Fisheries Re fIorts No. 57, VoL 4 FRm/R57.4 (Tri) PFOc: EINGS F THE WORLD SCIENTIFIC CONFERENCE ON THE PIOLOGY AN CULTURE OF AND ACTES D LA CONFERENCE SCIENTIFIQUE MONDIALE SUR LA BIOLOGIE ET L'ÉLEVAGE DES CREVETTES ACTAS DE LA CONFERENCIA CIENTIFICA MUNDIAL SOBRE OLOGIA Y CULTWO DE CAMARONES Y GAMBAS Mexico City, Mexico, 12-21 June 1967 Mexico (Mexique), 12-21 juin 1967 Ciudad de México, México, 12-21 junio 1967

j V

A FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS F4 0 ORGANISATION DES NATIONS UNIES POUR L'ALIMENTATION ET L'AGRICULTURE ORGANIZACION DE LAS NACIONES UNIDAS PARA LA AGRICULTURA Y LA ALIMENTACION 47 ? ROME, 1970 1499 FAO LIBRARY AN: 115086

FEm/5l03 FAO Fisheries Synopsis NoJ03 SÄST -

SYNOPSIS OF BIOLOGICAL DATA ON THE PINK SHRIMP Penaeue duoraruia duorarum Burkern'oad, 1939 poa6 synopticiue sur la biologia de Ponaeue duoraruxn duorarum Burkenroad, 1939 Sinopsis sobro la bio1oga dei Penaous duorarum duorarum Burkenroad, 1939

prepared by

P,J, COSTELLO and DONALD M. ALLEN Bureau of Commercial Fisheries Tropical Atlantic Biological Laboratory 75 Virginia Beach Drive Miami, Florida 33149, U.S.A.

Contribution No.249, Bureau of Commercial Fisheries Biological Laboratory, Galveston, Texas 77550, U.S.A. - 1501 -

/3103Penaeus d. duoraruin

No.

ID14TITY 1 i

1,1 omeno1ature 1

1011Valid nano 1 1,12Objeotive 1

1.2Taxono 1

1.2]. Affinities i 1.22Taxonomlo status i 1.23 Subspecies 1 1.24 Standard common names, vernacular flames 1,2

1.3 Morpho1oy 2

1,3]. External inorpholo 2 1,32 Uyto_morphology* 1,33 Protein specificity 2

2 DISTRJBtJTION 2*1

2,1 Total area

2,2Differential distribution

2,2]. $pawn, larvae and juveniles 2.22Adults

2.3 Determinants of distribution ohanRe 2:3

2.4 bridization*

3 BIONOMICS AND LIFE HISTORY 3,1 3.1 Reproduction i 3,11 Sexuality 3.12 Maturity 3.13Mating 3.3.4 3'erti1ization 3.15 Gonads 3.lb Spawning 3,17 Spawn 3:2

3.2Pro-adult hase 2

3,21 1knbryonic phase 2 3.22 Larval phase 2 3,23 Adolescent phase 2

3 3 Adult phase 3:4 3.31 Longevity 4 3,32 Hardiness 4 3,33 Competitors 4 3.34 Predators 4 3,35 Parasites, diseases, injuries and abnormalities 4

This synopsis has been prepared according to OutlineVersion No. 2 (H. Room Jr., FAO Fish.Synops., (1) Rev,1, 1965), -1502 - ii FBm/S103 Penaeus d., duorarum

PageZIo, 3.4Îutrition and growth 3*5

3.41Feeding 5 3.42 Food. S 3,43Growth rate 36 3.44Metabolism 6

3.5Behaviour 6

3,51Migrations and local movements 6 3.52 Schooling 7 3,53 Responses to stimuli 7

4 POPULATION 4*1

4.]. Structure i

4,11 Sen ratio 1 4.12Age composition i 4.13 Size oomposition i

4.2Abundance and density of population 1 4.21Average abund,a,nco* 4,22 Changes in abundance 1 4.23Average donsity* 4.24Changes in density 1 4.3 Zlatality and, recruitment 4:4 4.31 Reprod.uotion ratos 4 4.32Factors affecting reprodnotion 4 4,33 Reoruitmen-L 4 4,4 MortalIty and morbidity 4

4,41 Mortality rates 4 4.42Factors causing or affeotng mortality 4 4.43 Faotors affecting morbidity 4 4.44Relation of morbidity to mortality rates* 4,5Dynamics of population (as a rho1e)*

4.6 The population in the community and the ecosystem 4

5 ]ULOITATION 5*1

5,]. Fishinß equipment 1

5.11 Gears 1 5,12 ,Uoats 1

5.2Fishing areas 1

5,21General geographic distribution i 5,22 Geographic rangez 512 5,23Depth ranges 5.24Conditions of the grounds

Fishing seasons 5.3 2

5,3]. General pattern of oeacon 2 5.32 Dateof beginning, poak and end of seaeons* 5,33 Variation in date or duration of seasons* -1503 -

künJSì.O3 enaes 1. duorarurn iii Page Ido, 5.4Fishing operations anl resuJ.s 5; 2 5.41Effort anAl intensity 2 5.42 Selectivity 3 Catches 5.43 3 6 PROTXTION AlD MANAGEMENT 6i1

6.]. Regulatory (legislative) measures i 6.11Limitation or reduction of total oatoh i 6,12Protection of portions of population i 6.2 Control or alteration of physioal features of the environment'

6,21Regulation of flow 6,22Control of water levels 6,23Conirol of erosion and silting 6.24Fishways at artificial and natural obstruotions 6.25Fish screens 6.26 Improvement of spawning grounds 6,27Habitat improvement 6.3Control or alteration of chemical features of the environment'

6,31 Water pollution control 6.32 Salinity control 6,33Artificial fertilization of waters

6.4 Control or alternations of the biolocal features of the environment *

6.41 Control of aquatic vegetation 6,42Introduction of fish foods (plant, invertebrate, forage fishes) 6,43Control of parasites and disease 6,44 Control of predation and competition 6,45Population manipulation 6.5Artificial atockin*

6.51Maintenance stocking 6,52Praiisplantat±on introduction

7 P0ID FISHCIJIÌrtJRE 7;].

7.1Procurement of stocks 1

7.2Genetic seleotion of stooks* 7.3 Spawning (arfic.a1; inducea.;. natural) 7,4Holding of stock

7.5Pond management (fertilization; aquatic plant control, etc.)' 7.6Foods; feeding 7.7Disease and. parasite control' 7,8Rarveat' 7.9 Trmnapor-t'

8 REFEREMCES 'Ano information was available to the authors, these subjects are not listed in the text. -1504 -

eus d. duor 181 arthrobranoh on somite XIII;mastigobranchias IDENTITY on somites VII to XII.Zrgooardiao ossido consisting of a principal tooth followed. by a 1,1Nomenclature longitudinal row of smaller teeth which often 1,11 Valid name end in a cluster of minute teeth.Body glab- rous.(After Dall, 1957, slightly modified ivy Pérez-Farfante.) duorrìrijm Burkenroad. 1939 Spooi±'io 1.12 Objective eyxlonomy The holotypo and syntypes of Penaeuduo- Periaeus arum "Form A", Burkenroad, rarum (1 male and. 2 females) aro in the Dinghani 1939, Buli.Binghain oeeanogr.Coli., Vol.6, Oceanographic Collection of laie Bniversity, New p,1-.62, 36 text-rigs. Raven, Oonneoticiit, U.S.A., catalogued. B4O,Q. 255 and 256.Those specimens were taken off 1.2Taxonom' Alabama, U.S.A., ATLAIPPIS Station 23l3 in 37ni of water on March 20, 1937 (Pérez-Farfante, 1.2]. Affinities 1969). Suprageneric(to family after Waterman and Chaos, 1960) The following diagnosis of P. duorarum is from Williams (1965):"Female with thelyoum Phylum Arthropoda composed of two broad lateral plates, and. a me- Glass Crustacea dian plate.Posteromedian part of median plate Subclass Malacostraom of adult with s. well-developed, short, longi- Series Eumalacostraca tudinal carina extending anteriorly toward. Superorder Eucarida roughly esinioircular, concave anterior portion. Order Lateral platos produced medially to meet in mid- Suborder Na-tantia line, exCept variably divergent a-b anteromediaxi Section Ponaojdea oornera, thus exposing carina of median plate. Family Subfamily Penasinne .Petssma of male with distal ends of distoventral lobes curved medially, not project- ing free of distolateral lobos;external edge Gemelo of distoventral lobes with a serios of 2 to 12, Genus Penaeus Fabrioius, 1798, Suppl usually4to7,small spinulsmedian or at-. Ent.Syst:385,4ö8Type , by selection taobeci edge of distoventral lobes witha com- b3' Latreille, 1810, Consid,gén.Anim,Cruat. pact group of 6 iso 16 large, long, sharp, curved 2ncbri.Ina.2 102,422: Fabrioium, spines;fold of d.istolateral lobo rather small £798)Suppl.Ent.Syat,:408. Gandenmasculins, and armed inconspicuously if at all." Definition For a key to the western Atiantio apoden of the genus Penaous, see Pérez-Farfante (1969). Rostrum toothed dorsally and ventrally. Ocr&cc without longitudinal or transverse 1.22 Taxonomic sttus ouuaea cervical and orblto-axitenna]. amici rind. ciitannal oarlaao always present.Hepaisio Ponaeus duorerum is one of about 28 spo- incj oenna1 spines pronounced, piserygostomial oies of the genus Penaeus,P, duorarum and, -the aiJrounded.Telson with deep median sulcue, closely related. Penaeus brasilierisis Latreille without fixed subapical spines, with or without and. Penmeus a.ztecus Ives wore considered. a lateral movable spines.First antennuia.r sag-. single species (P. brasilienaim) befare the tan- meo ithou. a spine ori ventral diajomodian anomie revision based on morphological differ- borcJir.Antemnular f lngella shorter than cara- onces by Burkenroad (1939). paco,Nau!llulary paip with 2 or 3 seente, usually 3 Basial somos on let and. 2nd parejo- 1,23Subspecies poda;exoods on ist 4 perelopods, usually prusonon 5hPetasen symmetrical, pod-like Pérez-Farfante (1967) divided P, âuorrtrum rí.hhín medIan lobes with or without distal into the subspecies P, duorarum duorarum and. protuberanoes;lateral lobes often with thiczen- Pduorarum notialis.The basis for the dvj- ed ventral margin.Appondix masculina with die- sien was given by Pérez-Farfan6o (1969)anfol.-. tal eoent subtriangular or ovoid, bearing lows:"P, duorarum d.uorarum ,5roru the north- numerous spines,Thelyonni usually with an an- western Atlaìtio and the Gulf of Mexio27 has the icrior urocoss, varaabla in shape, lying be- dorsolateral mulcas narrower than P. duorarum ttree.i he coarto of 4th perelopods;with or with- notialis from the Caribbean Sea, the ALlantlo uni lo.Lernl platas on sternite XXV.Pleuxo- Coast of South America, and Africa,J3urkottroad. branohlrto on soml-Los IX to XXVia rudimentary (1939) was the first to point out thin difference ahrchranoh osomite 1111, and. a posterior between the two and called the former 'Form A', - 1505 -

1.2 FRm/Sl03 Penaeus cl. duoraruai and the latter'Form B'.Biometric studies "AbdomenwiÌisegments four to six cari-. have indicated a statisticallysignificantdif- nate, carina of sixth ending posteriorly in a ference in the ratio(K/S)of K (height of the spine and flanked on each side by a narrow keel) to S (width of the sulcus) between those groove. Telson with deep median groove s-rid populations."K/Srangesfrom about 2.5 to acuminate tip," 15,5(mode4,5)inP,d.duorarurn,and from 0.25 to 3.0 (mode 1,75) in P.d.notialis. In addition, The color of pink shrimp varies with lo- the third pereiopod "is proportionately shorter cality, diel period and age. Shrimp in coas- in P. d.duors-rumthan in P, d. noti.aijn" tal areas of Florida are green, brown, or rod- (Prez-Farfrate, 1969). he noted further that dish; on the offshore Tortugas (Florida) end "overlapping is very limited" and the two sub.-. Campeche (Mexico) grounds, colors range Zrcm species "do not seem tomixacross the Gulf pale rose to deep pink (Idyll, 1964).Along Stream" (P6rez-Farfante, 1967). the northern coast of the Gulf of Mexico, le- mon yellow is common (Anderson, 1962),Dey- Pérez-Farfante, therefore, equated P,d, night color differences were observedin south duorarum with Form AandP,.notialis with Texas; at sunset, shrimp became bright red FormB. In thissynopsis,the discussion is re- {jloese stai,, 1968).Juveniles and y,-wa& stricted to P.d,duorarum of thenorthwestern adults are gray, reddish brown, or bluh c'°y, Atlantic and Gulf of Mexico, Information con- and older individuals are red, pinkish13 o cerning P, duorarum, or Form A,from these geo- gray,or almost white (Williams, 1965). J graphic areas is considered referable to P.d. spot (gray, blue, purple, rod, or brown) mey duo rarum. or may not be present at the pleuraljuncture of the third and fourth abdominal segments 1,24 Standard common names,verne- (Anderson, 1962;Williams, 1965), cular names The morphology of pink shrimp changes Standard common during growth. Several parts and appendaGos names, U.S.A., pink shrimp become shorter in proportion to the oai-apacc' Mexico. oamar6n rosado length;lthis break is distinct in females Vernacular s: blue-tailed shrimp, brown about 108 mm total length2endin males at potted shrimp, rhannel shrimp, grass shrimp, about 96 mm total length (Drucker, 1960). green shrimp,grooved ehriinr, hopper, jumbo hop- per, pink grooved shrimp, pink night shrimp, Williams (1965) reported the rostrum to be pink spotted shrimp, red shrimp spot shrimp, "relatively shorter anddeeperin old indivi- spotted shrimp, skipper (U.S,A.5. duals than in young Ones, It extends to the end of -the basai antennular article inaverage-sized The common name "pink shrimp" in this syn- adults." opsis refers only to the subspecies P.d.duora- rum. (See also 1.21, 1.23, and 3,11.) 1,3 1,33 Protein specificity

1,31 External morphology Leone and Pryor (1952), who made aerologi-. cal comparisons of the blood proteins of three Figure 1 is a lateral view of an adult pink species of shrimp -pink, brown (Penaeusaziecus shrimp. The following description is from Ives), and white (Penaeus setiferus (Linnaeue)) Williams (1965). "Integument thin, polished, -from1'crth CaroliiTa,placed pink and. brown translucent. Carapace with a median carina con- shrimp closer to each other than either is to tinuous anteriorly with rostrum andextending the white shrimp. Serological differences s-up- nearly to posterior border of carapace flanked port the theory that these three speciesare on each aide by a broad, roirnded groovepos- distinct but closely related, terior half of carina with a median longitu- dinal groove; anterior half arcuate, highest above orbit s-nd with9or 10 sharp teeth; pos- 3-Carapace length is a head-length measurement tenor tooth remote frein others, anterior 6 or takenfrom the orbital notch inside the or- 7on rostrum proper,Lower margin of rostrum hits-1 spine, in a lineparallel to the with 2 to3teeth (occasionally 1); tip slen- lateral rostral sulcus, to the posterior der, horizontal br directed slightly downward, margin of the cophalothorax. unarmed, Anterior margin of carapace with strongantennal spine on carina extending back- 2Total length is measurement taken from the ? ward nearly to well-developed hepatio spine, of the rostrum to the tip of the telmoni' Cervioal groove extending halfway from hepatio abdominal segments extended in a stralght spine to dorsal carina, A subhorizontal sutura line. below hepatio spina, and a groove extending from near hepatic spine to near baso of ocular pa- dunole, An.orbital ridge behind ayo. - 1506 -

PlimJSlO3Penaeud.. duorarum 1s3

Fig0 i.Penaeus duorarwn thiorarum Burkernoad, 1939 -1507 -

FRinJS1O3 Penamus d. duorarum 2:1

2 DISTRIBUTION 1965; Oopeland and Truitt, 1966; Christmas, Gunter and Musgrave, 1966; Williams, 1969). 2.1 Total area In North Carolina, for example, Williams found that postlarvae usually enter the es- The world distribution of P.d.duorarum tuaries from Nay through November. (Fig.2) is restricted to the northwestern At-- lantic and the Gulf of Mexico (Atlantic, NW. - ANW; Atlantic S.L - ASW) - see Perez- Juvenile pink shrimp inhabit coastal Farfante (3.969). This ubspecos inhabits bays and also estuaries that penetrate coastal waters of land areas 240, 230, and deeply into the mainland (Williams, l955a 311 of the FAO distribution code (Holtbuis Tabb, Dubrow and Jonee, 1962; Joyce, 1965). and. Rosa, 1965). P.d..duorarum is found in They also have been found in the Gulf of BorTiniâa and, along the Atlantic coast of North Mexico west of Key West, Florida, in the America from lower Chesapeake Bay to sóuthorn grassy shallows near the Marquesas Keys Florida, and into the Gulf of Mexico. The (Ingle et al,,, 1959), According to Eldred range extends along the Gulf coast into Mexi- (1962), "Very small individuals prefer the can coastal waters and terminates slightly very shallow, protected, relatively oalm south of Cabo Ca-boche at Isla Mujeres (Pérez- areas near the shore...," They occur year Farfanto, 1969). P.d,d.ucrarum, as Form A, around. in Tampa Bay, Florida, where numbers was also found at the Doubleheaded. Shot Gays are greatest in the summer and autumn in the Straits of Florida (Burkenroad, 1939). (Eldred et al., 1961), A similar seasonal dietribution was noted. in Florida Bay P.d..duorarum is most plentiful whore the (Costello, Allen and Hudson, MS), and in North Carolina estuaries (Williams, l955a). Continental Shelf is broad and. shallow, arid where coastlines are characterized by shallow As juvenile pink shrimp grow, they bays arid estuaries. Highest concentrations move gradually into deep water (Iversen and. are in the eastern and southeastern Quit' of Idyll, 1960), Mexico, adjacent to the Florida and Yuoatd.n Juveniles and, young adulta (58 to 101 mm total length) marked and Peninsulas, released in northeastern Florida Bay in 2 m of water appeared., within a few months, as 2,2Differential d.ìetribution adults (87 to 155 mm total length) on the Tortugas grounds in depths of 17 to 44 m 2,21 Spawn, larvae and. juveniles (Allen and Costello, 1966),

Pink shrimp spawn offshore, The (See also 3,23,) demersal eggs were found from about 10 to 64 km off the west coast of Florida in 4- 2,22 Adults to 38- s depths (Eldred et al., 1965). The distribution of eggs on the Tortugas Shelf A few pink shrimp remain in mouth (Florida) was not observed directly, but the Florida estuaries after they become adults, spawning grounds, determined from larval but most move offshore (Iversen and Idyll, distribution, ranged. from 15 to 48 n in 1960; Allen and Costello, 1966). The aime depth (Jones et al., in press). Spawning of shrimp on the Tortugas grounds increases was year around and the center of activity with depth (Iversen, Jones and Idyll, 1960) and intensity varied seasonally, and the greatest concentrations are between 9 and 45 m (Kutkuhn, l962a), These shrimp After hatching, the larvae are planktonio are rare at depths greater than 51 m and move shoreward, apparently following an (Hildebrand, 1955), but a few have been indirect route determined by currents (Mucre, taken at depths to at least 110 m and Jones and Dimitrion, 1968).Newly hatched specimens of P. duorarum, subspecies larvae had their greatest abundance in the unknown, were caught at depths as great as eDawning area on the Tortugas Shelf, whereas 366 m off the western edge of the Great postlarvae were abundant near the coast Bahama Bank (Anon., 1961; 1962), (Jones et al., in press), Postlarvao enter south Florida estuaries throughout the year; The biomasm of Tortugas pink shrimp, the greatest numbers were taken in spring, generally, is least in the spring and summer, and autumn (Roossler, Jones and Munro, sunmer and greatest in the autumn and. early 1969; Allen, Hudson and Costello, MS). In winter (Kutkuhn, l962a), Annual fishable other areas studied, the annual influx of biomass from 1956 to 1959 was fairly postlarv'ae was similar to that in south uniform, but with a slight downward trend., Florida, but generally more restricted (Bearden, 1961; Joyce, 1965; ßldred eai., (See also 4.22, 4.24, and 5.31) 35N.

25

Diatr.bution Major estuarire fihLng grounds 20 Major offshore fishing grounds

Linit of atatitica1 ares South CarolrnaNorth UN L TED STATES Hisisippi Alabama Georgia Caroltni \ Chesapeake Bay Texas ATLAS T IC OCIAN Bermuda GULF 0F MEXICO Doubleheaded ffyucatn j Cabo Cstoche QSouth Atlantic STATISTICAL AREAS lOO w. o 95 T Tabasco ¡Caapecoe 80 Shot Cays Fig 2 tibuiozi arid 'jghjri areas for Penaeuß duorarum duorarum QSanibel-Tortugee QApalachicola QPensacola to Miss. River QMise. River to Texee QMexico-West of Long. 94 Texaa W. L 75 QMexico-Obrego7t-Campeche 70 ot: -1509-

FRm/5103 Penaeus d. duorarum 2.3

2.3 Determinants of distribution changes Young pink shrimp tolerate low salinities (Buricenroad, 1939) but not for extended Thedistribution of pink shrimp may be periods of time (Joyce, 1965), They were limited by salinity, temperature, and substrate most abundant in Texas bays at salinitiem (Williams, 1965). Details of these and add- higher than l8°/oo (Gunter, Christmas and itional factors are given here. Killebrew, 1964) and in Florida Bay at salinities from 30 to 50°/co (Tabb, MS). Temperature: Spawning temperatures (see Observations ooncerning distribution wore 3,16) are apparently critical and may deter- summarized by Hildebrand (1955) who con- mine area of spawning and subsequent distribu- cluded that juvenile pink shrimp "prefer" tion of shrimp populations (Eldred. et al,, salinities of 200/00 or more. 1961). In North Carolina estuaries, the majority of samples containing postlarvae Values of temperature and salinity were collected at water temperatures between should be considered together in evaluations 23° and 28° C (Williams and Deubler, 1968), of the effects of either factor on pink shrimp. Williams (1960) reported that the Temperatures required for survival of survival of shrimp at low temperatures is pink shrimp may vary with locality. In North best at moderate to high salinities, Joyce Carolina estuaries, juveniles have been (1965) noted that "The ability of shrimp collected in water temperatures ranging from to withstand low salinities appears to 4° to 340 C (Williams, l955a). These shrimp depend upon several factors.,." which became narcotized by the cold at 100 C include temperature and perhaps the oon- (Williams, 1955),,and in severe winters most oentration of calcium ion. of the shrimp population in estuaries may die (Williams, 1965), Death caused by low Currents: The distribution of pink temperatures may be relativelyunimportant shrimp may be controlled to a great extent in the warmest parts of the species range by currents. Concerning larvae spawned on because none has been reported. In Florida, the Tortugas grounds, Nuoro, Jones and shrimp have been collected within the range Dimitriou (1968) reported: "dispersal may 100 to 3505° C; tlij were narcotized at 13,3° be effected primarily by the current which C (Eldred et al., 1961), These authors stated leaves the Tortugas area through Rebecca that juveniles move into deep water withthe Channel, and enters the Florida Current in onset of cold weather. the Florida Straits. Entryinto the Florida Current would result in rapid transit to Mortalities attributed to high temper- the area adjacent to Florida Bay." The atures have not been reported, movement of postlarvae to estuarine waters is effeoted by inflowing currents (Idyll and. Salinity: Salinity requirements or Jones, 1965; Copeland and Truitt, 1966; preferences vary with geographic area and Hughes, 1969). Juvenile pink shrimpin shrimp size, Minimum salinities associated North Carolina estuaries concentrato near with pink shrimp, (irrespective of size) are inlets where tidal ourrents are moderate as follows: "no measurable salt (using stand- (Williams, l955a), In south Florida waters, ard hydrometers)" in the Florida Bay area some juveniles "move seaward on the ebb (Pabb, Dubrow and Jones, 1962); less than tides and back into the bays on subsequent 1,0°/oc in the Caloosahatohee ßstuary, Florida flood tides" (Tabb, Dubrow and Jones, 1962). (Gunter and Hall, 1965); 0.64°/oo in a north- Juveniles migrating to sea are carried by east Florida estuary (Joyce, 1965); and 2.7°/oc ebbing currents (Burkenroad, 1949; Copeland, in Mesquite Bay, Texas (Hoese, 1960). Maximum 1965- Beardeley and Iversen, 1966; Hughes, salinities recorded for waters with pink shrimp l969,, are600/co in the Upper Laguna Madre, Texas (Simmons, 1957), and 70°/oo in Florida Bay Substrate: The distribution of pink (Tabb, MS), Tabb, Dubrow and Jones (1962) shrimp follows closely that of sand, shell- caught postlarvae in the Florida Ba area sand or coral mud substrato (Williams, at saliriities ranging from 12 to 43 /oo, l965, The importance of substrate type, juveniles at salinities of about O to 47°/oo however, as related to food, cover, or some and adulte at salinities of 25 to 450/co, less obvious factor has not been established. Where adults are concentrated on the Tortugas williams (1958) reported that young pink grounds salinities range from about 36 to shrimp in North Carolina estuaries ar 38°/oo Iversen and Idyll, 1960), "onfined largely to areas near the sea where the bottom is composed of coarser Juvenile pink shrimp can survive in materials." He noted that pink shrimp waters having a wide salinity range,although oan burrow into extremely coarse substrates, optimum salinitiem may be relatively high as and in laboratory experiments, learned that compared with those of white and brown shrimp subadulte prefer she1l-and and boom peat (Ganter, Christmas and Killebrew, 1964). substrates. Adults prefer calcareous -1510 -

FRm 510, Penasusti.duoraruin sedirnnt but do ocour on hard sand bottoms that may serve aifood. for shrimp The geo- "particularly in non-turbid waters" (Hildebrand, graphie distribution of pink shrimp may follow 1955) closely the distribution of seagrassem. ac- cording to Hoese and. Jones(1963),who noted Seagrasees: The assooiatíon of juvenile that there are no pink shrimp fisheries near pink ahrimp with seagrams has been noted by areas where seagramses are rare or absent, mamy veetigators, including Hildebrand(1955), 14ourn et al,(l957) and Tabb, Dubrow and Environmental changes can alter growth of Mamig(i96, Eldred. (1962) reported that meagramsea and afeot associated organisme, in inehore waters, email juveniles were near whioh include young pink shrimp (TabbDubrow shore in bede of shoal grass, Diplanthera and. Hanning,1962), Destruction of seagrass ritii, and, the large juveniles were in beds by dredge-and-fill operations ham elim- deeper waters in turtle grams, Thalaemia inated nursery areas for pink shrimp in Tampa testudinum. Moore (1963) observed that turtle Bay, Florida (Saloman,l965), grams is a suitable habitat for many organisms FRrn/S103 Penmeus d. duorarum

3 BIONOMICS AND LIFE HISTORY 3.13Mating

3.1 Reproduotion No observations describing copulation are published, but it may occur several 3.11Sexuality times during the life cf a female (Idyll, 1964);this multiple copulation would Pink shrimp are dioeoious, Hermaphro- indicate that pink shrimp are promiscuous. ditism ham not been reported. Males aro Sperm transfer probably is between a hard-- distinguished from females by the presence shelled male and a recently molted female of the male sex organ, the potassa, between (cummings,1961), the let pair of pleopods. On the female, the distinguishing external organ, the thelycum, 3.14Fertilization lies between the 4th and. 5th pairs of pereio- pods. Figure3illustrates the petasma and Idyll(1964)described fertilization in tbelyoum, pink shrimp. "Fertilization is external, The male attaches a epermatophore to the Female Bhrimp attain a greater size than underside of the abdomen of the female.,,. males, Williams (1955) reported: "There is a sex size disparity in adult P. duorarum "Females bearing spermatophores are in which is statistically significant at mean various stages of sexual development. The total lengths which exceed 100 mm." The spormatophores are shed at ocdysis, largest female reported was 280 mm total length (Eldred,1958);the largest male was "The genital pores of the female open 43mm carapace length or about197mm total at the bases of the third pair of pereio- length (Iversen, Jones and Idyll,1960).In pode. After the eggs pass through these large pink shrimp, females have a greater pores they are fertilized by the sperma- oarapaoe length than males of the same total tozoa etorcl in the thelycum." length (Kutkuhn,1966). 3,15Gonads 3.12 Maturity cummings'(1961)report that ovary Femsle pink shrimp may become sexually ight increased with shrimp size suggests mature at85mm total length and males at that large females produce more eggs than 74mm (Eld.red et al.,1961), At these lengths, smaller ones, The number of eggs produced estimated woightsKutkuhn,1966)are about per spawning is unknown, but a female of a 5.2g for females and3.5g for males. related speoies--a white shrimp 172 mm total length--contained about860,000 The degree of sexual maturity, in female eggs (Anderson, King and Lindner,1949), pink shrimp has been described by Cummings Pink shrimp probably produce a similar (1961) as follows: number of eggs during one spawning, and Cummings(1961)indicated 'that they may Undeveloped stage: Ovaries flaccid, spawn more than once during their lifetime, small and translucent, 3,16 Spawning Developing stage: Ovaries flaccid, larger than in The ratio and distribution of sexes undeveloped stage, of adult pink shrimp on the spawning and white to palo grounds may vary seasonally (Broad,1951; orange buff colour. Iversen, Jones and Idyll,1960).

Nearly ripe stage: Ovaries slightly The location of the spawning ground turgid, large, and on the Tortugas Shelf (Florida) was given glauoous oolor. by Jones et al, (in press),The center of spawning shifted with the movement of adult Ripe stage: Ovarios same as shrimp from shallow to deep water am winter nearly ripe stage, approached (Munro, Jones and Dimitriou, ova with peripheral 1968), Eldred et al.(1965)reported that rod-like bodies. offshore from Tampa Bay, Florida, the ]ocation of intensive spawning varied from Males "with joined endopods and sperma- one year to the next, tophoros with spermatozoa" are considored. sexually mature (Eldred et al,,1961), Several environmental factors may influenoe spawning, Pink shrimp spawn (See also section 4.12.) throughout the year on the Tortugas Shelf - 1512 -

FRui/S103 Penneus d, duorarum bu îarval abundance im highest in the spring, against the membrane with the first antennaa summer and, autumn and lowest in the winter Energence from the egg requires 2 to3min- (Jones et ai. in press), In the more northerly parte of the range, the spawning season may be shorter than in the southern 3.22 Larval phase portion (WIlliams, 1965; Joyoe, 1965), Cummings (1961) noted a positive correlation The larvae of pink shrimp were des- btwen temperature of bottom water and the cribed by Dobkin (1961) who noted5 occurrence of ripe shrimp on the Tortugas nauplial stages, 3 protozoeal stages, groando., Shrimp spawn on the Tortugas grounds and 3 mysis stages. Figuro 4, taken from between 19,60 and 30.6° C (Jones et al., in Dobkirì, shows a 4th nauplius, He reported. preme), activity im maximum during the period size of' larvae irr total length as follows: of highest bottom water temperatures (Munro, nauplil, 0,35 to 0,61 mm; protozoeae, 0,86 Jones and Dimitriou, 1968), Eldred et al, to 2,7 mm; and myses, 2.9 to 4.4 mm, (1965), however, found evidence that pink shrimp spawn at temperatures as low as 16.10 C Development tise of the various stages and stated that "a rising temperature im the in the laboratory was reported by Ewald important factor in triggering spawniug ...." (1965), He noted that the number of' myeis On the Tortugas grounds, most spawning was stages and the length of larval life varied turing the waning moon (Munro, Jones and with water temperature and observed that Dimitrion, 1968). Spawning in the laboratory "Pink shrimp reared at 260 C took a minimum was "always in the very early hours of the of' 15 days to metamorphose whereas those morning" (Ewald, 1965). at 210 C took25 days,,,."

The female shrimp releases the fertilized Nauplii subsist on yolk granules in eggs freely into the water where they sink their bodies, and active feeding begins in slowly to the bottom (Idyll, 1964), the let protozoeal stage (Dobkin, 1961; Ewald, 1965), In the laboratory, larvae Joyoe (1965) showed that where pink, were fed, a mixture of unicellular algae white and brown shrimp ocourred together off and marine yeast supplemented, in the 3rd the northeast coast of Florida, spawning times mysis stage, with nauplil of brine shrimp overlapped during the warm months of the year. (Arteniia), Brown shrimp begin to spawn in February, white shrimp in April, and pink shrimp in May. Larvae reared in the laboratory survived bettor in Gulf Stream water than 3,17 Spawn in water from Bisoayne Bay, Florida (Ewald, 1965). Survival ratos of larvae on the Viable eggs measure from 0.31 to 0.33 mm Tortugas Shelf vary little and average 83 In diameter (Dobkin, 1961). They are opaque percent per day (Munro, Jones and Dimitriou, and yellowbrown and, the ohorion shows a 1968). blue hue under oertain light reflections, 3,23 Adolesoent phase The specifio gravity of fertilized eggs is slightly greater than water (Idyll, 1964), The postlarvas of pink shrimp were described by Dobkin (1961); methods for 3.2 Preadult phase their identification were provided by Williams(1959),Ringo and Zamora (1968), 3.21 Embryonio phase and Chuensrl (1968), The smallest pomtlarva reporten from plankton byPobkinwas 3,8 sIm The embryos develop in the fertilized total length; the smallest reared in the eggs thai are extruded in the water. There laboratory by Ewald (1965) was 2,9 mm, is no parental care. Ewald reported. that the poetlarvao reared at the lowest temperatures were smallest, Dobkin (1961) provided a photomicrograph of eggs in several stages of development, and Early postlarvao are planktonic in noted that development of pink shrimp may offshore waters (Jones et al,, in press), be similar to that of' Penasus iaponicua as Th more advanced ones enter the estuaries summarized by Pearson'i'T at about 8 mm total length (Copeland and Tr,itt, l966Allen, Hudson and Costello, Dobkin (1961) described hatching as MS), although size at entrance varios with fol1ows '1Prior to ite emergence, the season (Nattas, 1966) and area (Tabb, Dubror nauplius .vee its appendages convulsively and. Jones, 1962), They become bnthi at at short intervals, After the furcal spines about 10 mm and concentrate in shallow puncture the egg membrane, the nauplius water where cover Is adequate, Here, they emerges, posterior half first, by pushing develop into juveniles, and density of pjJ$Q3PIMuL ¿, duorexu

o i 2

MILLIMETERS

1ig. 3Sex organp of Penaeuo duorarum duornrum Petaeeof oale, 125 mm total length (zight later&1 view) Thelyouo 'f tomaIe, 125 mm total length (ventral view)

Fig. 4A ventral view of e 4th nau1iue of Penaeuo duorarue duors.rua. (?roa Dobkin, 1961) - 1514

FRa 510Ponaeus d, duoraruin dlvíUuals may exceed 32 per ,,2 (Costello, by the different speoims in respect to mizo Al1ei arid Hudson, MS), and season probably reduce oompetition, Offshore, pink and white shrimp sometimes Juvenile shrimp are more robust than oocur at the same depths,Differences in pcvlarvae and have proportionately shorter substrate preference, food, and dici 6114 ehdominal segments,Methods for identify- behaviour, however, probably reduce compe- iojuveniles were given by Williams (1953). tition there almo (Hildebrand, 1955). P.nk shrimp, as juveniles and adults, 3,34Predators marr r ,r.j from about 2 to at least 6 mo in the rh iree'yareas (Costello and Allen, Fishes that prey upon pink shrimp ¿ì ut as size mareases, they move grad- undor natural oonditionm include: snook eally toward deep waters,When shrimp migrate Centropomus undeoimalis (Marshall, l958S; from the estuaries to the offshore waters, spotted seatrout, Cynoscion nebulomum they average about 95 to loo mm total length (Moody, 1950; Tabb, 1961; Stewart, l6l); (Joyce, 1965),Size at emigration, however1 mangrove or gray snapper, Lutjanus ßrisous varies with season and area (Iversen and Idyll, (Croker, 1962); toadfish, Opmanus beta 1960; Eldred et al,, 1961; Tabb, Dubrow and (Woodburn et al,, 1957); red drum, Soiaenops Jones, 1962; Copeland, 1965), 2ila.a (Yokel, 1966); and. blue oroaker, Bairdiella batabana (Robins and Tabb, 1965). (See also 2,3,) We have found pink shrimp in the stomaoh of king mackerel, Sooniberomorus cavalla, 3,3Adult phase and mutton snapper, Lut janus analis (unpublished observations),Costello and 3.31Longevity Allen (1962), in tank experiments, found mangrove snapper, red. grouper (Epinephelus (See 4.12,) morto)and black grouper (Mycteroperca bonaoi5 predacious on pink shrimp,Undoubt- 3,32Hardiness edly, many fish not listed here prey upon pink shrimp, Juvenile and adult pink shrimp are hardy capable of to1eratinrough handling Birds, mammals, and reptiles that inhab- without excessive mortality (Costello and. it shallow estuaries are probably predators Allen, personal observations),They may be of pink shrimp, but this matter is poorly taken in trawis from depths of 9 to 35 is, documented, h1d in tanks at the surface, and returned. to the bottom without undue loss,Costello We have observed that pink shrimp react (1964) noted, houever, the importance of in two ways when attacked by predators:(1) maintaining water temperatures below about backward movement to avoid contact is ano- 24,0° C when ho1din,hpnt in tanks, omplished by rapid contractions of the abdominal muscles and sweeping of the uropode When pink chrimp are transported in and (2) thshrimp seek shelter of any kind, iruoks in the live bait indu.mtry, large volumes of air are pumped into the water, 335Parasites, diseases, injuries Undo,- thcsa conditions, they do not perish and abnormalities ro,t rie oabol:m due to eXQCsbubbles in the 't»r thcn huJd in bait tanks, pink shrimp The following parasites and. diseases va j'o'c h'.ruy than white shrimp and. less have been recorded in pink shrimp: hç- I,n bro,v' shrimp (De Sylva, 1954), Class Eu.bacteriae:Pseudomonas sp, Like other , pink shrimp are Pseudomonae was listed byThn 1964) delicate imrnsdiutaly following ecdyeis, as a parasite in hrip along the Florida west coast, 333 Competitors Clase Telosporidea:Cha1o1obu Burkenroad (1939) suggested that pink iaeus Kruse, 1959,This gregarine was shrimp are in interspecific cornptition with isolated from the digestive tract of shrimp brown shrimp and Caribbean brown shrimp, (Kruse1959),Kruse found 20,8 pexoeifl of u un brasiliensis,He proposed that jur- a sample infected with this parasite,It je 1uuO of these species had optimum salinity found in r2uuiercus Florida lacaiitiee (Hutton, to which determined species doreinanoe l964',

Nomatc ois penaeue Sprae, 1954.Th ornpotition between pink shrimp and other young forme of this gregarias were isola-led speotee bas been diacaeed by Williams from the intoutine of shrimp (Kruse, l99),, (1JJ)Hildebrand (1955), and Joyce (l96 It occurs in numerous Florida looali.tjan avestigators uoncluded -that in the (Hutton, 1964), varinijoce inhCOi%ats occupied 1515

510Penaeus d. duorarum

Class Cnidosporidea: Thelohania sp. This Parachristianella dimogacantha Kruse, parasite 000urs in muscles and body organs, 1959, This parasite, recovored from the Kruse(1959)reported1.9percent infection digestive gland, oonare infroquontly in 784 pink shrimp examined, and considered (Kruse, 1959), but it is found in numerous this organism to be a cause of the "cotton" Florida localities (Hutton, or "milk" condition which makes shrimp 1964), uriaooeptable for marketing in the United Clase Nematoda: Contracaecw, habana States, Found in Alligator Harbor s.ndApalachi- (Linton,1900)Linton,1934, Immature forms cola Ray, Florida, were taken from the hepato-pancreas, pylorio gland, oephalothoraoio musculature, testes, Thelohania duorara Ivermon and Manning, and ovaries, Incidence of infection was 11 1959, Body muscle fibers and legs of shrimp percent (Hutton, Ball and Eldreà,1962). aro infeoted by this organism which causes It is found in numerous Florida localities a milky color, The incidence of infection (Hutton,1964), is low (Ivoreen and Manning, l959) but it is found in numerous Florida looalities (Hutton, Pink shrimp aro intermediate hosts for 1964). the trematodes, cestodes, and nematodee listed above, Nosema nelsoni Sprague, 1950, This organism, isolated from the body muscle tissue Class Jiexapoda (Ineecta): An unidenti- causes a cotton ox' milky appearanoe of shrimp fied larva of the family Sarcophagidae was flesh, It has been reported in shrimp from noted in the terminal ampoule of one Boca Ciega Bay, Florida (Kutton et al.,1959), epecimen caught in Tampa Bay, Florida (Hutton and Eldred,1958), Epistylis ap, This ciliate was Isolated from shrimp in numerous Florida localities 3,4 Nutrition and row-th (Hutton,1964), 3.41Feeding Class Trematoda: Parorchis sp, (tentative identification). A metaoercarja was taken Feeding habits of pink shrimp wore frosi one shrimp from Tampa Bay, Florida (Hutton determined from analyses of stomach oontonts et1,, 1959), by Eldred et al,(1961), Pink shrimp are bottom feeders, and juveniles and, young ?icrophallus gmaeus (Levinsen,1881) adults feed, primarily in shallow waters Baer, 1943, Metacercariae of this trematodo :here marine plants grow, Although most were found in shrimp from numerous Florida feeding is at night, shrimp may feed in the localities (Hutton,1964). daytime when the water is turbid,

eooelodes fimbriatue (Linton,1934), Feeding varies seasonally, Sastrakus- Metacercarias were found in the hepato=-panoream, umah(1970)reported that feeding activity onads and other soft tissues of shrimp of juveniles in south Florida was minimal (Hutton et al,,1959). This trematoda occurs in the late winter and summer, and highest in numerous Florida localities (Hutton,1964), in September, Eldred et al, (1961) suggested that shrimp feed more in the Class Cestada: Poypocephalus sp, summer than in the winter, (rParataonia sp,),This cestode is found in shrimp from numerous Florida localities (Hutton 3.42Food 1964), Idyll(1964)reported that pink shrimp Prochristianella penae Kruse,1959, are omnivorous and aleo ingest inorganic This organimm was isolated from the digestive detritus, gland and tissues that surround the digestive gland and stomaoh, Phis is probably the moat Ewald(1965)found that mioroplankton common helminth parasite of pink shrimp, In was suitable food for early larval etagen a sample of 137shrimp, Kruse(1959)reported reared in the laboratory, Advanced larval 97percent infected, It je found in numerous stages and postlarvae were fed naupili of Florida localities (Hutton,1964), brine shrimp (Artemia), and mioroplankton,

Parachx'im ianella monomegaoantha Kruse, Scotrakusumah(1970)reported that 1959, Thin parasite was isolated from the orustaccanc and. polychaetes are the maim digestive gland, The incidence of infection foode of juvenile pink shrimp in south is ortroreely low (Kruse, 1959),but it i Florida waters, He detected no difference found In numeroun Florida localities (Hutton, in diet seasonally, or between small and 1964), large speoimenm, - 1516 -

FRis SiG Penasus d, duorarum

Eidrcd et (1961) recorded, the follow- , 3,5 Behaviour ii iorrc from ,omahof juvenile and adult uh'itip '.orn tho Tampa Bay, Florida, arear sand, 3,51 Migrations and local debve L.lgae, iletoins, seagrso particles, movements dJ,fl&grllatee, forarniniforaim. nematodoci po1yuhaeou, ost:oodn, ccpopod, riyeilu, The larvae of pink shrimp are plank- ioopcdu, amphipou, caridean shrimps, oaridean tonic and move vertically in the water eggs, mollusks and fish scales, column, generally ascending at night and descending in the daytime (Jones et al, 17.illiama (1955) and Broad (1965) listed in press), As age increases, the larvae Thodo in shrimp stomachs, but these lists move farther from the bottom, inojudo items ingested by several species of Panneun. For migrations of larvae as related to currents, mee section 2.3, 3,43 Growth rate Migrations of postlarvam also are Growth rates have been calculated ox' Cffeoted by currents, In south Tetnc, eo'tirnated by a number of authors. These rates most postlarvae migrate into the bayo arc .'apovted nc .Lncreases in weight or length "when the bays are being refilled after por uiii imo, Length frequently te given as previous seasons of low water" (Copeland either onrapaos length or total length. Growth and Truitt, 1966), Migration into the of larvaeandpoetlarvae was observed under nursery grounds of south Florida is on th laboratory conditions, Estimates of growth flood tides (Tabb, Dubrow and Jones, 1962; rates for juvenile and adult shrimp are based Idyll and Jones, 1965), According to on marking (releaserecovery) data, and on Hughes (l969a), tidal transport of postlarvae lengthfrequency distributions, For conver- may be initiated by the organisms' response sions of various size measurements, see4.13. to tidal salinity changes,

Rates of growth, am measured by length In the Everglades National Park, juven- increases, vary with size and sex (Iversen and iles migrate usually at night on the ebbing Jones, 1961) and water temperature (Williams, tides (Idyll, Iverson and Yokel, 1964). 1955; Teinsongrusmeo, 1965), As larvae, pink Further observations in the sanie area were ohriem inorease their total length from about made by Idyll, Ivermen and Yokel (1966), 038 rna (nanolli) to 4,1 mm (postlarvas) in who reported, "periode of high relativo 2 J3ik (Dobkìn9 1961; Ewald, 1965), Estimat- abundance occur in spring, summer and early ou of juvenile grorth range from about 7 to fall, but ,,. interyear variation is 52 mm tots.l length per mo (F!i1l1am,l955 considerable." These authors noted that Cou'tello and 1lon, 1959; Costello and Allen, the smallest shrimp usually appear when l61 F!dvurì et al,. 1961; TabbDubTow 3n relative abundanoC is highest, Jcunoon 1962)n ihoroao month!y incromointm of mubadult ond adult shrimp range from O to Several factors may cause migration about 22 mm (Costello and Allen, 1960; of pink shrimp, Tabb, Dubrow and. Jonso ui'rrjen end Idyll, 1960; Iversen and Jones, (1962) saggested that juveniles move to l96l Cootello, 1963; Ku.tkuhn, 1966; Knight, escapo unfavourable conditions anali rio 9í6; TioCoy nd Brown, 1967), sudden reductions in water temperature and salinity, Extensive movements from the 3,44 Metabolism estuaries to the offshore grounds take place as maturity approaches. These move- Stood and Copoland (1967) reported that ments apparently are breeding migration the metabolic rate of pink shrimp in mea (Broad, 1951; Allen, 1966), Tabb and an meocured by oxygen consumption, Jones (1962) suggested that a hurricane io lower then thai of brown shrimp. When caused shrimp to move from south Florida the anImals itero exposed to petrochemical estuaries to ths offshore grounds earlier waste, however, the metabolic rate of pink and at a smaller size than usual, shrimp inoreased, and that of brown shrimp deorased, Juvenile shrimp from the Everglades National Park migrate to the offshore Oumotic regulation in pink shrimp is Tortugas grounds (Iverson and Idyll, 1960; influenced by temperature (Williams, 1960). Cote110 and Allen, 1966), Obmervatjons on Pink shrimp adjust rapidly to salinity changes the timing and direction of pink shrimp in a normal range of temperatures, but at migrations from mouth Florida estuaries to lower temperatures, regulation is more the Tortugas and Sanibel grounls were made difficult, by Costello and Allen (1966), who reported 1517 -

FR,n/5103 Penaeus d. duorarum that juveniles spend from 2 to6mo in the Ogron,1966), In the laboratory, they nursery areas before migrating offshore, 'l'hese have buried in sand to a depth of15.3cm investigators noted that some marked shrimp (Eldred et al,,1961), The depth of traveled at least 277 km before recovery on burrowing may be influenced by shrimp size the offshore grounds, and although migration (Fuss, 1964),and bottom type (Williams, routes were broad, shrimp that left particular 1958), sections of the nursery grounds demonstrated distinot distributional patterns on the The response of pink shrimp to light offshore grounds0 Once they reached the may vary with shrimp size or with light offshore grounds, the shrimp generally ooritinued intensity (Fuss and Ogren,1966), Larvae to move into deep water (Iversen and Idyll, held in aquaria are positively phototrophio 1960;Costello and Allen,1966). (Ewald,1965), In a natural environment, however, the larvae migrate vertioally Migration of pink shrimp in North Carolina away from the water surfaoe during the waters was reported by Mccoy and Brown(1967), day (mee3,51). Juveniles respond posi- Movement out of the estuaries apparently was tively to moonlight by moving to the water influenced by tide. In the croan, migration surface on the ebb tide (Beardsley, in prese). was southward along the coast0 Ordinarily, adults are buried in the nib- strate during daylight and are activo at night (See also 2.21 and 3,23.) (various authors), but juveniles and adults may be active by day under conditions of 3.52 Sohooling low light level (Eldred et al,, 1961), Juvenilem, moreover, tend to be more active The movements and catches of the shrimping than adults in the daytime (Eldred et al,, fleet indicate that, at times, pink shrimp 1961;1'uss and Ogren 1966; Hughes,l9), may sohool schooling may be by size, Fuss and. ogren(1966)considered burrowing aotivity a direct effot of light at the Stookm of pink shrimp probably mix time of the burrowing', Wickham(1967)and frequently. Costello and Allen(1966) Hughes(1968),however, presented evidence reported some overlap between stocks of the that diurnal activity of juveniles and Tortugas and Sanibel grounds, Florida, young adults i influenced partly by They stated, however, that "Apparently, previous light-intensity experience, Tortugas shrimp do not migrate to the Sanibel grounds and migration t'von the Sanibel to the Under laborntory oonditions juvenile Tortugas grounds is minimal," shrimp showed nocturnal activity peaks that probably were related to tidal rhythm Pink shrimp are taken frequently in trawl (Wiokham,1967), Activity of juveniles' catches with several other species of shrimp, also may be affected by water movement arid Williams (1955)observed that in North water height (Wickham,1967),and that of Carolina, the seasons during which white juveniles and adults by feeding (Hughes, shrimp, brown shrimp and pink shrimp appear 1968). Hughes(l969a)found that salinity in the nursery grounds are fairly distinct; affected the tide-associated movements of hówver, there im some mixing of postlarvas, postlarval and juvenile pink shrimp. Adults mi wherever they occur in the same general area, Tabb, Dubrow and Jones(1962)observed that the oxygen depletion and buildup of Vrioal movements of pink shrimp were hydrogen sulphide that followed a hurricane reported by Costello and Allen(1968),who caused pink shrimp to swim at the water observed shrimp in dense mohools "near the surface in full sunlight, surface at night," Concerning night-time vertical movements of migrating juveniles, See2,3for responses to salinity and Beardsloy and Iversen (1966) stated: "Ari currents, and2,3and3,16for effects of' average of about91%of the shrimp oaught temperature, on full moon tides were taken on or close to the surfaoeonly about75%were oaught in the Artificial stimuli surfaos layers during new and quarter moon tides," Burrowed pink shrimp responded to mechanical stimulation of their dorsal 3,53 Responses to stimuli body surfaces by burrowing deeper into the substrate (Fuss,1964), Continued agitation vironnìental stimuli of' the dorsal surfaces or stimulation of the animals' sides caused -the shrimp to hop Pink shrimp burrow into the substrate as vertically out of the substrate, "a protective measure against predators and adverse environmental conditions" (Fuss and Photo-orientation of juvenile and young 1518 -

FRis 810Penaeu6duorarum aduii; chrimp was studied by Jachoweki and. Eleotrioal stimuli Myrbc'(1968).The shrimp were attracted ThOi'& cquent1y toward light than toward Pink shrinip9 mostly adolt, subjected to arknees, and toward higher intensities and interrupted direct current9 responded by ìoner wavelengths than toward lower intonsi- flipping, tail first1, to9ard the anode ties ic1 shorteí' wavelengths.Matimum (Rigman, 1956).This galvanotaxic reaction, arfjcl hotoactjvatjon of juveniles and affeoted by shrimp size and position9 pulse you wan during the full moon9 mm- width and water temperature, causes pink ims 'u the flew moon (Aaron and Wisby9 shrimp to hop from their burrows (Ioscler9 1964), hoe.oivation and photoaotio 1965),The intensity of the responso lessens drive varied oith shrimp elze buriot with when thshrimp is shocked repeaodly (Nelson, sezo 1962), 1519

FRm/SJ.0 Penaeua d. duorarum 1

4 POPULATION April and the largest ones (about 140 mm total length) in September, 4.1 Structure The size composition of captured pink 4.11 Sex ratio shrimp is affected by the time of day that fishing is done, In Tampa Bay, the largest Saloman(1965)reported a sex ratio of shrimp were caught at night (Eldred et al., about 1:1 after examining 11,695 juvenile 1961; Saloman, 1968), pink shrimp from Tampa Bay, Florida, Idyll (1964) also reported a sex ratio of about Conversions of size measurements were lU, but he noted (as did Broad, 1951) that given by Kutkuhn (1962),Kutkuhn (1966), the ratio may vary with season and size of and Fontaine and Neal (1968), Formulas shrimp, for the total length-total weight relation- ship in pink shrimp are as follows: 4,12 Age oompomition female, w 5,06 z Age composition of' the commercial catch male, w 4.49 z lo623.13(Kutkuhn, 1966), varies with recruitment, migration, mortality (see 3,11,3,12 and 4.12) aM praotioes of the fishing fleet, 4.2 Abundance and densjtj of Age at various life atages can be estima- population ted. with size-ago conversions provided by Kutkuhn (1966). The youngest pink shrimp 4,22 Changes in abundanoe for commercial use are captured in the bait shrimp industry, Theme shrimp are about Fiehing effort and practices are 6 wk old, sinos Saloman (1965) has reported variab1, but in certain areas, uoh as the bait shrimp marketed as small as 47 mm total Tortugas grounds, landings may reflect length, Male and female shrimp that reach actual changes in abundance, Nonth1 sexual maturity at 75 and 85 mm total length, changes in abundance by area (Fig. 5), respectively (Eldred et al,, 1961), may be were compiled from Fishery StaticUom of' 9 or 10 wk old, Kutkuhnl966) gave 15 wk the United Stat (United Staboz Bureau of (107 mm total length) as the age when shrimp Commercial Fisheries) and Kuthuhn, l962n. are first recruited to the Tortugas fishery, Jnnual landings by United Statem vessolo wiere they have a fishable life spari of about are listed in Pable I, For areas of capture, 68 wk, Halmo estimated 83 wk as the see Fig, 2. average maximum age, Absolute maximum age is not known, but F1dred et ai. (1961) suggested Seasonal and geographic changes in that specimens 200 mm total length may be abundance as well as annual fluctuations 2 yr old or more, may result from varying oceanographio conditions, (See 2,3, 3,16 and 4,33,) 4.13Size composition The exploitation of newly recruited Size composition varies similarly to age shrimp may affect annual yields in the oomposition, (See 4,12,) Tortugas fishery (Kutkuhn, l962a),

It is generally true that small, immature Annual landings from the Tortugas pink shrimp live near shore, and largo, mature grounds showed "No markmd changes or trends individuals offshore. Size increases with ,.," from 1951 to1965,although they depth of water (Iversen, Jones, and Idyll, fluctuated from year to year (Berry, 1966). 1960), Shrimp caught In Tampa Bay, Florida, ranged from 11 (Eldred et al,, 1961) to 144 mm 4,24 Changes in density total length (Saloman, 196g.On the Tortugas grounds, however, shrimp ranged from about Kutkuhn (1962a) found that oatch-psr- 49 to 230 mm total length (Iversen, Jones unit intensity values were highest in the and Idyll, 1960), Tortugas fishery in the late autumn and lowest in the spring, Fie noted a downward Size composition also varies seasonally, trend in the Tortugas annual population In Tampa Bay, the smallest average size (about from 1956 to 1959, but a population buildup 31 mm total length) was in the summer and the from 1956 to 1958 for the Apalaohioola largest (about 82 mm total length) in the (Florida) area, followed by a decline in lato spring (Eldred et al., 1961).Length- 1959, frequency data from the Tortugas fishery (Wlteelor, Benton mM Hudson, 1965) showed On the Tortugas grounds, shrimp are that shrimp with the smallest average sizes most dense in the hnl1oweet wators (about 95 mm total length) were oaught in least dense in the deepest waters (Lindner, 1966), 1520

FRa 310, Penseus d duorar

) ) ) SOUTH ATLANTIC SANIBEL- TORTUGAS APA LAC H ICOLA PENSACOLA- MISSISSIPPI RIVER OBREGON- CAMPECHE 1,400-

1,200- \ ¡N N u-1-j IO 'I 3 \i

800- \ I-O

600- w z 400- I uI- o 200

Fig.5 jor United States pink shrimp fisheries (seasonal abundanoe se indicated by average monthly landings 1957-59). $tates vessels and area of ual landings (metric tons, ihole weight) o,f pink shrimp by United sot available for all areas capture, 1951-661/. TA.BLß I Data for earlier years Year Carolina North Georgia-FloridaSouth Carolina east coast Sanibel- Mtxico, 1/ 1951-55 from Idyll (1964); 1956-66 compiled from Fishery Statistics of the United States, Tortugas Apalachicola MississippiPensacola to River MississippiRiver to Texas Texas vest of94°Wlong. Obregon-Mexico, 1951 publiahed annually by the United States ßureau of Coimiiercial Fisheries. Campeche 1952 1953 1954 1955 -- - 4,952.47,529.38,314.2 7,893.9 8,916.3 - -- --

------3,550.6 13,961.3 9,931.2 10,748,6 1956 -_ -- 9,246.9 577.9 345.5 1.1 26.9 2.1 10,601.510,498.4 19611960195919581957 556.3934.8368.8961.0792.8 17.4 0.34.7 0 10,670.110,665.3 8,070.86,011.27,209.9 1,080.5 343.0388.2904.6 151.84.8 121.4380.9 49.182.0 0.03 7.24.48.20.06 30.347.354.6 6.93.0 3.14.36.9 5,306.8 0 8,192.47,998.97,085.89,193.4 19661965196419631962 1,018.0 240.1251.4765.3878.5 0.85.21.0 0 9,649.09,306.09,013.57,733.76,391.9 1,408.8 1,304.81,176.8 453.9682.9 154.6 36.429.070,045.6 1.23.13.50.30.04 23.210.929.4 8.67.7 4.62.92.46.28.0 2,772.67,896.28,361.17,840.78,753.1 1522 -

4j FEin SlOPenaews d. duorarun 43 Natality and recruitment 4,42Factors causing or affecting 4,31Reproduction rates mortalIty For survival raten of larvae, neo 3.22 Causes cf mortality ax's varied,Period- and 4.41. ically, large numbers of pink shrimp perie as a result of hurricanes (Tabb and Jones, 4,32Factors affecting reproduction 1962),Occasionally, red tide (mani»e water which is toxic due to exoessiv (See 2,3 and 3,16,) concentrations of the d.inoflageliate, nium breve) causes mass mortality of 433Recruitment pink shrimp rountor, Smith and WiiiJ.eme, 1947), For recruitment of poetlarvac to the eetuaries, moe 2,21, 4,43Factors affecting morbidity The relative abundance of juveniles (See 2.3 arid 3.35.) emrating from Evorgladec National 'ark oan be pocitively correlated i:ith the subsequent 4.6The population In thin communfl oonimeroial catoh on the Tortugac grounds a cos st em (Yokel, Ivorsen and Idyll, 1969), The optimum habitat for pink ehx'imp Migration studies (Costello and Allan, varies with shrimp size (Eldrcd, 1962), 1966) indicate that young adult shrimp aro Young shrimp prefer quiet, clear, shallow recruited to the Tortugas fishery year around, waters and firm substrates, with protective Maximum recruitment is in the spring and growth such as turtle grasa (Viosca, l957) autumn (Berry, 1966),In more northerly pink According to Hoese and Jones (1963), shrimp fisheries, recruitment io seasonally charactenistio associates of juvenile shrimp more restricted (Fig5), in a south Texas turtle grass community were rainwater killifish1 Lucaniparva; For factors that influence recruitment, mojarra, Cerros embreas; pinfish, gdcn see 2,3,3,16, 3.34, 3.43, 3.51 and 4,42, rhcmboides; goby, Gobiosa nobuotum; nrnd crab, zanopc texena; end granr hrimp 4,4Mortalitand morbidi Palaernonetes puio, 4.41Mortality rates Adult shrimp typ1al1y inhabit offshore waters that have no soagrassee,,The faunal Mortality rates for the larvae of pink associates of adult shrimp on thCampahe shrimp have been estimated as a nearly copstant grounds in the 11.- to 29-m depth tholuded 17 percent per day (Munro, Jonce and Dimiirtou, the followingsfighting ccnch, Stronibus 1968),only a small percentage of the larvae alatus; West Indian chunk, Xonou iclatua; produced survive to become juveni1es pontunici crab, Portunuc sc5.nisonunmn,oeth cd grunt,athyn cmxi aarojinoe rísior Estimates of mortality in adult populations and porgy,tonoomue eaynuc hdcbcod-, are based on data from mark-recovery experi 1955),, (See oleo 2,2 and mento and, in crie study (Berry, in presa), on the de]ne in abundanos of age groups.In the eatiniatse in Table II, derived from expon- mento performed in waters off south Florida, Z te the instantaneous total mortality coefficient; F in, the instantaneous coefficient of mortality caused by fishing; and X is the Luj-ntaneoucoefficient of other 1osse i ho marking theory, including true natural riora11ty (M) plus losses of individuals hoh for any reason become unavailable for recapture. stimatos of mortality rates of pink shrimp in Florida waters TABLE II Z F X Rates of

Iversen (1962) 0.02393 0.05998 Fishing 1-week period Other losses BerryKutkuhn (1967) (1966) 0.22-0.27 1.51 0.160-0.227 0.96 0.024-0.061 0.55 14.8-20.37. 627. 2.4-5.97. 62 Berry (in rsss) 0.10-0.16 (males) 0.07-0.12(females) Rates of reduction per Costello andAllen (1968) 0.3570.233 0.13850.0689 0.20.1644 185 13.1% 6.8°ì 2-week period 19.7%14.87. -1524 -

FR ' Penas us d duorarum

5 EXPLOITATION Fishing for pink shrimp is usually at night, because these animals typically are 5,1 Fishing equi buried in the bottom during the day, Recently, trawling gear was developed 5.11 Gears that employs an electrical field whioh causes pink shrimp to leave their burrows The fishing gears used in estuaries or (Pease and Seidel, 1967) and enables in relatively shallow near-shore waters to fishermen to take shrimp in daylight, capture young pink shrimp for bait or food vary regionally, In North Carolina, fishermen Descriptions of various types of use otter trawle (Broad, 1951a) and channel shrisping gear were given by Dumont and nets (Guthrie, 1966). Another type of channel Sundstrom (1961), net is used in Florida (Higman, 1952). Aleo used in Florida are pushnets (De Sylva, 1954), 5,12 Boats dipucts (Iversen and Van Meter, 1964; Joyce, 1965), bridge nets (Higrnnn, 1952), and roller Shrimp boats range from 4-m skif fo frame trawis (Woodburn et al,, 1957; Tabb, powered by outboard motors to 34-m vessels 1958), On the Yuoatán Poninsula, Mexican with a gross weight of 118 metric tons fishermen uso beaoh seines and traps (Idyll, powered by 360 hp diesel engines, 1964), Most vessels used in shallow waters In the shallow water United States have wooden hulls and shallow draft and are fisheries, most pink shrimp are caught with less than 15nilong. They are powered by roller frame traw1n, These trawls usually are engines that burn either diesel or gasoline constructed of iron pipe and rods and. have fuel; most engines are 140 hp ox' loss, rectangular openings 1,8 to 3.0niwide and 61 to 77 orn deep. The bottom parts of the Typical shrimp trawlers used offshore frames servo as attachments for trawl flots are 18 to 21 m long, are powered by a diesel that terminate in ood ends similar to those engine of 200 to 300 hp, and have gross used on small otter trawls, This gear is weight of about 61 metric tons, Hulls are effeotive in grassy areas in shallow water, usually wood, but now tee1 hulls are being The rollers (about 127 mm in diameter) move built and are recommended espeoially for over the seagrass without uprooting it, and boats longer than about 18 m, These boato rakes, built on the frames, prevent large generally have a singlo propeller and in- objects from entering the frame mouths. corporate a 3-drum winch with a power (Otter trawls used in grassy areas soon fill takeoff from the main engine to handle -the rii.h upr000dgrass and are ineffective,) otter trawls and trynet, The development iol1er frama traiils used to catch live shrimp of large vessels with more power has been 'oí hait nro omptied about every 10 to 15 min. necessary to acoommodate the double-rig trawls and large otter trawls now in unie, Otter trawls are the principal gear fished in deep water, The trawls used in the Most shrimp trawlers are of conventional United States have been described by BulliB hull design, although at least one catamaran (195l)n Knake, Murdook and Cating (1958), and (double hull) shrimp trawling vessel, 21 in Robas (1959), The mesh size of the cod end long, was launched and used in the Gulf of of he raw1ing gear used inshore is typically Mexico. Aluminum hulls and fibreglaso 251 o 41,4 mm stretoh measlu'e, whereas the reinforced hulls may be used in the near meith sine of that used offshore is usually 50.8 futuro (Captiva, 1967), to 57.2 mm, Thread sizes range from about 36 -o 48 in the cod ends, and from 12 t.o 18 Ringhaver (1960) described the design in the body and wings of the trawls, Doors and rigging arrangements of shrimp trawling of the otter trawl vary with trawl size and vessels, vooscl horsepower, Until about 1957, single oo' raw1tc with openings 24 to 31niwere 5,2 Fishing areas uoocl In hc Uniad States, but at present, mos voosols fich two otter trawls, eaoh 5,21 General geographio oponlng 12 o 14 m, Also, a third, small distribution oìux' rowl-- the trynet-- is fished to determine the density and size of shrimp at Pink shrimp are fished throughout moot o paiou1ar location. If it produces good of' their range, Williams (1965) observed, ontohus, thu largo nets are lowered. Trawls however, that pink shrimp are not uniformly aro dragged cboui 3 h at a speed of 2 to 3 kn abundant and that fisheries are "oonoentraod (307 o 505 los/x) and then taken aboard, smp- at diverso points," cud reao, 1525 -

'5,2 FRm/S103Pea?u d cluorarum Fishing areas are ehown in Fig. 2. Major Fishing seasons vary with the type of fisheries for bait or food shrimp in estuaries fishery and with geographic location, In orhal1ow inshore waters are in North Carolina, North Carolina estuaries, juvenile pink Florida and Campeche,Mexico (land areas 238 shrimp are caught for food during May, June and 3115 (Burkenroad, 1949; Woodburn et al., and July (Burkenroad, 1949). In south Florida 1957; Idyll 1964; Joyce, 1965; Costello and estuaries, such as Biaoayne Bay, juveniles Allen, 19665. The major offshore fisheries are taken for live bait throughout the year are off North Carolina, the Florida west coast, (Saloman, Allen and Costello, 1968), Here, Alabama, Mississippi, and Campeche (land areas peak catches are in the winter and spring 238, 235, and 311) (Broad, 1951; Kutkuhn, (Weloh, 1965; Jones and Smith, 1966). 1962a), By far the largest pink shrimp fisher- ies are the Tortugas fishery off the southwest Seasonal variations in offshore coast of Florida and the Campeche fishery off fisheries for adult pink shrimp ar, shown the west coast of the Yuoat&n Peninsula (Kutkuhn, in Fig, 5, 0ff North Carolina, South 1962a), Carolina, Georgia, and northeast Florida (South Atlantic fishery), shrimp are taken 5,22 Geographic ranges from April to December; catches are greatest in June and October, Off southwest Florida The distance of fishing areas from the (Sanibel.-Tortugas grounds), the fishing ooat varies throughout the pink shrimp range season is year around, and catches are and with the type of fishery. Bait shrimping highest in the winter and spring. The often takes place a few meters from shore, fishery near Apalaohicola, Florida, is from The Tortugas fishing grounds, however, extend March to August with greatest catches made from about 83 to 222 km from the mainland in Nay, In the Pensacola-Mississippi Hiver coast (Costello and Allen, 1968), area, shrimp are taken from April to August arid peak landings are in June, The Obregon- 5.23 Depth ranges Campeche grounds produce large quantities of shrimp throughout the year with peaks Depths in which pink shrimp are caught in the winter and early mummer. vary regionally and with the type of fishery. Bait shrimp ax's taken in waters as shallow 5,4 Fishing operations and results as 0.5 m and most adult shrimp arc caught in waters between 9 and 60 m. 5.41 Effort and intensity

(See 2.22.) The unit cf effort in United States otter trawl fisheries for pink shrimp im 5.24 Conditions of the grounds designated "days fished", This unit, representing 24 h of fishing effort, i Offshore shrimp grounds are usually used by the United States Bureau of areas of smooth bottom euitable for trawling. Commercial Fisheries in their statistical In the eastern Gulf of Mexico, they "are reports for uniformity in tabulation of characterized by sand, shell and coral gravel; data, and by live coral overlying white, gritty, oaloareous mud" (Springer and Bullis, 1954). Fishing effort also is reported by Typically, the bottom on the Tortugas grounds the United States Bureau of Commercial im silt and mud, with 20 to 25 percent white Fisheries as the number of trips made by coral sand (Idyll, 1964). The Tortugas and oomrneroial fishing vessels in a given time Campeche grounds have dead. ehell and patches period. Landings pa:' uot of 'ishiig effort of coral and sponge (Springer and Bullie, may be calculated from U.S. "tatistloal imt 1954; Hildebrand., 1955). because landings and effort are repu:' for specific areas. A few reports dealwith For conditions of the estuarine and catches per unit arec, or both, inmhore fishing grounds, see 2,3. The following tabulation, abridged from 5.3 Fishing seasons Kutkuhn (l962a), shows offshore fishing effort in the Gulf of Mexico primarily for 5,31 General pattern of seasons pink shrimp, The units represent the numbers of 24 h periods fished,

Area Year

1956 1957 1958 1959

Sanibel-Tortugas 17,519 17,356 20,690 17,098

Obregon-Campeohe 22,236 21 491 16,899 l9709 1526

/5103 Penaeu d, duoraruin 5:3

Fishing effort and intensity vary because lY2 or 2- in [38.1- or 50.8_mmJ mesh of shrimp availability, size of shrimp, market nets...trawls with largo meshes throughout price, weathor conditions, vessel efficiency, will permit some marketable shrimp to escape mciregulatory measures (Iversen and Idyll, and therefore ooulcì profitably be used only 1959;Iverson, 1962). In addition, effort in certain situations." is affected by: (i) seasonal changos in the number of hours of darkness; (2) changes in Market preference affects the sizes of moon chase, i.e., fishermen report poorer pink shrimp retained by fishermen in same omiohes when the moon is full than when it is areas. On the Tortugas grounds in 1963, waning or waxing; and (3) changes in amount approximately il peroent of the total catch of jellyfish (Aurelia sp.) or algae, i.e., (by weight) was not landed because the shrimp excessive amounts clog noto (Iverson and Idyll, were too small for the market (Wheeler, 1959). Benton and Hudson, 1965), Discarding declined from 1963 to 1966, apparently be- During the period l95O.59, fishing cause of the increased value cf the small effort increased on the Tortugas grounds but shrimp (Berry and Benton, 1969), total catch "remained fairly (Ivereen and Jones, 1961). Available 5,43 Catches information mugesto a declino in thcatch per unit of effort during this period. Monthly and annual reoordm of catches of bait shrimp (mostly pink shrimp), although 5,42 Selectivity not complete, are given in Summarof Florida Commercial Marine Landiri e, which is publish- The selectivity of various sizes of trawl ed annually by the Flori. a Board of tìonser mesh in catching pink shrimp was reported vatioua., by Megan, Idyll and Iversen (1957).They reported escapement of pink shrimp through Total annual yields of pink shrimp in aoci ends of varying mesh sizes as shown in the offshore waters of the United Stetes Table III, are given in Fishery Statistics of the United States, published by the United Berry and Hervey (1965) ,5-tudied the States Bureau of Commeroial Fisheries, selectivity of various mesh (stretched) sizes Table I lists annual landings of pink as applied to Gulf of Mexico shrimp (species shrimp by United State vessels, A com- not given) and suggested that, because large parison of yields from different areasis mesh increases fishing power, "the oatoh of showu inFig. 5, large chrimp by nets with 2/2 and 3-in E63,8- and (602-mn meshes should be approximately 15 cud 20 peroent greater than the catch by - 1527 -

18103 Ponaour I 4 d. duor

PABLE III

ooapement of pink shrimp through cod end. of varying mesh sizes

Mesh size' in Shrimp size Escapement cod end (mm) (number of tails through cod per pound2) end (percent)

44.5 110 10 83 5 51 0

50.8 110 50 83 10 51 0

57.2 83 30 51 10 33 0

63.8 83 40 51 15 33 5

Measurement between the midpoints of knots (stretched mesh)

2 1 pound equals 0.45 kg 1528

F1tnjSlQ, P nasue dduorarwa 6i

6 PROTECTION liND M'INAGEMENT of regulations applying to pinkhr1mp anh recommended that the elzelimit law be 6.,. Regulatory (legislativemeasures repealed., He observed that "The sise distribution of our shrimp on the fishing 6.,l]. Limitation or reilu.ction of total catoh grounds is such that sise limits oauee more waste than benefit.," The efficiency of gear fishing for pink shisp bas been oontrolled in oertain Florida Regulations specifying a sanctuary for ficheries., Limits are placed on the size of small pink shrimp were applied in a section tatding gear used to take live bait ehrimp. of the Tortugas grounds in 1957, The area Thí rgnlation limits the total catch in is olosed to trawling when it is determined sews that shrimp smaller than50to the pound (o45kg), heads off9 predominate In the area., 6,12 Protection of portions of population In the United States, regulations that apply to commercial shrimp close oertain The usefulness of certain laws to protect areas at speoified seasons in States bordering small shrimp Is the subject of some contro- the Gulf of Mexico, and in Georgia and the versy. For example, a1956Florida regulation Carolinas., placed limitations on the mesh size of ood enm ofrawis used on the Tortugas grounds Generally, no restrictions are placed but the regulation was later withdrawn (Idyll9 on the use of pink shrimp., For the fishery 1957)., Another Florida regulation Is a "size- In Biscayne Bay, Florida, however9m. regul- limit« law designed to prohibit the taking ation specifies that shrimp are io be taken of undersized pink shrimp in the Tortugas only for uso as livebait0 area, Lindner (1966) reviewed the problem -1529 -

510

POND CULTURE 7,4 Holding of stock

Pond cultivation of pink shrimp is just Ewald(1965)held larval pink shrimp beginning and information on the culturo of individually in oompaz'tmented plastic boxes, this species is soarce. Information on culture Survival was better in Gulf Stream water of related shrimp species is provided by Cook (oceanic) than in Biecayne Bay water and Lindner(1970),and Lindner and Cook (estuarine). Survival was beet end devel- (1970). opment time least at a water temperature of 26° C. 7.1 Procurement of stocks Costello and Allen(1959)held. pink Ewald (1965) optured large, ripe female shrimp in a small salt-water pond with a pink shrimp on the Tortugas grounds with a sand-mud bottom for3Y2 sowhile the shrimp trynet to obtain spawning stock. The shrimp grew from juvenile to adult size. were transported to the laboratory in plastio garbage cans. In a later experiment, juvenile shrimp were held in the same pond at. water tempera- 7.3 Spawning (artificial, induced, tures ranging from26° to 3300 and salin- natural) ity from27 to 36°/oo(Costello and Allen, 1961), Laboratory spawning of shrimp captured on the Tortugas grounds was described by 7.6Foodsfeedjng Ewald(1965) "At the laboratory, individual shrimp were placed in 15-gallon running Ewald(1965)provided a mixture of seawater aquaria. Spawning, if it occurred, several unicellular algae and a marine yeast was within two days after return to the as food for early larval stages of pink laboratory and always in the very early hours shrimp. The last mysis stage and early of the morning, Water temperature in aquaria postlarvae were fed nauplil of brine shrimp at times of spawning was27°and29°C, The (Art ernia), eggs, being demersal, settled to the bottom of the aquaria. They were present in such groat numbers that they were visible to the naked eye and appeared like fine, white powder," 1530 -

'Re/Si0 Penneus d. duorarum 8,1 e IIEFEREhCES

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D.MO and T.J. Costello, Releases and recoveries of marked pink shrimp, Penmeus duorarum 1966 Burkenroad, in south Florida waters1958-64, Data Rop.Fish Wildl.Sorv.U.S., (ii)s 2 microfiches

Allen. D.N., J.H. Hudson and T.J, Costello, Postlarval pink shrimp, Penaeus duoraruinduorarum, (MS) in the Florida Keys. Miami, Bureau of Commercial Fisheries, Tropical Atlantic Biological Laboratory,20 p.

Allen, J.A., The rhythms and population dynamics of docapod Cruetacea.Ooeanogr.mar.Biol.., 1966 4x247-65 Anderson, W.W., Recognizing important shrimps of the south.Fisher Leafl.Fish Wildl.Serv.U.S., 1962 (536):5p. (Rev.) Axderson, W.W, J.E. King and M.J. Lindner, Early stages in the life history of the common 1949 shrimp Penaeus sotiforus (Linnasus). Biol.Bull,sar.biol.Lab., Woods Hole,96(2), 168-72

arden, C.M., Notes on the postlarvee of commercial shrimp (Ponaeus) in South Carolina. 1961 Contr.Bears Bluff Labe,(33)8 p.

Beardsley, G,L., Jr., Distribution of migrating juvenile pink shrimp, Penasus d. duore.rwn (in prose) Burkenroad, in :Buttonwood. Canal, Everglades National Park, Florida. Traxis.Am.Fish. Soo., Vol. 99 Beardsley, G.L,, Jr. and ILS. Iversen, Studiès on the distribution of migrating pink shrimp in 1966 Buttonwood Canal, Everglades National Park. (Abstract) Proo,Gulf Caribb.Fimh.Inst,, 18,17 Berry, R.J., Shrimp population studies, Circ,Fish Wildl,Serv, Wash.,(246)246 1966

Dynamics of the Tortugas (Florida) pink shrimp population.Thesin, University l97 of Rhode Island, Kingston, 160 p. Probable rangs of total mortality ratos for penaeid shrimp, Proo,Gulf Qaribb, press 22 Barry, R,J, ano ILC. Donou, Disoarding practicas ín parts of the Gulf of Mexico . 1969 ?A0h,I.9(57)Vol.3983-99 rry, R,J. and J.B. Hervey, Meuh selectivity utudies Ciro.Fish Wildl.Sorv. Wa,(230)4l-4 1965 Broad, A.C,, Resulta of shrimp research in Morth Carolina,Proo,Gulf Caribb.Fish,Inat.,327-35 151

The shrimps in 1orth Carolina, In Survey of marine fisheries of North Carolina, l951a ditsd by H.P. ayior, chapel Hill, University of North Carolina Press,pp.191-204 Environmental requirements of shrimp. Puhl,Hlth Serve Wash.,(999-WP-25)s 6-91

Bul.o, LI.! jr., Gulf of Mexico shrimp trawl designa. Fishery Leafl,Fish Wildl.Serv.0 S 95 (394)l6p. Burkenroaì, MAD., I'urthor obeswations on Penaeidae cf the northern Gulf of Mexico.Bu11, 1939 JJinr!h ao,Co1lri, 6(b)l-62 1531-

8:2 5].03 Penasue d, duorr

Burkenroad, M.D., Oocurrenoe and life histories of commercial shrimp. Science, N.Y., 110 1949 (2869)s688-9

Captiva, P.3., Ie in. shrimp trawler design and construction over the pant five decades. 1967 Proo,Oulf Caribb.Fish,Innt,, 19:23-30

Christmas, J.Y., G. Gunter, and P. Muagrave, Studien of annual abundance of poatlarval penasid 1966 shrimp in the ostuarine waters of Mississippi an ro1atd to subsequent commoroisi oatohea. Gulf Res.Rep. 2(2)s177-212

Chuansri, C., A morphomotric and merintic study of postlarvul brown shrimp, Penaeuc asteono 1968 Ives, pink shrimp, P. duorarum Burkenroad, and uhito shrimp, P. cotifoz'ua (Linnneuo), Thesis, University of Miami, Coral Gables, 108 p,

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Survival of stained, tagged and unmarked shrimp in the presence of predators. 1962 Proc,Gulf Caribb,Fish,Inst., 14:16-20

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Mortality rates in populations of pink shrimp, Penaeus duorarusi, of the Sanibel 1968 and Tortugas grounds, Florida, Fishery Bull,Fish Wildl.Sorv.U.S,, 66(3):491-502

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FErn $103 Ponneus d. duorarum 83

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HiCuan, J.B., Preliminary investigation of the live bait shrimp fishery of Florida Bay and the 1952 Keys. Miami, University of Miami, Marine Laboratory.Report to Florida State Board of Conservation, (ML 3789):8 p.

The behavior of the pink grooved shrimp, Ponaous duoraruin Burkenroad, in a direót 1956 current electrical field. Tech.Ser.Fla St.Bd Conserv., (l6):23 p.

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Hoese, H.D., Biotic changes in a bay associated with the end of a drought. Limnoi.Oceanogr,, 1960 5(3),326-.36

Hoese, H.D. and R.S. Jones, Seasonality of larger animals in a Texas turtle grass oommunity, 1963 Pubis Inst.mar.Sci.Univ.Tex,, 9z37-47

Hoses,H.D. , ,Fauna of the Aransas Pass Inlet, Texas, 3.Dialand seasonal variations in 1968 trawlablo organisms of the adjacent area. Tex,J.Soi,, 20(l):33- ,0 - 1533 - 84 FRm/S].03 Penasus d. duoraru Hoithuim, L.B. and B. Rosa, Jr.,List of species of shrimps and prawns of eoonomio value. 1965 FAO Fish.tech.Pap., (52):21 p. Hughes, LA.,Factors oontrolling emergence of pink shrimp (Penaous duoraruxn) from the substrate. 1968 Biol.Bull,mar.biol,Lab., Woods Hole, l34(l):48-59 On the meohaniems underlying tide-associated movements of Penasue duorarum. 1969 FAO Fish.Rop., (57)Vol,3,867-74 Responses to salinity ohange as a tidal transport mechanism of pink shrimp, 1969a Penaeus duorarum.Biol.Bull,mar.biol.Lab,, Woods Hole, 136(1):43-53 Hutton, R.F..,A second list of parasites from marine arid coastal animals of Florida.Prans.Ain. 1964 mioroso.Soc., 83(4):439-47 Hutton, R.F. arid B. Hidred,A saroopkiagid larva (Diptera) from the terminal ampoule of the 1958 pink shrimp, Penaeus duora.rum Burkenroad,J.Parasit., 44(1): 27 Hutton, R.F., T. Ball and B. Eldrod,Immature nematodes of the genus Contracaeoum Railliet and 1962 Henry, 1912, from shrimps.J.Parasit,, 48(2)s327-32 Hutton, R.F, et al.,Investigations on the parasites and diseases of saltwater shrimps 1959 (Penaeidao) of sports and commeroial importance to Florida.(Prelim. Rep.)Tech. Ser.Fla St.Bd Conserv., (26):38 p., Idyll, C.P,.,The commercial shrimp industry of Florida.Eduo.Sar.Fla St.Bd Conserv., (6):28 p. 1957 (ML 17114) A summary of information on the pink shrimp, Penaeus duorarum. Paper presented 1964 to the CSA Specialist meeting on Crustaceans, Zanzibar, April 19-26, (64)lOa:22 p. Idyll, C,P, and A.C. Jones,Abundance and distribution cf pink Bhrimp larvae and postlarvas 1965 in southwest Florida waters,Circ.Fish Wildl.Serv., Wash., (230)s25-7 Idyll, O,P,, LS. Iversen and B. Yokel,Movements of juvenile pink shrimp in the Everglades 1964 National Park, Florida,(Abstraot).Proo.Gulf Caribb.Fish.Inst., 16:51-2 Abundance of juvenile pink shrimp on the Everglades National Park nursery grounds. 1966 Circ.Fish Wildl.Serv., Wash., (246):19-20 Ingle, R.M. et al.,Preliminary analysis of Tortugas shrimp sampling data 1957-58.Tech.Ser. 1959 Fla St.Bd Conserv., (32):45 p. Iversen, E.S.,Estimating a population of shrimp by the use of catch per effort and tagging 1962 data,Bull,mar,Soi.Gulf Caribb., 12(3): 350-98 Iversen, LS. and C.P. Idyll,The Tortugas shrimp fishery:the fishing fleet arid its method of 1959 operation.Tecb.Ser.Fla St.Bd Conserv., (29):35 p. Aspects of the biology of' the Tortugas pink shrimp, Penaeum duorarum.Tranm.Am. 1960 Fish,Soc., 89(1)l-8 Iversen, E.S. and A.C. Jones,Growth and migrations of the Tortugas pink shrimp, Penaeus 1961 duorarum, and changes in the catch per unit of effort of the fishery.Teoh.Ser.F1a St.Bd Conserv., (34):30 p. Ivereen, E.S, and R.B. Manning,A new miorosporidian parasite from the pink shrimp (Penaeuu 1959 duorarum). rana.Am,Fish.Soc., 88(2)':130-2 Iversen, LS. arid N.M. van Meter,A record of the microsporidian, Thelohania duorara, para- 1964 sitizing the shrimp, Penaeum brasilienmis.Bull.rnar,Soi.Gulf Caribb., 14(4):549-53 Iversan, LS., AJ. Jones and C.P. Idyll,Size distribution of pink shrimp, Penaeus duorarum, 1960 and fleet conoentrations on the Tortugas fishing grounds.Spec.scient.Rep.U.S.Fieh Wildl.,Serv.(Fisb.), (356):62 p. -1534 -

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Jona, A.O, etal., Distribution of early developmentalstages of pinkshrimp, Penaeus duorarum, (inpraae)in Florida waters,, Bull,mar,Sci.

Jones, IL, and J, Smith, Swnmary of Florida commercial marine landings, 1965.Tallahassee, Florida 1966 Board of Conservation, Salt Water Fisheries Division, Marine Fisheries Research, 61 p.

Joyce, E.A., Jr., The commeroial shrimps of the northeast coast of Florida. Prof.Pap,Ser.mar. 1965 Lab,Fla,(6):224 p.

Kessler, LW., Electrical threshold responsos of pink shrimp, Penasus duorarum Burkenroad. 1965 Bull,mar,Scl., 15(4):885-95

Knake, B4O., J,F. Murdock and J.P.Cating, Double-rig shrimp trawling in the Gulf of Moxioo. 1958 FisherLeafl.FishWildl,Serv,TJ,S., (470):].l p.

Knight, C,E., Mark-recapture experiments. Circ.U.S.Fish Wildl,Serv., (246)t21.-3 l96 Kruse, D.N., Parasitos of the commercial shrimps, Penaeus aztecus Ives, P. duorarum Burkenroad, 1959 and P. setiferus (Linnaeue). Tulane Stud.Zool., 7(4):l23-44

Kutkuhn, J.H., Conversion of "whole" and "headless" weights in commercial Gulf of Mexicoshrimps. 1962 Spoc,scient.Rep,Ij,S.Fish Wildl.Serv,(Fish.), (409):7 p,

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Moore, D.R., Distribution of the sea grass, Thalassia, in the United States. Bull,mar.Sci. 1963 Gulf Caribb,, l3(2)329-42

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Nelson, D,R., .n evaluation of the Columbia obstruction method for studying the behavior of the 1962 pink shrimp Penaeus duorarwn Burkenroad. Thesis, University of Miami, Coral Gables, 77 p.

Pearson, J.C., The early life histories of some American Penaeidae, chiefly tho commercial 1939 shrimp, Penaous setiferus (Lino.). Bull,Bur.Fish., Wash,,49(3)73 p,

Pease, N.L. and. W,R. Seidel, Development of the oleotro-shrimp trawl system.Cosmi Fish.Rev., 1967 29(8-9)58-63

Pr z-Farfante, I. (Isabel C, Canot), A new species and two ne'.r subspecies of shrimp of the 1967 genus Penmeus from the western Atlantic, Proo.biol,Soc,Wash,,80(14):83-lOO Western Atlantic shrimps of the genus Penaeus. Fishery Bull,U.S,Fish Wildl,Serv., 67(3) 461-591

Regan, J., C.P. Idyll and E,S, Iverse, Mesh size regulations as a possible method of managing 1957 the Tortugas shrimp fishery. Proc,Guif Caribb.Fish,Inst,,9,18-.22

Ringhaver, L,C., Design and mass production of shrimp trawlers,In Fishing boats of the world, 1960 oò.ited by Jan-Olof Traung, London, Fishing News (BooksLtd.,Vol.2s6l5-23

Ringo, R,D. and G. Zamora, sr., A panasid postlarval character of taxonomie value.Bull,mar, 1968 Soi., 18(2)*471-6

Robas, J.S,, Shrimp trawling gear as used in the Gulf of Mexico, In Modern fishing gear of the 1959 world, edited by filmar Kristjonsson, London, Fishing News (Books) Ltd.,pp.311-16

Robins, C.R. and D.C. Pabb, Biological andtaxonomie notes on thecroaker, Bairdiella batabana. 1965 Bull.mar.Soi.,15(2)495-5l1

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Saloman, C.H., Bait shrimp (Penaeus duorarum) in Tampa Bay, Florida - biology, fishery economico, 1965 and changing habitat. Speo.soient,Rep,U.S,Fish Wildl.Serv,(Fish,),(52O)l6 p.

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Stewart, IC,W.P Contributions to the biology of -the spotted seatrout (ynosoion nebulosus) in 1961 Rverglados National Park. Thesis, University of Miami, Coral Gables,103 p.

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Tabb, D.C., A contribution to the biology of the spotted soatrout Cynosoion nebuloeue (Cuvier) 1961 of east-central Florida, Tech.Ser.Fla St.Bd Conserv., (35J:22 p.

Distribution and occurrence of fishes, crustaceans, echinoderms and mollusks in Florida Bay and the Ten Thousand Islands. Miami, Instituts of Marine Science, Univer- ity of Miami.

Tabb, D.C. and A.C. Jones, Effect of hurricane Donna on the aquatio fauna of north Florida Bay. 1962 Trans,Am.Fish,Soo,, 9l(4)3758

Tabb, D.C., D,L. Dubrow and A.E. Jones, Studies on the biology of the pink shrimp, Penaeus 1962 duorarum Burkenroad, in Everglades National Park, Florida.Tech.Sor.Fla St.Bd Conserv,, (37):30 p.

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Teinsongrusmee, B., The effect of temperature on growth of postlarval pink shrimp, Penaeus 1965 duoraruin Burkenroad. Thesis, University of Miami, Coral Gables, 66 p,

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Waterman, T.H. and F.A. Chace, Jr., General biology. In The physiology of Crustaooa, 1960 edited by T.H. Waterman, New York, Academic Press, Vol.1:1-33

Webb, E., Summary of' Florida commercial marine landings, 1964, Talishassee, Florida Board of 1965 (Jonservation,Salt Water Fisheries Division, Marine Fisheries Research,77p. Wheeler, R.S,, R.C. Benton and J.H, Hudson, The Tortugas pink shrimp fishery. Ciro,Fish Wildl. 1965 Nerv., Wash,, (230):36-7

Wiokham, D.A., Oboervations on the activity patterns of juveniles of the pink shrimp, Penaeus 1967 duorarum, Bull,mar.Sci., l7(4):769-86

Williams, A.B,, Identification of juvenile shrimp (Penaeidae) in North Carolina,J,Elisa 1953 Mitchell scient.Soo,, 69(2)15660

A oontribution to the life histories of commercial shrimps (Penaeidae) in North 1955 Carolina, Bull.mar,Soi,Gulf Caribb., 5(2):ll6-46

A survey of the North Carolina shrimp nursery grounds.J. Elisha Mitchell scient. l955a Soo., 7l(2):200-7

Substrates as a factor in shrimp distribution.Limnol,Oceanogr,, 3(3):283-90 1958

Spotted and brown shrimp postlarvas (Penaeus) in North Carolina. Bull.mar.Soi. 1959 Gulf Caribb., 9(3):28l-90

The influence of temperature on osmotic regulation in two species of estuaririe 1960 shrimps (Ponaeus). Biol.Bul].mar,biol.Lab. Woods Hole, 1l9(3)560-7l

Marine decapod crustaceans of the Carolinas.Fishery Bull.Fish Wildl.Serv.U,S,, 1965 65(l):l-298

A ten-year study of meroplankton in North Carolina estuaries: cycles of occur- 1969 renco among penaeidean shrimps. Chesapeake Sci,, l0(l):36-47

Williams, A.B, and E.E, Deubler, A ten-year study of meroplankton in North Carolina estuaries: 1968 assessment of environmental factors and sampling success among bothid. flounders and pertaeid shrimps, Chesapeake Sci., 9(l):2(-41 Woodburn, K.D. et al,, The live bait shrimp industry of the west coast of Florida (Cedar Key to 1957 Naples3T Tech,Ser.Fla St.Bd Conserv., (21):33 p. - 1537 - 88 */slO3 Penasus d. duoz'sr

Yokel, B.J,, A contribution to the biology and distribution of the red drum, Soiaenops ooellata. 1966 Thesis,University of Miami, Coral Gables, 160 p.

Yokel, B.J0, LS. Iversen and C,P. Idyll, Prediction of the suocess of commercial shrimp 1969 fishing on the Tortugas Grounds based on enumeration of emigrants from the Evergiade National Park Estuary. FAO Fish.Rep.,(57) Vol.3:1027-39

Anon,, Extensive survey made off Florida east coast for stocks of shrimp and soallopii. 1961 1(/V SILVER BY Cruise26. Cosmi Fisb,Rov.,23(1):32-4

Exploratory fishing for shrimp, scallops and small snappers inouth Atiantio, 1962 E/Y SILVER BAY Cruise34.Cosmi Fisb.Rev., 24(l)29-31