Fisheries Re fIorts No. 57, VoL 4 FRm/R57.4 (Tri) PFOc: EINGS F THE WORLD SCIENTIFIC CONFERENCE ON THE PIOLOGY AN CULTURE OF SHRIMPS AND PRAWNS 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

- 1439 - FAO LIBRARY AN: 115084

FRm/S101 FAO Fisheries Synopsis No.101 SAST - Shrimp

SYNOPSIS 0F BIOLOGICAL DATA ON THE WHITH SHRIMP Penaeue aetiferue (Linnaeus)1767

Expose synoptique sur la biologie de Penasum setiferus (Linnaeus)1767

Sinopsis sobre la biologia dei Ponaeus setiforus (Linnaeus)1767

prepared by

MILTON J. LINDNER andHARRYL. COOK Bureau of Commercial Fisheries Biological Laboratory Galveston, Texas77550,U.S.A.

Contribution No.240, Bureau of Commercial Fisheries Biological Laboratory, Galveston, Texas77550,U.S.A.

- 1441 -

FRm/$lOi PenaeusBtifO'?,5 i

CONTENTSil

Pago No,

3. IDENTITY 1*1

1.1Nomenclature i

1.11 Valid naine i 1.12 Objective synonymy i

1.2 Taxonomy i

1.21Affinities i 1,22 Taxonomic status 2 1.23 Subspecies 2 1.24 Standard common names, vernacular names 2

1.3 Morphology 2

1.31External morphology 2 1,32 Cyto-morphology * 1.33Protein specificity 2

2 DISTRIJTION 2s].

2.1 Total area i

2.2 Differential distribution i

2,21 Spawn, larvae and juveniles i 2.22 Adults

2,3Determinants of distribution changes i

2.4Hybridization

3 BIONOMICS AND LIFE HISTORY 3 si

3.1 Reproduction i

3,11 Sexuality i 3,12 Maturity i 3,13 Mating i 3.14 Fertilization i 3.15 Gonads i 3.16 Spawning 2 3,17 Spawn

3.2 Pre-adultjhase 2 3.21 Embryonio phase 2 3,22 Larval phase 3.23 Adolescent phase 3 3 3.3Adult phase 4 3.31 Longevity 3.32 Hardiness 4 5 3.33 Competitors 3. 34 Predators .5 5 3.35 Parasites, diseases, injuries and abnormalities 6 i Thissynopsishasbeen prepared according to Outline Version No, 2 (M. Rosa Jr., FAO Fish, Synops.,, (i) Rev,1,1965). -1442

Page No, 34Nutrition arid owth 3:6

341Feedthg 6 3.42Food 6 3.43Growth rate 7 Metabolism 3.44 7

3.5 haviour 7

3.51 Migrations and ].00a]. movements 7 3.52Schooling 9 3.53 Responses to stimuli. 9 4 POPULATION 4:1 4.1 Structure i 4.11 Sex ratio i 4.12Age composition i 4.13Size composition i 4.2 Jthundnrice and densit of o ulation i 4,21Average abundance 4,22 Changes in abundamos i 4.23Average density i 4.24Changes in deneity i

403Natality and recruitment 2

4.31Reproduction ratee * 4.32Factors affecting reproduction * 4.33Recruitment 2

4.4Mortality and morbidit 2 Mortality ratee 4.41 2 4.42Factors causing or affecting mortality 2 Faotors affecting morbidity 4.43 2 4.44Relation of morbidity to norta1ity ratee *

4,5Dynamics of o ulationas a whole *

4,6Thepopulation in the community and, the ecosystem 2 5 EXPLOITATION 5:1

5.1 Fishing ectuipment 1 5.11Gears 5.12Boats

5.2FishinR area8 i 5.21General geographic distribution I. 5.22Geographic ranges i 5.23Depth rangos i 5.24Conditione of the grounds 1

5,3Fjshjn seasons 1 5,31 General pattern of season(s) * 5.32Dates of beginning, peak and end of soason() * 5.33Variation in date or duration of season * - 1443 -

FRni/S1O1 Penaeus setiforus iii

Page No. 5.4Fishing operations and results 5d

5.41 Effort and intensity 1 5.42 Selectivity 3 5,43 Catches 3

6 PROTECTION AND MANAGEMENT 6:1

6.1 Regulatory (leßislative) measures 1

6,li Limitation or reduction of total catoh 6,12 Protection of portions of population *

6,2 Control or alteration of h sical features of the environment 1

6,21Regulation of flow * 6.22 Control of water levels * 6,23 Control of erosion and silting * 6,24Fishways at artificial and natural obstructions * 6,25Fish screens * 6,26 Improvement of spawning grounds * 6.27Habitat improvement *

6.3Control or alteration of chemical features of the environment *

6,31Water pollution control 6,32 Salinity oontrol 6,33Artificial fertilization of waters

6,4Control or alterations of the biological features of the environment *

6,41Control of aquatic vegetation 6,42 Introduction of fish foods (plant, invertebrate, forage fishes) 6,43 Control of parasites and disease 6,44Control of predation and competition 6,45Population manipulation

6,5Artificial stoc *

6,51 Maintenanoe stocking 6.52Transplantation; introduction

7 POND FISH CULTURE 7t1

8 REFERENCES 8sl

As no information was available to the author, these items have been omitted from the textO 1444 -

FR» 5101 Penaeusotiferus 1 1

IDENTITY Definition

1,1 Nomenclature Rostrum toothed dcrsally arid ventrally, 1.11 Valid name Carapace without longitudinal or transverse suture s; cervical and orbito-ante nnal subi Penaeus setliferus (Linnemue) 17o7, Syst. and antennal carmas always present. Hopatio Na%,(od.12) 1:1054 and antennal spines pronounced, pierygoctomial angle rounded. Telson with deep med&n cul- The controverey over tho use of the cue, without fixed subapical spines, with or ucMio setiforus for the present species or for without lateral movable spines. First on- ochmiti has been definitely decided in tennular segment without a spine on vo,irn.l favor of thfirst alternative by the inter- diatoiosdian border. Aatennujar fli1a naiona1 Commission on Zoologica]. Nomonclatur shorter than carapace. Naxillulary pair with in 'helr Opinion 817 (1967, Bull. cool. 2 or 3 segments, usually 3, Bastai cpiiice on No:icjnol. 24(3)* 151). let and 2nd pereiopods exopode on let 4 per- eiopcde, usually present on 5th. Petasma 1,12 Objective synonymy symmetrical, pod-like with thin median lobes with or without distal protuberancee; lateral Astacue fluviatilie, Americanus, Seba, lobes often with thickened ventral margin, Appendix masculina with distal segment sub- 1759 triangular or ovoid, bearing numerous epinon. Cancer setiterue Linnaeue, 1767 Thelycum usually with an anterior prococo, variable in shape9 lying between thecoxee of Aetaous eetiferus (Linnaeus) Olivier 4th pereiopodm; with owithout lateral plates 1791 on eternite XIV. Pleuobrancbjae on omite IX to XIV: a rudimentary arthrobrancb on Cancer (Oamrearel1us) aetiferus semite VII, and a postarior arthrobranch on (Linnaeus) Herbst, 1793 somito XIII; mastigobranchias on mosites VII to XII, Zygocardlao osio1e oonreieting of Penaeus fluvjatilie Say, 1818 (n a principal tooth folloned by a longitudinal objective synonym of Cancer setii'eruo row of smaller teeth which often end in a L., 1767, through the type selection cluster of minute teeth, Body glabrous, by }Iolthui, 1964, Bull. zool, Noreenol,, (After Dall, 1957, eliglitly modified by Proz- 21(3): 233). Farfante), Specific Penaeus setifer (Linnaeus) L. Agasmiz, 149 Identity of type specimens

1.2 Taxonomy The specimen figured by Seba (17599 Locu- plet. Her, nat. Thee, 3: pl. 17 fig, 2) c 1.21 Affinities holotype, or if the original description nao based on more material, the lectotyps of Cd.cor Suprageneric (to family after Watermm eetiferus L. (by the eelection by Hoithuic, d Ohace, l9bO) 1962, Gulf Research Rep. l(3):1l5). The original typo material is lost and a neotype Phylum Arthopoda has been designated by Lurkenroad (1939, Bull, C1ae Cruetaoea Bingham ooeanogr, Coil. 6(6):l7). The neotypo Subclase Malacoetraoa is a male froni Matanzas Inlet, Florida (O-10 Series Eurnalacostraca fathoms, otter-trawl 2 April 1934, N.B, Bishop) Superorder Eucarida in the Bingbam Oceanographic Col1ection of Ya)o Order Docapoda University, New Haven, Conn, U,SA,, prasoroO Suborder Natantia under no. B.O.C, 237, Phis neotype eo1o'jon Section Penaeidea has been validated by the International Cee- Family Penaeidae mission on Zoological Nomenclature in their Subfamily Penne mas Opinion 817 (1967, Bull, cool. No:nencl, 24(3)* 151), Generic Type locality Genus Penaeus Fabricius, 1798, Suppl. Eflt.Syet,35,4OT Type species, by selection "In Indue" (Linnaeus, 1767), restricted by Latreille, l8lo, Consid.gdn.Anim.Crust. to "Amerloa" (Seba, 1759) by the leototypo Araohn,Ins,:lO2,422: Penaeus rnoncdon Fab- selection (Holthuio, 1962) and still í'iiibo' rioius, 1798, Supp1,Ent,Syst.4O, Gender: restricted to Matanzas Inlet, Floricn., U,S.t. masouljne, by the neotype selection (Burkenroad, 1939), The authors wish to express oir appreoiatíon to Dr, Hol-thuis who provided theinformation in this section, - 1445 -

1:2 FRin/S101 Ponaeus sotifrua

Diagnosis 1.3 Morphology

Adroetral carina not reaching behind 1.31 External morphology middle of oarapaoe; gastrofrontal carina absent; hepatic carina present; rostral Young (1959) described in detail the teeth usually 9 dorsal, 2 ventral; an- skeletal and muscle systems and discussed the terior portion of the ventral margin of the nervous, circulatory, digestive, excretory, and piouron on the ist pleonic soznite almost reproductive systems. Burkenroad (1934, 1939), straight; thelyouxn of female with open Rioja (1939, 1940, 1941), and King (1948) seminai receptiole; posterior portion desoribed minuto details of the external and of 14th sternite of female with conspic- internal reproductive systems. uous pair of fleshy protuberances; disto- ventral lobule of distornedian lobe of petasma Williams (1965, p.18) described the color rounded and not projeoting; petasma of maie "Body translucent, bluish white with dusky bands with diagonal ridge aoross ventral face of and patches composed of scattered black specks; distoventrai lobe (Burkenroad, 1934, 1936; rostrum and sides tinged with pink; blades of wald, 1965), pleopods marked with dark rod; antennae dark brown; uropode with tips of blades dark brown- Subjective synonymy ish purple with narrow stripe of yellowish green along margin." Penaeus orbignyanus Latreilie, 1817 (see Burkenroad, 1939, p.19). Burkenroad (1934) reported that specimens from the Atlantic coast of the United States Penaeusraoilirostris Thallwitz, 1891 bave longer rostrums than those from the Gulf (see Burkenroad, 1939, p.17). cf Mexico, but Lindner and Anderson (1956) be- lieved Burkenroad was comparing immature Atlan- Key to species tic specimens with mature Gulf shrimp. Pérez- Farfante (personal communication) detected no For key to species of the western differences between the Atlantic and Gulf of Atlantic see section 1.21 of synopsis on Mexioo populations. Penaeus sohniitti by Isabel Pérez-Farfante (1970). Mature shrimp have a shorter rostrum than immature shrimp of the same total length. Lind- 1.22 Taxonomie status ner and Anderson (1956, Fig.30) showed the rela- tion between rostral length and total length for Penaeus setiferus is one of the about immature and mature female shrimp from Louisiana, 28 species of the genus Penaeus, and the first to ever get a valid scientific name. The bodies of older shrimp tend to thicken, It is very olose to Penaeus schmitti. Burken- and their weights are greater in proportion to road (see Speoies Synopsis io) and was only length than are those of younger shrimp (Ander- distinguished from that species as late as son and Lindner, 1958; Vioca, 1920). 1936. (See also 3.11) 1.23 Subspecies 1,33 Protein specificity No subspecies are currently recognized in this species, Serological comparisons of saline-hemooya- nm filtrates of P. setiferus, P. azteous and 1.24. Standard common naines P. duoraruin from North Carolina by Leone and Pryor (1952, p.31) ".,. place P, aztecus and U.S.A.: white shrimp P. duorarum olosor to each other than either is Mexico: camar6n blanco to P. setiferus. , ." and "P. setiferus is more similar to P. duorarum than it is to P. azta- Vernacular names cus." These authors also stated, "The sero- logical differences are significant, and support U.S.A.: common shrimp, archaic the theory that these organisms aro three dis- green shrimp (Southport, N.Carolina tinct, but closely related, species". (Jacksonville, Florida gray shrimp (N.Carolina and Florida green-tailed shrimp (Painlico Sound, N. Carolina blue-tailed shrimp (Ocracoke, N. Carolina) lake shrimp (Louisiana) - 1446 -

na u tifon 21

2 oIS'rRIBUTION waters more than i mile (1.9 km) from ahora, although some spawning may actually occur in 2,1 Total ares inside waters in this area,"

According to Pérez-Farfante (personal Off Lousiana, most spawning was in depths communication), P. setiferuis limited to the between 9 and 31ni(Lindner and Anderson, east coast of the United States and the Gulf 1956). The greatest depth at which Pearson of Mexico, land areas 235, 237, 238,and. 311 (1939) encountered larvae was about 55 ni. of FAO distribution code (Hoithuis and Rosa, 1965). It has been reported from Fire Island, The eggs ere denieraal and the larval New York (Burkenroad, 1939), south to St. stages planktonic. Occasionally, eggs and Lude Inlet, Florida (Gunter, 1963), with the larvae are carried. into the estuaries (Pearson, center of abundance off Georgia and north- 1939), but normally the young shrimp enter the eastern Florida,It was generally thought to estuarine nurseries as postlarvas at a total bc absent between St. Lucio Inlet and length (tip of rostrum to tip of telson) of 6 Och1 ocknee River, Florida (Joyce and Eldred, to7mm (Weymouth, Lindner and Anderson, 1933; 1966), but Springer and Bullis (1952), re- Pearson, 1939; Anderson, King and Lindner, corded one specimen from about 46 m of water 1949' Bearden, 1961; Baxter ond Renfro, at iat.24°46' N, and long. 82°59' W. From l966, On reaching the estuarios, the post- Apalaôhicola Bay, Florida, a concentration ex- larvae become benthic (Anderson, Lindner and tending westerly and southerly to northeastern King, 1949; Williams, 1955), Habitata crc Mexico ha s center of abundance in Louisiana, characterized by shallow water, muddy or peaty

,. tiurd concentration in the southern part of bottoms high in organic detritus, abundance ahe Gulf of Campeche extends northeasterly marsh grasses, moderato to low salinities, end along the coast from about Tupilco, Tabasco widely fluctuating temperatures (Weyaouth, to Campeche, Campeche (Lindner, unpublished Lindner and Anderson, 1933; Anderson, King records). Bullis end Thompson (1965) caught and Lindner, 1949; Williams, 1955, 1955a; P. aetiferus at lat,2O°17' N, long. 91°35' i, Loeech, 1965; Mock, MS). which appears to be near the limit of its oc- currcnca in thia area. The greatest depth re- As the shrimp grow, they move gradually corded for P, aetiferus is about 82 m in the from the shallow marginal areca to the deeper Gulf of Mexico near the mouth of the parts of the estuarims and then out to the Mississippi River (Bullia and Thompson, 1965, open sea, They first appear on the eatuarine OREGON station 1495, ist. 28°48' N, long. fishing grounds in June or July, depending on 89°26' W; 45 fa (82 m)), locality, at a modal size of between 80 and lOO mm total length. Ninimun, length im abou The species is most abundant in areas 50 mm (Weyaouth, Lindner and Anderson,1933; characterized by extensive inland, brackish Gunter, 1950; Lindner and Anderson, 1956), marshes connected by passes to shallow, off- Usually within a month of their appearance on shore areas of relatively high salinity and the estuarine fishing grounds, some of the mud or clay bottoms (Anderson, King and young have reached the open aea at a modal Lindner, 1949). In the bays and shallow length of between 100 and 120 mai, This sea- weterm along the coasts of Georgia and the ward movement continues throughout summer and northwestern Gulf of Mexico, it is most is heightened in fall and winter by lowering frequently taken by otter trawle in associe- temperatures (Lindner and Anderson, 1956), tion with fishes of the Family Sciaenidae (Anderson, personal communication; Gunier, 2,22 Adulte 1945), Seo sections 2.t, 3.51, and 5.31. 2.2 Differential distribution 2,3 Determinanta of distribution chan em 2.21 Spawn, larvae, and juveniles The determinants of distribution changos Most, and possibly all, spawning of P. are largely unknown. Generally, it ja presumed 3a1forn5is at sea. Off Louisiana, spawning that postlarvae enter estuaries for food and probobly does not occur in depths less than shelter, Lindner and Anderson (1956) specula- about 9ni(Pearson, 1939; Lindner and ted that the movement from estuarine areas to Anderson, 1956). At Port Aranss, TexaS, the open sea was associated with approaching Heegaard (1953, p. 78) believe it "spawns maturity, They and Gunter (1950) agree that immediately outside the coast at a distance this movement is intensified. by falling tem of 5 to 6 milos(9,3to 1101 km) from shore." peraturos during the latter part of the year, Soiio spawning along a portion of the Atlantic coast may occur near shore. Pearson (1939) Migrations along the Atlantic coast and collected eggs on 5 occasions at St. Augwstine offahore-onshoremovements in the northern Inlet, Florida, and Lindner end Anderson Gulf of Mexico during fall and winter were (1956, p, 616) on the basis of ripe shrimp attributed to temperature changes by Lindner taken in trawis atated, ". moat spawning and Aederson (1956), in Georgia and northern Florida is in outside - 1447 -

IPRm/S1O1 Penseus setiferus 3:1

3 BIONOM lOS AND LIFE HISTORY shrimp in the population exceeded 100 mm, the mean total lengths of the females were consist- 3.1 Reproduction ently larger than those of the males.

3,11 Sexuality 3.12 Maturity

Heterosexual. Sexes aro easily dieting- Broad (1965), intopreting length-frequen- uishedz ma]s with endopod of the let pair of cy plots of Louisiana white shrimp published pleopods modified to form a copulatory organ, by Lindner and. Anderson (1956), determined the petasma; female with open type thelycum that sexual maturity is reached at a total situated between the 3rd, 4th and 5th perelo- length cf 140 mm. Burkenroad (1934) reported pods. that females become sexually mature at a carapace length of abcut 35 mm (about 165 mm Rioja (1939, 1940) described the intern- total length) and that ripe sperm first appear al or medial antennular flagellum of adult at 25 mm carapace length (about 119 mm total males as being thicker and more robust than length). that of adult Lemales; and having 16 of its segments each bearing 1 uniformly-large Distinctive ohàngos in color and size central and 2 small lateral spines. The later- accompany maturation of the ovary (King, 1946). al spines ars lacking on some of the segments Using these changes, King differentiated the however, especially on the distal ones. The following phases of development which aro spines are absent in females, which have a easily identified in the field, tuft of bristles near the base of the external border much longer than in males, Rioja (1939) Undeveloped: ovary small and transparent. believed this antennular organ of the males to be sensorial and intimately related with Developingt ovary opaque; scattered sexual functions, melanophores over the surface.

Four male white shrimp (143 to 180 mm Yellows ovary yellow to yellow-orange. total length) from the Texas coast that were examined by us differed from Rioja's descript- Ripes ovary drab, olive-brown colored; ion, which presumably was based on shrimp from very distended. the Gulf of Campeche. Three of the shrimp had 26 of their antennular segments bearing a large Recently spent: ovary collapsed and not central spine, and the 4th had 24. There was as deeply colored as when ripe, only 1 lateral spine per segment and not all segments with large central spines had lateral 3.13 Mating spines. The number varied from 12 to 17. Dall (1957) said in his description of the There is no record that mating has been genus Penaeua that the 3rd. maxilliped. is observed, but it is believed. that P. setiferus sexually dimorphic. Rioja (1948) who studied is promiscuous. The male attaches a spermato- in detail the 3rd. maxilliped. of P. californien- phore to the female before spawning. The sis Holmes, P. stylirostris Stinlpson, spermatophore is placed externally between the P. vannamei Boone, and P. setiferus, stated 3rd, 4th, and 5th pairs of pereiopods (King, that the 3rd ma.xilliped of P. californiensis 1948). was sexually dimorphio.He did not describe, however, sexual dimorphism in the other species. 3.14 Fertilization F'om this statement we conclude that Rioja found no difforenoe between the 3rd maxilliped Generally believed to be external; upon of P. setiferus males and. females. We examin- emission, the eggs are fertilized. by sperm ed 2 females (150 and 191 mm total length) from the spermatophore. Cook, however, (un- and 1 malo (163 mm) from the northwestern published records) reported that recently Gulf of Mexico and could find no sex-linked oaptured gravid females, without external differences between proportions of each seg- spermatophores, spawned viable eggs in the ment of the 3rd maxilliped against dactylus laboratory These eggs hatched into apparent- or between the total length of the shrimp ly normal larvae which were reared to subadult against length of 3rd maxilliped.. In all 3 size, The sex ratio of the juveniles was Isi. specimens, the 3rd maxiiliped extended to the distolateral edge cf the antennular pedunole, 3.15 Gonads

Weymouth, Lindner nd Anderson (1933) No detailed investigations have been noted a difference which increased with age in made on the relation of gonad size and number mean total lengths of the sexes. This dispar- of eggs to body length, weight, or age. It ity was later studied by Williams (1955) who has been estimated, however, that from 500,000 conoludod that after the mean size of the to 1,000,000 eggs are produced by one 1448

3i2 FRni/S1O1 Psnauuaetiferus vidual at a spawning.Anderson, King and to be moro olosely associated with rising and Lindn' (1949) oountod about 860,000 eggs in falling temperatures than with absoluto tempe- th:'ar les of a ripe female 172 mm total rature... The comparatively abrupt rise in 1en«.Burkenroad (1934) reported that the the spring temperature ooinoides rather close- opermatophot'ee of large mature males wore no ly with the beginning of the spawning season larger than those from smaller mature malos, and may, indeed, initiate it, while the season Ciog (1948), who examined the entire gonads of seems to terminate as soon as the temperatures moiee1 specImens of both sexes, reported them begin to decline rapidly in the fall, even to be homogenous in structure and etage of though they are at that time appreciably high- deveopment of the germ cells, er than those which evidently induced spawning in the spring," 3.16Spawning Data from studies in areas where the 3 The number of times a white shrimp spawne species P. setiferus, Pd.uorarun and Pazte- in n single year is unknown.Most shrimp cus occur together suggest that erawning sea-. bIologists agree that these shrimp may spawn sons of the first 2 probably coincide,That repeatedly in a single season.Lindner and of P. aztecus overlaps the others but its peak Anderson (1956) speculated that a single of spawning activity is either later or earlier shrimp could spawn as many as 4 times in a (Williams, 1955;Joyce, 1965; Christmas, Gun- season and indicated that a few females probab-. ter and Musgrave, 1966; Renfro and Brusher, ly survived to spawn during a socond season, MS),Temple and Fischer (1968) indicated that in the northwestern Gulf of Mexico peak spaw- Weymouth et al, (1933, p. 24) stated that, ning of P, aztecus occurs later than that of "P. setiferue spawns from March or April to P. setiferus andthat the larvae or postlarvae, August or September . ,."With consider-. or both, of P. aztecus overwinter in waters of ably more data than those available to the continental shelf Weymouth et aL, Lindner and Axid.erson (1956, p.616) concluded, "When we oonaider sampling The eggs ere spawned directly into the technique arid errors, it appears probable that water and there im no nesting or reproductive there is little, if any, difference between isolation, spawning seasons in any of the localities we covered between South Carolina and Texas, Spawning activity within a season appears There probably is not more than about 2 to increase to a single peak and then decline, difference in the beginning of spawning between but tho young produced do not follow this pat- any of the localities,Spawning may start tern,Depending upon the looalityd two or thr later in South Carolina and Georgia than it broods of young shrimp may appear on the -fichi s does in Florida, Louisiana, and Texas, but in grounds each season (Ounter, 1950;Lindner oíd all the localities it probably begins either Anderson, 1956), during the latter part of March or early in Aprii and may possibly contiOue on into 3.17Spawn november, though probably it is completed by the end of September," The eggs are demorsal, nonadhesve and spherical.Live eggs measure 0.28 a;' Each egg Lindner and Anderson (1956) placed the has a thin, transparent sembrano, or c0rion, pcc of spawning in Juno or July,More rooent that gives a purplish-blue coloration in rel'leot- thc1os on ovarian development In the north- ed light Cnder the microscope (Pearson, 1939), ouLe.'Gulf of Mexico by Renfro and Brusher (ss) and on 000urrence of larvae in plankton 3.2Pre-adult phaas collections by Templs and Fischer (1968) substantiate the findings of Lindner and 3,21Embryonic phase Anderson (1956), In late developmental stages, the embryo Spawning in laboratory aquariums has is well-differentiated almost filling the egg, oocurrcd only at night.On 2 occasions, ant leaving only a narrow parivitallise space, j ietiforu. spawned between 2230 and 2300 The length of the embryo iabout 0,26 nm Th Cotral Stsrtdard Time (Cook, unpublished dark brown embryo is inveeted by a thin, parent, and colorless embryonic membrau., Whrc pairs of appendages, oorrespondi.cg to d;uoh Spawning is e.sooiateç1 with sise and 2nd antennae anc mandiblee, can h@ oe'c'olcld probbly alec. with age because the largest cloSely to the bodi.Several hours bc:re í'esals ain first (Weymouth et al, ,1933; hatohing the embryo miL'ces convulsive movements Lndoor aiAnderson, 1956). within the egg.The ombryonio membrane is broken by the antennae,The egg rneabranç is Spawning also appears to be affected by normally ruptured soon after and thlarve temperature. Lindner and Anderson (1956, emerges.Actual emergence from the egg takes a. 620) stated, "Spawning in Louisiana appears onLy a few seconds (Pearson, 1939), - 1449 -

FRmJS1OI Penasueetiferu 3*3 3.22Larval phase TABLE I Size of egg, larvae and first postlaru-ae White shrimp larvae iere described by of white shrimp (Pearson,1939) Pearson (1939) fron specimens taken from the plankton.He encountered 5 nairplial sub- stages, 3 protozoeal oubstages, and 2 mysis eubetageri.Heegaard (1953) also reported 2 Stage of development Length (mm) mysis substagee but an editorial footnote suggests more than 2.Three mysis eubetagea Egg o 28 are present in laboratory-reared larvae (Cook, unpublished records).It is probable that Nauplius I 0. 30-0. 34 Pearson did not encounter the 2nd. mysie sub- stage. Nauplius II 0. 32-0. 34 Johnson and. Fielding (1956) reported that Nauplius Xli 0.36-0.40 the length of larval life was 10 to 12 days. They said, however, that the metamorphic period Nauplius XV 0.38-0,44 is not of fixed duration, but depends to some extent on local food and habitat.When reared. Nauplius V 0.46-0.56 in the laboratory at 300 C, larvae grew to ist poatlaryas in 9 to 13 days; those reared at Protozoea I o, 801 14 22° C required. 15 to 24 days (Cook, unpublished records). Protozoea II 1.3 '-1.7 Nanplii do not feed., but utilize yolk Protozoea III 2.2 2.6 granules whioh fill their body,Feeding starts at the 1t protozeal substage.In the labora- Mysis I 3.2 3,8 tory, they are fed. sucoesefully with microsco- * pio green algae and brine shrimp nauplil, Mysiin II 4.0 4,4 First protozea feed, while swimming; movements of the maxfllae and maxillipeda produce a feed- Postlarva I 4.0 -5,0 ing current (Pearson, 1939). The sizes of the various larval stages are * Probably !ltysis III. given in Table I. 3.23Adolescent phase Williams (1955,1955a), who studied estuarino habitats, notad that bottom areas of miring the early postlarval stages, the mud. or fibrous peat are preferred by young young shrimp are still planktonio in the off- P. setiferus, but that they occasionally occur shore waters.At a total length of about 7 mm on bottoms of sand or clay.He also concluded they enter the estuarine nursery grounds and that the depth of water over these ai-Gas did take up a benthio existence, undergoing no not affect the numbers of shrimp, and that striking morphological changes as they continue preferred areas usually had abundant oover, development.Williams (1953, 1959) has given consisting primarily of organic litter and characters which allow separation of certain living vegetation.In controlled laboratory size ranges of poetlarval white shrimp from experiments, juvenile P. setiferus occurred postlarval P. azteouand P. duorarum of more frequently on loose peat, sandy mud, arid. comparable size. muddy sand than on shell sand arid beach sand (Williams, 1958). Various estimates have been made of growth during the adolescent phase.Viosca (1920) and In Mobile Bay, Alabama, postlarval and Ounter (1950) estimated growth at about 1.0 mm juvenile P. setiferum (15 to 70 mm total length) per day; Williams (1955)at 1.2 mm per day; and were most abundant in water less than 0.6 m Lindner and Anderson (1956) at 80 mm in 2 mo or deep where the bottom contained large amounts 1.3 ins per day.Loesch (1965) suggested that of organic detritus (Loesoh, 1965).Mook (MS), "very young white shrimp may grow as much as who sampled pomtlarval and juvenile white shrimp 65 mm per month" or about 2.2 mm per day. in an eatuary at the chore and at a distance Johnson and Fielding (1956) estimated that of about 15 a an30 a from shore, encountered captive postlarvae grow at a rate of 2,1 mm per 98 percent of the white shrimp at the shore day for 24 days and 1.7 mm per day for the next station,Phis shore area was oharaterized by 28 days.B. S. Wheeler (personal communication) anorganic mat or peat bottom (composed of more grew postlarvaefrom 12,2 mm to 92,9 mm in than 40 percent organica) bordered by emergent 5 wk during the summer in a pond at Galveston, vegetation that was under water at high tide. Texas; growth rata was 2.3 min per day. - 1450

Rm/S101 Penanus eetiîe

Vj-ioumme t igatore, inoluding Ouater Zoula Zin-Eldin and George Griffith (195û 96), Ganter, Christmas and. Killebrew (personal communication) believe that the period (1964), WiJiians(1955a),and.Joyce(1965)have of acclimation is extremely important in deter- reported that postlarvau and juveniles of mIning the extremes of salinity and temperature P. -irerue, P, aztecus, and P. duorarum at which Ponmeus epp. can survive. psn r1.ato:a.rd. fresh water in the order given. ce.c'istmas and. Killebrew (1964) recorded Temperature extremes are not documented. as epo ,.1in, L,y limite of 0.42°/oo for P. asti- well as salinity extremes. Weymouth et al. Lr' d.&)°/co for P. aztecue and 2.50O/?or (1933)encountered P. eetiferu in temperatures run in the northern Gulf of Moiao, On ranging from9°to 31° C. Both P. setiferue ro hast oat of Florida the lower sail- sud P, aztecus have survived a temperature of utt i Lijwer.e P. setiferus, 0.260/00; 36° 'ina pond In North Caroline (Lunz,1956), etoua, O.46°/o; and. P. duoraruin, 0.64°/ao The total length ranged between5.1and9.5 cm (ou ¿565). for P, setiferus and between 11.4 and 17.8 cm for P. aztecus. The same author(1958)stated. In Galveston Bay, Toxa, Lee Promt and. that shrimp (presumably P. setiferus) in ponds Jauk Pui1n (personal communication) on 23 April survived temperatures exceedTC. 1966, oaught9P. azteous, rangïng in-total lnngthetwen 40 and 75 mm, in bottom salinity Trent and Puilen (personal oommunication), of 0.10/ao, The bottom temperature was28.00C. hfter examining data from studies in Galveston On 12 July 1966they captured 70 P. setiferus, Bay, reportad the catch of ilive but immobile 35to 65 mm long, in a salinity ofo7 and P. setiforus(65mm total length)on28 January a temperature of29,00C. (There is some ques- 1963,where bottom temperature was2,60C and tion an to the accuraoy of the above low sali- salinity12.80/00, Their records also show a nity records because they were determined from catch of531P. setiferus (mean total length ohiorinity analyses.) 85 msi) at bottom temperature and. salinity of 35.5°C and 24,00/ao on26August1963, At the other extreme, Ganter (1961) report» ed. P. estiferus from a salinity of 41.3°/ao end Rapidly decreasing temperature is known to Hildebrand(T958)caught them, from 23 to 76mm kill P. setíferus, The rap:Ldty of the ohange long, in Laguna Madre, Nexioo in salinity of appears to be more important than the actual 47,96°/oo. Hildebrand (personal communication) temperature (Ganter, 1941; Gunter and Hildebrand., siated "In Laguna Macire of Texas, I have taken 1951 Lindner and Anderson,1956; lung, 1958). several specimens at a salinity of 62,40/00, lethal temperatures are nab known but Ganter However, this vas in South Bay where the eali.- (1941)re-ported surface water in Copano Bay, muynight vary greatly with wind shifts and. Texas, to have reached a low of 40 C durìng the tidal patterns ovar pelatively short periods of 1940kill and Lana (1958) mentioned4050 C. time. I aleo took it in eal:inities of56.8°/aa in the Upper Laguna Madre near South Bird. Island. Weyinouth et aI,(1933)end Gunter(1950) In general, ï would agree with Glanions(1957) obtained a positive correlation between salinity that it is generally raro at salinitiss grealer and sise of white shrimp, hu.t Lindner and than 400/co end that the few tbs.t occur at high- Anderson(1956)demonstrated that th size re1ai. er ealinities are strays or particularly hardy ion was with loonlity rather than salinity. inctividualowhich have survived a gradual in- orGane in salinity," Guntsr and Hildebrand developed a relatiow between Texan rainfall and. Texas catch of P, set' In general, reaLte of laboratory expuri- fru, The highest correlation nao between 'ho mox,s c'La tentiate field ohservaí&oae, catch of thyear aoci the avorage annual raa (:it cad ZainE.id.in (pereonal communication) fall of that year and the two prev:Lous years, 80 percent survival of' P. aetiferus with high catches a000iated with high rainfall '.o's exposure to salinitisa of 4.0 and. values, They believed this relation was caused. 5..(i°/oo. These speoimens were betwoen 50 arid. by salinity changes in the bays (Hildebrand. and hCi ¿c oc1 longththey were held. at a eon- Gunter, 1953 Ganter and. Hildebrand,1954). temperature of2500 and cere gradually aeïeated to the final salimities over a period 3,3 of 3. 31 liongevity î: :ct-her sxperisent, when ea':iap morsas- oct ±. a'ye..c a total length from 6 to 50 mm in No method. han been developed to determine 20 cayo 0 peroent of the experimental popì.- reliably the age of an individosl shrimp, L-:-.tunn. and allnitio of 2.0 and 40.0°/ca. Lindner 1953)developed aeohnique for estim- u.:-,rf n were held oanct.ant at250C and. ating age for restricted periods by mark-recap- t:..a period was48h, ture experiments, and Lindner and Anderson(1956) - 1451 - PRw/S1O1 Penaaun istiferus 3L demonstrated Its uufulnesu in interpreting 3 species occur off Galveston, Texas, length-frequency distributions,Neither thin however, the adults occupy quite distinct technique, howevor, nor thu von,Bert1anffy areas.Adult P. setiferus are most abundant growth formula, lt L (i_l_t_to)), be on mud or sandy mud bottoms shoreward of 27 ni, used as a reliable indicator of shrimp age when and adult P. aztecas are on similar bottoms the constant K changes an it does in P. enti- seaward of 27 ¡n.Adult P. duorarum occur ¡noei ferus between warm andcold periods.The me- abundantly on shell or sandy shell bottoms, thod of Ursin (1963) may be an alternative. On the Galveston Bay nursery grounds, Using information gained fron maturity the bulk of P. aztecus young are leaving or studies and length frquency plots, Lindner have left these grounds before most poetlarval and Anderson (1956) demonotroted that P. neti- P. duorarum and P. setiferus have arrived. ferue lives at least 16 mo, but owing to high When the postlarvae occur together on the same mortality, the number that live 2 yr or mora nursery grounds, those of P. setiferue are is small with respect to the total population. closer to shore than those of P. aztecas (Loesch, 1965; Mock, Ms).Poetlarvae of 3,32Hardiness P. duorarum seem to prefer grass flats. Williams (1955a) reportad seasonal, size, and According to de Sylva (1954), p, seti- areal differences in the estuarine distribut- 'erus is the least hardy, P. aztecus is the ion of poetlarvaa and juveniles of the 3 spec- most hardy, and P. duorarum is intermediate. ies in North Carolina. Field observations by the senior author while trawling, confirmed by David Harrington (per- 3.34Predators sonal communication)9 indicate P. duorarum to be the most hardy of the 3 species and P. No exhaustive study of white shrimp pred- metiferus the least,Butler (1962) observed ators has been undertaken.Predators on larvae, that under laboratory conditions P. aztecas undoubtedly numerous, are unknown.Robert is more hardy than P. setjferus, Temple (personal communication) has observed Penaeua mysis in the digestive tracts of arrow On the basis of size distribution in wint- worms (Chaotognatha), but he was unable to er, Lindner and Anderson (1956) believed that determine whether the arrow worms had, been juvenile and subadult P. setiferus are better feeding on the larval shrimp or whether they able to withstand cold than the adults,Con- had accidently ingested the larvae while in firmation of this relation by experimental the plankton net or while dying from the pre- evidence is lacking. servat ive, Shrimp are extremely vulnerable to agricul-' Knapp (1949) examined 10,016 stomachs of turai pesticides (Chin and Allen, 1957; Butler, 34 species of fishes from coastal waters of 1962, 1963, 1966; Butler and Springer, 1963), Texas between 7 June and 31 August 1948, Postiarval and smal] shrimp are generally more Shrimp, principally P. aztecus, occurred in sensitive to pesticides than large shrimp; 61,8 percent of the 5,946 stomachs containing large white shrimp are less tolerant to some food,He reported shrimp as being the moot pesticides than large brown shrimp (Butler, widely used food between June and August, the 1962).Butler and. Springer (1963, pp. 380, season when P. aztecas is at peak biomass 381) stated, "Because shrimp, crabs, and other (Kutkuhn, 19x2). crustaceans aro arthropods, like the insects, they understandably are sensitive to many of The following fish predators of Panaeus the chemical insecticides.Concentrations of shrimp have been reported from the United heptachior, encirin, and liridano in the range of States (areas 235 and 238 by Ganter (1945), 0.3-0,4 p.p.b. killed or immobilized half of Knapp (1949), Darnall (1958): the adult commercial brown and pink shrimp exposed in 48-h laboratory tests.Other chlor- Carcharhinus leucam Coryphaena hippurus inated hydrocarbons, including DDT, chlorclane, Dasyatis sabina Bairdiella chrymura toxapherie, and dieldrin, showed similar effects Elops sauras Cynoscion arenarius at 1-6 p.p.b.In the laboratory, paralyzed Msgalops atlanticus Cynosoion nebulosas individuals may live for days or even weeks5 Anohoa mitchilli Micropogon undulatum but in the struggle for survival in the sea Bagre marinas Menticirrhus sp. this condition results almost immediately in Galeichthys felis Pogonias creíais death," Iota] urus furcatus Soiaenopa ocellata Roocuri mIrisîsnippieflnBScomberomorus cavalla 3,33Competitors Lut,janus blrickfordi Scosberomorus maculatus Pomatomus saltatrix Istiophorus (Ìakaira) Little is known about the competition Rachyentron canadum Parsliohthya lethostigma among the 3 species of Penaeus. P. setiferus Caranx hippos and P. astecus may oe competitors during most or all of their lives (Broad, 1965),Where the \4u believe the above list of predators is 1452

3s6 FR!js1O1 Ponaeue setiforus incomplete.To us, Gunter's (1957, p. 102) Class Cestada:P-roohristianellaenaei remarks seem entirely appropriate:"It in a Kruse (1959).Found by Kru7T59in general observation of everyone working with "digestive gland arid tissues surrounding shrimp that they are eaten by alaot everything digestive gland and stomaoh, blastocysta of that is large enough to eat them, all the time, larvae frequently penetrating wall of digest-. They are eaten by everything that lives around, ive gland."Pleroceroi occur in Penaeu.e, in, or above the sea, including the lowly jelly-. adults in the ray Dasyatis sabinaTAldToh, fish,They are not only eaten but they are 1965),Kruse (1959) stated,haisbc north- preferred." eastern Gulf of Mexico 944 pci'CS1Tlof P..metl feruswere infected with an average of 14.7 The defense reaction of Penaeus is a rapid plerocerci,Aldrich (1965) reported 50 percent flexing of the abdomen that caUses the animal of the white shrimp in Galveston Bay, Texco, to to dart backward.In large specimens the dis- be infected.Both incidence and. intensity of tance may be1 a or more in less than 1seo infection increase with the sise of the shr)mtz (observations of the authors)0 up to a size of about 14 mm carapace length (Aldrich, 1965), 3.35Parasites, diseases, injuries, and abnormalities Claes Cestoda;Pol, socs phalus ap. (.. Parataenia sp.).Hutton194 listed1do The following parasites have been recorded cestode as infecting P. setiferus as annLer- from P. aetiferue:Class Telosporideaz mediate host.He reported it to bpresenifl Wmmaiopsls naeus Sprague, 1954.Found in numerous localities in Florida, intestinal tract of host (Sprague, 1954); oocurs as trophozoites, sporonts, and. gameto- Clase Nematoda:Contraoaeum sp. Kruse oystm (Kruse, 1959).Kruse (1959) reported (1959) found juveniles "In digestive gland and 100 percent infection in Alligator Harbor and tissues eurrounding digestive gland end, stomach, Apalachicola Bay, Florida,Shrimp have to be not encysted,"The same author reported 5.5 continually reinfectsd, or they lose the infec- percent infection in Alligator Harbor and. tians (Sprague, 1954; Kruse, 1959).Intestinal Apalaohicola Bay, Florida, and an average of epitholium can be appreciably damaged (Sprague, 4 parasitessr shrimp,Hutton, Ball and 1954). Eldred (1962) encountered 15,5 percent infest- ation and as many as 22 parasites iii onz speci- Class Cnidoeporidea:Tholohania penaei men. Sprague, 1950,Found in reproductive organs of male and female (Sprague, 1950); occurs as Dawson (1957) found maturing Balanus on 4 spores an1 sporonts (sprague, 1950).Viosca white shrimp. (1943) stated that in 1919 about 90 percent of P, aetirerusin Louisiana were infected with 3,4Nutrition ind1rth a protozoan that destroyed their reproduotiye organs. 3.41Feeding Clase Cnidooporidea:Thelohanie. sp. Kruse, From observations on Penasus feeding in 1959.Found mainly in muscles, it also occurs aquariums, the senior author is convinced that in other organs as sporonte, pansporoblasts, they are selective and particulate feeders, and spores.Kruse (1959) recorded 10 percent Shrimp search through the san'l grains with their infeotion in Alligator Harbor end Apalachicola, peroiopods and pase toward the manddbles only Florida, and considered this organism to be the that which they choose,They appear to be cause of th"cotton" or "milk" condition in particularly fond. of polychaste worms, P. setiferus, P, aztecus and P. duoravum, "Cottonhrimp," for esthetic reasons, are not 3.42Food acceptable for marketing in the United States, Studies on the food habits of shrimp hcir Class Trematoda;Opecoeloides fìmhriatus been hindered because their stomach contss.c (Linton, 1934).Hutton et al, (1959, p,19) are usually macerated and d±fficult to identii'y, found this parasite "Enoysted in the hepato-. Viosca (1920) and Weymouth et ai, (1933) report- pancreas, the gonads, in the soft tissues of ed P. setiferus to be omnivorous,In reviewing the head, around the mouth, around the stomach the literature on the food of the 3 species of and in the soft tjsuea under the exoskeleton," Penaeus of the United Staten, Darnsll (1958) It occurs as metacercaria in Penaeus, adulte in and Broad (1965) reached thsazie conclusion. various epecies of fishes of' the Family Cannibalism in captive P, setiferus was observ- Sciasniclas (Kruse, 1959; Hutton et al. ,1959), ed by Weymouth et al, (T935).h'il1'isa (1955), Hutton et al, (1959) found a single specimen in discussing the food of the 3 species in in white shrimp from Apalachicola Bay, Florida, North Carolina estuaries, stated that stomache Kruse (1959), who worked in the Satze area, usually were full during surmaur, full or half- encountered it only in P. luorarum, filled during fall, and nearly always empty in winter, - 1453 -

FRm/S101 Penaeus setiferus 37

3,43 Growth rate Lova and Thompson (1966) studied the amino acid concentration in abdoméns of P. estiferus Lindner and Anderson (1956) demonstrated, and P. aztecas. The amino acids they found are by means of markrecapture exporimenta, that shown in Table II. Nitrogen content did not growth in P. setlferus along the coasts of the vary seasonally in brown shrimp tails or offal, United States was much loss when the temper- but certain amino acids did. Concentrations of ature was below 200 C than when it was above. a number of amino aoid differed uignificantly They developed Walford (1946) growth lines from between P, azteous and P. setiforus, but the successivo 10-day intervals for the period of variation between analyses of P. aztecus on rapid growth. These data, oonverted to 30day different dates indicates that the differences lires, had the following oharacteri.stics between species may not be truly significant.

Males Y - 45.8 + 0.7427 X L 178.0 mm In an earlier study, Jones, Noeller and Gersdorff (1925) reported tryptophane in the Females Y - 56.4 + 0.7225 X Li,, - 203.0 mm muscle of P, setiferus.

Combined Y - 51,00 + 0.7322 X L, - 190.45 mm Boroughs, Chipman and Rico (1957, p. 82) stated that P. setiferus "accumulated strontium Klima (1963, and personal communication) rapidly from sea water," calculated von Bertalanffy growth parameters from two markreoapture experiments. For 3.5 Behaviour 7day periods these parameters were 3.51 Migrations and local movements (Personal 1963 communication) The young shrimp aro planktonic in the offshore waters during their larval life and to -0.6 -0,2 enter the estuaries as postlarvas about7mm total length. It is generally believed that L 224 mm 214 mm these younger stages are not capable of exten- sive movement and that to reach the inside wo 87 g 75.1 g waters they must encounter favorable currents. Once in the estuaries, they make their way to K 0.06 0.09 the less saline, shallower areas, presumably by utilizing tidal currents. After growing rapidly 3,44 Metabolism for 3 or 4 mo, the young move into the deeper waters of the rivers and bays and then to sea. Johnson and Fielding (1956) fed fresh fish Movement out of the estuaries appears to depend to 2 groups of small white shrimp at a rate of on stage of maturity and temperature. 10 percent of their initial body weight per day. One group was held at 340/00 salinity and th Lindner and Anderson (1956) described the other at 18.5°/oo for 2 wk. Their results are general migration of adult white shrimp from summarised in the table given below, shrimp tagged with Petersen discs, Klima (1963) reported on 2 experiments in Louisiana in wt,ich P. setiferus are able to regulate total biological stains were used to mark white shrimp body fluid concentrations. Body fluide aro and found the same pattern of movement as des- byporosmotio in brackish water and hyposmotic cribed by Lindner and Anderson for tagged shrimp in hyperealino environments. Regulation app- in that area. ears to be more effeotive in lowsalinity water. Also, the shrimp regulate the jonio The following account of white shrimp concentration of Na, Cl, K, Ca and Ng in their migrations has been taken from Lindner and serum (McFarland and Lee, 1963). Anderson (1956), The longest migration rEecordd

Salinity Initial Tormína]. Gain Gain Food Conversion weigh t weight consumed ratio 0/00 (g) (g) (g) (%) (g)

34.0 32.7 37,3 4.6 14.1 19,2 4.09:1

18,5 29.4 32.7 3.3 11.2 17,4 5.27:1 1454

3 8 FRm/S101 Penaeuo aetifarus

TABLE II

Average amino acid concentrations in tail meats of Gulf of Nexioo Shrimp (adapted from Love and. Thompson, 1966) Signifionail, upecioc vera amino acid. ooncentrtio difference botvoen o aoids eamplos talcen on 20/2/64

»olesJm" N 3,70 3.64 4.26 4.36 Arginino 3,16 3,81 3.60 3,81 Aspartic acid. 3,90 4,14 4.64 5,57 0yiin/2 0,33 0,36 0,09 0,38 0ltamic acid. 4,90 4097 5,64 6,22

Glycthe 6,35 6.19 6.45 6.96 Histiô.ine 0,67 0,79 0,76 o,86 Hydroxylysine 0,05 0,03 0,01 0,04

FLydroxyproliae 0,09 0,11 0,03 0,01 Icoleucine 1,86 2,02 2,13 259 3,35 3,59 3.87 4,32 2.82 3,52 3.1]. 3.46 Nethloniria 0.87 1,04 1,06 1,25 Ornjthjne 0,13 0,10 012 0,11 Phenylalanino 1,11 1,20 1,30 1,42 Proline 2.11 2,29 2,46 2.47 Serine 1,92 1,96 2.33 2,40 Taurine 0,47 0,31 0,57 0,65 Thi'eoniae 1,70 1,77 1.93 2.16 0,39 0.45 0,77 0,85 0,74 0,89 Urea 0,07 0,08 0.16 2,06 2,32 2.57 2,80 indicates difl'erence -1455 -

FRin/S1O1 Penaeus setiferus 3t9

was360mi (667 km) in 95 days, and the great- There should also be a corresponding northward est time between marking and recapture was migration out of Mexico in the spring. 257days. 3,52 Schooling After reaching outside waters, movements vary with size, locality, and season. Shrimp Hildebrand(1954)stated that white less than130mm, total length, did not migrate shrimp school more extensively than related extensively, but remained adjacent to their spe oies. nursery areas. Shrimp larger than 130 mm exhibited distinct migrations that varied with The senior author believes that white loo al i. ty. shrimp of about the same age or length tend to aggregate in groups but that these aggregations Atlantic coastt The white shrimp in this do not possess the consistency through timo area migrate southward during the autumn and gensrai]y attributed to a school of fish. early winter and then move northward again in late winter and early spring. These movements 3,53Responses to stimuli parallel the coast, and do not extend into deep- er water far from the coast. n'ing late Thermal spring and summer movement is not pronounced. See section3.16for effect of temperature Northern Gulf east of the MississipiiAll on spawning and 4.42 for lethal temperatures. that is known of movement in this area is that during autumn and winter the shrimp move into Lindner and. Anderson (1956) found that deeper water and toward the mouth of the P. sotiferus moved from 8hßllow to deeper wat- Mississipi River. er in respiss to unusual cold spells. They also uncovered evidence that the smaller shrimp Northern Gulf west of the Mississipi Rivers moved back into shallower water as the water The only definite movements in this area are temperature increased. offshore and onshore movements. The large shrimp move offshore and disperse during autumn Optical and winter, Apparently the Mississippi River acts as a natural barrier, Temple and Fischer(1965)demonstrated that penaeid larvae showed vertical diurnal Central Texas and northern Mexico: The migrations when the water column was vertical- migration in this part of the Gulf appears ly stable. Larvae moved into the surface similar to that on the Atlantic coast, bu-t the layers in darkness and. into deeper watei during evidence is not conclusive. Indications ars daylight. that the shrimp move from central Texas into northern Mexico dnrïng autumn and early winter. 1456

I.w/S1O Penaeus setiferue 41

4 POPULATION Lindner and. King, 1949). Between1945and 1948,production of white shrimp deolinec' and. 4,1 Structure has not approached. the high levels of the early1940'ssince then, During the years 4,11 Sex ratio for which we have reliable statistics(1957 to19659Table III), the catoh has varied The eon ratto is 131, from14,441metric tone in 1957 to36,188in 1963, The oauso of these fluctuations is not 4,12 Ageoompomition known,

Ph mortality rate is high, and, while TABLE III some white shrimp maylive more than 2 yr, United States white shrimp landini, the percentage is insignificant, Age corn- in metric tons, whole weight position of the catch oonmequently varies di. roctly with recruitment and movemenof the Past coast, Gulf coast, maturing ehrinip into offshore watere., YearNorth Carolina Florida Total to Florida to Texas The young enter the inshore fishery at an age of about2mo and the offshore fishery 1957 6,673 7,774 14,447 at about3 or 4mo., 1958 5,032 17,980 23,012 4,13 Size composition 1959 5,816 17,167 22,983 Size composition varies in the same man- ner as age composition, 1960 8,522 19,826 28,348 The young enter the inshore fishery at a 1961 6,365 10,093 16,458 minimum total length of about50mm and, the offohore fishery at a minimum total length of 1962 59504 16,181 21,685 about 100 mm, Lindner and Anderson(1956) give an L,,, of203mm for females and. an L,, of 1963 3,297 32,891 36,185 178 mml'or males (see also3,43), 1964 3,683 30,720 34,403 The weights of large shrimp are greater im relation to length than are those ofsmall 1965 7,395 23,4'O 30,865 ones, There is no appreciable difference in length-weight relations between sexes,. Chin (1960) expressed the length-weight relation In 1940, the white shrimp pcpuiaion in with the following equation: South Carolina aM Georgia was decimated by a severe cold. spell, la1941,the return of Log W= 5,289 + (3,075)(Log L) la.ndinge to normal indicated thei a normal crop may be produced by relativrlj few speun- This equation agrees with the length. ers (Lindner and Anderson, l956) Kutkuhn weight relation given by Anderson and Lindner (1962)attributedthe low 1957landings of (1958)until the shrimp reacha totallength white shrimp in Louisiana to the effects of of 1,43 mm, Above this size, weights at come- e-vere torma 'hioh otruok th coast while the pl lengths given by Anderson end. Lindner yosr1g of 'che spring opawning class were in the wore greater than those of Chin(1960), oeurioe, Ageth, the fishery mein a good re- idereon and Lindner found a difference in oov®ry oî'mr 1 yr., length-weight re la ions between matura and. immature aIrnp naurohriep more 4,23 Avere e denaity . t"eao2llength, Chin be 1io't, tbt thic J'ao.or nhi oLplin bhc d&f'orcnces be- Kuthuija (1952) indicated the 4-r .rund te-m he 2 mate of data, (195G to i')59) is hionasu o' P, 'crue in the iorthcrn Onif ci' ,f,:joo,qj icn 4,2Abundance ,,neity of poprl c'ce ing too ec.t cl' tin4smr'ci'p1, flico aOi "ccrece,n .c -1m 4.22Changos in abundanoe 4q24Changes in dønsity Whi. 9'fimpproduction 'esked er1y L94C - Production in 1940e cimu.od There are 2 periods of increased eity t hc'o been r.out 659000 metric onrm, uhole on the offshore fishing grounds, one i lnt wot&.:, At thic tnenoive fshnb for fall and, the other in spring (Weymo ar,' o.nl: shriep baci mot etavcdn and. 1933'Lindner and Anderson, 1956 a th1to nThcimp usro oaima,od to account for95 19625, peroent or more of the total catch (Anderson, - 1457 -

4*2 FRm/S101 Pennous setiforu 4.3Natality and. recruitment 4,43Factors affecting morbidity 4,33Recruitment See sections3.23, 3.32, and 3.35

Rate of recruitment has not been determine&. 4.6 The porulation in the community and With some looal variation, inahore recruitment the ecosystem starts in June and continues until the onset of cold weather (Lindner and Anderson,1956). After making an extensivo survey of macro fauna on the white shrimp grounds in the Gulf 4.4Mortality and morbidity of Mexico, Hildebrand.(1954)listed the domin- ant organisms to bet White shrimp, Penaeum seti- 4.41 Mortality rates feras; blue crab, Callinectee donne; sea pansy, Ranilla mullen; TexasVOflUBpPitar texasiana; Klima(1963)gave an instantaneous total silver seatrout, Cynoecion nothus; Atlantic mortality rate of0.92for white shrimp off croaker, Mioropogon undulatus; Atlantic thread-' Louisiana from September through !ovember. Lin, Polydactylus octonemus; and sea catfish, This is a 14day rate of decline cf60peroent Oaloiothys fells.

4.42 Factors oauming or affecting mortality

See sections3,34and 4.22 1458

Wr/o 101 xaeue sotiferus 51

XfL)ITATION 52 Fishin areas

5,1 I?ishiag equipment 5,21 General geographic distribution

5,11 Gears P setiferus is fished along the Atlantic coast from Pamlico Sound, North Carolina to LbS various garre used in the Inshore about Vero Beach, Florida (land areas237 and fishery for h1te shrimp rango from the modern 238) and in the Gulf of Mexico from Apalachiocia, tï to primItive eire and include trynets, Florida to about let, 23° N on thø M-:.- can coast or ce framo travis, lift or drop nets (land areas 235, 237, and 311), 2i ri ial, ridc. frame raule, Seines, oast nets sud dip 230 N and Tupilco, Tabasno, Mexico, 1Jro is no nst In the (Jolted States some shrimp are fiehery From Tupilco, however, thfishery tkr, in emiunries for sale as bait, largely extends eastward to about Champoton, Coapeohe, wIth a elogie3-to 11-m otter trawl with a The3centers of abundance are in Georgia, trtohod mesh elze of 3 to5cm, If the Louisiana, and. between Alvaro Obregdn, Tabasco, shrimp are to be kept alive, they are placed and isla dol Carmen, Campeche (Hiidebrend, in a tank on the boat equipped with running 1955; Carranza, 1959; Anderson and Luire, ì9G5 sea eater, or in a live box which is towed and Lindner, personal observations), behind the boat, On shore, the live shrimp are kept usually in larga bait boxes suspended in 5,22 Geographic ranges the water or in large holding tanks through whioh sea water is pumped (Inglis and Chin, The accompanying map (Fig i) adapted from revised by Baxter, 1966), Hildebrand (1954), Lindner and Anderson(1956), Anderson and Timm (1965), and Lindner (personal Most vessels fishing in offshore waters observations), dol locates the primary fishing use a pair of 12io 15-m flat or balloon otter areas, trade, but some use only one, Almost all boats uso trynots, which aro miniature trawls, 523 Depth rangos to locate flhablo concentrations of shrimp, A detailed description of trawling gear was The white shrimp has beenrecorded fros given by Robas (1959). depths down to 82 n, but it is most abundant on bottoms shorewerd. of27 n(see sections 21 nui! In the early fishery in the United States, 3,33), fishing was primarily with haul seines and. cast nets, In Mexico, fixed traps predominated, 5,24Condition of grounds Between 1912 and 1915, the otter trawl was introduced and by 1917, ii had become the stand See seoticri 21 aa'd commercial gear Practically all shrimp boats on the Atlantic coast were equipped with 53 Fis'n!neansona power winches by the mid-tentios, but with the sxcøptlon of Texas, winches were noi iotrodUo White shì'isp er's hnrrau td ajl ed into the Gulf until th late thirties peak production is in the autunu Be:ian 1956 (Johnson ana Lindner, 1934 Anderson, Liîc1ndr and1959, thegreateot moth1ycatch in Texan and King, 1949) In the Gulf of Mexico, doubic- waters was taken in September, about 6 pccnt rig trawling was tried first in1955 and, was of the catch was made during September aol. noon adopted by most offshore boats (Knako, October and about 84per000t from Augunt through urdook and Cating, 1958) November, The only ol:.h.sr month of high prod.uoi' ion was May (Gunter, 1962) Phci greatest catch 512 Boats offtheeast coast of ».orid is made froc Septemberthrough Jono...rrand tlic psk io in VassoIo of varied design, usually n9t December (Joyce, 1965) exceeding9to 14 n in length, are used in the inshore fishory Power im supplied by eninse 5,4Fish1nro:a.i;o that burn gaso:tine or dietillate The boats ere equipped usually with power winches and rope or 541 Fffot arid intentï wtre towlines (us Fish and Pildilfe SerVice,

1958), Fl,sbereso frsquent1: I id nccl ci nhec cf brown, white arid. pirhrih .I-n -r 1ïaessl in the offshore fishery ara en, effort isroportsirt o it.y r h... driii riant predominantly Florida-typo trawlers, which are speciem and not by ludir :uni poc oo s u deecwl'oed detail by Cook and Lindner (l97O) resait there 'nave be ro rai: cala s urdnn b. t require effort information on tnctiv:Idua:( -1459- FRm/S1O1 Penaeua setifBrtuj

FIRE IS.

GENERAL DISTRIBUTION L MAJOR FISHING GROUNDS

N. C PAMLICO SOLJNO

o C

o GA ALA. ' Oip, ISS. LA. !I

T EX.

VERO BEACII

ST. LIJCjE INLET

T4MPJCO

L,

1. Ditri1mtton and major f i&iing areas for Fenaeusetiferu '1460

/S1( rlaoueeetlferw3

542.1tivty pok abunlence of one speo1oeboatefroquent- ly oonoentz'ate in relatively emall areas, n cvitoui cra aro available for leaving other areas aoci.peotos unfiebsd At otho aeleotive 3roperties of other tì5e of the year, the fleet le diepere. or u'oe of ohangeein ineheizee buta ed over the entire fiehing range 7vu!hly -1a'cUee onheother 2epoclee 5.,43Catohes triepricee of all cornrnoroial pnaeldo Seeoection 4,22 jç1 Statoe are haecd On i3iOe blie U1fl1 OOrnIflOd ho hLcvopri.ce Table IV givee Uolted Storj oh'1hrn r-ocJ U)IÍ1Uh 'O(OOl,Y Oh in aocu3 that yir)lci uet,on bhrouh 1940Q:t. Je ooir'ud that !C&L UhCLhiO IJ p'O.'U1jii0O to a'tci ihoo vhi.o ehrrnp oonI3titui.id floro than 95 poroet ivy uriull uhilrnp0Uhon 1arc quant1iiu oÌ u!1ïee landins0 !u1le içd Si.bor bge: niul ohrirup oci le riot uneorneon for oo!letii, moro drtilc ritaioIc,u on hu t, ho didudolthoruoeueohoy io not Guluoaeloic inoo 1956 cudxncnì'dhie uic diuut uiu o ct1onuoroo1rie }io u'ra;o i.e tho Atiuni.io coast fLcbo,v in i95î Luh'iOi (.O!ut lL'fl to bohoritli thnn, rLablo LL (ooton 4,22) gie thu Unitod Sac,o vhite shtip lanc1ine e1u 195'? These U\7J on ri1lLah1 ClOu tuo OluMtOtl by l.nuinge do toi. inolude produoion Eroe tito dIiduuL otatoo co'dhof'e bui;, i'isherieo, 'thioh In sorno .treae lu eubttant.. ia During964, ror examric, imre i.hnu OtTohoro oh.p ]oubo tu:e hijbly wobilo9 85OoOo lb (about 400000 it) of chrirnp vero cof L3.VU ì1OV!h t,cC iiuu11y ioiu oho lotion or bait in 0a1eu i.on i2ayTe'cc.c (Inglie to ìouii c.hindanoo uI tho 3 clioroia1tr and Ohin:oiood byaxov,96G),'boi' iwtt'.nt epeui ÜIi1uhJI1f poiothi ei' 65 poi'oûnoi 250,000 tg voe vhlo cbnp (tettoth huter pt-.ocl muiort thu) 1461 5,4 FRm/S101 Penaous sotiferiis

TABLE IV

United States shrimp landings through1940 in metric tons, whol,, weight (Power, 1960)

East coast, Gulf coast, North Carolina to Florida Florida to Texas Total

1889 337 3,767 4,104

1890 337 3,380 3,717 1897 284 3,081 3,365 1902 1,729 5,609 7,338 1908 2,584 5,835 8,419 1918 7,101 14,672 21,773 1923 10,761 21,371 32,132 1927 13,604 31,242 44,846 1928 15,109 37,271 52,380 1929 13,984 35,254 49,238 19 G 11,886 28,083 39,969 1931 11,564 32,185 43,749 1932 10,189 29,846 40,035 1934 11,776 42,346 54,122 1936 15,226 38,348 53,574

1937 12,445 51,636 64,081 1938 12,503 51,068 63,571 J1939 12,558 54,605 67,163 L1940 10,297 57,986 68,283 1462 FPUiì/S1Oi Penasus tfen" 61

6 pOrpF(T:O ]) ' iii) limitations on oatoh 'ininside waters; (iv) permanent and temporary closure of insIde watere (ir) temporaryolosure of outside waters In the U. .. ,..h of 'th rr )r'oc1uç within territorial 1imiia 1LLoo lico c oro 1J o r'r) ''0 .1 ou ouco'or rxu Ou 62 Coni;rol or alrc,!ooLl o$ p1iylocl u on1 b.,'o o1'-' 1) irru; o 'i cod Leatucoc o'the envronrncn c0000rcr cc oro $ JC mL ,cri c:i',: () s )i!dIfco' iuc, Hook (MS) demonstratedthat the abundance !unor1957) of poetlarval and juveniie shrimp is reduced greatly along bulkheaded portions of ostuarine In 'the UOSOAO, regu1eto o borders0 He has aleoobtaineddata (personal Lor,ly 1)0iTOCJit StO 00 did 0000 t rc0O0yjii oommunioation) whioh showìtaí £',con ho rou itior2 ui hi u. S 'Co'o'L]o\7ing dredging and spoil bcn1a 1:tlJo oubmerod VeGOt- '/10C O iG,U1iOry OO"OO 0 C at!on Submorod veotat5.oi .o a pro°crrod 00000 haul tat of youngonauuon and auho vngotat- tor divappoara fron co aroa tho u'iober of () Limitations on eio and tyorri P doorecceo gfeLr&) used; (ii) limitations on cias of to be lcnded -1463 -

YRm/S101 Penaeus setiferus 7cl

7 POND FISH CULTURE avere total length of 112 mm, Survival was 84 percent. Wheeler did not add supplemental In general, P. setiforus ie regarded am foodj instead ho fertilized the pond water with being well suited for pond culturo. Attempts chicken manure. During the first35days, the at culturing have been experimental, however, shrimp grew at a rate of 2.33 mm per day, but and there are no commercial pond oulturo oper-. during the remainder of the experiment average ations at the present time, growth woe reduced to only 0.23 mm per day,

In one of the earlier culture experimonts bins(1967)reported that white shrimp oonduoted by Johnson arid Fielding(1956),sex- stocked in ponds as young adults in the spring ually mature P, setiferus that had been reared reached sexual maturity the following spring, or held in dirt ponds spawned viable eggs when but they did not spawn. Personnel at the Bears plaood in a 21 x9 x12 m concrete tank filled Bluff Laboratories, Wadmalaw Island, South with sea wator. The resulting larvae were rear- Carolina, have also succeeded in rearing white e1 to poetlarvae in 10 to 12 days. The shrimp shrimp postlarvae from eggs spawned in the wore reared to a total length of 50 mm in the laboratory. Lunz (1967) said that some eggs tanks in whioh they were hatched. Water in the hatched and 2 percent of the resulting larvae tank was fertilized with a oornrneroial inorganio were reared to postlarvae. fertilizer(7parts elemental nitrogen9parts phosphoric aoldj O parts poreh) to promote Johnson and Fjsldir (1956)were troubled algal growth for larval fooc. during early by excessive mortalitlee cc have been subse- development. Later stages were fed daily on quent workers, One ofhe causeo of mortality ground fish. has been the presence of predc000uo fich, Lune and ardon(1963)found that rotonono9 e opec- The work of Johnson and Fielding has not ifio poison for fish, does not kill olirimp and been repeated and subsequent workers have had other dooarod crustaceans when applied in prop- to rely on poetlarvae taken from the estuaries er ooncentrations. They uoed 1.4 kg(3lb) cf to etook their ponds, Recently, however, Cook powdered cubo root with o 5-porcont-rotenono (1969)developed a method for culturing white content for each 1,233m3of îator, iving e shrimp larvae under laboratory conditions, rotonono concentration of about1.5 parts por Although this method needs further modification million, Saponin,a poison contained in toc before large-scale culture becomes praotioal, seed cake, is used for fish control in Mien sufficient numbers of postlarae were reared to but liznz and a.arden(1963)found it,o be Loo stock a 0,05-ha (1/8-acre) pond. After the expensive for general use, oven though it laboratory-reared. shrimp had been in the pond worked satisfaotorily. 117 days, Ray Wheeler (personaloommunioation) harvested 32 kg (o lb) of shrimp with an - 1464 FSm 8101 Penaeum setiferue 61

8 REFERENCES

Agassia, L., Twelve lectures on comparative embriology, delivered before the Lowell 1849 Institute, in Boston, December and January1848-9. Redding and Co., Boston, 104 p.

Aldrich, LV., Observations on the eco1ogr and life oycle of Proohristianella penaei 1965 Kruse (CestodaTrypanorhyncha), J,Paraait,,51(3)37O-6 Anderson, W.W. and M,J. Lindner, Length-weight relation in the oomnon or white shrimp, 1958 Penneus setiferue. Spoo,soient.Rep.U.S.Fish Wildl.Serv.(Fish.),(256)l3 p.

Anderson, W.W. and 0.R. Luna, Southern shrimp, a valuable regional rosouroe, Leafi, 1965 Atlant.St.mar.Fish,Çomurn, (4)6p.

Anderson, W.W., J.E, King and M.J. Lindner, Early stages in the life history of' the 1949 common marine shrimp, Penasus sotiferue (Linnaeus).Biol,Bull,mar,biol,Lab,, Woods Hole,96(2):l68-72

Anderson, W,W., M.J. Lindner and J.E. King, The shrimp fishery of the southern United 1949 States. Commi Fish.Rev,, 11(2):l-.l7, Also issued as Fishery Leafl.U,S.Fish Wildl,Serv.,(36B)17 p.

Baxter,K.N.and W.C. Renfro, Seasonal occurrence and size distribution of poatlarval 1966 brown and white shrimp near Galveston, Texas, with notea on species identi- fjtiatjon, Fisher Bull,U,S.FishWildl,Sorv,66(1):14958 arden, C.M., Notes on postlarvae of commercial shrimp (Penaous) in South Carolina. 1961 Contr,Bears Bluff Labs., (33):8 p.

Boroughs, H., W.A. Chipman and T.R, Rico, Laboratory experiments on the uptake aoou- 1957 mulation and loss of radlonuolides by marine organisms, In The effsct8 of atomic radiation on oceanography and fisheries, Pubis natn,Res.Goun. Wash., (55l)8o-.7 Broad, A,C,, Environmental requirements of shrimp. Publs ubl,Hlth Serv., Wash.,(999- 1965 WP-25):86-91 - Bullis, E.R,, Jr. and J.R, Thompson, Collections bythefishing vessels OREGON, SILVER 1965 BP.Y, COMBAT and PELICAN made during1956-1960 inthe southwesternnorth àtlantio. S ec,ecient,Re ,U.S,Fish Wildl.Serv,(Fish,),(5lO)l3O p. Burkenroad, LD., The Penoeidao of Lou.ìFlianawith a discussion of their world relation- 1934 ships. Bul1.Am.!us.nab,his.,.68(2)361--143

, A newspeciesof paeu from the American ktlantic, Anata Aoad,brno l93C 8(4)3i5-8 Further observations on Penneidas of the northern Gulf of Rsxioo, 1939 Bgham ppsano r.Coll,,6(6)31-62

Butler, P.A., t'feotm on commercial fisheries. 171Effects of pesticides on fish and 1962 rildlife in 1960. Crc,Fisb Wi1dl,Serr, Wash., (143)2O.-4

Commercial fisheries L1tg'tions. InPesticide-wildlife studio, a 1963 rswlowof Fish cnd tid1ifo Sr'ice investigations during1961 and 1962. Ciro,Fiah lIildl cir.,Uach.. (l67)l1-25 Fixation of DDT in estuaries, 'Trans.N.Am.W±ldl,Conf. 313184-9 - 1465 - 82 FRm/S1Oi Pepasus setifere Butler, P,A. and P,F. Springer,Pesticides - a new factor in coastal environments. 1963 Trans,N,Ain.Wildi,Conf,, 28:378-90 Carranza9 J.,Pesca y reoursoepesqueros,In Loa recursos naturales dei sureste y su 1959 aprovechamiento, Mexico, D.F.,Instituto Mexicano de Recursos Naturales Renovables, 3:149-238 Chin, E,,The bait shrimp fishery of Galveston Bay, Texas,Trane.Am.Fieh.Soc, 89(2): 1960 135-41 Chin, E. and D.M. Allen,Toxicity of an insecticide to two species of shrimp, Fenaeus 1957 aztecus and Penasus setiferus,Tex,J,Sci,, 9(3):270-8 Christmas, J.T,, G. Ouater and P, Muegrave,Studies of annual abundance of' postlarval 1966 penaeid shrimp in the estuarine waters of Mississippi as related to subse- quent commercial catchen,Gulf Res,Rep, 2(2):177-212

Cook, H.L,,A method of fearing penaold shrimp larvae for experimental studies. FAO 1969 Fish,Ro, (57) Vol.3:709-15

Cook, E,L, and M,J0, Lindner,Synopsis of biological data on the brown shrimp Penaeus 1970 aziocus aztecus Frez-Farfante,FAO Fish.Rep., (57) Vol.4:1471-97

Dall, W,,A revision of th3 Australian species of Penaeinae (Crustacea Deoapoda: 1957 Penaeidao),Aust,J.mar,Froshwat,Res,, 8(2) :136-231 Darnell, ILM,,Food habits of fishes and larger invertebrates of Lake Poutchartrain, 1958 Louisiana, an estuarine community,Publa,Inst.mar,Sci,Univ,Tex,, 5:353-415 Dawson, C,E,,Balanus fouling of shrimp Penaeus0Science, N.Y., 126(3282):l068 1957

De Syiva, D,P,,The live bait shrimp fishery of the northeast coast of Florida,Tooh, 1954 Sor,Fla.St,Bd Conaerv,, 11:35 p, Ewald, J.J,,Investigaciones sobre la biologia dei oamarn comercial en el ocoidente 1965 de Venezuela0 Infine a.Fondo nac.Investnes agropeo., Caracas, 2:139-47 Gunter, G.,Deatiiof fishes due to coldon Texas coast, January, 1940, 1941 22:203-8 Q2!&' Studies on marine fishes of Texas,Publs,Inst,mar,Sci Univ,Tex,, i(i) 1945 l-190 Seasonal population changes and distributions as related to salinity,of certain invertebrates of the Texas coast including the commeroial shrimp, Publs,Inst,niar,Scj,jjniv,Tox,, l(2):l-.51 Principles of shrimp fishery management,Proc,Gulf Carjbb Fish Inst 1957 8:99-106 Habitat of juvenile shrimp (Family Penaoidae).Ecology, 42,598-600 1961 Shrimp landings and production of the state of Texas for the period 1956- 1962 1959, with a comparison with other Gulf States.Pubis Inat.mar.Sci,Univ. Tez,, 8:216-26 The proper zoological name of the North American white shrimp.Proc. 1963 Gulf Caribb,Fieh.Inet,,, 15,103-10 - 1466 -

/SiOi Ponaous setiferue

Gunter, G. ond U.H, Hildebrand, Destruction of fishes and other organisms on the oouth 1951 Toao coast by the cold wove of Jantiry 28-February 3, 1951. E000v,32(4)i 731-6

The reict&on of ,ou1 iina1l ofho oto.o and catch of the marine 1954 olirimp (Ponuouo oclforu) o Toxuo fatoro.Buil,mar.Soi,Gulf Caribb., 4(2): 95-103

Gunter, G., 3,Y, Chriatmam and R. Killebrew, Soma relations of salinity to population 1964 distributions of motilo estuarine organisms, with speoial reference to penaoid shrimp.Eoology, 45(1)l8l-5

Hoogaard, P.E., Obeervationo on spawning and larval history of the shrimp Penasue 1953 setifarus (L). Pubis lnst.mar.Sci,Univ,Tex,, 3(l):73l05

Herbst, J.F,W.,, Versuob einer Naturgeschichte der Krabben und Krebse nebst einer system- 1793 atisohen Beschreibung ihrer versobiedonen Arten, Berlin, Vol.2, 225 p.

Hildebrand, H,H,, A study of the fauna of the brown shrimp (Penasus zteouo Ive) 1954 grounds in the western Gulf of Mexico,Publo Ins par.Sci..UnivTex 3(2) 231-366

A study of the fauna of the pink shrimp (Penaous duorarum Burkenroa4) 1955 grounds in the Gulf of Campeche. Pub1,Ine-tomar,5cj vTa,.4(l) 169-232

Estudios biol6 icos prelininaros sobre la Laguna Madro deTamaulipas, l95BCiencia, Mix,, 177-9):l5l.-73

Hildebrand, H.H, and G.. Guntor, Correlation of rainfall with the Texas catch of white 1953 shrimp, Penaous setiforas (Linnaeus), 82:1515

Holthuis, L,B,, On he nemes of Penasue setiforus (L,) and Penaeus Burkonroad, 1962 3eo.Re,, l(3):1l5-

Cancer Lforug Linnaous, 1767 (Crustacea, Deoapoda): proposed valida;.. 1964 tion of neotype celootion under the plenarypowers Z,N,(S,) 1617, Bull, zool,Nom0, 21(3):227-9

Holthuis, L,B.. and H, Roea, Jr.., List of species of shrimps and prawns of economic 1965 yalue, Q_Fish.tea, (52):2l P.

Hu.ton R0N0: A second list of parasites from marine and coastal animals of Florida, i96Z Trns,P,mjc-oso,.Soc,, 83(4) :439-47

Hutton, R,F,, T. Ball and B. Eldred,Immature nematodes of the genus caecun 1962 Railliot and Henry, 1912 from shrimps, J. Parasit,, 48(2):327-.32

Hutton, R,F,, et al. Investigations on the parasites and diseases of salt water 1959 shrimps (Penaeidae) of sports and commercial importanoe to Florida, Teoh,Ser,Fla SBd Consery 26:38 p.

Inglis, A. and E. Chin, (roiced by I1.1, Ba:er), The Bait shrimp industry of the 1966 Gulf of Mexico,Fielorry Lsaf1,U.S.iish Wildl.Sox'v., (582)l0 p. Also issued as Fishory L3afl,U,S.Pumh iIlldl,Serv., (480)14 p, (1959)

Johnson,F.F.and M.J, Lindner, Shrimp induetry of the South Atlantic and Gulf states, 1934 Investi Rep,U,S,Bur.Fish,, 2l83 p.

Johnson, M.0 and J,R, Fielding, Propagation of the white shrimp, Penaeus setiforus 1956 (Linn.), in oaptivity, Tulane Stud.Zool,, 4(6):l73.-90 -1467

8t4 FRm/S101 Penasue setiferue

Jonee, D.B., 0. Moeller and C.E.F. Gersdorff,The nitrogen distribution and peroontase 1925 of some amino acIds in the muscleof the shrimp, Ponaoue aetiferus (L). J.biol.Chem.,65(1):59-66

Joyce, LA., Jr., The commercial shrimps of the northeast coast of Florida.Prof.Pap. 1965 Ser.mar.Lab.Fla.,6224 p.

Joyce, LA., Jr. and B, Eldred, The Florida shrimping industry. Eduo.Ser,Fla St,Bd 1966 Conserv.,15:47 p.

King, J.E., A study of the reproductive organs of the common marine shrimpPonaeue 1948 aetiferus (Linnaeus). Biol,Bull,mar.biol,Lab,1Woods Hole,94(3244-62

Klima, E.F,, Mark-recapture experiments with brown and white shrimp In the northern 1963 Gulf of Mexioo, Froc,Gulf Caribb.Fish.Inst.,l652-64

Knake, B.O., J.F. Murdook and J.P. Cating.Double-rig shrimp trawling in the Gulf of 1958 Mexico. Fishery Leaf.l.U,S.Fish Wildl,Serv., (470)dlp.

Knapp, F.T.,Menhaden utilization in relation to the conservation of food and game 1949 fishes of the Texas Gulf ooast. Thana.Am.Fiah.Soc.,79:137-44

Kruse, O.K., Parasites of the commercial shrimps, Penasus azteoua Ives, P. duorarum 1959 Burken.road, and P. setiferus (Linnaeus). Tulane Stud.Zool.,TÇ4)l23-44 Kutkuhn, J.H., Gulf of Mexico commercial shrimp populations - trends and oharaotor- 1962 istics,1956-59,Fishery B:1l.U,S.Fish Wildl.Serv.,62(212343-4O2

Leone, C,A, and C.W. Pryor, Serologioa]. correspondence amone three species of penasid 1952 orustaoea. J. Elisha Mitchell soient,Soo.,68(l):27-3l

Lindner, M.J., Estimation of growth rate in animals by marking experiments.Fishery 1953 Bull.U.S.Fish Wildl,Serv.,54(78),65-9

Survey of shrimp fisheries of Central and South America.Sec.soient. 1957 Rep.U.S.Fish Wildl.Serv.(Fiah.), (235):l66p.

Lindner, M.J. and W.W. Anderson, Growth, migration, spawning and size distribution of 1956 shrimp Penaeus setiferue. Fisher Bull.IJ.S,Fiah Wildl,Serv.,56(106)s555-645

Loesoh, H., Distribution and growth of penaeid shrimp in Mobile Bay, Alabama. Pubis 1965 Inst,mar.Sci.Univ.Tex.,10:41-58

Love, T.D, and N.H. Thompson, Annual report Bureau of Commercial Fisheries Technolo- 1966 gical Laboratory, PascagoulaMississippi, Fiscal Toar1965, Ciro. Fish ildl.Serv.,Wash., (25lj:25p,

Lunz, G.E., Harvest from an experimental ono-acre salt-water pond at Bears Bluff Labora- 1956 torios, South Carolina.Progve Fish Cult, 18:92-4 Pond cultivation of shrimp in South Carolina, Proo,Gulf Caribb.Fioh, 1958 Inst.,10:44-8

Annual report1965-1966, Bears Bluff Laboratories,Wadmalaw Island, 1967 South Carolina, Contr,Boare Bluff Labs.,44:11 p,

Lunz, G.E, and G.M. Bearden, Control of predaceous fishes in shrimp farming in South 1963 Carolina. Contr,Bears Bluff Labs,,36:9 p.

McFarland, %.N, and B.D. Lee, Osmotic and ionic concentrations cf ponaoidean shrimps 1963 of the Texas coast, Bull.mar.Soi.Gulf Caribb,13(3):39i-417 - 1468 -

FRo/S101 Penasus setiferue 8:5

Mock, 0.5., Natural and altored estuarios habitats, Galveston, Texas, Bureau of MS Commercial Fisheries Biological Laboratory, 18 p,

Olivier, G,A., Ecrevissa, Astaoue, In Enoyciopddie methodique.Histoire naturelle. 1791 Insectos, Parie, Vol.6:327-49

Pearson, J.C., The early life histories of some American Penaoidae, chiefly the corn- 1939 marcial shrimp, Penseus setiferus (Lion.). Bull.Bur,Fish.jjaeh.,49(3):73p.

P&ez-Farfante, I,, Western Atiantio shrimps of the genus Penaeus. Fish.Bull,U,S.Fish 1969 Wildl.Serv,, 67(3):46).-591

Sinopsis de datos biologicos sobre el oamarón blanco Penaeus schmitti 3.970 Burkenroad, 1936, FAO Físh.Rep., (57) Vol.4:1417-38

Power, E.A., Fishery statistics of the United States, 1958. Statist,Di'.U.S,B'ish Wildi. 1960 Serir., 43:424 p.

Henfro, WC, and H,A. Brusher, Seasonal abundance, size distribution, and spawning of MS penaeid shrimp in the northwestern Gulf ofMexioo,

Rioja, E., Estudios oarcinológioos. 1, Caracteres sexuales secundarios de los 1939 Penaeidae - El órgano antennular de los machos,An.Inst,Biol.Univ,Méx., 10 313-9

Estudios carcinológicos. 4, Observaciones sobre las antenulas d al- 1940 gunas especias dei gónero Penseus, An.Inet.Biol.iJniv,Néc,, ll:26773

Estudios carciriológicos. 8. Contribución a la morfologZs e inter- 1941 pretación del pstasrna de los Penaeidae (Crust. Doopcdos),!.n.Inst,Biol,Univ, Mdx., 12:199-221

Estudios carcinológicos. 20, Valor dìagn6stco dei tercer saxilpedo 1948 y de sus cerdas on algunas especies del gdnero Penanus de la costa de Móxico, kn.Irtst,Biol.Univ,Móx., 19:453-67

Robas 3.5,, Shrimp trawlin..g gear as used in the Gulf of Mexico. In Modern fishing 1959 gear of the world, edited by H. Kriatjonsson.London, Fishing News (Books) Ltd., pp. 311-6

Say, T,, An account of the Crustacoa of the United States, J.Aoad.nat.Sci,Philad. i, 1818 235-53

Sba, A,, Locupletissisi rerun naturalium thesauri accurata descriptio, et ioonibu 1759 artificiosiseimie expressio, per universs physices historian. Anni;alaedami, H.D. Gaubium, Vol.3, 212 p,

Simmons, E.G., An ecological survey of upper Laguna Madre of Texas.Pub1Inst,sar. 1957 Scj.Unjv,Tex,, 4(2)156-2O0

Sprague, V,, Notes ca three zicrosporidian parasites of deospod orustacea of Louisiana 1950 coastal waters. Occ,Pap.msr.Lab,La St.Unlv., (5)sl-8

_, Protozoa. in Gulf of Mexico, its origin, waters and marine 1if, 1954 Fishery Bull,U.S,Fih Wildl.Serv., 55(89):243.-56

Springor, Sand H,B. Builis, plortory shrimp fishing in the Gulf of Mexico, 1950W 1952 51. Fishery 1.U.S.Fish Wiidl,Serv., (406)34 p.

Temple, R.F, and C.C. Fisher, VertIcal distribution of the planktcnio stages cf penseid 1965 shrimp. Pubis Inst,mar.Sci.Univ,Tex,, l0:9-67 - 1469 - 8:6 FRm/S101 Penaeus aetiforus

Temple, R.F. and C.C. Fisher, Seasonal distribution and relative abundance of planktonio- 1968 stage shrimp (Penaous app.) in the northwestern Gulf of Mexico,1961. ery Bull,U.S.Fish Wildl,Serv,,66(2):323-34 United States Fish and Wildlife Service, Survey of the United States shrimp industry0 1958 Vol.1. poo.scient.Rep.U.S,Fish Wildl.Serv,(Finh,),(277)2311 p.

Urein, E,, On the incorporation of temperature in the von Bertalanffy growth equation. 1963 Meddr.Danm,Fiak.-o Havundors,4(1)21-16

Viosoa, P,, Louisiana greatest in the production of shrimp, Penaeuo sotiforue. Biorin. 1920 Rop.La Dep,,Conserv,,41106-30

A critical analysis of practices in the management cf warm-water fish 1943 with a view to greater food production. Trana,An.Fish,Soo.,732274-83

Welford, L.A., A new graphic method of describing the growth of animals. Biol,Bull, 1946 mar ,biol,Lab. ,Woods Hole,90(2):l41-7

Waterman, P,E, and F,A, Chaco, Jr., General crustacean biology, In The physiology 1960 of orustaooa, editad by T.H, Waterman, Now York2 Academic Proas, Vol.1: 1-33

Weymouth, F.W., M.J, Lindner and W.W. Anderson, Preliminary report on the life hietory 1933 of the common shrimp Penaeus sotiferue (Lizui.), Bull,Bur.Fish,,Wash,,48 1-26

Williams, A.B.., Identification o± juvenile shrimp (Penaeid.ae) in North Carolina.. 1953 J,Elisha Mitchell sciont,Soo.,69(2):156-6O A contribution to the- life histories of commercial shrimps (Ponaeidae) 1955 in North Carolina.. Bull mar,Sci Gulf Carib,5(2):1l7-46 A survey of North Carolina shrimp aursery grounds. ,Elisha Mitchell 1955e soient.Soc,,71(2)2200.-7

Substrates as a factor in shrimp distribution.. Limno].,Oceanogr.., 3(3): 195 283-90

Spotted and brown shrimp poatlarvee (Penaeus) in North Carolina. Bull, 1959 mar.Sci.Gulf 0aribb, 9(3):28l-90 Marine docapod crustaceans of the Carolinas. Fishery Bull,U.S.Fimh,Wildl. 1965 Serv,,65(1)1298 p.

Young, J.H., Morphology of the white shrimp Penaous sotiforus (Linnaeus1758).. Fish, 1959 Bull,U.S,Fish Wildl.Sorv.,59(145)2168 p, Anon,, Canoer setiferus Linnaous,1767(Crustacea, Decapoda): validation of the 1967 neotype under the plenary powers, Bull.zool.Nom,,24(3)1151-2