UNITED STATES DEPARTMENT OF THE INTERIOR Fred A. Seaton, Secretary

FISH AND WILDLIFE SERVICE Arnie J. Suomela. Commissioner

NATURAL HISTORYOF THE SANDBAR SHARK

EULAMIA MILBERTI

By STEWART SPRINGER

FISHERY BULLETIN 178

From Fishery Bulletin of the Fish and Wildlife Service

VOLUME 61

PUBLISHED BY U.S. FISH AND WILDLIFE SERVICE • WASHINGTON • 1960

PRINTED BY U.S. GOVERNMENT PRINTING OFFICE, WASHINGTON, D.C.

For sale by the Superintendent of Documents, U.S. Government Prlntln~Office, WashlnAton 25. D.C.Price 35 cents

The series, Fishery Bulletin of the Fish and Wildlife Service, is catalogedas follows:

U. S. F·i8h and TV-ildlife 8el'l'i.ce.Fishery bulletin. v. 1-

·Washington, U. S. Govt. Print.. Off., 1881-19

v. in illus., maps (part fold.) 23-28 em.

Some vols. issued in the congressional series as Senate or Housedocuments.

Bulletins composing v 47- also numbered 1-Title varies: v. 1--49, Bulletin.Vols. 1--49 issued by Bureau of Fisheries (called Fish Commission,

v. 1-23)

1. Fisheries-V. S. 2. Fish-culture-V. S. I. Title.

II

SHll.A25

Library of Congress

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t59r55b1]

9-35239*

CONTENTSPall'e

Introduction_______________________________________________________________ 1~omenclature______________________________________________________________ 2Populations of Eulam-ia milberti____ _____ _____ _____ ______ ___ _ 3Ecological and systematic relationships of the genus ElIlam-ia.__ ___________________ 4Specimens examined ._ ______ _____ __ ______ ____ ____ 5Field recognition ____________________________________________________________ 6Distribution of E'ltlamia milbertL _____________________________________________ 10

General nature of distribution_ ___________________________________________ 10Factors affecting distribution_ ____________________________________________ 12Nursery grounds and distribution of young_________________________________ 13Distribution off Atlantic coast of the United States and in eastern Gulf of

]dexico______________________________________________________________ 14Distribution in Bahamas and West Indies___ ___ __ ___ _____ __ _ 16Distribution in western Gulf of ]dexico and western Caribbean _______________ 17

~igration---------------------------------------------_____________________ 19Eleproduction_______________________________________________________________ 20

Courtship and mating___________________________________________________ 20Time of mating____ __ ___ _ ______ __ ___ __ __ _____ __ _ _ 21Development of the embryo___ ___ _____ __ _ ___ __ __ __ 22Number of young in litter . ___ ___ ___ _____ __ ___ ____ ___ _ 23Length of young at birth ________________________________________________ 23Abnormal embryos___ ____ _____________ ______ __ _______ _______ __ _____ ___ __ 25

Sex ratios__________________________________________________________________ 25Growth and size at maturity _________________________________________________ 26Food and feeding habits _____________________________________________________ 29Abundance_________________________________________________________________ 32Enemies ~____ 32Summary__________________________________________________________________ 34Literature cited___ _________ __ ___ __ ______ _____ __ _________ _____ ___ 36

ill

IV

ABSTRACT

Populations of l'andbar sharks of the eastern and western parts of the AtlanticOcean are defined and general problems of nomenclature, the ecology of largecarcharhinid sharks, and field recognition of sandbar sharks are discussed.

A more-detailed account of observat,ions on Eulumiu milberti, restrir.t,ed t,o thepopulation of the western North Atlant,ir., is given, out.lining distribution of adultsand young, migrations, development, and behavior, based on observations fromthe commerr.ial shark fishery which operat,ed from r.enters in the SoutheasternStates from 1935 to 1950 and supplemented by dat,a from reseal'r.h vessels operatingafter 1950.

Comparisons wit,h other species in the area, lists of large species-of sharks takenat cel't,ain times off Salerno, Florida; Bimini, Bahamas; t,he mouth of the Missis­sippi River, Louisiana; and the Caribbean coast of Nicaragua-Costa Rir.a, as wellas disr.ussiollS of interspecies r.ompet,ition. are inr.luded.

NATURAL HISTORY OF THE SANDBAR SHARK, EULAMIA MILBERTI

By STEWART SPRINGER, Fishery Methods and Equipment Specialist

BUREAU OF COMMERCIAL FISHERIES

This account of the sandbar sha,rk, Eulamiam;ilbe~l'tl (Miillerand Henle), is an attempt to bringtogether aU the significant information on onekind of common and moderately large shark.Sharks have been studied because they are occa­sionally dangerous to man, often a nuisance tofishermen and, in the past at least, have beenvaluable as a source for food, leather, vitamin A,fish meal, and some specialty products. A rathercomprehensive body of knowledge exists aboutsome of the smaller species, such as the compara­tively valuable soupfin shark of the coast ofNorth America (Ripley, 1946) and the schoolshark of Australia (Olsen, 1954), both species ofGaleol'h.i-nus, nnd the common spiny dogfish,Squaltl8 (Ford, 1921; Hickling, 1930; Temple­man, 1944). Information on the natural historyof the larger species is fragmentary. This is tobe expected, because large species not only aredifficult to cntch and handle, but also are far­ranging and require observation over a wide geo­graphical area.

The sharks, together with their relatives, theskates, rays, and chimaeroids, form a class ofvertebrates that is sharply set off from the classeswhich contain the fishes, amphibinns, reptiles,birds, and mammals. The sharks and other mem­bers of the class Chondrichthyes have cartilagi­nous skeletons, and while elements of the sharkskeleton may become calcified, no true bone isformed. This is the basis for the definition thatis generally used to distinguish the Chondrich­thyes from the higher vertebrates. But there areother differences in the chemistry and physiologythat are very likely of great importnnce but arelittle understood. The evolutionary connectionsof the modern sha.J'ks and their allies with othermodern vertebrates are obscure and of great an­tiquity.

NOTE.-Approved for publl~8tlon. October 27. 19118. FisberyBulletIn 178.

SIUH'ks occupy a place in nature at the top ofthe food chain. As predators they compete withman, but it is by no means established that theirpredatory activities are always harmful. Theyare a nuisance or are harmful to fishermen chieflybecause of the damage they do to nets or to fishthat have been caught on setlines. In some locali­ties, in England and Australia, for example,sharks are utilized and are consequently of somevalue. In the United States, landings at presentare of no great importance.

Sharks may be chUlgerous to man through at­tacks on swimmers and survivors of marine dis­asters, but Etdamia '1nilbe'l'ti is· not a speciesimplicated in well-documented records of attacks.There may be sevel'ltl reasons for this. E. mHbe1'tiordinarily stays away from beaches and does notoften feed at the surface. It usually seeks smallprey. During the summer, when the female sand­bar sharks come inshore near the heavily popu­lated centers from New York southward alongthe Atlantic coast to give birth to their young,it is not t.heir habit to seek food. The large malesdo not come inshore. So the sandbar shark, whilela,rge enough to be dangerous and perhaps themost common of the larger sharks in shore waterssouthward from New York, is isolated by itshabits from encounters with man. Nevertheless,the sandbar shark is potentially dangerous to manand might become a more, serious danger withminor shifts in the environmental situation.

The most annoying aspect of my work withsharks, prior to the publicat.ion in 1948 of thefirst volume of Fishes of the 'Western North At­lantic on sharks by Dr. He.nry B. Bigelow and'Villiam C. Schroeder, was that many westernAtlantic sharks could not be identified with con­fidence because of It scattered literature of vary­ing quality. It is appropriate that I acknowledgethe importance of this excellent general work tome, because without it I\nd without the encourage-

1

2 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

ment of its authors, I would not have attemptedpreparation of t.his report. Dr. Richard H.Backus of the Woods Hole Oceanographic Institu­tion, Dr. Giles W. Mead of the U.S. Fish and'Vildlife Service, lwd Dr. Leonard P. Schultz ofthe U.S. National Museum made many helpfulsuggestions during the preparation of this report.Captain B. ,V. "Tinkler was especially helpful inkeeping fishing logs and measurements of about1,300 large sharks he took off the Bahama Banksand off Nicaragua. Records obtained while I ,vasemployed by the Shark Industries Division ofthe Borden Company and while I was aboard theexploratory fishing vessel Oregon of the Fish and'Wildlife Service comprise. the basic data usedhere. Special assishtnce was given me also by theLerner Marine La.boratory of the American Mu­seum of Natural History, by permitting 2 monthsof field study at Bimini, Bahamas, in the summerof 1948. In all my work with sharks, I have beengiven the most generous help by my associates incommercial shark fishing and aboard exploratoryfishing vesse.Is.

NOMENCLATURE

This report is not int.ended to set.tle problemsof nomenc.lature and taxonomy, but. to be usefulit is necessary t.o name the sharks under discus­sion and t.o define the names used. My choice ofa name for t.he sandblu' shark is Enlmnia, ndlbe'l'ti(MUller and Henle) 1841. Use of Eula1nia fol­lows my partial revision of the carcharhinids(Springer, 1950). For the specific name milbe'l'fiI follow Bigelow and Schroeder (1948) who notethat, if it is finally proved t.hat the Mediterraneanform is ident.ical with t.he American, the nameplum,beu8 Nardo 1827, must. be used for the com­bined species in place of'rnilbe1·t-i.

J disagree, however, with Fowler (1936), witht.he preceding stat.ement by Bigelow and Schroe­der, and with Tortonese (1951,1956) t.hat Nardo'sdescription is valid. The description by Nardowould apply to almost any cal'charhinid amI thespecific mention of the rounded snout 1 wouldapply better to some other carcharhinids than tothe sandbar shark. Because t.here is no type andbecause Nardo's descript.ion would apply to al-

1 The total description and diagnosis of Sqrwl;'8 pll/ll1 bello h~'

Nardo, 1827. p. 35, Is as follows: "Speciel secllndae cOII\·p.nitexacte Sqllal. Glallell8. Hloc..1 color excil;pretur pt formarostri quae in exemplarl nostro rotunda est."

most any carcharhinid if applied liberally butto none if applied strictly, I regard Sq1talJu81)lum,beu8 as It nomen nude'ln.

I am also unable to accept Nardo's descriptionas specifically applicable to the sandbar sharkbased on the argument that t.he. sandbar shark isthe most common large carcharhinid of theAdriatic.

A most extraordinary snarl has developed overthe years in the determination of the scientificname to be applied to the sandbar shark. Theorigin of t.his complication probably lies in thepeculiarities of the distribution of species ofcarcharhinid sharks along the Atlantic coast ofthe United States. Mistakes in identification ofspecimens have been frequent, probably becausethe descriptive accounts of the early authors werevery brief and did not select truly diagnosticfeatures for emphasis. Systematists had too fewspecimens ilnd too little datlt on distribution tonote that. segregation of the sexes and segregationof the adults and young characterized thesesharks at some seasons.

In the latitudes from New York to ChesapeakeBa.y at depths within easy rench of collectors orfishermen, two common large carcharhinid sharksoccur, the sandbar shark, E. m,ilbe1'ti (MUller andHenle), and the bull shark, Oa.1'charhimt8 leucas(MUller and Henle). The sandbar sha.rk is rep­resented in this area by adult females and by theyoung of both s~xes, but ra.rely by adult. males inthe observable elements of the population. Thebull shark is represented usually by adult males,but females and young are also present sporadi­cally.2

The ranges of the sandbar shark and the bullshark will be discussed later a·s well us the ap­pa.rent competition between these species. Aneffect of the occurrence of the two species to-

• Large male Ct,rchm'hilil/o lellca8 were reported from theChesapeake Bay area by Schwartz (1958) ; one was taken fromthe Patuxent River In 1957 and another at Flag Pond in thesummer of 1958. This Is apparently the first published reportof the sllecies from Chesapeake Bay. Specimens of large sharkscame to the attention of Edgar H. Hollis. of thp. Mar~'land De­partment of Tidewater Fisheries in 1957, because Chesapeakefishermen regarded them as rarities. Photographs of the speci­m",ns sent me b~' Mr. Hollis were sufficient to permit identifica­tion as C. lel/Ca8. Nichols (1918) and Nichols and Breder(1927) reported a. lel/Ca8 from the south shore of Long Island,noting that these specimens were huge males. Attention Iscalled to this parenthetically because the appearance of adultmales at the perlpher~' or In the cooler parts of ranges ofcarcharhlnld sharks iM frequent.

NATURAL HISTORY OF THE SANDBAR SHARK 3

gether, howe,ver, has been to foster confusion inthe nomenclature. There appears to have beena tacit. assumption by some nat.uralists that sexualdimorphism accounted for differences in sandbarsharks and bull sharks despit.e recognition of theexist.ence of both species.

Superficially, the sandbar sha,rk and the bullshark resemble one anot.her but, as will be shownlater, sandbar sharks can easily and positively beseparated from bull sharks on t.he basis of severalanat.omical characteristics. Identification ofspecimens from t.he Middle Atlantic States, par­ticularly the Atlantic coast from Cape Cod toChesapeake Bay, presents an added difficult.y be­cause of t.he several very similar offshore specieswhich may be caught occasionally, but probably .rarely in inshore waters: Eulmnia. obsc/u'u (Le.Sueur), E. fa.leifo·r·mls (MUller and Henle), E.ftorida'lla (Bigelow, Schroeder, and Springer),and E. allilna Springer. R.ecent. unpublishedrecords of the occurrence of E. 11lRberti youngand of t.he occurrence of '(!al'ch.arM'I1US leucasadults are rather numerous, and following publi­c!ltion of t.he first. volume of Fishes of t.he W'est.­ern North Atlantic (Bigelow and Schroeder,1948) t.here appears to be lit.tle confusion of thetwo species.

Belaboring the point t.hat. descriptive accountsof carcharhinids must. be det.ailed and selective tohave any meaning seems necessary t.o affect t.heint.renched misconcept.ions about. E. 11l.ilbertl t.hat.can be derived from t.he literature. Fowler's(1936) description of E. plttjnbe1ls (plum.beu.s =

11l.ilber!i), which was based on American MiddleSt.ates examples, alt.hough in a report on 'VestAfrican marine fishes, is not. unique in confusingE. 11lRb&rti wit.h anot.her species,S but it is de­t.a.Hed enough to be especially vulnerable t.o criti­cism. Mon." elements of his description fit. thebull shark, (!aJ·eh.a.rhIn1ls leu<Jas, t.ha.n E. milberti,but additional confusion is int.roduced by theprobabilit.y t.hnt juvenile and adult charact.eristicsof bot.h spedes are mixed. There is no selectionof diagnostically useful characteristics for em­phasis. The result. is a, plausible literary syn-

3 Garman's illustration of Carr.lla·rhinus platllodon (1913. pI. 3.IIgs, oj, and 6) appears to bE" ver~' well drawn frolll a specimenof Eulamia milbl'rti. Bigelow and Schroeder (1948) note thatGarman's illustration Is mI81I1b('l('d, Th(' accompanying illustra­tion of the teeth In Garman's plat(' 3. IIgur(' 5. appears to hav('be('n drawn from the t('('th of platllodon, Oa·rclla·r/linlts pla.tll­odo'n (Poeyl Is a synon~'rn of C', Il'lIcas (Milller and HNlle),

thesis that is a. haza.rd to one attempting to fit areal shark to a position in zoological classification.

POPULATIONS OF EULAMIA MILBERTISandbar sharks, Eulmnia m.ilbe-rti, occur in

portions of the te~nperate and tropical Atlant.ic,the Caribbean, t.he Gulf of Mexico, and the Medi­t.erranean. Our data in this st.udy primarily coverthe population inhabiting t.he western NorthAtlantic, the Gulf of Mexico, and the east.ernCaribbean. Lat.er, the normal movements anddistribut.ion of this population will be discussed.We need first to consider the relationships of thevarious populat.ions. In addition to the popula­tion of the western North Atlantic two othersmay be roughly defined. One occurs along thecoast. of South America from Trinidad east.wardamI southward. The other is found along thl'west coast. of Africa and is presumed to be con­tjnuous with the st.ock entering. t.he Mediter­ranean.

The population occurring on the coast of SouthAmerica appears to be a minor one. The specieshas been reported and figured by R.ibiero (1923)from the coast of Brazil.

'While engaged in commercial shark fishing inApril and May 1949, I made shipboard examina­tions (Springer, 1949) of a series of sandbarsharks from the north and east coasts of Trinidadand identified them in error as E. plu.m.belM. Inow believe that differences between the Trinidadspecimens and typical '11l.ilbe1'fi from the Atlanticcoast of t.he United States are insufficient to war­rnnt. recognition of separate species, and that. the­conservative course, pending accumulation of newdata, is to look upon the various Atlantic sandbarsharks as representing a single species. At. theTrinidad locations, adult males and fpmales aswell as young of all sizes from 4 feet upwardwere taken on single setlines. Although COln­mercial shark fishing was carried on throughout1949 from the const"of French Guiana westwardto the Gulf of Venezuela" sandha.r sharks werereported only from the north and east coasts ofTrinirlnd and chiefly from depths of 5 to 20fathoms near Galera Point.

There are no records of snndbar sharks fromthe 'West Indies north of Trinidad except. fromthe north coast. of Cuba and from the westernpart of the Bahama Ba.nks. This is not., of

4 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

course, conclusive evidence of their absence, froma region sO poorly known ichthyologically as theWest Indies. Nevertheless, all of the evidencepoints to a discontinuous distribution with noregular contact betwee·n the population knownfrom Cape Cod to Costa Ricll and the SouthAmeriean population known from Trinidad andthe ellst coast of South America south of theAmll7.0n.

In connection with possible future work withthe sandbar sharks, it should be noted that theE. mHbertl from Trinidad were taken in eddiesof the very strong, westerly current flowing be­tween Trinidad and Tobago; and that recruiL­ment for this stock could take place in part bytransport by the Equatorial Current of the grow­ing young to Trinidad from shore waters of theAfrican coast.

The stock of Eu1amia milberti in the easternAtlantic. is known from scattered records fromthe Mediterranean and the west coast of NorthAfrica as sumnmrized .by Tortonese ( H)56) .These records cover a long period of time andalthough critically reviewed by Tortonese andunquestionably aceurate, they give. little basis foran estimate of the abundance of the slUldbarshark in rehttion to the abundance of other largespecies of the area. For our purposes here, thatis, to estim.ate the relative. importance lmd abun­dance in comparison with other large sharks ofthe area, reports by Cadenat (1950, 1957) onE. 'milbe1'fl and other species from the coast ofSenega.! ltre quite informative. Cadenat has beenable to make observat.ions on fresh material froma fishery taking relatively large numbers of thelarger species of sharks. His reports suggest thatthe stock of E. milberti off northwest Africa isa strong one.

The list of species of large sh~rks reported byCadenat is quite similar to lists of large sharksfrom the southeastern coast of the United St.ateR.The endemic species of both areas are the smallersharks.

Precisely the same factors of prevailing windand surface currents that make the southerncrossing from the North African coast to Trini­dad easier for man when it is from east to westmay be expected to operate for sharks. Simi­larly. for a more northerly crossing, the one fromwest to east is more easily followed. The postu-

late that such contacts as exist between the stocksof the western Atlantic. and the stoc.k of theeastern Atlantic result frqm exchanges followingthis general clockwise eirculation is a reasonableone. No actual evidence of regular contacts be­tween the three stocks exists, however, and thereis substantial reason for the, belief that move­ments of individual slutrks from one stock to an­other are relntively infrequent occurrences.

Knowledge of the distribution of large sharksin oceanic situations at conside,rable distancefrom land was ext.remely meager until very re­cently when data from oceanographic vessels andexploratory fishing vessels became available. Themost comprehensive study c.overs sharks of theCentral Pacific (Strasburg, 1958) in which dat.ashowing patterns of dist.ribution of some of thelarger species is given. Before the appearance ofthat study and of a less comprehensive ac.countof Atlantic pelagic sharks (Backus et aI., 1956),questions of shark distribution seaward of thecont.inental shelves were unanswerable.

Now, while it. is known that neritic species oflarge sharks are capable of moving over greatdistances of open ocean, the.re is increasing evi­dence that they rarely do.

ECOLOGICAL AND SYSTEMATIC RELA­TIONSHIPS OF THE GENUS EULAMIA

The genus E1tkunia, nItty be divided into twogroups on the basis of tIl£' st.ructure and arrange­ment of the dermal denticles. The group towhich E. 'm-ilbedi' belongs is characterized bynonimbricate denticles as contrasted with theother group whic.h has denticles with overlappingedges or points. The mHbel'ti group includescomparatively few species. Probably Eul<!,miadU8811:m.iel'i (l\:Hi.lle.r and Henle) and E. japo'niC'/18(Schlegel) of the western Pacific belong here. Inthe Atlantic, the group is represented by a deep­water species, Eulam.ia, aUima. Springer, which isquite different. from milberti, l.lOt only in its mor­phology but in its habitat. Aside from E. Idti'111.a,the only Atlantic representative of the genusE'ltlamia (or any carcharhinid genus) withwidely spaced, nonimbricate denticles is E1tlmniamilbati. the sandbar shark.

E. mJlberti has the shoalest range and occupiesthe most inshore habitnt of any of the 5 or 6species of Eul<lmi{l, of the Atlantic. coast of North

NATURAL HISTORY OF THE SANDBAR SHARK 5

America. Although 1nilbel't-i may be in competi­tion with other spec.ies of E~da1nia for food insome parts of its range, it does not comiJete withother species of Enhnnia for nursery grounds.

Of the other carcharhinids of the northwestenlAtlantic, the genera Prio'nace and Pte1'olamiopsare pelagic surface dwellers; Hypoprion is con­fined to waters generally deeper than 1.00 fathomsnear shelves or banks; Negaprion, .A. priondon,ScoUodon, and Oa1'cha?'hinu,s are shallow watersharks that spend at leust some part of their livesin shallow lagoons, river mouths, or estuaries,and venture into deeper water rarely except fort.ransitory movements; the species of E~tlanllia aresharks of" the continental shelves, oceanic banks,and islund terraces, although some species extendtheir ranges well inshore and a.Iso for consider­able distances beyond the limits of the Conti­nental Shelf. The only other western North At­lantic carcharhinid genus, Galeoce·rdo, is repre­sented by a single spec.ies in subtropical andt.ropica.I waters ont. to depths of at least 200fathomfl. It does not exhibit the specialize.dschooling habits of the other carcharhinids, showsno strong migratory tendencies, and is less re­stricted in habitat choice than the others. There.seems to be a tendency to greater variation in thenumber of young produced as well as a greaternumber per litter in Galeocm'do and P1io'nanoeand possibly also in Pte'l'olam;iops than in othernorthwestern AtJantic carcharhinids. Insofar asis known, there are no very important differencesin the general outlines of the life history patternsof Negapl'io-n, .A.p1'ionodon, ScoliodO'n, Oa.rchm·­Mnlls, and E-u.lamia" although there appear to bemany differences in deta.il.

Barriers which may restrict the movements ofthe larger sharks including E~tl((.1ni(L milbe1'ti arenot readily apparent. Occasional captures ofsharks outside areas of normal c.oncentration ofthe species prove that they can and do wander.The remarkable thing is that la.rge sharks tend torema.in within definable habitats and geographi­cal ranges.

Since species of Eu,lam.ia a.re, in general, les!';dependent on land masses than (!arcna1'Mnlls andextend their activities regularly to surface watersof the open ocean beyond the Continenta.I Shelf~

it would not be surprising to find that somespecies have a very wide distribution in temperate

552508 0-60--2

and tropical seas. Eu.lam-ia fioridana (Bigelow,Schroeder, and Springer) may be an example ofsuch a distribution (see Strasburg, 1958). Thosespecies of Eula'In'ia, such as 1nilbe1,ti, which aretied to shallow-water habitats are presumablysubject to a greater degree of isolation.

SPECIMENS EXAMINED

Spec.imens, records, and field observations forthis report have been assembled over a period ofabout 25 years during which. time I have exam­ined seve.rnl thousand sandbar sharks. Availablerecords of the commercial shark fishery covermore than 100,000 adult Eul(J;nl-ia mUbe1'ti. Abouthalf of these sharks were measured at the pointof landing. Earlier records of the stations in­clnded specimens of Eulmnia altlm..a and EUl(l;m.iafiorida;Jl(t under the heading sandbar sharks.Since I visited most of the stations frequently,llnd during part of the period between 1935 and1950 supervised rec.ording procedures, I saw rela­t.ively large numbers of sandbar sharks. Speei­mens which appeared unusual to stat.ion employ­ees were retained when practicable for my inspec­tion. :Most of my observations were made alongthe coasts of southern Florida. Adequate num­bers of specimens for some purposes have beenexamined from the eastern and northern parts ofthe Gulf of Mexico, the Atlantic coast of theUnited States south of Cape Cod, and from theCa.ribbean coast. of Nicarllgua lmd Costa Rica.The available material in several museum col­lee-tions in the United States was studied, but t.hisconsisted chiefly of preserved embryos or veryyoung sharks and dried jaws.

The collection of data in the shark fishery suf­fered from inte~'ruptions and was assembled toaid ItIl industrial operation rather than for abiologic.al study. The difficulty in handling speei­mens, averaging nearly 7 feet in length as adultswith an average weight of about 135 pounds, hasmade it necessary to select different series orsamples for diffe·rent object.ives: one sample forlength-weight relations; another for tooth counts,and so on.

I was unable to find spirit-preserved specimensof eastern Atlantic or :Mediterranean origin re­ferable to eit.her E. lJlumbetls or E. mHbe1'ti dur­ing II hasty examination of catalogs and specimensat the :Museum d'Historie. Naturelle in Paris or in

6 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

the British Museum of Natural History, althoughspecimens from the western Atlantic were pres­ent. Tortonese noted (1938) that there are twosppcimens in the Musee di Trieste collected in1869 and 1871 and, after exa,mination of speci­mens labeled milbe'l'ti from the western Atlanticin the Museum at Paris and the British Museum,he indicated (in 1951) that he regards 1nilbe1'tias a synonym of plU'lnbeus.

FIELD RECOGNITION

Eulamia milbert'i is commonplace in appear­lwce. It has neither unusual color markings norspectacula l' stTuctural features. The length ofthe shark at maturity, 7 feet, makes the speciestoo large for the biological collector and too smallto interest the journalist. It is necessary tosearch for distinguishing featUl'es (fig. 1). Theopportunity for comparison of series under onli­nary circumst-ances is negligible, and almost allidentificat-ions of the larger sharks are necessarilymade in the field. The suggestion that many ofthe presently recognized species of carcharhinidsharks are not in fact separable from one anotherbut should be regarded as unidentifiable parts ofa species complex has been advanced in specula-

tive conversation by some of my friends who areichthyologists. This view may easily developfrom unsatisfactory attempts to make identifica­tions with methods which are quite adequate andsuccessful in application to teleosts but fal] shortwhen applied to sharks, and particularly to car­charhinid sharks. E. 1nilbeTti, in waters off theUnited States, is readily defined and problemsconcerning it are not complicated by the existencp.of geographic or environmental races or sub­species, insofar as the available evidence shows.This is apparently not true of some of the otherca.rcharhinids where separate populations may bedefined on the basis of morphological differencesshown in the analysis of adequate series fromdifferent a,reas.

The keys and descriptions given by Bigelow andSchroeder (Hl48) are adequate for the identifica­tion of the carcharhinid sharks of the westernNorth Atlantic excepting Eula1nia altil1UL, whichwas described (Springer, 1950) after publicationof this work. Nevertheless, identifications needto be made carefully because of the general struc­tural similarity of the species which look alike onsuperficial examination. Sharks of the genusEulamia in the !aldfm'mis-sp1"in,qeri group are

FlOUR!;; l.-E/llQ1n?·a, millJel"li in un exhibition tank. TIl(' high, triangular first dorsal fin, nonfalcatE' pectoral fins, andrrlativrly hip;h srcond dorsal and anal fins, ncarly cqual to one anothcr in area, arc chantct('ristic of the species_ (Pho­togl'llph courtesy of Marinc Studios, Marincland, St.. AUll;ustine, Fla.)

NATURAL HISTORY OF THE SANDBAR SHARK 7

not well known and possibly are incompletely de­fined. Minor differences between Atlantic andGulf of Mexico populations of E. obse'll,Ta needfurther study. There is no difficulty, however, indistinguishing E. 1nllbeJ'ti from these species orfrom other species of sharks ordinarily foundwithin its geographical range.

The importance of determining the presence orabsence of a middorsal ridge (a low ridge in theskin extending for all or a part of the distancebetween the first and second dorsal fins) for theidentification of carcharhinid sharks cannot beoveremphasized. This minor structural featureis certainly nonadaptive and its usefulness as anindicator of probable relationships should begreat (see Springer, 1950: p. 1, and Backus,Springer, and Arnold, 1956: p. 180, for discus­sion). The first mention of this characteristic inpublished work was by Nichols and Breder(1927), but correct identifications of the commonlarge ground sharks of the east coast of theUnited States were made by Nichols and by Rad­cliffe independently before 1916.

In one of the more valuable papers on sharks,Radcliffe (1916) made the first general use of thestructure of the dermal denticles to show differ­ences in western North Atlantic carcharhinidspecies; and his illustrations show clearly thedistin~tive denticle type and arrangement whichsets E. 'I1l.ilbert·i. off from other carcharhinidswithin its range, except. for the newly describedE. al#ma. Both E. lIlHberti and E. altima differfrom all other North American carcharhinidsharks in having nonimbricate denticles withoutstrongly projecting points; however, the denticlesof E. altima are much smaller than those ofE. lIlJlbeJ'ti.

Commercial shark fishermen at Salerno and KeyWest, Fla., recognized althna as distinct from thesandbar shark and called it the bignose shark orKnopp's shark before it. received a scientificname. The diagnosis given with the original de­scription of E. altima. (Springer. 1950) shouldbe adequate for the determination of specimensof all sizes. All of the known examples of E.althna have been taken at depths of 50 to 150fathoms off Salerno. Florida, in the Straits ofFlorida, in the northeastern Gulf of Mexico, andin the Dragon's Mouth between Trinidad andVenezueill. Its vertical range overlaps that of

the shllllower water E. lnilbe-di in the Straits ofFlorida area and extends well into the nighttimerange of the night shark, HypQprio·n signa-tUBPoey. The geographical range of E. altima maybe quite extensive, but it is unknown becausecomparatively little fishing has been carried outat the depths where this species might be expectedto occur. Such fishing as has been done in mid­water and just beyond the edges of the Conti­nental Shelf by commercial shark fishermen indi­cates that the species is relatively common.

Probably many more E. altima would havebeen taken by the commercial fishery were it notfor the fact that in the Florida-Caribbean regionthe liver oil of altima is characteristically lowerin vitamin-A content than is that of any of theother species of E'Ukunia or of H ypoprion in thatarea.

In a large measure, the confusion in the nomen­clature of the Ia.rger American carcharhinids thatexisted before the publication of the 1948 workby Bigelow and Schroeder would undoubtedlyhave been avoided if descriptive literature hadincluded information on the presence or absenceof the middorsal ridge. A fine replica of a shark,which in the light of the better descriptions nowavailable can easily be identified as Cal'chal'h:inu-sle1tcaS (Mimer ~nd Henle), a species without. amiddorsal ridge, is shown in an illust.ration in aninformative n,rt.icle (Rockwell, H116: p. 161)under the caption Oa.1'cn..a1'hi11ll.tIJ obscu/ru8 (Eula­n!-ia ObSC1bra.), a species with a middorsal ridge.Determination of the presence or absence of theridge is sometimes difficult, particularly for mu­seum specimens or for specimens that have beenexposed to the sun for a long period. Althoughidentifications can be made without reference tothe ridge, they are likely to be difficult and useof all of the available differentiating character­istics is desirable.

The confusion of the sandbar shark with thebull shark extends to the Pacific. Referenceshave frequently been made to the sandbar shark,Eula:m.ia 1n-ilberti, as occurring on the Pacificcoast of Panama. There is no evidence of thisand the species probably is not found there. Gar­man's (191:l) synonymy of m.ilbert-i includedEulfUnia nic(lra{/ue'l/IJiil Gill and Bransford, thefresh-water bull shark. The two bull sharks,nical'ague'1lsis and leucas are so similar to one

8 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

another that their separation is doubtful. Thereis a Pacific species so similar to nicaragnensis andlenca~ that commercial shark fishermen who fishedon both coasts claimed they were una·ble to dis­'tinguish one from the other except by area ofcapture. I do not know the scientific name forthe form if it has a name, but whatever thespecies, it is like nicm·agu.l!'l18is or le1tca.s and notlike mHbeJ'ti. Meek and Hildebrand seeminglyhad difficulties with this one in Fishes of Panama(1923), whe.rein they discuss a Pacific specieg('aJ'charh.inuB azw'e'll-8 (Gilbert and Starks) as asynonym of m.ilbeJ'ti. But Meek and Hildebranddid not see a specimen from the area which theythemselves could identify as m.ilberti and thesignificant sentence in their treatment states-

We certainly must regard the present arrangement astentative only. for more specimens must be comparedbefore the true affinities of the specimens from the op­posite coasts can be established.

A paper by R.osenblatt and Baldwin (1958) onsome of the carcharhinids of the eastern Pacificpresents for the first time information on thepresence or absence of the middorsal ridge inPacific species. This is an impOitant contributionand includes more comprehensive descriptionsthan have hitherto been available for sharks ofthe eastern tropical Pacific. These authors findthe separation of E-u.him-ia from Oarcnarldnu.sunacceptable for Pacific species. In support ofthis an unfortunate choice of illustrative argu­ment is used. They say (!. altima, for example,has a definite dermal ridge but teeth which are asl1!trrow as those of any member of the smooth­backed group (Springer, 1950). This is an error.The teeth of althna in the upper jaw are similarin general shape to the teeth of the other speciesof Eulmnia. These authors logically ca.ll atten­tion to the ill-assOl:ted group left in the genus(!arcnaJ'M-nus by my 1950 revision. mentioningwu.cas and ·velo.l! as examples. I am in completeagreement wit.h this but find no cogent argumentfor the elimination of Eulmnla, since the speciesof E-ulOln-ia as restricted are remarkably similarto one another in all of their morphological f('a­ttIres. The sharks allied to the genus C(u"chaJ'­Mn1l8 are far too widespread and numerous andthere is far too little known about them for anadequate study of thp. entire group. Additionalrevisions of the group are needed.

Differences between ndults of E. altbna. andm.ilberti are quite apparent in field examinationwhen the two are seen side by side. The snout ofalthna is longer and notably thicker dorsoven­trally. Furthermore, the first dorsal fin in E.altima looks quite different because it is not quiteso far forward as in E. 'mRbeJ'ti and is neither soerect nor so high. The high and erect dorsal finof E. m./lbeJ'ti in a forward position (fig. 2) is areliable and adequate character for field recogni­tion of adults in the water, if the size. of theshark is taken into consideration.

Gill (1862) based his c.lassification of the car­c.harhinid sharks almost entirely on the structureof the tp.eth. His arrangement of genera was notsatisfactory and it is apparent that short. descrip­tions of shark teeth are inadequate and leq,d toconfusion even though the number and form ofthe teeth show comparatively little variationwithin species and are of considerable diagnostic.value. The persistenc.e of essentially similarshape and structure in the suc.cessively largerteeth appearing in some· carcharhinid sharks asthey grow has been fairly well established by ob­servation. In E. m:ilbeJ'ti, at least, this appearsto be true, although this is neither universalamong sha.rks nor adequately demonstrated formany speCIes.

To obtain some verification of the extent ofvariat.ion in the number of tooth rows in car­charhinid sharks, I took advantage of a situation.requiring the preparation of several hundreddean, dry shark jaws for a commercial order. Icarefully identified the sha.rks and tagged thejaws of a series of 110 E. 'fIz.ilbe-rti together withaU other sharks appearing at the same time onthe dock at Salerno, Fla. All of the rnilberti andmost of the other sharks were adults; sex was notnoted. After the jaws were clea,ned I c.ountedand recorded the number of tooth rows (table 1).To the extent tn:at this sample represents t.hepopulation of m.ilbertl, the counts of rows ofteeth indicate that variation is small in tha.tspecies.

The shape and the relative position of the finsin carcharhinid sharks are reasonably usefulcharacteristic.s for identification. Small differ­ences in the size of fins or even in their positions,however, are of comparatively little value becauseof differential growth and the diverse trends this

NATURAL F.USTORY OF THE SANDBAR SHARK 9

FI: HI~ 2.-E/llamia wilbert?' turning in front of the camera at 1\1urinel:lJld, Fla. Note that the pectoral fins arc quitebroad at their base., relatively pointed and not strongly concave on thoir trailing edges. \\"ith the exception ofthe caudal fin, all fins function as rudders or stabilizers and CHunot be used indeppudently for locomotion TIll'I)('ctornls provide lift to ofT~l't the lift of the asymmetrical caudal since without a forward lift· the shark would tl'ndto somersllult. The large stiff fius iu forwllrd positions reducl' thl' ability of this spccips to roll :lI\d twist but maybe expected to increase the precision of it,s forwarrl swoops at creatures on tIlt' se:L bottom. (Photograph courtesyof Marine Studio.', Marineland, Fla.)

growth may take in dif:l'erent species. Data tosho,,, adequately the differential grmvth in car­charhinids are lacking. But one example willsuffice to show how unreliable proportional meas­urements can be for comparisons between speciesin whieh speeimens of different sizes an 1 agesarc involved, and in which the growth patternsof the species being conl])ared are unknown. ]nt.hree CXHlllples of yOllllg 1,,-'. 'II1ilb('l'ti, Ci8!). (iSO,

and (i;3!) 111m. long, from the vicinity of "Tood.Hole, Mass., the lengths of pectoral fins (meas­ured on their outer margins or leading edges)are 16.2, ] 5.!), and 15.0 percent of n1e total lengthof the sharks. In three adult rnilberti from off

Englewood, Fla., 2,210, 2,070, and 2,240 mm. long,the pectoral fin lengths are 21.3, 21.5, and 21.0percent of the total length. Let us compare theseproportions with measurements of pectoral linleno'ths of the whitetip shark, P/CI'olam.i.ops

~ .lon,qi.mmll/8 (Poe,)'). ~\ latc-l'ndJI'Yo \"hitet ip 5~()

mm. long, takcn 1~5 miles off New Smyrna, Fht.,11<1,-; a pectoral fin length 22.G percent of the tobilength; :l yOllng whiletip ],020 mm. IOllg, takenofl' Tampico, ]\Jexico, has a pectoral fin 25.5 per­cent of the total length; and an adult whitetip,2.310 mm. long, from the central Caribbean, hasa pectoral fin 22.0 percent of the total length.The figures ind icate a proportionately longer

10 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

TABLE l.-Tooth-ro!O counts in carcharhinid sharks taken off Salerno, Fla., summer oj 1947

SpeciesNumber of specimens bavlng toothrow counts I of-

-----------------------1------------------------

1 3 _I _3

41 29 116 13 23

12 7 28 1 1

Eulamia milberti (110 specimens):Upper jaw_. • • . .• .___ 3 37 55 11 4 • . _Lower jaw_. •• .. .. __ .• • . __ 12 20 44 30 3 1 • _

E. jloTidana (48 speclmens): . .Upper Jaw ....• •• __ . • .------------- ---- __ ._ ---- --._ ---- . . - _Lower Jaw • • •. • .. . • • . 1

E. ooBcuTa (39 specimens):Upper jaw ._•..• . -. -- . ._._ 3 15Lower jaw__ • • - . ._ 1 3 15 10

CaTc:IlaThinuB I,uras (24 specimens):Upper jaw. . . .. .__ 2 5 15 1 1 • . _Lower . . . . • . ._ . 18 3 3 . • _

C. limbatuB (31 specimens):Upper jaw .. . .__ 1 0 4 14 11 0 1 _Lower jaw . . . . 1 25 5 . _

C. maculipinniB (13 specimens):Upper jaw ._. . ._____ 1 1 2 4 3 2Lower jaw . • . . . . 2 1 9 1 _

Gakocerdo cuoier (21 specimens):Upper jaw ._. . • .______ 4 3 12 2 . • __

Neg~~ t~~~~~t;~~;(~;~~~~~~~~;;~;;;;;;;;;;;;;;;;;;;;;;;;;;;;; ;;;;;;;;;;; ::~: ::~: :;~: ::~: ::~: ;;;; ;;;; ;;;; ::~: ---. --~- ::~: ;;;; ;;;; ;;;; ;;;; ;;;;I All counts made from cleaned jaws from which all membranous sheathing had been removed to permit accurate counts whether or not teeth of the

functional row were missing.

pectoral fin in adults than in young for E. 7nil­b&r#, but an entirely different condition in P.lo-ngimanus.

The sandbar sharks available to me were re­markably uniform in general appearance and inthose features that I could measure, count, orcompare. In an attempt to learn something frommorphometrics, a considerable number of 7nilbe1'tiand other species were mea.sured carefully and indetail. However, the principal value that I de­rived from this excessively laborious task was inthe deliberate examination of specimens enforcedby measurement of detail and in the notes madl:'to accompany t.he measurements. The exerciseserved also to impress upon me the difficultiesattending att.empts to get adequate series to showgrowth patterns among some of the species oflarge sharks which are not only migratory butprobably short lived.

A characteristic of great import.ance for fieldrecognition of specimens of carcharhinid sharksis the tot.al length of the specimen considered inconnection with its sex and maturity (fig. 3).The mammalogists and ornithologists have longconsidered total length important in identifica­tion because mammals and birds have determi­nate growth patterns. As will be shown later.E. 'I1tilbertl has growth characteristics which re­sult in adults of predictable size. Furthermore,

the size range of adults within the known seg­ments of the population falls within limits whichare narrow enough to facilitate field identifica­tion by process of elimination. Thus, an adultEula·m.ia more thttll 92 inches in tota.} length isprobably not -m.llbe1·ti, and adult males less than70 inches or adult. females less than 72 inches intotal length are unknown.

DISTRIBUTION OF EULAMIA MILBERTIGeneral nature of distribution

The distribution of EulQlrnia milberti is diffi­cult to treat adequately bec~use, even thoughfurther discussions will be limited to the popuht­tion of the weste.rn North A.tlantie, the distribu­tion patterns are extremely complex. The adultssegregate by sex and to some degree have differ­ent vertical ranges. The nursery areas occupiedby the very young sharks are free of adults ex­cept when the females come inshore to give birthto their young.

The migratory patterns of young and adultsdiffer greatly. Finally there is a well-definedprincipal range oceupied by at least nine-t.enthsof the western North Athmtie population and anaccessory rnnge of uncertnin importance. It isquite possible that the populntion occupying theaccessory range is not self-sust.aining and exist.sonly beeause there is continuous but quite acci-

NATURAL HISTORY OF THE SAND'BAR SHARK 11

Carcharias taurus

Ga/eocerdo cuvier

Negaprion brevirostris

Aprionodon isodon

Carcharhinus ·/imbatus*

Carcharhinus maculipinnis

Carcharhinus /eucas

Eu/amia milberti

Eu/amia altima

Eu/amia f/oridana

Eu/amia obscura

Sphyrna sp.**

........ ......... ... ..... 1IIIIIIIII1IIIIIIB1II!IIIIl1llllliill15.... ?~ ~~~::::: ~ ~~~~~~~~~~78

:~ ~ ~ ~ ~ :::~ ~ ..... :::: :~ ~: ~ : ~ ~ :.~i~6.., ~

....••••.•••••• :lIlIIII2o

~:::~ ~

.. .; -- .......... ... . &i!iJii.................. _12

::: ••••••~:::: :~:::::::: •• :: ~.~142····················-.n5• • • • • • • .. . • • • • • • • • • • .1IIIIIII!lIIIII899

··:;.·.·.·.:~.·.:~.·.·.·.·.·.·.·.·.·.·.·.·.·.·.·.~15

.. '?:: ~ ::::::: ~ '::::.~34~ ~ •••••••••••• ~ •••••• ::: ••••••••••••••••••••••••~ ~ ~ ••~150

~ ~ ~ ... -. -. ~ ~ ~ ~ ~ ...-.-. -. -. -...~... -. ~ -. -. -. ~ -. -. -. -. -.~l!3i!6 ilMil!la!lil!l!l!!ili!!ilJ!}@!aESl1i!&l1i!O\i\i&i&52

I I I I I I I

o 10 20 30 40 50 60 70 80 90 100 110 120 130 140 150 160 170 180 190

_ =. siZE! range of embryos ~ = size of adults

FIGURE 3.-Comparative sizes of adults and young of common large sharks found within the geographical range of E.milberti. The figures at the right indicate the number of specimens in the sample used to determine size range. Thesize ranges for embryos are estimates based on maximum observed lengths of embryos and minimun lengths of free­swimming young observed in Florida collections. (* Size range in Florida-Antillean specimens. Western Gulf ofMexico and Central American coast specimens are smaller and produce smaller young. The western stock mayprove to be distinct nnd if so should take the name Carcharhinus natator Meek and Hildebrand. ** The greathammerhead of the West Indian region, following Bigelow and Schroeder (1948). The nomenclature is now unsettled.The name Sphyrna tudes is not available for the great hammerhead and probably should be replaced by Sphyrnamokarran (Ruppell)')

dental recruitment from the principal population.Within the expected vertical and geographicalrange of the species are some areas which appearto be avoided. It is well to mention again thatthe sandbar shark, like other large sharks, is notprevented by well-defined barriers from wander­ing out of its normal range.

The limits of distribution are therefore notsharply defined. Following the traditional pat­tern in descriptions of distribution it may be saidthat the sandbar shark, as represented by thewestern North Atla.ntie population, is eommonin summer off the Atlantic coast from Cape Codto "Vest Palm Beach, Fla., and in winter from

the coast of the Carolinas around the tip ofFlorida to the gulf coast of Florida as far northas Tarpon Springs. It occurs uncommonly in thewestern part of the Gulf of Mexico and along thecontinental shores southward to Costa Riea. Itis a casual visitor on the northern coast of Cubaand the western edges of the Bahama Banks. Itsvertical range is from the shC'reline out to H35fathoms. It. enters bay mouths but is not foundin fresh waters.

The principal source of information on thedistribution of the sandbar shark c.omes from the.C0Jl1lUe.l"C.i111 shark fishery. Atlant.ic coast sharkfishermen used bottolU setlines more often than

12 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

any other genr. Each unit of gear consisted of It

main line of chain or wire rope a half-mile ormore in length. This was set on the bottom:anchored at both ends with the anchors riggedwith buoys so that the lines could be retrieved.Short, branch lines made of chain, each with abaited hook, were spaced at intervals of 20 to 40feet along the central part of the main line. Thetypical unit had 100 hooks. Floating lines andtmchored gill nets were also used occasionallyin the fishery.

Positive information from the shark fishery onthe distribution of E. m.ilberti is quite volumi­nous and detailed. Systems of payments to fish­ermen required detailed records involving iden­tification and measurement of all sharks landedby vessels of the principal fishing company. Al­together, records of landings of more than100,000 milberti during a period of 15 years havebeen pxamined.

Information on the absence of E. 11IHberti fromspecific areas has been difficult to assemble, buthere also the records of the shark fishery supplymost of the data. The species was first reportedin the Florida area from correctly identifiedspe.cimens after the eommercial shark fisherybega.n (Springer, 1938), so the earlier scientifieliterature has been useless in the establish­ment of the ra.nge of the spedes in the Floridaarea southward. Offshore records, from areaswhere water depths a,re more than 500 fltt,homs,are exclusively from catehes made, on tuna.longlines used by the explortttory fishing vesselOregon (for descriptions of this gear seeBullis, 1955, and Captiva, 1955). Some informa­tion on the distribution of the young was obtainedfrom otter-trawl eatehes made by the exploratoryfishing vessel. Dela·ware off the coast of NorthCarolina. Additional scattered records werepicked up from accidental catches made by com­mercial and sport fishermen who used varioustypes of gear, from catches made by collectorsfishing for aquarium specimens, and from biolo­gists who captured specimens incidental to othereolleeting activities.

The area of greatest uncertainty is in the off­shore and midwater range. Reeent marine ex­ploration has sho"'n that substantial populationsof large sharks, fishes, and invertebrates live insubsurface waters beyond the Continental Shelf

where they have escaped the attention of natu­ralists. On June 11, 1954, the first E. '1TIilbe'1'tiknown from waters beyond the Continental Shelfwas taken on a tuna longline hook at USFWSOregon station 1099, 85 miles off the coast ofTexas where the depth was approximately 600fathoms. Since the hook was set to fish at about30 fathoms, this shark, an adult male, was cruis­ing in midwater. Throughout. the second half of1954, all of 1955, and the first part of 1956, long­line fishing was cn.rried on in the offshore watersof the Gulf of Mexico by the M/V OregO'n and afew commercial vessels. Large numbers of sharkswere talten, ehiefly spec.ies known to be partly orentirely pelagic. No addit.ional mHbati weretaken until early February 1955, when a CODl­

mercial vessel eaught two adult females about 50miles off the nort.hern edge of t.he CampecheBank where dept.hs were est.imated to be morethan 1,000 -fathoms. These sharks were caughton longlines with hooks fishing not more than50 fathoms deep. These three captures, outsidethe princ.ipal I'ange, appe.ar to have little signifi­cance in the general picture of t.he distributionof E-ulam-ia mRbel'ti.

Factors affecting distribution

It may be assumed that water temperature andsalinity are import.ant in limiting the distributionof the sandbar shark and that there are otherfactors clen.rly influencing the movements anddistribution of the species. The reaction of sand·bar sharks to ocean currents, the availability offood, the reln.tion between the growth rate or thereproduetive pattern and the migratory move­ments, all appear to be important in forcing thespecies into a part.icular range. No data areavailable, however, to show the relative st.rengthsof these conditions as determinant.s of the rangeof the sandbar shark.

The facts, from superficial examination at least,do not support t.he thesis that competition withother species is a powerful influence in the selec­tion of a partieular range. Young E1l1mnla, m.il­bel'tl, for example, apparently cannot long sur­vive wlwre large (!al'f'hul'ldnU8 leu.cas in propor­tion to mHbedi arc relatively nbundant.. Thepresence of largE:' numbers of large C. le'Ucas inthe vicinity of the mout.h of the Missi~Bippi Riwl'seemingly does not. deter gravid female milbert-ifrom moving into the area to give birth to young.

NATURAL HISTORY OF THE SANDBAR SHARK 13

Whatever the particular reason or reasons maybe, the general absence of young 'l1l-ilbe'rti. in theGulf of Mexico shows that conditions are un­favorable for them. Circumstantial evidencesuggests that interspecies competition is respon­sible, because large O. le.ucas eat young milberti.

No explanation is apparent for the commonoccurrence of E. 11'liilberti along the continentalshor~s and its absence from most of the WestIndian shallow waters except that the speciesseems to have preferences for certain types ofbottom. E. ,milbe-r# is ordinarily not common inareas of coral reefs or where the bottom is rough.Since it is chiefly a bottom-dwelling species, it isnot surprising that it would exhibit preferencefor one type of bottom over another. In itsmigratory passages around the southern tip ofFlorida and the Florida Keys, however, thereappears to be active avoidance of the fringingreef. Here the migrating adults leave the rela­tively shallow areas they inhabit on both the eastand west coasts of Florida and temporarily enterand feed in much deeper water.

Nursery Arounds and distribution of young

The principal nursery grounds of the westernNorth Atlantic population of E1tlamia milbertilie in. relatively shallow water alon,g the Atlanticcoast of the United States from Long Island toCape Canaveral, Fla. This range may be ex­tended slightly at its northern end to the southside of Cape Cod in favorable years but thesouthern limit is more definitely fixed. Not oneyoung mRberti has been taken south of CapeCanaveral, around the tip of Florida, or in theeastern Gulf of Mexico. On the east coast ofFlorida, south of Cape Canaveral, a few sexuallyimmature 1nilbert-i of almost adult size have beentaken; but in this area adult milberti are com­mon. A great quantity and variety of fishingeffort has been concentrated south of CapeCanaveral on the Florida coast. The total ab­sence of young milberti here is remarkable inview of the somewhat indefinite range limits ofthe adults.

A secondary nursery range apparently lies inthe northwestern pa.rt of the Gulf of Mexico. Itis indicated only by the capture of a few femaleswith near full-term embryos near the mouth ofthe Mississippi River, the capture of a large

1152li08 Q.-..6O-3

m.ilbe1,ti with nearly full-term embryos off theTexas coast (Henry Hildebrand, 1954), and aspecimen 747 mm. (nearly 30 inches) from theTexas coast (Bigelow an~ Schroeder, 1948).

It is probable that gravid females wanderingaway from the principal range of the species givebirth to young along the Mexican and CentralAmerican coast, but no records of the capture ofyqung in this area. have been found. Shark fish­ing on a small scale has been carried out overmost of this area and catches have been examinedat various points from the mouth of the RioGrande River to Costa Rica; but excepting theGulf of Campeche, no young 'l1Iilbe'rt-i appeared.

The female EuJawlt m,ilbe1·ti, which move intothe principal nursery areas to give birth to theiryoung, do not remain there long and do not feedactively while there. This may explain thescarcity of records of captures of adult E. mil­be'rti along the Atlantic coast. Great South Bay,Long Island, is one of the nursery areas of E'.'l1l,ilberti and accounts of the appearance of fe­males in the bay and birth of the young are givenby Thorne (1916). Additional mention of theappearance of E. milberti in the Great SouthBay area is made by Nichols (1918), who notesthat the interesting fact about them is that theadults of the two sexes of the same species arealmost never taken together near Long Island.Here the adult females are E. milberti and theadult males are CarcharMntUJ l.eu·CaB.

Records of young sharks from Chesapeake Bayshow that E. m.flbe-rti gives birth to young in thebay in summer. William Massmann, of the Vir­ginia Fisheries Laboratory, has kindly given me(in correspondence) records of E. milbert-i fromthe lower Chesapeake Bay. He says-

Although young of this species are probably the mostabundant shark In the Bay In summer, I would not saythat It is numerous. • •• It Is commonly caught byanglers and probably rather generally distributed In thelower Bay. I have not seen an adult In the Bay or anyIndividual more than three and a half feet long..

After a comparatively brief period in shallowwater or in the mouths of bays, perhaps at thebeginning of cool weather, young m.ilberti appearto move offshore. The only area from whichyoung 1nllbe1'ti are known in the winter seasonlies off the coasts of the Carolinas at depths outto 75 fathoms.

14 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Distribution oft Atlantic coast of the United States andin eastern Gulf of Mexico

Th~ northern limit of th,e range of the sandbarshark is easily established.. ': There are no reliablerecords of its capture from the Gulf of Maine(Bigelow and Schroeder, 1953), but SO~lt.h ofCape Cod it has been taken frequently but Irregu­larly at. Woods Hole, Mass. Numerous recordsof the spec.ies along the Atlant.ic coast of theUnited States are summarized by Bigelow andSchroeder (1948). Sandbnr sharks may be saidto be common in summer along the Atlantic coastof the United St.ates from Long Island to the tipof Florida and in winter n.Iong the waters of theContinental Shelf off the Carolinas southward tothe southern tip of Floridn, in water of moderatedepths in the Florida. Straits and along the ~est

coast of Florida northward to Tarpon Sprmgsor the Middle Grounds (the rough bottom areasouth of Cnpe San Blas, Fla.). This area is theprincipal known range, but the species hilS alsobeen taken in small numbers from the northernand western Gulf of Mexico, the western bordersof the Bahnma Bank, the northern coast of Cuba,and the Caribbean c~~st of Nicaragua and nort,h­ern Costa Rica (fig. 4).

The hypothesis is advanced here that the sand­bar sharks of the eastern and southern sides ofthe Gulf Stream in the Straits of Florida arecasunl visitors to those areas and that the stockof the northern and western parts of the Gulf ofMexico is a breeding stock which is not self­sustaining, but is recruited in part from migra­tory adtilts moving to the northern and westernparts of t.he Gulf by mistake or through error inorientation and navigat.ion during the regularwint.er migration of the principal stock.

TIlP- sandbnr sluu'k has been taken frol11 theshnllows a.Iong beaches out. to a depth of about135 fathoms. The young have been t.aken most.often in shallow waters to depths of about. 5 to25 fathoms in summer, but in wint.er- they moveoffshore t.o warmer wnt.er and depths ns great. as75 fathoms off the Ca.rolinn const..

The sandbar shnrk is known only from theshallower part, of the Continentn.I Shelf in thewnrmer months in the extreme northern part ofits range. Probably the adults are more commonoff beaches Hum in major bays or inlets. Hilde­brand and Schroeder (19~S) found the SpeCleg

rather rare in Chesapeake Bay although morecommon than nny other shark except the spinydogfish. Radcliffe (1916) st.ates t.hat t.he speciesappears to be rare in the Beaufort, North Caro­lina, region. However, it was regarded as com­mon in bays on t.he ocean side of Long Islandfrom mid-june to mid-September, by Nichols andBreder (1927).

The apparent scarcity of adtilt E. milbertinoted by Hildebrand and Schroeder and by Rad­cliffe is' easily explained. It is possible that E.mHbe.·rti ente~s the mouths of bays t.o give bh·thto young more frequently than records suggest.Fema.Ie Carcha:rhIn:u.s le.1wa.s and Negapri(rn b·re·"i­rostris move inshore and stop feeding for a shortperiod at t.he time of the birth of their youug,and immediately aft.er the young are born t~e

females move into comparatively de.epe.r water.This may be a common habit among carchar­hinids al~d certainly a very use.fnl one to providefor survival of the species. The Long Islandre.cords are to a large extent based on harpoonedspecimens, and adult females should probably notbe. expected to be easily availahle. t.o capture onbaited hooks in areas where the young are born.

The best fishing depth for adult E. 'milbertifrom the Carolinas south to Miami was found bythe commercial shark fishermen to be 15 "to 30fathoms. On this stretch of coast. it was rarelyif ever tilken beyond 50 bthoms on bottom set­lines and made up less than 5 percent of the catchon floatlines set beyond the 100~fathom curve.

Sout.hward from Miami, m./lberti was rareamong the. keys, in HawkChannel, or along t.heshallower portions of the reefs south of tl.leFlorida Keys. In the. winter, a few appeared III

catches made in the Northwest Ship Chmmel but.,in general, the,se waters were le,ft to other species.The sandbar sha,rks, however, were the common­est sharks on the bottom be,yOIi.d the fringing reefout t.o depths of 50 fathoms an,d made up sub­stantial portions of catches out. to 100 fathoms.They wel~ appreciably nwre numerous in catchesoff the lower kevs where currents were not sostrong. NOl1.hw;rd from the keys along t.he westFlorida coast as far as Tampa, E. m1lbe.rti wasfound t.o be most abundant. in depths of less tha~

30 fnt,11onls. . .. ";, .."Shark-fishing vessels operated out ·of Salerno,

Fla., almost e,very day that weather permitted

NATURAL mSTORY OF THE SANDBAR SHARK 15

PRIMARY WINTER RANGEOF ADULTS

•

...

.',,•

..

o····~ .".•.. '.'"C , • I I IE'. S E ,

NORT. AMERICA

IILF till IEIICD

FIGURE 4.-Geographical distribution of the western North Atlantic population of the sandbar shark, Eulamia milberti.

from 1936 to 1950, except for parts of 1939 and1940 when activities were suspended due to over­production. These vessels landed .their catchesdaily at Salerno where the sharks were identified,measured, and recorded. Cate-hes included in theSalerno landings were made from Bethel Shoal,north of Fort Pierce, Fla., to the offings of

Jupiter Light. Some details of these landings,showing catch per unit of effort, were reported inan earlier publication (Springer, .1951). Thesedata show that at Salerno the highest averagerate of catch of E. mHbe'rt-i occurred annually inthe month of Februa.ry and the lowest averagerate of catch in September. These data concern

16 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

only adult 'lldlberti and the comparatively fewsexually immature m.ilbertl of adult size occur­ring with the adults.

On the west coast of Florida, shark-fishingvessels caught no E. m.ilbert·i at all from Maythrough November and reported their largestcatches from January through March in each ofseveral years for which data are available.

At Key "Vest, Fla., high catch rates were ob­tained for mHbe'l'tl from deepwater sets made inthe winter and early spring but catches at otherseasons were poor.

High catch rates of adult E. mHbe'rti were ob­tained by shark-fishing vessels off the Carolinasin September, and from 1946 to 1949 one or twoof the' more able shark-fishing vessels followedthe fishing for adult m.ilberti southward, arrivingoff Salerno in January or February.

Scouting by shark-fishing vessels showed thatsome adult milberti were present in each monthof the year along the Atlantic coast between thelatitudes of Charleston, South Carolina, andMiami, Florida. Although E. milbedi was theprincipal species sought in this area, other speciescontributed variously to the value of the totalcatch. Table 2 shows the comparative availa­bility of large sharks to the kinds of fishing gearemployed in the Salerno area in the late spring.

Distribution in Bahamas and West Indies

E. mllberti is common only on the western sideof the. Gulf 8tre.am. On the eastern side of thestream, it is replaced by E. falelformJs as thecommon inshore EU}(J.,mi.a. The wandering ofmilbe-rtl into the Antillean area may be quitelimited. There are no records of "111iilberti. east­ward through the West Indies nor from thesouthern shores of Cuba. A shark fishing opera­tion o~ ·the eastern part of the Bahama, Bank inthe period from 1947 to 1949 did not take mll­be-rtC ..

.'. E. falc-ifol'mis was reported by Evermann andMarsh (1902) from. Puerto Rico; by Be.ebe andTee Va.n (1928) from Port-au-Prince Bay, Haiti;by Nichols (1929) from Puerto Rico; and byBackus (1957) from open sea situations east ofthe Gulf Stream. None of these authors notedthe presence of E. m,ilbe-rti.

Frank Mather III has told me in correspond­ence of the capture of E. fakifo-rmi~ and E. flo-ri-

aa:na from the vicinity of St. Croix and St. Thomasin the Virgin Islands. Mather's fishing operationsrovered the depth range in which E. m.-ilbe·"tiwould be expeet.ed if its geographical range ex­tends through the West Indies and if the absenceof milbert-i follows the usual pattern in the WestIndies.

TABLE 2.-Sharks taken by commercial.fishing I.essels in theSalerno-Fort Pierce area (Bethel Shoal to Jupiter Light)and landed at Salerno, Fla., in May and June 1945-46

[I to 3 vessels; only sh...ks with hide length of 55 in. or more included; ftshingdepths from 18 to 40 fathoms~

Number of sharks

Species

1945 1946

Eulamla mllbtTtI (MiilIer and Henle), sandbar shark_ _ 1,515 987Sph/ITua sp.,' hammerheads .___________________ 323 268Elliamia ob8cuTa (LeSueur), dusky sh...k______________ 269 94Eulamia (lOT/dana (Bigelow, Schroeder, and Springer),silky sh...k ._____________________________________ 220 191GalroctTdo cuvitT (LeSueur), tiger sh...k______ ___ _ lIS 64CaTchaTh/uusltllcas (Muller and Henle), bull shark____ 61 41Net/apTian bTeviro'/T/s (Poe.y), lemon shark___________ __ 20 19CarrhaThi-llus sp.,' blacktips____________________________ 3 16Ginglgmos/oma ciTTa/um cOmeliu), nurse shark_ _ 2 6CaTchaTodon raTrhaTias (Linnaeus), great white shark__ 2 5l,uTlII OIg,/nehus Raflnesque, mako ._____ 0 2Unidentified ._____ 120 2

I 3 species; station records do not distinguish kind.'2 species; station records do not distinguish kind.

That the Gulf Stream is not itself a barrier toE. mHbe:l'tl is apparent from the occasional cap­tures along the Bahama Banks and off the north­ern coast of Cuba. It is possible t.hat large nUlll­bel'S of migratory sha.rks may wander away fromnormal migratory routes at t.imes when unusualcondit.ions prevail. Cert.ainly a few 1nilbe-l'tlcrosg the stream.

From May 18, 1948, to July 8, 1948, I under­took a program of e.xpe.rimental shark fishingalong the western edge of the Bahama Bank fromRiding Rock nortllwRl'd. Fishing operationswere carried out from a base a.t. the Lerner Ma­rine Laboratory of the American Museum ofNat.ura:l History at Bimini, using a fishing vesseland gear provided by t.he Shark Industries Divi­sion of the Borden Co. Fishing was carried onchiefly by bottom setJines at. dept.hs from 10 to200 fathoms, but some, float.ing lines were used t.oassure coIled-ion of as wide a variety of sharks aspossible. Sets included some made at variouslevels along the ext.remely precipitous slope of thebank, which drops off abruptly from about. 20fathoms down to the floor of t.he Gulf Streamchannel where depths are more than 150 fathoms.

NATURAL HISTORY OF THE SANDBAR SHARK'" 17

Number of sharkstaken in-

I Antillean form.• Nomenclatorial status of this species not determined.

TABLE 3.-Sharks taken in, exploratory fishing from theDusky along edges of northwestern Bahama Banks,May 18 to July 8, 19J,.8

During the entire fishing period along the Ba­hama Banks only 14 E. mHbel'ti were caught inthe lot of 447 sharks. Of t·he tot.al, 197 werereef sharks, Eulmnia sp1,tngel'i (BigelO\y andSchroeder) .

at Snlerno. This is a, poor fishing spot, however,so the commercial vessels rarely caught springel'i.

Distribution in western Gulf of Mexico and westernCaribbean

Posit.ive knowledge of the distribution ofE'u,{anl-ia 'lnilbe'l't'i in the western part of the Gulfof Mexico and along the Caribbean coast of Cen­tral America is based on records of 8 gravid fe­males off the. Mississippi River delta, 1 gra,vidfemale and 1 young male from t.he coast of Texas,1 adult male and 2 adult females from the south­central part of the Gulf of Mexico over deepwat.er, and 51 adult.s from the Caribbean c,oast ofNicaragua and Costa Rica.

From the mouth of the Mississippi River east­ward and southward around the Gulf to thevicinity of Tarpon Springs on t.he Florida westcoast., m-ilbel'ti appears to be abseJlt or at leastrare from inshore waters out to 30 fathoms. Nocatches were reported by shark fishermen and nospecimens were seen. I should note t.hat whileemployed in the shark fishery, seasonal shark­fishing stations were maintained at various timesat all of the fishing ports of any consequencefrom the mouth of the Mississippi River aroundthe tip of Florida to the Carolinas. I visited aUof these stations frequently and sometimes par­t.icipated in fishing operations. The presence ofmilbel'ti in appreciable quantity would almostcertainly have been noted in catches from the sta­tions at Panama City and Carrabelle, Fla., hadspecimens been taken.

It is not possible to present a meaningful ac­count of the sandbar shark, Eu,z<lt111>ia -milbel'ti,without frequent reference to the bull shark,Ca'l'Cnarldll:lf8 leu,caJj, and its fresh-water l'el>l"e­sent.ative, (!a,rchal'himM niCf.f.l·agtte'lIsiJJ (Gill andBransford). The sandbar shark has been con­fused with the bull shark because of the peculiarmanner in which their ranges overlap. In addi­tion, bull sharks appear to be the most importantof the predators on young sandbar sharks andthe primary factor t.hat prevents E. milbel'ti frome.xtending its nursery range into otherwise suita­ble areas in tropical seas.

As has already been not.ed; bull sharks arefound along the Atlantic coast as far north asLong Island but increase in numbers somewhatin the latitude of Salerno, Fla., where, as noted intable 2, they were sixth in number of large sharks

Bimini Walkerarea Key

(20-200 0\)-100fathoms) fathoms)

10 18760 1346 52 18

17 2

15 010 412 211 32 35 23 0

3' 11 20 31 01 00 11 01 0

Enlamia ,prlngeri (Bigelow and Schroeder) ,reef shark_Ga~ocerdo cuvl" (I.eSueur), tiger shark • _Eulamia ob,cura (LeSueur" dusky shark • _Ntgaprion brtpirostri8 (Poe)-), lemon shark • _Eulamia altima Springer, I,ignose shark. _Fulamia florldana <Bigelow, Schroeder, and Springer),silky shark • • • _Eulamia milberti (MUller and Henle), sandbar shark _Hupoprion aig1l4t,'I Poey, night shark. _Scoliodoll ttrrlHlovae (Richardson), sharpnose shark _Oi'lll/lpm08loma tirralnm (Omelinl, nurse shark _Sphprna ltwilli Griffith, southern hammerhead. _Herallthu, sp. (not H. grlseu,), little cowshark _. _Cartharhillu, limbatns (MUller' and Henlel,' littlehlacktlp . • • •• _Sphprna Sp.•f great hammerhead .. _Carrharhlnu,lellrtu (Milller and Henle), bull shark _Mna/eltte calli, (Mitchl1l), common smooth dogshark _Pttrolamiop, IOllgimailllS (Poeyl, whitl>tip • . _Carrbarbil1'" maculIpil11li, (Poey), big blarkUp _Carcharbhl"s acronotus l.Poeyl, blacknose shark _Eulamia sp., undetermined • • _

Species

During the same season of the year in whichthe exploratory fishing was done, a great. numberof sharks were landed across the Gulf Stream atSalerno, Fla., about. 80 miles from t.he northernend of our Bahama fishing area. But more thanhalf of t.hem were E. 1'nilbert-i, and no E. falci­fo-rmi,s was landed. Results of the Bahama fish­ing are summarized in table 3, which showscatches made in two areas off t.he Bahama Banks.For purposes of camparison, catches made in theSalerno-Fort Pierce area in May and June 1945and 1946 are shown in table 2. The two fishingoperations are not exactly comparable, of course,not only because they we~ carri~d on in differ­ent years but because the Bahama fishing wasessentially exploratory while the Salerno-FortPierce fishing was a part of a continuing COlll­

mercial operation concentrated in limited depthsand locations. Exploratory fishing with sets scat­tered in different depths and locations off Salernowould presumably produce a fe.w E. spl'inge'ribecause there are normally a few to be foundalong inshore reefs adjacent to the St.. Lucie Inlet

18 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

-------------_.--------

I Also taken but not recorded becallSt' of small size: several Scoliodonlerra·nOllae (Richardson), Carcharhinus porosus Ranzani, and Alus/tlus canis(MitchiJIl, and one young Gingl/lmos/o·ma clrralltm (Ornelin).

, Continental form-typical of western Gul! of Mexico.

TABLE 4.-Large sharks taken from the Joe Leckich onbJltom lilies set in .Ii to 35 fathoms oJ! the mouth of theMississippi River, JunR- 25 to July 29, 1947

landed by the shark fishery. The abundance ofbull sharks at various loeations in the Gulf ofMexico varies seasonally, but fro111 the vicinityof Apalachicola, Fla., westward along the north­ern coast. of the Gulf of Mexic.o and southwardalong the coasts of Central and South Americaas far at least as Frenc.h Guia.na, bull sharks formthe major part of catches of large sharks made byshallow-water setlim.'s in some seasons. Table 4:shows the comparative frequence of capture oflarge sharks during a test-fishing period off themouth of the Mississippi River.

These two species are also reported by Backus(1957) from the A.tlantic beyond ContinentalShelf limits.

A lot of 51 adult sandbar sharks was taken byCaptain B. W. Winkler from off the Caribbeancoast of Ni.caragua and Costlt Rica. This groupincluded all of the sandbar sharks in a collectionof 854 sharks of all species which Captain Wink­ler measured and recorded for me from Septem­ber through December, 1948. The most inter­esting feature of this collection of E. milbertlwas that it was made up of approximately equalnumbers of adult males Itnd females (26 malesand 25 females).

The shark fauna of the area as represented byCaptain 'Vinkler's collections included a largenumber of species that are predatory on sharksof the size of young E. milbe:rti. The followingshltrks wei'£>' taken:

A few days' fishing with bottom longlinesalong the outer edge of the Continental Shelf offnorthern Nicaragua and on Serrana l\l1d SeranillaBanks in Februal'Y 1949 failed to produce E.mRbert-i or any adult sharks, but. moderate num­bers of young E. fiO'l'ida·na were taken.

As a grader and buyer, I eXllmined several lotsof dried shark fins said to have been taken off thecOitsts of Colombia and Venezuela. E. 'milbM'tifins were not noticed although it is possible. that

Specie8: NumberC'areharhill us,' chott,os or bull sharks ._ _ 421Galeocerdo elwier (LeSueur), t,iger shark_________ 85'Eulamia. obseura (LeSueur), dusky shark _______ _ 76Eulamia jloridana (Bigelow, Schroeder, and

Springer), silky shark ._ _ 70C'areharhillus limbatus (Milller and HenlE') ,2

little blacktip_ __ ___ _______________________ 54

Eulam ia milberti (Mtiller and Henle), sandbltrshark____________________________________ 51

Sphyrn.a sp. (not determined), hammerhE'adshark____________________________________ 27

Eulamia Sp.,8 reef sharks_____________________ 25Seoliodon sp., sharpnose shark ._ _____________ 15Hexallchus Sp.,4 cow sharks___________________ 13Ginglymostoma cirratu.m (Gmelin), nurse shark_ _ 3Eulamia altima Springer, bignose shark_________ 2Negaprion brelJirostris (Poey), lemon shark______ 1Undetermined 6 " ._ _ _ _ _ _ _ _ _ _ _ 4

I Either C. Iwcas (Miiller and Henle) or C. niraragl/ensis (Gill and Brans'ford l or both.

, Continental form.• Probably F. sprl-ngeri (Bigelow and Schroedrrl or F. 'a/riformis (Miiller

and Henle).I IllrJudes two species.• Possibly a sm..1l spe.cies or Galeorhill-u8.

3853

129

3491314262012

164

Av~r8ge

weightof liverlib.)

Z758356

47988743

Numberof

sharksSpecies I

The eight E. m.ilbej'ti were adult females andfive of them were gravid. Since '11-dlberti hasliver oil of comparatively higher potency thanthe oil· from otlll;'r spe.cies in the area, a speeialeffort was made to catch them in subsequent,larger scale fish~ng efforts. Nevertheless, catchesof milherti were not made later during 1947 andcommercial fishermen operating in the. area re­ported sandbar sharks absent in 1948 and 1949.The few records of E. 111ilbedi from offshoreGulf waters wOlild not be important except thatthey serve to show the species can move into andacross deep areas of the ocean. Off t.he AtlanticStates there has been comparatively little long­line fishing be.yond the Continental Shelf andthere are no records of '/nilbert-i from deep water.There are, however"'l'ecords of catches of EulamJiafal-dfQl'/nis (MUller and Henle) and E. obscu-rafrom longline sets made by t.he exploratory fish­ing vessel Dela-wa.re beyond the limits of theContinental Shelf off the Middle A.tlantic States.

("'archarhinns lencas (MiiIler and Henle), bull shark. _Fnlamia abscllra (LeSueur), dusky shark _Galeocerdo envier (LeSueur), tiger shark. _Careharhinlts limbo/Its (MiiIler and Henle), littleblooktip ' .. _Sphyrna sp.• great hammerhead .. . _Careharhi"us maeulipinnis, big bJacktip _Fltlamla millvr/i (MiiIler and Henlel, sandbar shark __(·'archarias/au",s Rafinesque, sand shark_. _Negaprion bWliros/ris (Poeyl, lemonshark. _SphyrnlJ I,wini Oriffith, southern hamml'rhead. _Eltlamia floridana (Bigelow, Schroeder, and Springer),silky shark. . __ . . • . _FUlamia spring..i (Bigelow and Schroeder) . __ . _

NATURAL HISTORY OF THE SANDBAR SHARK 19

a, few might have been overlooked. The fins ofE. milberti adults (pectorals, first dorsal, andlower caudal lobe) are more desirable for com­D.1ercial purposes than the. fins of some otherspecies because t.hey are thickeI' and have a rela­tively large proport.ion of t.he material used forshark-fin soup. Shark fishermen with whom Italked and who would recognize 11liilbe1·a also re­port.ed the absence of t.he species from t.heircat.ches made off Colombia and Venezuela. Thus,evidence for t.he occurrence of E. m.ilberti in thesouthwestern Caribbean, while not very sat.isfac­tory, is negative.

MIGRATION

It is probable that t.he migrations of E. nI,il­berN are of two kinds. One is simply the gradualwithdrawal of t.he sharks from waters that be­come too cold or too warm-a movement that isaccompanied by normal feeding activities and ischaracteristic of immat.ure sharks. The other isa movement, generally, over a greater dist.ancethat may· or may not be induced by temperaturechanges. The general patterns of the majormovements of adult. sandbar sharks suggest thatoc.ean currents grently influence the. direction andext.ent of the movements.

It is necessary t.o consider the migratory move­ments of the adult. male, the adult female, andthe immature Eu,l.mnia mHbert-i separately. Wemay look upon the Atlantic coast from the vi­cinit.y of Charleston, South Carolina, t.o thenort.hern part. of Florida as the core· of dist.ribu­tion of the principal stock of the western NorthAtlantic population because it is only in this areathat all three groups are known to be found.This may mean merely that in this area there isoverlapping distribution. 'Ve know t.hat theadult females go as fa.r north as Long Island togive birth to young in summer and in some yearseven farther, to the vicinity of Cape Cod. Thereare no data to show whether the adult males ornongravid females move northward int.o the por­tion of the species' range lying north of the Caro­linas. All that.. is known of the dist.ribut.ion ofthe young is that the young are born in water ofmoderate salinit.y from Cape Canaveral to Ca.peCod and that some of t.hem move in winter intothe comparatively warmer offshore water foundat. depths of 50 to 75 fathoms on the Carolina.

coast.. Until the young sharks reach adult sizethey do not t.ake part. in the long sout.hern migra­tion characteristic of adult.s or move south ofCape Canaveral. One capture of a lot. of nearly200 young off North Carolina indicates that theyoung occur in schools of both sexes and of mixedSIzes.

Migratory movements of adult E. lnilbertisouth of Cape Canaveral are more dearly out­lined from the data available from the sharkfishery. The annual southward movement ap­pears to be coincidental with the beginning ofcooler weather and to be accelerated by' -coldsnaps.

A very much larger number' of adult femalesandbar sharks were taken t.han adult males bythe Florida shark fishermen. This and the tend­ency to segregation by sex, will be discussed laterin connection with reproduction. But it should bepointed out here that no adult males are recordedfrom inshore nursery grounds and probably occurthere but rarely. In the Florida shark fishery,adult males of most species brought more moneythan adult females and were particularly soughtby fishermen. The fishermen were convinced byobservation that. adult males of most species in­cluding lnilberti were usually in deeper and coolerwater than the females, that they usually pre­ceded the females in migration, and that theyusually were to be found in more compact aggre­~ations so that fishing was best where males couldbe found. Such figures as are. available bear thisout. A rationale for this condition is suggestedby the thermal sensitivity (decreased fertilitywith applicat.ion of heat.) of the male germ plasmamong vertebrat.es in general (see Cowles, 1945).

The sandbar shark is properly considered a"ground shark" and is rarely t.o be seen from thesurface except when it comes int.o shallow water.An e.xception t.o this occurs off Salerno immedi­ately following periods of especially cold winterweather north of that. area. At such times, whenweather and wat.er surface condit.ions permittedobservations from a boat, large schools of n~il­

berti were t.o be seen headed south, swimming atabout 3 to 5 knots 5 to 10 feet below the surface,but where water depths were about 20 fat.homs ormore. Shark fishermen have. told me, and myown experience bears t.his out, that. it is uselessto try to follow these sharks or to try to divert

20 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

them by chumming or by setting lines ahead ofthem. The appearance of southbound schools atSalerno was generally accepted as a harbinger ofbetter fishing a few days later.

The southbound migratory movements of E.mRbe1·tl at Salerno were inshore and within thesouthbound eddy of the Gulf Stream; north­bound movements were not observed. It is sug­gested that these movements were either offshoremovements or slower movements of more diffuseaggregations. Northbound movements offshorewould be aided by the Gulf Stream. An hypothe­sis which may. be more convenient than signifi­cant is that E. milberti tends to follow currentsin migration and if the currents are strong doesnot go against them. Of course, sharks wouldnot make appreciable he.adway against the.. cur­rent at the surface near the axis of the GulfStream without vigoro.ufi and persistent effort.The sandbar shark does appear to take advantageof eddies or countereurrents and the fishingplans of some of the more successful shark fisher­men were based on an assumption that the shark'sseasonal movements would follow the currentsavailable at the time.

The distance traveled by various segments ofthe population probably does not extend from oneend of the geographical range of the species tothe other. From the southern end of the nurseryrange of the principal stock at Cape Canaveral,a seasonal gradient of availability was shown bycatch per unit-of-effort data. This availabilitydecreased in summer southward and around thetip of Florida to the west coast of Florida wherethe species was completely absent from summercatches. Thus, the minimum migratory travel ofthe part of the stock reaching the vicinity ofTampa would be approximately 600 mile.s.Catches of E. m.ilbeJ'ti throughout the area be­yond the southern end of the nursery range reachtheir highest peak in midwinter. Catch per unit­of-effort data pl'eviously published (Springer,1951) show the catch of E. 'lIlllberti at Salerno,Florida, as decreasing from 4.8 fish per 100 hooksfor FebrUllry to a low of 1.1 per 100 hooks forSeptember. A cold upwelling over the narrowContinental Shelf immediately north of JupiterLight usually occurs in June or July. It is prob­ably of brief duration but annually stuns greatquantities of fish, a.ItllOugh the sharks are not

affected. This phenomenon coincides with spec­tacularly good shark fishing and possibly alsowith considerable mating activity on the part ofE. 1nilbe·l'ti. This may give some bias to Salernocatch per unit-of-'effort figures for early summer.

REPRODUCTIONCourtship and mating

I have seen neither the courtship nor matbIg ofEulam,ia. milbe·I't-i. The general pattern may beconstructed, however, from fragments of infor­mation and from inferences based on the fewfacts known about related "large. sharks. Thecomparative morphology of the secondary sexualapparatus of male sharks has been given compre­hensive discussion by Leigh-Sharpe in 11 papers.The functions ascribed by Leigh-Sharpe in threeof .these papers (1920,1921, and 1924) to the vari­ous parts of the apparatus in carcharhinid sharksa~"e in general accord with my observations onGa.leoeeJ'do, the tiger shark. The courtship pat­erns in Galeoce:rdo, Eu.lmnl-a m.ilbM·ti, and otherlarge carcharhinids probably do not differgreatly.

A brief outline of the mechanics of fertilizationin the carcharhinid sharks is included here toorient the reader in following some of the infer­ences made in later discussion of differentialdeath rates in the sexes. ·Carcharhinid sharks areborn alive and fertilization is internal. Pairedintromittent organs of the male known as dasp­ers are supported by cart.ilages. Immediatelyfollowing the rapid enlargement of the testes,which oc,curs at maturity, layers of calcificationappear at the surface on the principal daspercart.ilages. At this t.ime the claspers becomesemirigid except at the basal area of attachmentof the claspe.rs to the base of the pelvic fin adja­cent to the doaca. The tip of each clasper, how­ever, is expandible. 'When expanded, the carti­lages of t.he tip are transverse to the ma.in axisof the clasper and' open as the ribs of a fan. Theexpanded tips are thought to serve both to hold theoviducts of the female open and to prevent wit.h­drawal of the claspers because of the rigid cart.i­lages in a transverse position. The very largedasper siphons are a distinguishing and peculiarfeltture of the apparatus in male carcharhinids.These siphons are a pair of sepa.rate sacs lyingjust under the skin of the belly on either side of

NATURAL HISTORY OF THE SANDBAR SHARK 21

the midline and extending from the pelvic to thepectoral areas. The,y function as reservoirs forthe sea water used to flush the male sex cells fromthe bases of the claspers into the oviducts of thefemale during mating. The siphons may hold alarge amount of sea water, as much as 2 gallonsin Galeoct3l'do. The siphons do not ordinarilycontain the sea water which is presumed to enterthe siphons during the period immediately pre­eeding mating.

During the mating season the area at the basesof the claspers of the huger carcharhinids exhibitextraordinary vascular congestion. Charaete,ris­tically, 'in lilale (l-a.leocerd(), a mass of very softspongy tissue appears around the cloaca. This ispresent to a lesser degree in the smaller carchar­hinids such as E. milbel'ti. Unusual congestion,edema, and subdermal hemorrhage at the base ofthe claspers are evidences of courtship activity onthe part of the male.

Large, sharks are not highly maneuverable andCtl'nnot swim backward, so it is necessary for theclaspers to rotate and point forward during mat­ing. Since the musde system in the typicalcarcharhinid clasper seemed functionally inade­quate or feeble, I carried out an experiment.which incidentally revealed the probable methodby which the clasper siphons are filled with seawater. I obtained an adult male OOol'chal'hinU8lim bOofus about 5 feet long and evidently in mat­ing condition. The choice of species was dic­tated by circumstances, one of which was the factthat a 5-foot shark was as large as I could man­age. By injecting a, considerable quantity of anisotonic solution into the caudal ve.in, I was ableto induce the claspers to assume the normal mat­ing position. This action caused the claspers torevolve inward and forward. As the claspersmoved into a forward pointing position, a fllnnel,formed by a membrane supported by rods ofcartilage, opened at the base of each clasper. Themouth of the funnel was also directed forwardand the constricted end led i~to the siphon. TheclHIdal vein was plugged experimentally to holdthe claspers and funnel in position and the sharkwas moved forward as rapidly as possiblethrough the water. This caused the claspersiphons to fill with water. Application of addi­tional pressure to the caudal vein resulted in com­plete expansion of the fanlike tip of each clasper.

The course of courtship and mating in all ofthe, larger carcharhinids including E. mJlbertiprobably follows the pattern in which the malepersistently follows and occasionally bites thefemale on the back until she swims upside down.Both claspe,rs probably function at the same time,one entering each oviduct of the female by wayof the lateral opening from the cloaca. The C011­

tact of the two shal'ks may be presumed to forcethe sperm-Iadened sea wate,r from the siphonsinto the oviducts.

The mating pattern has been given in some de­tnil to emphasize the point that mating is verycomplicated in carcharhinids, and that the me­chanical difficulties are' compounded among thelarger species by their greater weight and lessermaneuverability.

Time of mating

The approximate period of the mating seasonis established by the appearance of males withenlarged testes and also with some evidence ofvascular congestion of the pelvic-fin area and bythe appearance of females with eggs of full sizein the ovaries (about 1 to ltA, indIes in diameterin E. m.ilbe1'fi). In the. vicinity of Salemo, Fla.,mating of E. m:ilbe-rti. evidently takes place in thespring or early SUllllner. Males appear commonlyin inshore c,atches after the first of February butremain segregated for some time. Catches ofboth sexes indicating mixed schools are more fre­quently made in April and May than in othermonths. After May~ male E. '1l'dlbe1'ti are rela­tively 'lcarce in Salerno catches. Among car­charhinids, the males may stop eltting during thecourtship period. This is an inference drawnpartly from the general reduction of catches ofmales on baited. hooks during mating seasons,partly from the observed smaller size of thelivers of males immediately following the matingseason, and partly from observations on the mat­ing activity of Galeoce·l'do.

Fertilized eggs and the smallest detectable em­bryos were observed. first in Salemo catches fromthe first part of .July to the first part of August.The time of matiilg can be established with moreprecision, however, by observation of the time ofappenrance of the fresh courtship scars on thefemales. Scnrs, tooth marks, and small openwounds produced by shark teeth are commonly

22 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

found on adult female carcharhinids and are gen­erally restricted to the dorsal surfaces betweenthe two dorsal fins. These are never present onmales or immature females and are obviouslyproduced during court.ship. Scars or wounds arenot always present on gravid females or at leastare not always detectable hut were found onabout half the gravid E. 'lnilbm'ti taken atSalerno. The coexistence. of. old and completelyhealed scars with fresh scars on some females isone bit. of evidence that female E. 'lIl:ilbeJ,ti pro­duce more than one litter of pups in a lifetime.,

All available evidence points to the month ofJune as the time of maximum mating activity ofE. milbel'tl in southeastern Florida waters. Ithas alrea.dy been pointed out that males wererarely taken during the month of .Tune when

.mating act.ivity is assumed to he at a peal\:. Thereis some evidence from catches that the males werepresent in suhstantia,} llumbers at that time.From 3 to 5 percent of the catches of the betterfishermen at Salel'l1o, who keptt.heir hooks verysharp, were- snagged sharks; that is, the sharkswere caught by hooks in the fins or tails 01' occa­sionally in other parts of the body but not in themout.h. More males were caught. in this way dur­ing .June thnn were hooked by mouth.

Development of the embryo

In E. m./lbe'J'fi. as in other carcharhinids, it ispresumed that fel'tiliztttion occurs after the largeegg leaves the single functional ovary. It is alsopresumed .that fertilization occurs before the egghas been moved through a shell gland. Shellglands are located near the anterior end of eachof the two functional oviducts. In passingthrough the shell-gland area of an oviduct, asingle egg is enveloped by a diaphanous tubelikeshell capable of great expansion to acconllllodat.ethe growt.h of the embryo to a very hU'ge size.The nutrient material from the egg yollt is suffi­ciellt only to provide for early growth of theembryo and t.o supplement nutrient materialsnecessary for intermediate growth. The meansby which nonrishment is supplied t.o the growillgembryos probably varies in different species ofcarcharhinids, hut. in species of E:ulmnia, threeprincipal methods appear to he involved. Inaddition to t.hat. supplied by the yolk some ab­sorption of nutri~nt material from fluids in the

oviducts may he assumed to take place. Thiswould appear to be necessary to provide sufficientmaterial to carry the embryo to a length of about12 inches at which length the pseudoplacenta isformed from the yolk sae.

My observations on the embryology of the sand­bar shark are limited to general notes on the ex­ternal appearance of the eggs and embryos atseve-ral stages during development.

The spherical, unfertilized eggs in the singlefunctional ova,ry reach a diameter of 1 to 1%inches. In winter and el\rly spring. large, num­bers of a.c1ult felllltles not earrying embryos werefound to have developing eggs % to % inch indiameter in the ovaries. In a few installces, fe­males taken in July and August were found witheggs of nmximum size in the ovaries a.s well asfertilized eggs in the oviducts. In the greatestdispa.rity of development noted, theore were twolarge yellow eggs remaining in the functionalova.ry while embryos in the oviducts ranged fromless than 6 nun. to 10 mm. in length.

A female m:ilbedi. 6 feet 7' inches long collectedoff Salerno on .Tuly 2, 1948, in 25 fathoms, wastypical of It series taken in early July of thatyear. This female contained 10 egg cases, 5 ineach oviduct; no large. eggs remained in the. flUlc.- .tiona'! ovary. Each st.ringlike egg case was aboutl~O centimeters long, with thin membranous,a.mber-colored. transparent walls. A single yolkwas contained in one expanded oval section ofeltch egg case. The expanded section, approxi­mately 6 cm. long, was located a.bout 10 to 12 cm.from one end of the egg case. This section alsocontained a clear fluid in each of 8 of the eggcases thnt. had developing embryos 9 to 13 mm.IOllg. The remaining 2 egg cases, one anteriorlyin each oviduct, contained milky fluid and therewas no evidence of fertilization nor developmentof the single egg yolk eontained in each. Thesect.ion of the egg cn.se occupied by t.he embryo,spherical yolk, and dear fluid was held in shapeby two longitudinal folds and by folded eonstric­tions of the egg-case membra.ne at either end.The egg case could, however', be unfolded and ex­panded with rell\tively light internal pressure.

The egg case surrounds the embryo until birthand unfolds or stretches to accommodate the de­veloping embryo. 1Vhen the embryo reaches fullterm the pseudoplacental mass extends outside of

NATURAL HISTORY OF THE SANDBAR SHARK 23

the eggshell membrane. but my notes do not imli­cate nt what point the pseudoplacentn functionsoutside of the sheIl membrane. The amotUlt ofclear fluid within the shell increases as the em­bryo grows. It is present at. the time the embryoreaches the approximate size at which the youngare born-24 inches in the Florida area. Exter­iutl gills were noted on embryos up to about 4inches but were absent on 41/~- to 5-inch embryos.Some yolk was found remaining in the .pseudo­placental apparatus in embryos that had reacheda size. of 12 inches, but yolk material was entirelyabsent from the well-developed, yolk-sac placen­tal Rpparatus of embryos 15 inches long. As theembryo grows, the yolk or attachment of thepseudoplRcenta lengthens. In E. m.ilbertl and inother western North Atlantic Eula:lnla the yolkstalk has no structural embellishments at. anystage in its development insofar as I have. beenable to discover. Structures of this kind occurin some species of hammerhe.ad. 8phyrna., butdiffer in the various species. When embryonicE. '/nllberti are near full term the stalk (pseudo­placental attachment) is easily broken at thepoint of attachment to the embryo. The scar ofthis attachment remains clearly visible until theyoung shark has attained an inch or more ofpostnatal growth.

Normally thp. large eggs in the ovary of E.milberti are bright yellow; but white eggs, sug­gesting some pathological condition, were founda few times in the fall. White eggs were noticedless freque.ntly in E. m.ilberti, however, than inEulmnia obsC11tl'a, ('a:l'chal'hIm.lsleueas, or SphYl'na.sp. In these last three species, white eggswere noticed in e.xC(>.pt.ionally large sharks (E.obscu,l'!l. all(~ 8ph1'Y'na sp.) and deformed sharks(l-ell('a.s). No excessively large deformed orobviously diseased m.ilbedi were Seen in ex­aminations of lllany hundreds of Rdult fe­males. Dead embryos were noted oc.casionaUybut. not frequently in E. ·milbe·rtl and these weregenerally 10 to 15 inches long, that is, at aboutthe size at which the egg yolk would be com­pletely absorbed. The de.ad embryos sometimesappeared t.o be dehydrated but there was nonoticeable putrefactive decomposition.

Number of young in litter

In an earlier publicat.ion (Springer, 1940), Ireported the collection of 13 litters of Eulmnla

milbertl pups from Englewood, Fla. In this lot59 were males alld 63 were females. Anotherseries of 28 litters from the west coast of Floridahad 130 males and 130 females. A t.hird seriesof 24 litters from the east coast of Florida in­cluded 116 males and 112 females. The numberof embryos in .each litter varied. from 1 to 14 butthe modal number was 10 and the average num­ber was 9.

The available data do not show an increase inthe numbet' of young with increased size of themother as reported for some other genera(Bac.kus et al., 1956). In Eulamla milberti,

however, the size range of adult females is notgreat and such a correlation, if it exists, wouldbe diffic.ult to demonstrate.

Spedes of Eula1nia taken in the Florida regionother tha.n m.lJberti all frequently carry 10 em­bryos to full t.erm. The average number of em­bryos to the litter for these other species is lessthan 9, which is the average litter number forE. m./lberti. All of the other Florida species ofEu..lamd.a are somewhat larger than milberti andthe largest species, EuJam.ia OOS(JlI.ra, has thesmallest average number of embryos to the litter.In E. obs(:U,ra, pups were often found only in oneof the oviducts. This suggests the possibilitythat fertilization was effected only through oneoviduct. Various other observRtions on t.he loca­tion and numbers of embryos in E. obscwra andother Florida Eulmn·ia. le.ad to the conclusionthat the normal maximum [usual] number ofyoung in edc.h species is 10 plus or minus 2, butthat act.ualnumbers are progressively smaller thelarger the mother.

Length of young at birth

Consideration of all of the available dataplaces the time of birt.h from March to earlyAugust. and the size at birth from 17 to 25 inches.The length at birth of 24 inches seems, however,to be the best estimate for the young born innorthern Florida'waters. Smaller young and asomewhat later birth dRte. may be characteristic.of the part of the population in cooler waters.An estimRte of the gestation 'period ba.sed onsouthern Florida specimens is 9 months with lim­its of the estimate 8 to 12 mont.hs. Some vltria­tion might reasonably be expeeted to result fromdifferences in water temperature during devel­opment.

24 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

TABLE 5.-Embryos of F.ulamia milberti, by size group and month, from females collected off th~ southeast coast of Floridabetween Fort Pierce and Tor/ugas, 1946-49

Number of littersAveragc litter length

__________I._J_Ul__y_~~I~~~~~ Ma~~~~I inch and under______________ 5 7 . _2inches_______________________ 5 _3 inches________________________ 4 • _4 inches_______________________ 6 • _5 inches________________________ 1 1 , _

~ :~~g~~::::::::::::::::::::::: :::::::::: --------~- :::::::::: :::::::::: :::::::::: :::::::::: :::::::::: :::::::::: :::::::::: :::::::::: :::::::::: ::::::::::8inches_______________________ 2 2 _9 inches_______________________ I • _10Inc)les______________________ I _11 Inches______________________ 1 _12Inches______________________ I I . • • _13Inches______________________ I • _14lnches______________________ 2 3 _15Inches______________________ _ 2 1 27 _16Inches______________________ I I 1 11 _17Inches______________________ 6 I • _18Inches______________________ 7 4 I . _l\linches______________________ 4 6 3 1 • _20 Inches______________________ 3 1 • _21Inches .____ 1 _22IIlches . ':.. . 5 _~ ~ __ . ~ _

~ :~~~:~:::::::::::::::::::::: :::::::::: :::::::::: :::::::::: :::::::::: :::::::::: :::::::::: :::::::::: ::::::::::1 ~ --------~- ~ 1

Table 5 shows the length distribution bymonths of emhryos of E. m.il.be'l'fi from off south­east.ern Floridn. It is possihle that some of thevariation in length of embryos neal' full term isdue to inclusion in the sample of measurementsfrom females of widely differing geographicalorigin, for example northern Florida and Chesa,­peake Bay.

At Salerno, Fla., 5- to 10-mm. sandbar sharkembryos were found commonly in July and Au­gnst bnt no large embryos were seen duringthese months. In.Jnne only a few among thelarge numbers of adult. female Eo -m.ilbe1't-i landedat the Salerno dock contained embryos and thefew that were found were 24 inches long. Sinceno free-swimming young were taken at Salernoor around southern Florida, est.imation of thesize at birth in this area depends entirely ondetermination of the maximum length of em­hryos. R.ecords of young E. m.llbeJ'li. from CapeCa.nnvern.l, about 60 miles north of Salerno, north-·wa.rd are common. Twenty-four-inch embryosfrom south Florida. were found to be most C01U­

mon in May but substantial numbers were seen inApril, and one set of 24-inch embryos wn..s takenfrom a female collected off l\farquesas Island inthe Lower Florida Keys in Marph. Eight adultfemale sandbar sharks were tnken on July 2, nearthe mouth of the Mississippi River. Of these,five contained 24-inch embryos and one had 25­inch embryos. Two had evidently ghoen. birth toyoung just before they were caught.

Hildebrand and Schroeder (1928) report sixE. milbe1'ti. in their colle.ction from Chesapea,keBay 17%, to 25% inches long. Presumably thesewere not embryos. Records furnished by WilliamH. Massmann (in correspondence) for youngChesapeake Bay mllbedl include specimens from21% to 28 inches that were taken in 10 collectionsfrom June 7 to October 7. Massmann'8 series isnot large enough to establish progressively largersize with later dates.

Some young E. m.ilbeJ'ti may be born prema­turely at lengths of less than 20 inches either be­cause of crowding, in large litters, or extraordi­nary activity on the part of the mother. Cap­hIres of very small young in otter trawls, forexample, might result from the entry of themot.her into the. net followed by a successfulstruggle to escape. This could bring about pre­mature birth of one or more young which mightbe left in the net.

Nichols and. BredeI' (1927) note that femalesca.rrying young were taken in Great South Bay,Long Island, from .June 22 to August 5. Alsothey state, when relea.sed the young were about22 inches long and weighed 2% pounds. One ofa.bout 3 feet seen in Sandy Hook Bay as early asJune 9 may have been of the preceding year. InSeptember 1924, five young ranged from 24% to26 inches in total length. Bigelow and Schroeder(1948) summarized the indications of size atbirth from various Atlantic. coast records ·northof Florida in approximately the same way. It is

NATURAL HISTORY OF THE SANDBAR SHARK 25

reasonable to expect that E. ndlbe1# at birth lllaybe somewhat smaller in the northern part of itsrange than in the southern pttrt and that thegrowth mte in eooler waters is slower.

Abnormal embryos

In the l:tte spring of Hl42 I collected at Salerno,Fla., a series of Enla-nda m.ilbel'ti embryos ofboth sexes and near full term which were a,p­parently perfect exeept for having the eyes onthe lower side of the snout, almost in contactwith one another and just posterior to the nos­trils, and having no trace of a,n opening in theskin for the mouth, although the jaw eartilageswere apparently normal. The speeimens werepreserved in formalin but dried out during thefollowing years and were discnrded. Again ill1946 similar embryos were collected, and abouthalf a barrel were preserved; but all were, lost illa hurricane which destroyed a dock building.

No abnormal young were found in the rela­tively large series of litters examined in 1948 andI hud no late.r opportunity to see substantialnumbers of E. milbel'ti embryos. All litter matesexhibited the same abnormal condition and wereremarkably uniform structurally.' A very roughestimate of the frequency of occurrence is one setof abnormal young in 500 to 1,000 sets of ap­parently normal pups.

SEX RATIOS

It. was the general observation that landingsof adult Euhun';((, mRberti at Salerno were in theratio of 5 females to 1 male.. A simila.r sex l'lttiowas estimated for Salerno landings of E. obscu:raand for Bahama landings of adult E. spl'in!!el'i.A disproportionately large number of female E.florida-1w, were landed at Salerno' but the recordsdo not furnish an adequate basis for an estimate.An insufficient numbe.r of E. altiJ1/'(~ or E. spring­e/'j, were recorded for estimate. Murphy andNichols (1916). say that the commonest largesharks in the waters about New York are theground sharks (0a'rcnar'ldlJl.uJ5), and also thatmales of these fishes are mrelyseen but towardmidsummer many' of the females eMer our bayswhere they give birth. to their young. Theyfurther state that the commonest ground shark'is Oal'cnm·h.imf.s mHbe·rtl.; The Ol)ly record sug­gesting equality in the number of adults of thesexes of E. m.-ilbedi is Captain 'Vinkler's I'ecord

of the capture of 26 males and 25 females otT theCttribbenn coasts of Nicaragua, and Costa Ricain the fall.

Data on the sex ratio in young E. m-ilbe1# islimited to a series of 203 young from 26 to 50inches long collected by otter trawl in February1958 otT North Carolina. There were 91 malesand 112 females in this collection.

Florida shark fishery records of carcharhinidsother thnn those of the genus E'Illmnia show localsegregation by sex and size but in no other car­charhinid nor in the IUlI1unerheads is there anydenr indicntion from available records fromFlorida of great imbalance in the sex mtios ofadults.

TI~e unavailabilit.y of male m.:ilbe·rti to baitedhooks during the mating season may explain inpart. t.he smaller number of males in the landings.However. because Florida commercial sluu,l\: fish­ing after IH46 wa·s carried on out to dept.hsgrent.er than t.he maximum knowll depth range ofthe species, and because the males brought thefishermen a higher price t.han the femal~s. it iscertain that there was no intentional selection offemnles. The fishermen believed that the schoolsof males, if found, were easier to catch in htrgenumbers. Females were far more abundant thanmnles in the deeper water catches made off theFlorida Keys in the lat.e fall and early spring,and n much greater abunda.nce of females chnr­acterized the wint.er catches on the west coastof Florida.

It has already been shown t.hat. approximatelyequal numbers of male nnd female mllboofi nrehorn. The evidence that there are subst.antiallymore females than males in the adult populationis very strong, if our information adequatelycovers the geographical range of the. species. Al­though it is quite possible that segments of theadult population have been entirely overlookedin the otTshore and midwater depths in the north­ern part of its range, the short.age of ma.Ies in thepopulation around southern Florida is remark­able.

There is some indirect evidence nlso of n short­age of males in the breeding population. If thefemnles bear pups in alternate years. 50 percentof the adult females would be expected t.o begravid in winter. I have previously reported(Springer, 1940) that. only about. 17 percent of the

26 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

lldult females taken in winter off Englewood, onthe west CO:1st of :Florida, were gravid. I sawlarge numbers of E. milbel't{ in 1942 and againfrom 1946 through 1949 at Salerno, and my notesinclude seyernl estimates of the proportion ofgravid to nongruvid lIclult females seen on thedock. For the late-wint.er :ll1d early sprillgperiods, it was estimated that substantially lesst.han a third of the females were currying pups.In my sample of iW9 adult females taken forlength-frequency data, approximately' 18 percentwere grnyid.

Three interest.ing, if theoretical, explanat.ionsare suggest.ed to account for the apparent differ­ences in the number of male,s and females inm.ilbertl and in other Eulamia.

The mating pnttern appears to be particularlydangerous to the male,s, since mating oc('.urs whenthe females nre in a. feeding cycle while the malesare not. That is~ during court.ship males may nipor slash to :,;ome extent but. do not take l:wgebites. The fema.les have no such inhibitions ex­cept at. the time the young are bol'l1, and fatalaccidents to males may be frequent during court­ship. However, if this expla,nution is to be ac­ceptable, some further speeulatiOll is needed tosuggest why aaleo('t~}'do with a similar matingpattern is not represented in the adult populationby a preponderance of fema.les.

Geiser (1924) summarized a. variety of reportson the higher death rate for males in some mam­mals, fishes, and invertebrates, and suggests thatthere' is a genetic bn·sis for this in certain caseswhere the possession of two sex chromosomes bythe females * .., * ensures a greater longevity ofthe female by "canceling out" possible mutationsin the. x-chromosome. especially associated lethals.while in the ma.le there is no such "cance.lingout:'

The third explanation is that the males occupywider geographical and vertical ranges than thefemales. remain in the cooler parts of the ranges,and exhibit a grenter tendency to wander t.hanthe fema.les. Thus, greater numbers of malesthan females are lost to the breeding populntionby wandering and death in unfnvorable environ­ments.

'\Vhntever the explanation for the unequal sexratio, the smaller number of: males is not a suffi-

cient handicap to prevent E. mHberti from beingone of the commoner sharks.

GROWTH AND SIZE AT MATURITY

In Florida catches. adult male Eulmnia m.il­berN average 4.2 inches shorter than the avemgendnlt femille nnd weigh 32 ponnds less than theaverage nongravid adult female. The smallersize of the adult male. is characteristic of all ofthe western North Atlantic carcharhinids al­though the size of males and females at birth isapproximately the same. For about 20 years Imaintained n close watch on landings of oneor more commercial vessels and saw no m.ilbe'l'tifemales longer than 9:3 inches and no males longerthan 89 inches among the thousands that wereexnminec1.

The smallest sexually mature male in the ma­terial exnmined wns 71 inches and the smallestsexually mntnre female was 72 inches in totallength. SexlHlI maturity in the male is easilynnd positively determined because enlnrgement ofthe t.est.es to functionnl size is immediately fol­lowed by the nppearance of a ring of calcium atthe surface of the major clasper cartilage. Thisring is easily seen in cross section but since it.seffect. is to stiffen the segments of the clasper,seet,ioning is unnecessary for positive determina­tions. Determinat.ion of sexual maturity in fe­ma.le speeime.ns whel'e the specimens we,ee 11on­gravid or had no courtship scars wa·s made byexamination of the OVIUY and -the oviducts. Thefemales were regarded ~s sexually iinmature ifnone of the eggs in the ovary had begun to incrensein size and if the oviducts were· smaller in diame­ter than is chal'actel'istic of the fully contractedoviducts in females following parturition. Itmay be noted in table 6 that, while nt len:o;t ~

femllle mRbel'ti were. mature at a length of 72inches, 5 immature females of greater length werecollected. Obviously the length at. which the fe­males may hecome sexually mature. varie.s morethnn 4: inches.

The left skew of the length-frequency polygonsshown in figure 5 may he the result. of a,ny of se.\'­eral varin,bles including the length nt whiehmat.urity is re.aehed.·

A t.otal of 51?, adult sandbar shnrks was select­ed from southeastern Florida. entches for meas­urement of total length and for comparison of

NATURAL HISTORY OF THE SANDBAR SHARK 27

\\ ,

recorded or none were recorded. Selection wasaffected by the scarcit.y of males. The compara­tively large sample of fall females at Salerno hadto be me.asured to get. any catches that includedmnJes which rarely appeared there at tlutt sea·son.In the sample t.aken from the lower east coast of:Florida, 10 shM'ks, 5 males and 5 females, werefound to be immature. These immature sharkswere e.xcluded from the calculations of meanlengt.hs· of adults, but the sizes together with thedates of capture ai'e given in footnotes to table 6,which shows the mean length of the sample lots.

By its migratory movements and its restrictionto limited nursery areas, the North Americanpopulation of E. m.ilbeJ'ti appea.rs to be subjectto constant mixing. It does not seem reasonableto expect a rigid segregation by area of origin ofthos~ mRlJeJ'ti mating off southern Florida. Thismay be one factor in the apparent homogeneityof the population.

In some of the other carcharhinids, environ­mental or racial factors appear to affect the sizeat which the species becomes mature. For ex­amplE'. t.he avera.ge size of the bull shark, 00.1'­

(:h(r.J'hInu.~ 7eneas, from the vicinity of Trinidadis appreciably less than t.he average size of adult.sof the. same species from the Gulf of Mexico.Important difference.s in the size at. maturity aswell as the size at birth se.parate t.he Texas andsouthern Florida populations of the little black-

10I------::.,:..,,........-------~:__------\-__I

801----~-------_+----t-------~

d. = ----

~r=-701-----'----------,f-----1~-----

601-------·----+-------<r-------j

901---------------=:---------__�

n M ~ ~ ~ ~ ~ ~ ~ ~

lOTAL LENGTH IN INCHES

II:\oJm~z 301--------I~...:.,,::_--------___\---__I

" \\

201-------,1--/------'\---------\,..--__1

IL

o 401-----------1----------\----__1

Ul

'"~ 501----------+--------+------/ill

FIGURE 5.-Length-frequency polygons for adult male andfemale Eulamia milberti from southern Florida.

length frequency wit.h the length frequenciesof 76 shark-s from off Fort Myers on the west·coast. of Florida, and of 51 sharks from the Carib­bean coast of southern Nicaragua and nort.hernCosta Rica (table 6). To reduce bias, all sharksof all species in any catch were measured and

TABLE 5.-.Mean length, number measured, and length range of adult Eulamia milberti, by sex, area, and season of collection

[.Lengths ill inches; length range in samples in parentheses)

January-March April-June July-September October-December Combined data

AreaNumber Mean

in lengt.hsample

Numberin

sample

Meanlength

Numberin

sample

Meanlength

Nnmberin

sample

Meanlength

Numberin

sample

Meanlength

-----------------------------------------------

71

26

399

75.0171-841

48

10

.';9

79.9(72--89)

16

47

Gravid (from I school>. _

Nongmvid (from I school>' _. _

78.1 36 78.4 5 78.8 5 78.8 114 78.7(71-891 (72--861 (7fHl2) (73-82) (7\-89)79.0 . -- _

(73-841Nicaragua-Costa Rlca .. _ 78.4(7J-8\l1

82.9(72-921

83.:1173-88182.8 _

(75-88)83.8 _

(75-87183.3 -- _

m~Jo)Nicaragua-Cost.a Rica: Nongravid_____________ 7 82.6 18 83.9 25 83.5

(78-88) (76-90) (76-90)

Southwestern Florida (from 2 schoolsl- _

Southwestcm Florida: Nongravid _

FEMALES:Sont.heastern Florlda.__________________________ III 82.8 63 82.9 1)4 83.0 161 82.9

(72-91) (76-92) <74-91) (73-90)Gravid· • _

MALES:Sout.heastern Florida , _

I 5 immatnre males were collected wit.h t.his sample bnt exduded from tabulation and from calculations of mean length: I specimen 64 inches long collected inMarch, 2 specimens each 66 inches long collected In Augnst, 1specimen 59 inches long collected in October, and I speCimen 6S inehes long rolle.cted in November.

• 5 Immature females were collected with this sample but excluded frolll tabnlation and from calculation of mean lengths: 1 specimen 72 Inches long collectedin January, I specimen 76 Inches long collected in April, I specimen 73 inches long colle·cted in July, and :l specimens collected in October of which 1 was 75inches long and the other 76 inches.

• From preceding collections.

28 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

tip shark;, O"arcnadl.lnus limbatus, but whetherdifferent 'species, subspecies, or races are involvedremains to be determined.

In the last week of February 1958, I was ableto get evidence that the edge of the ContinentalShe.lf off the Carolinas is indeed an importantwintering ground for immature milbel'ti. Ase.ries of 25 tows with a modified Number 41 ottertrawl made by the Bureau of Commercial Fish­eries exploratory vessel De1'«(·lNtl'e while I wasaboard took 203 immature m.ilbel'ti between theoffings of Cape Hatteras and Cape Lookout. Inone tow in 52 fathoms, 197 '/n.ilb&l'ti from 27 to 43inches long were taken and in three other tows atdepths of about 50 fathoms 4 m.ilbel'fi werepicked up. Tows made at. ot.her depths from ~o

t.o 250 fathoms failed to eat.eh mRbel'ti. Thewater t.emperature at the bot.tom at. t.his time. wasabout. 64° F. in 50 fathoms, but temperatures wereltppre.c,iably eooler, ltbtmt [.7 0 F., in both 80fathoms and 80 fathoms.

It is suggested that the 197 young m.ilbeditaken in February 19[,8, we·re young of the year,born in the summer or early fall of 1957, and thattheir average length of 34 inches representedgrowth from birth of about 12 inches in approxi­mately 7 months. However, if it is correct thatall of these young we.re born in 1957, some. mayhave grown as little as 5 inches and some morethan 20 inches. Such an irregular growth pat­tern would help to explain some other facts ofthe m:ilbel'fi. life history, for example, the sea­sonal differences in the vitamin-A potency ofadults taken by the shark fishery.

In .January and February 1945, I obtained twoboatloads of sharks from O. E. Holley, an expertshark fisherman at Fort Myers, Fla. Of the 133sharks brought in, 73 were adult E. mHberti andnone of the female E. m.ilbel'ti were gra vid. The27 males in the sample ranged from 74 to 84inches in total length (average, 79 inches) andweighed from 94 to 130 pounds (average, 113.7pounds) . The 46 females ranged from 72 to 90inches in length (average, 82.3 inches) nudweighed from 102 to 184 pounds (average, 145.1pounds). The specimens were weighed and metlS­ured about 36 hours afte.r capture and the stom­achs of all of the sharks were empty or nearlyempty. 'Veight among sharks of the same length\Taried considerably. Thus, three 7S-inch males

weighed 94, 115, and 1~2 pounds, while e.ight 83­inch females weighed 12g, 127, 141, 14~, 146. 148,161, and 162 pounds, and six 7S-inch femalesweighed 102, 1:36, 147, 147, 1M, and 164 pounds.'Veights of livers, other viscera, fins, and eviscer­ated carcasses were obtained, but unfortnnatelywhe.n t.hese. weights were recorded they were tiedin solely with the totnl we.ight of each shark.Analysis of the figures gives some indication thatliver weights in female E. m.ilbf1'tl contributeddisproportionately to total weights. Livers ofmales weighed from 7 to 14 pounds and livers offemales from 7.25 to 31 i)OlUlds. The a,verageweight of 40 livers from females was 18.5 pounds.

It. would he convenient and would fit most. oft.he facts to assume that the growth of Eulamiam.ilbe'l'fi. is very rapid, but. there. is merely pre­sumptive evidence for this. It is hy no means theonly view that could be reasonably advanced andit would be especially weak for milbedi forwhich the immature part of the popultltion is notwell known. Nevertheless, on the basis of littlereal evidence I suggest that it is probable thatgrowth from birth to mat.llrity ta.kes about 2 yeltrS,occltsionally 1 yenr, rarely 3 years.

The length uniformity indicat.ed in table 3 forboth male and female adult m.flberti from differ­ent seasons would be. expected from a species withdeterminate growth. Predictable adult size inrelatively narrow ra.nges nlso characterizes otherspecies of Eu,l{tm.ia. Negapl'io'n, and perhaps Om'­chaJ'M/I:Ils. On the other hand, it is presumedthat some individuals of Pl'iorwce and Pterola:nd­ops continue growing after maturity to reachnbnormally large sizes. Records of Galeoce'l'domore than 14 feet long are not uncommon in theliterature and some of these records apparentlynre relinble and I accept them as such. Never­theless, I hnve seen more than a thousand largeGa.leocerdo from the Florida-'West Indian regionand have metls1U'ed all of the excel>tionally largeones that I have seen. None were found to exceed14 feet in total lengt.h. Great. increa.se beyondthe usual adult size hns been noted, howewr, for8phy1'Wf sp.. the great. hammerhead. In mysample of 52 adult females, 5 were much largerthan would be expected. If, as indicated by mysample. the' nsual size for adult female grenthammerheads is 10 to 12 feet, the attainment. ofapproximately 15 feet by 10 percent. of the lot

NATURAL HISTORY OF THE SANDBAR SHARK 29

preeludes description of the growth as determi­nate, ·at. least for t.he great hammerhead. OneindividlHlI, measured but not included in thesample, was 18 feet long. Although t.he continua­tion of growth after maturity as a peculiarity ofthe female apparently does not. obtain in E. m:il­7/e1'1i. it is evidently the pattern in some speciesof Spnyrna., in Ptero7amIops, and occasionally in(;a7eoce·l'do.

FOOD AND FEEDING HABITS

Examination of stomachs of Eu7amla mHbedias a source of information on the food of thespecies has been disllppointing. Compnl'ison oft.he feeding habits of E. mHbe"l'fi wit.h the feedinghabits of other ctlrcharhinid species of theFlorida region, however, has proved to be moreilluminating. Various bits of evidence show thatE. mHbertl is a discriminat.ing feeder: that it is abottom feeder; that it feeds on small bottom fishand invertebrates rather than t.he larger ones;that it. prefers fresh fish t.o stale or decomposedfish; that it prefers fish to porpoise meat or tothe flesh of domestic animals; and that its feed­ing is remal'kably successful in comparison withsome of its Ia.rger eal'eharhinid relatives.

A very large proportion of the sharks in eom­mereiallandings were found to have empty stom­achs. An obvious explanation for this is thatmost of the sharks were examinee] after the. pas­sage of several hours wlien small and readilydigestible meals would presumably have beeneompletely hydrolized. Observations and exami­nat.ions of stomachs of sharks of many speeieshas led me t.o the opinion t.hat the larger speciesfind food less oft.en than the small ones, and t.hatthrough no cJlOice of their own, large sharks haveempty stomachs more oft.en than not.

In the st.omachs of Enlamla. m:ilbe'l'ti a few fishremains, usually not identifiable as to species,eephalopods. and crust.ncean remains were foundfrom time. to time. Goatfishes (Mullidae), snakeeels (Ophichthyidae). sea robins (Triglidae), andcusk eels (Ophidiidae). wel'e among the t.ypesmost. commonly found. A eoJIect.ion of 16i m./l­oerti, all with fish, octopns, or crabs in t.heirstomachs was reported in an earlier publicat.ion(~pringt'r, 1946). There were unusual cireum­stances about this catch, however. which sun'(l"est. 00

that the large amount of food in t.he stomachs

resulted from t.he sharks being taken in ltll area. of localized upwelling that had stunned large

numbers of fish a·nd bottom invertebrates. Veryfew instances were. noted in which shark remninswere found in 1II./lDertl stomachs and no evidencewas found that 'l1IHbedi commonly fed on largeturtles, porpoises, birds, ships' glubage, or sur­face material recognized by the inclusion ofSargassum weed or t.ypical surface-dwellingforms.

Some indication of the probable food prefer­ences of E. m:ilbel'ti can be found in the experi­ences of eomme.l,cial fishermen. It. was demon­strated to the satisfaction of Florida shark fisher­men that there were somewhnt different require­ments for bait depending on what species wassought.. It was also found that the increased costof very fresh bait. or bait. frozen when fresh, wasfully justified by the improved catches. All ofthe carcharhinid species of the Florida area, eventhose frequent.ly feeding on garbage, apparentlyprefer fresh bait. The bllit. most. universttlly ac­eepted by all species was fresh fish and E. ?nU­Dei'tl rarely was taken on a.ny other bait.. Somespecies. not.ably E'lilamia 008C1(.1'« and Gal.eoce1'do,took cut. porpoise readily, and one species, Oa1'­cha.rnimlB 7e·/l.cas, occasionally preferred pieces offresh shark. Cut bait was found to be very muchbetter than the entire fish of any species. Prob­ably the diffusion of juices from cut bait wasgreater than from fish in the rounel.

In t.he Salerno, Fla., area the best catches wereobt.nined by sets made in the late afternoon withpieknp of the lines stnrting the following morn­ing as soon as there was sufficient. light to locatethe buoys. The freshness of the bait may havehad some bearing on t.he applll'ent. better fishingat dusk and during the early part of the night.Catches made during the early morning nndthroughout the day were not infrequent, how­ever. and it seems probable t.hat the early partof the night is m~rely a period of increased feed­ing activity for E:u}:a:lIlia lIIHbel'tl.

It is the common habit of many species ofsharks, ineJuding Eulwnia m1ne/,ti, to nudge orhit. objects with theIr noses. I am convincedthat. they do this to test t.he object for ·juices.The edible qualit.ies of the object are thus deter­mined by the shark through sensory crypts whichare widely dispersed over the skin of the hE-ad

30 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

and body. These organs, whieh perform a fune­tion eorresponding to taste, have been described·in detail by BudkeI' (1938). Under ordinarydrcumstances the testing procedure is routine forobjects which are not moving or which are notpart of a series of similar-appearing objeets, oneof whieh has already proved edible. The pro­cedure may be omitted if there is eompetition forfood.

It seems improbable that the sensory eryptsof the head function in any othe.r way than asgustatory organs. To find suitable food or tofind an area having food in it, Eulmnia tnilbe'l'fias well -as many other species depends on olfac­tory organs. Experiments by Parker and Shel­don (1913) with llIuJJtelu8 CI1:1l.iJJ probably outlinethe general pattern of beha.vior of the eachar­hinids in their search for food.

Careharhinid sharks in general are opportun­ists in feeding and necessarily so. Without spe­cial techniques, developed in some spedes but notto my knowledge in E1tlam1a -mH.,berti, sharksare relatively ineffective at catching uninjuredfish in open wnter or even at finding such slow­moving objects as crabs. The shark's ability toexist on a regimen of feast and famine, imposedby its ineptness in eatehing food at will, is prob­ably made possible through its unique digestiveand fnt-storage organs. A general outline of theprocesses of digestion and fat storage by sharksis beyond the scope of this report. It is pertinentto point out that their digestion is rapid andthorough, and that the shark's liver with its highperce.ntage of oil is a good index of its meta­bolie well-being. The larger, fatter livers arefound in sharks in good condition while smalllivers with little oil are' frequently found insharks having severe injuries, sharks in obviouslypoor conditiqn, or in t.he males at the end of themating season.

The liy~n:_ of adult E." milberti typically re·l)l'e­sents beh~eEm 10 and Hi percent of Ow animal'stotal weight, rarely more than 18 percent andrarely less than 6 percent. The proportionalweight of the liver in a.dult E. mRbe-di males islower than in f~males, but the liver weight of thespecies is remarkably· uniform in comparison withlarger species of carcharhinids within its geo­graphical range. This is strong evidence thatE. -milberti is suecefisful in getting an adequate

supply of food regularly. Galeocerdo au-lIfer,Oarchm'kinlls l-ellca~, and Eu..lam;ja obsC1./-m, allspedes more than twiee as heavy as milbe'rt-i,frequently have livers 25 pereent or more of thetotal weight of the fish. On the other hand, theyoften have very small livers, as low as 3 pe.rcentof the total weight. The inferenee is reasonablethat the larger spedes have greater difficultiesfinding food, or more precisely, may have to waitlonger between meals. This inference from liverweight also is eonsistent with the observationthat the foods of the larger species are frequentlyof a less digestible type and may be taken as akind of desperation measure and certainly not asa first choice. Galeoc.e-rdo, for example, fre­quently fills its stomach with large horseshoeerabs, huge horse conchs complete with shell,or even old shoes and tin eans.

Bee-ause of eommercial interest in the vitamin­A eontent of shark-liver oils that existed for alllunber of years, a large amount of data is tobe found in the literature on the subject. Alsoavailable to me are data on the oil and vita­min-A content of livers of E. m:ilbe-l'ti taken atSalerno. Some determinations for Salerno spe­cies were given in an earlier publieation(Springer and Fre.nch. 1944). Better methods ofestimation of the vitamin-A eoneent-ration in liveroils eame into general use later. but these did notappreciably alter the general trends observed.

The tendency in each species to an inerease inthe vitamin-A coneentration, or poteney, of theliver oil with the inerease in size of that specieshas been noted by many workers (Pugsley, 1939;Broeklesby, 1941; Templeman, 1944; Ripley andBolomey, 1946, et c.atera). The liver-oil vitamin­A potency varies considerably. Eaeh species andlocality produces sharks having potencies that areapproximately predictable. Characteristically,in most. speeies the males produce oil of some­what higher vitamin-A poteney than that of t.hefemales, but they often have less oil in theirlivers.

An hypothesis whieh has general support isagain advanced here that within a given speciesand loeality, the. total amount of vitamin A inthe liver of a shark is roughly proportional t.othe llge of the shark. Thus, the older the sharkthe greater is the total amount of vitamin Ain its liver. The rate of increase in vitamin A

NATURAL HISTORY OF THE SANDBAR SHARK 31

may rise sharply as the shark becomes sexuallymature but this does not affect the general trendin the total amount of vitamin A stored. Fluc­tuations in the amounts of oil stored in the livermay normally accompany such events as youngbearing, nutting, .and periods of poor feeding,and these fluctuations apparently affect potency,but the total amount of vitamin A is subject tofluctuation to n lesser degree. It. has been shownfor Ga.leOJ·hlnus zyoptel'1.IS of the Pacific coast(Ripley und Bolomey, 1946) that the totalamount of vitamin A in the largest males andlargest females does not greatly differ.

Livers of Squalus 8uddeyi of the Pacific coasthave. a higher average vitamin-A pote.ncy thanlivers of Squ,a1118 (UJanthios of the Atllmtic. Thetwo species are morphologically so similar thnttaxonomists have had doubts about their sepa.ra­tion. Concerning the differences in liver oil po­tency, Templeman (1944) has the. following com­ment:

Whether the greater length of the mature Squa~us

8I1CJ..,Ui reflects a greater age or a greater growth ratethan those of Squallts aca.nthias is not known, but thisgreater size is possibly partly responsible for the highervltamin-A value of its liver oll.

The greater vitamin-A content of shark liversdoes not ulways occur in the larger individualsin a given species. For example, my own meas­urements and assays show that O. lencas off th~

mouth of the Mississippi River average appre­ciably longer and heavier than O. lcu.cas takenoff the mouth of the Orinoco River; but the liver­oil potency of the species from the vicinity ofthe Mississippi River mouth is extremely low,while that of the C. leucas from the vicinity ofthe Orinoco is high. These differences are quitegreat and reasonably constant. Adult male O.leucas from the Orinoco area have liver-oil po­tencies above 50,000 I.U./gm. (international unitsper gram uncorrected for irrelevant absorption),while adult males from the Mississippi area gen­erally have liver-oil potencies of 1,000 to 5,000I.U'/gm. Variation in liver-oil potency by areais perhaps greater in O. l.eueos than in most sharkspecies of the northwestern Atlantic, but the vari­ation is appreciable in all species.

A rough classification by regions of part of thenorthwestern Atlantic on the basis of shark-liver

vitamin-A potency from data assembled by theshark fishery is as follows:

Extremely low potencies-Gulf ()f Me.xico; lowpotencies-northern Bahamas and shallow-waterareas of the 1Vest Indies; intermediate potencies-Carolina COltst, east coast of Florida, and COlLstof Cuba; high potencies-southern Caribbean,coast of Costa. Rica to the Guiana coasts. Toa limited extent this classification is useful totrace probable origins of elements of migratorystocks. Potency differences among liver oils takenfrom single adult specimens of Eula'lnia '1nilbe'l'tiwere in the range from about 2,500 to 25,000

.I.U./gm. One exception' was noted in that oneassay (the only one available) on mixed E. mil­berN livers from the coast of Nicaragua-CostaRica was substantially higher, about 38,000 I.U./gm.

If E. mHbertl from the coast of Nicaragua­Costa Rica returned regularly to the species' At­lantic population center from Florida northward,some among the hundreds of livers of mllbel'tih\ken at Salerno, and assayed separately shouldhave had potencies greater than 38,000 I.U./gm.None had such a high potency.

The average potency of oil from livers of Eu­la:1nia 11lHberti was found to be substantiallylower from the southwest coast of Florida thanfrom the southeast coast although the extremesin jndividual potencies were about the same.Since both stocks have a common origin from theAtlantic coast north of Cape Canaveral we mightassume. that the difference in a.verages resultsfrom either the lower age of the. adult rnilbertireaching the west coast of Florida or a lowerrate of vitamin-A accumulation due to the periodspe.nt in the Gulf of Mexico, or from bot.h ofthese.

The liver-oil. potency of the eight adult femaleE. m.ilberfi from the mouth of the MississippiRiver was higher and the total vitamin A fromthese sharks was about the amount to be ex­pected from similar sharks. taken on the easteoast. of Florida. This poteney would be ex­pected only if these sharks originate outside ofthe Gulf of Mexico.

To the extent t.hat data on vitamin-A potencydeterminations have any validity for the deter­mination of age or the areas of origin of migra­tory stocks, they support. the general conclusions

32 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

that E. lIlilb('rti does not live to a great age andthat those individuals migrating a great distanceto the coast of Nicaragua-Costa Rica do not re­turn..

ABUNDANCE

The inshore range of the sandbar shark hasmade this species readily available to the sharkfishery. This was particularly noticeable tit Sa­lerno, where the range is restricted by a narrow­ing of the Continental Shelf and where specialconditions of current and temperature tend toproduce a narrow path inshore for the south­bound migrants. Approximately 58 percent oft.he sharks landed by the fishery at Salerno be­tween 1938 and 1946 were sandbar sharks. Al­though present throughout tlle year, they werecomparnt.ively more frequent in Salerno catchesfrom December through July as shown by rec­ords of the catch per unit of effort (Springer,1951). Except for winter fishing on the south­west. coast of Florida which produced 11 fe.whundred E. m:ilberti each winte.r, few sandbarsharks were ta.ken in the Gulf of Mexico.

It is estimated that during the period 1935 t.o1950 from 5,000 to 15,000 sandbar sharks weretaken yearly from the entire range of the fisheryand th:;lt these were nearly all adults with anavera.ge weight of about 130 pounds. The. yearlycatch might be estimated, therefore, as between650,000 and 1,950,000 pounds, a small qmmt.itycompared with yearly landings by the commer­cial fishery of many species of bony fishes. Dur­ing the years that the shark fishery operated, itwas prosecuted vigorously, and, alt.hough therewas no evidence that fishing pressure reduced thestock of sandbar sharks, it was found that thecatch per unit of effort was reduced by concen­trating too much gear in one area. Efforts toexpand production were successful chiefly by ex­t.ension of the arel1 of fishing into the ranges ofother species of sharks. A fluctuation in abnnd­Il.l1Ce with a· low in every third year was found inthe catch per uni.t of effort. at Salerno (Springer,19!11).

Although data are lacking to support such acontention except observations at sea, it. is esti­mated that several other species of E'll.lmn:ia, par­ticularly E. {loridana, occur in substantiallygreater species-mass around Florida than does E.

mHbn'fl, chiefly because of the greater area oftheir habitats.

It is generally recognized that marine animalpopulations are unstable and are subject to re­ma.rlmble changes in total numbers and occasionalshifts in geographical range. !3uch changes may,of course, occur in the Atlantic population ofE. mHbertl.

A concentration of E. m.llberti appeared for afew cla.ys in the late spring or early summer of1935 off Dog Keys Pass on t.he coast of Missis­sippi. These sharks were not only out of theirnormal range but. behaved in a way that is notnormal for mHbel'ti or perhaps for any shark.They struck at anyt.hing thrown into the water,fought one another over pieces of charcoal, foughtso vigorously that some were killed and eatenby others of the school. Sharks frequently fol­low shrimp boats such as we were using at thet.ime, and churn the surface of the water afterscrap fish thrown overboard; but the intensi.tyof this attack by out-of-range mHberti was muchstronger than any shark action I have seen since.

Alt.hough no great fluctuations were noted inthe general abundance of E. m.ilberti during theperiod from 1935 to 1950, its competitor, the bullshark O. leuNl8, appeared once to go through amajor but temporary shift in abundance. In1n37 so many bull sharks were caught off Salernothat catches exceeded the demand. The· companybuying shark livers and oil was forced to delaypa.yme.nts to fishermen for a period of severalmonths. This was not merely a matter of eco­nomic adjustment to be settled by a reduc.tion inprice but the supply of tanks and drums forstorage of liver oil was exhausted and a courtinjunction was finally issued to stop all sharklandings because shark carcasses were accumulat­ing along the shores. This was the only knownappearance of bull sharks in large numbers onthe east coast of Florida. It seems likely thatpersistence of great. numbers of bull sharks, es­pecially north of Cape Canavera.l, would haveadversely ltffected the population of E. m.i7berti.

ENEMIES

The principal predators on sharks are othershnrks of larger size. 'Vherever concent.rationsof mixed sizes occur predation by the largersharks on the. smaller ones is normal. The con-

NATURAL HISTORY OF THE SANDBAR SHARK 33

centration of sharks occurring near the mouth ofthe Mississippi River in the summer months maybe as great as anywhere in the world. Some idea,of the predation that may occur is furnished bythe results of' one line of 180 shark hooks set onthe night of July 22, 1947, off Pass a rOutre, La.This line caught 68 large sharks, whole and Ull­

damaged, but 40 of the hooks had only theheads of large sharks and 12 more had onlythe heads of small sharks. Almost all of thelarger sharks on the line contained small sharksor llieces of large ones. One tiger shark, Gltl,f?Q­

ee-rdo, had swallowed a medium-sized bull sharkwhich in turn had a major portion of a somewhatsmaller blacktip in its stomach. The blacktip,however, had driven the head of a M'!l~~teln8 ea'ni.sa.bout 4 feet long up the leader so it. would bereasonable to assume that the large, tiger sharkwas the fourtll shark to be taken in that onenight Oli one hook. The bottom conditions.depths, and temperatures where this line was setappeared to be similar to conditions on lnllbf?'1,tinursery grounds but predation would presumablyeliminate young mllbe'r# in the area.

Full-grown sandbar sharks probably are rarelysubject to successful shark attack by other spe­cies. The tiger shark (Galeoeel'do) , dusky shark(E-ulcunia oOs("l.(,m) , and bull shark (CarcharMnu.sle-ucas) , have all been found occasionally withpieces of full-grown sandbar sha.rk in their'stom­achs, but the sandbar sharks may have been onshark lines when. attacked. Perhaps none ofthese species are able to catch adult sandbarsharks under ordinary circumstances. Greatwhite sharks, (!a'1'charodo'n C(il'cnarias. have beenfound with adult milbedi in their stomachs andit is probable that the great white shark couldcatch them. The white shark is not commonenough, however, to be an important factor inpredation. At Salerno~ captures were about 27great white sharks per 190,000 of--all species.

All carcharhinids more -than 6 feet long mayoccasionally eat young E. n1iilberti, but circum­stances of seasonal and geographical distributionkeep most species from preying on them. Therenre two notable exceptions: the tiger shark andthe bull shark. These sharks feed on youngsharks or on small species regularly, and bothlllay be found at times within the known rangeof young m.ilbel'ti.

Tiger sharks, perhaps for reasons of poor speedand maneuverability associated with their smallerand lighter weight fins, catch young sharks lessfrequently than do bull sharks. Furthermore,tiger sharks are primarily nocturnal in forayl:illltO inshore waters and at such times newbornor small sharks retreat to shoaler water. On theother hand, bull sharks are not exclusively noc­turnal and they are more frequent in relativelyshallow water. Stomach contents show that theyare regularly predatory on small species such asCareha.l'hInus aCl'o-noht-8, C. porosus, 8colioilonterra.-'1l-lH'ae, and Apl'io-nodo'n i.sodon. The bullshRrk also is the only species in the range ofE. milbe'rti with a preference for shark as abait.

There is strong circumstantial evidence, derivedfrom an examination of the geographical rangesand nursery-ground preferences of the varioul:ispecies of Eulam.ia, that the bull shark is themost important predator on young sharks andactually restricts the distribution of mHberti. Thelife history of the bull shark is similar to thatof species of Eulmnia, but the bull shark is al­ways found in shallow water and its nurserygrounds are in bays or estuaries, even in brackishand fresh water. The nursery range of the bullshark is shoaler and less saline than the- nurserymnge of n~llb&rti, but adult bull sharks inhabitthe depth and salinity range normal to nurserygrounds of milberti. Young bull sharks remainin estuarine waters during t.he' early growingseason, and since the females do, not eat at thetime the young are born and gen~rally move outof the very shallow water inllnecliately after thebirth of the young, the young hull sharks areprotected to some degree from predation by thelarge members of their own species.

Bull sharks occur from Long Island southwardand are migratory but t.heir centers of abundanceare in the GuIf of Mexico and soutlnva;rct par­ticularly nea,r the mouths of large rivers. Alongthe Atlantic coast north of Florida, hull sharksare not common. but m,ay be subject in t.his areato great. fluctuation in abundance. Bull sharksare ext.remely common around the mouths of theMississippi and Orinoco Rivers and between thesepoints along the inshore, Continental Shelf. Thereis pract.ically no information about their occur­rence in the West Indies.

34 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

No Euhunia except milbel'ti is known to Ill\,venursery grounds within the geographical andhabitat range of the bull shark. The nurseryranges of E. althna. and E. obsc·/tra are well off­shore. in deeper water, E. jlOl-idana nursei'ygrounds are on offshore banks where bull sharksdo not go, and bull sharks are uncommon atleast, or normully absent around reefs which arenursery grounds for spl'ingeri. The sugges­tion is advanced he.re that predation by the bullshark is the chief reason, and perhaps the oniyreason, for the scarcity of E. milberti in the Gulfof Mexico and sout.hward in continental waters.The comparatively few bull sharks of the Atlan­tic coast may be presumed to act as a eheck onthe numbers of young E. 11I.ilberti, but withoiltdisastrous effects for that species.

Some interesting eomplications are associatedwith predation of large sharks upon smaller ones.Evidently all carcharhinid sharks will eat othersharks sometimes if not regularly. However, itis a shark fisherman's axiom that sharks left onthe line to spoil will burn out the fishing groundsand make it necessary to move away 5 miles orso to eontinue fishing. A rationale has beenpartly outlined in an earlier publication(Springer, 1954) covering experiments with 1111(.8­

tellfs in tanks. Feeding of lI/1fstelus was entirelyinhibited in the presence of shark flesh that hadbeen allowed to stand at room temperature for4 days.

Presumably sharks have no difficulty digestingshark flesh provided it is fresh. Although sharksappear to be able to digest the partially decolll­posed flesh of other vertebrates, their digestiveprocesses are slowe,d or stopped completely whenthey sWil.Ilow large quantities of decomposingsharks or rays.

The flesh of sha.rks becomes strongly alkalh1eduring decomposition. The continuous liberationof amlllonia through action of enzymes producedduring the course of ordinnry putrefactive de­composition on the uren that norma lIy occurs inshnrks (see Smith. HIS(1), seems effectively toblock, or at least to greatly retard, digesti0l1.The proteolytie enzymes of the shark's stomachare most active in an acid medium (Sprissler,1942). In my examinations of juices of sharkstomachs in the field, estimates were made of thehydrogen-ion concentration, as shown by indica-. .

tor solutions. .Juices from shark stomachs thatconta.ined huge amounts of shark flesh in obvi­ously decomposing condition were found to besubstantia.lly above pH 8.0, whereas juices fromshark stomachs conta.ining fish, tm'tles, birds, orsmall amounts of fresh shark were estimated atpH 4.0 or below. It is not dear whether ex­eessively large meals of decomposed shark fleshproduce any result more serious than delayeddigestion.

I have not seen evidence that internal parasitesever greatly damage E-u.la.·mJ((, mRberti. Round­worms in the st.omach, roundworllls and adulttapeworms in the scroll-type intestine, and cope­pods on the gills and external surfaces werecommonly seen. In comparison with other largeFlorida sharks,mRbeJ'fi seemed to he the leasttroubled by parasites. Occasionally, sharks takenfrom lines apparently set in areas of extraordin­nry abundance of II small isopod were 'found t.ohave a large proportion of the viscera eaten awayby swarms of the isopoch" which had entered tll('body cavity by way of the soft parts aroundthe a·1lUS. It has been assumed that. these iso­pods attack successfully only after the sharksdie on the lines or at least have been restrirt.edin movement.. Similar isopods attack living smallfishes and man (Springer, 1957) and it seemsquite possible that. they may a'!so attack livingsharks. If they successfully attack living sharksthey may be one of the principal enemies ofsharks in temperate and tropical waters of shal­low and modernte depth. I have seen evidenceof their work on shark catches made off SouthCarolina, Florida, Louisia.na., Cuba, and sout.hernCalifornia.

SUMMARY

The overlapping geographical and habitatranges and superficial resemblance of E'ulmnhtm.ilbedi, the sandbar shark. and (Ym'chal'hinU8leu.ca.~, the bull shark, ha.ve led to some, confu­sion and n tangled nomenclature. The recogni­tion of m.ilbeJ'ti of Muller and Henle as the spe­cies name for the sandbar shark is based on theopinion that plltlnbeu8 of Nardo is a nome'll'1I:lldf;~m,.

The sandbar shark differs from the bun sharkin the structure and spacing of the derma'! denti­des, in having a ridge in the skin of the backbetween the first and second dorsal fins, and in

NATURAL HISTORY OF THE SANDBAR SHARK 35

the smaller size and less robust body form. Thesandbar shark swimming in the open sea or inan exhibition tank may be distinguished readilyfrom the bull shark been-use of its comparativelymore erect and higher first dorsal fin placedslightly further forward, its some\vhat larger eye,its somewhat longer snout, and its genei'ally less

. robust body form.Although the sandbar shark inhabits both the

eastern and western parts of the At..lunt.ic, thisreport concerns only one popnlation of the spe-.cies centered along the southeastern coast of theUnited States.

Species of the genus Etdmnla usually are sharksof the continental shelves, oceanic banks, andisland terraces throughout their life cycles incontrast to most species of Oa1·cha.J'1d'ntl-8, whichaTe typically shallow-water forms having theirlHlrsery grounds in brackish or fresh-wllter estu­aries, river mouths, or along continental beaches,or in island lagoons. Eulmn-ia. m.ilberti occupiesthe shoalest habitat range of the [) or 6 species ofEulmnia. of the Atlantic coast of North AmericlLand enters est.tllll'ies t.o some extent to give birt.hto its young. Its habitat preference may thusbe said to be. intermediate between that typicalof Eulamia and t.hat typical of Oal'clW1'hi:nus.

The sllndbllr shark is clearly separable from allother species within it.s geographical range andexhibits little variation in form and in toothcount.

The sandbar shark, Eula'lnia. 'I1l:ilbe1'tl~ is foundin waters of suitable depth, southward from CapeCod a.long the Atlantic coast. It occurs in theGulf of Mexico and along the Caribbean coastof Cent.ral America to Costa Rica. It. occurscasually off the northern coast of Cuba and alongthe western edges of the Bn-hama Bank, but prin­cipal elements of the population are confined tothe western side. of the. Gulf Stream. Sandbarsharks occur as bottom dwellers out to depths of100 fatllOms (extreme record 135 fathoms) andmay occasionally move out in midwatel' to oceanicsituations. The principal part of t.he ra.nge ofadults, away from the nursery grounds, ha.s beenshown by commercial catches to be in depths of10 to 30 fnthoms.

The sandbar shark populations of the northernnnd western parts of the Gulf of Mexico and theCaribbean coast of Central America are small

and are probably not self-sustaining wit.hout re­cruit.ment from the main population by migra­tory wandering or failure of such orientationmechanisms as the species may possess.

The primary nursery range lies in shallowcoastal waters of less than 20 fathoms fromCape Cod, Mass., to Cape Canaveml, Fla.; anda secondary nursery range lies in the Gulf ofMexico west of Mobile Bay and north of the28th pn,rallel. Newborn sandbar sharks are notlmown outside the general geographical limits ofthe nursery ranges, but the young sharks mov~

offshore to deeper (and ,varmer) water duringthe winter.

The species is migratory with annual move­ments of some segments of the population ex­tending at lenst 600 miles. Sandbar sharks inmigratory pa.ssage around the southern tip ofFlorida hold to dept.hs of 50 fathoms or morein apparent avoid:Ulce of coral reef a.reas. Theirvertical distribution in the southern part of thespecies' geographical range, however, is soml:"­what deeper t.han in the northern part, suggest­ing thermal influence in the selection of habit.at.

There is reason to believe that femnJe sandbarsharks are inhibited from feeding at the time ofbirt.h of the young and for a short time t.here­aft.er. Nursery grounds are away from the nor­mal range of the males thus giving additionalprotection to newborn young. Feeding appears tobe inhibited in male sandbar sharks during peri­ods of active courtship.

Young of nearly uniform size, numbering from1 to 14 in each litter, are born in early summer(probably also in late spring) off nort.hern Flori­da. The average number in a litter is 9 andthe modal number 10. The gest.ation period isfrom 8 to 12 months' duration. with 9 monthsestimat.ed in the latitude of northern Florida.Both oviducts are functional, a single embryo de­velops in each shell membr~ne. and a pseudopla­centa with :t simple stalk (yolk-sac attachment)forms.

Lengt.h at. bil"th is approximately 24 to 25inches in the latitude of nort.hern Florida butmay be less in lligher latitudes. Young bornin the vicinity of Long Island, N. Y., have beenreported to be 22 to 23 inches long at birth, andeven smaller ·young hnve been reported fromChesapeake Bay.

36 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

Although approximately equal numbers ofmalel!! and females are born, catches indicate amuch larger proportion of adult females thanadult males except off the coast of Nicaragua,.

Females produce young no oftener than everyother year, but since only about 18 percent ofthe adult females taken in the fishery were foundto be gravid, this apparent low productivity maybe the ~esult of the low proportion of males inthe adult population.

In samples from Florida, adult males are from71 to 89 inches in total length and adult females.from 72 to 92 inches. Average weights of adultsin samples from southern Florida are ~or males,114 pounds, and for nongravid females, 145pounds.

The mean length and length-frequency dist,ri­bution (size range) by sex in catches believed tohave been made from single large schools closelya,pproximated the mean' length and size range bysex of all adults of the spe~ies from southernFlorida. No appreciable difference was noted inmean lengths a.nd size ranges of adults in sam­ples ta.ken at various seasons on the east coast ofFlorida, the west coast of Florida, and the Carib­bea.n coast of Nicara.gua and Costa Uica exceptthe constant difference of about 4.2 inches be­tween average lengths' of adult males and adultfemales. Differences in the average lengths ofgmvid and nongravicl females were small.

The rate of growth is not known. Indirectevidence indicates that it may be very rapid untilse.xual maturity is reached, after ·whic.h little.growth occurs'.

The sandbar shark feeds chiefly on small bot­tom-dwelling fishes, mollusks, and crustaceans.It. 'rarely swallows indigestible materials. Uni­formly plump livers suggest that milbe1,ti has littledifficulty in meeting its requirements for food.

From commercial catches it is estimated tha tthe population is small in species mass as com-pared with many teleosts. .

The only important predators on sharks areother sharks and not necessarilv sharks of otherspecies. Fuil-grown sandbar ~harlts, unless in­jured or ca'ught on a shark liile, appear rarelyto be eaten by other sharks. Young sandbtu'sharks are especially vulnerable to attack by

large shnrks, particularly by bull sharks, butalso to a lesser extent by tiger sharks, duskysharks, and full-grown slutrks of their ownspecies.

LITERATURE CITED

BACKUS, RICHARD H.1957. Notes on western North Atlantic sharks.

Copeia, No.3, pp. 246-248.BACKUS, RICHARD H., STEWART SPRINGER, and EDGAR L.

ARNOLD, JR.1956. A contribution to the natural history of the

white-tip shark, Pterolamiop8 longiman1t8 (Poey).Deep-Sea Research, vol. 3, pp. 178-188.

BEEBE, WILLIAM, and JOHN TEEVAN.1928. The fishes of Port au Prince Bay, Haiti.

Zoologica, vol. 10, No.1, pp. 1-279.BIGELOW, HENRY B., and WILLIAM C. SCHROEDER.

1948. Sharks. 111 Fishes of the western North At­lantic. Memoir No.1, Sears :Foundat.ion MarineResearch, pt. 1, pp. 59-576.

1953. Fishes of the Gulf of Maine. U.S. Fish andWildlife Service, Fishery Bullet.in 74, vol. 53, pp.1-577.

BROCKLESBY, H. N.1941. The chemistry and technology of marine

animal oils with part.icular reference to those ofCanada. Fisheries Research Board of Canada,Bull. 59, pp. 1-442.

BUDKER, PAUL.1938. Les c,ryptes sensorielles et les denticules cutan~s

des Plagiostomes. Annales de I'Inst,itut Oc~ano­

graphique, vol. 18, pp. 207-288, pI. 1.BULLIS, HARVEY R., JR.

1955. Preliminary report on exploratory longlinefishing for tuna in the Gulf of Mexico and theCaribbean Sea. Commercial Fisheries Review,vol. 17, No. 10, pp. 1-15.

CADENAT, JOHN.1950. Rapport sur Ies Selaciens des cotes du Sen~gal

et. plus specialement sur lea Requins. Bulletin del'Institut franliais d'Afrique noire, vol. 12, pp.944-975.

195i. Notes d'ichtyologie ouest-africaine. XVIIBiologie. Regime alimentaire. Bulletin de I'Institutfranliais d' Afrique noire, vol. 19, ser. A, No.1,pp. 274-294.

CAPTIVA, FRANCIS J.1955. Longline gear used in yellowfin t.una explora­

tion. Commercial Fisheries Review, vol. 17, No. 10,pp. 16--20.

COWLES, RAYMOND B.1945. Temperature induced st.erility and evolut.ion.

Science, vol. 101, No. 2618, pp. 221-222. .EVP;RMANN, BARTON W., and MILLARD C. MARSH.

1902. The fishes of Puerto Rico. Bulletin of theU.S. Fish Commission, vol. 20 (1900). pt.. 1, pp.51-350, pIs. 1-51.

NATURAL HISTORY OF THE SANDBAR SHARK 37

FORD, E.1921. A contribution to our knowledge of the life

histories of dogfishes landed at Plymouth. Journalof the Marine Biological Association of the UnitedKingdom, New Series, vol. 12, No.3, pp. 468-505.

FOWLER, HENRY 'V.1936. The marine fishes of west Africa. Bull.

A:merican Museum National History, vol. 70, pt. 1,pp. 1-605.

GARMAN, SAMUEL.1913. The Plagiostomia (sharks, skates, and rays).

Memoir Museum Comparative Zoology, vol. 36,pp. 1-518, pI. 1-77.

GEISER, SAMUEL WOOD.1924. Differential death-rate of the sexes among

animals, with a suggested explanation. Washingtonl;.Tniversity Studies, Science Series, vol. 12, No.1,pp.73-96.

GILL, THEODORE N.1862. Analytical synopsis of the order Squall.

Annals Lyceum Natural History, New York, vol. 7,pp.367-408.

HICKLING, C. F.1930. A contribution towards the life-history of the

spur-dog. Journal of the Marine Biological Asso­ciation of the United Kingdom, new series, vol. 16,No.2, pp. 529-576.

HILDEBRAND, HENRY H.1954. Shrimp (Penaeu8 aztecu8 Ives) grounds in the

western Gulf of Mexico. Publication of theInstitute of Marine Science University of Texas,vol. 3, No.2, pp. 233-366.

HILDEBRAND, SAMUEL F., and WILLIAM C. SCHROEDER.1928. Fishes of Chesapeake Bay. Bulletin of the

U.S. Bureau of Fisheries, vol. 43, pt. I, pp. 1-366.LEIGH-SHARPE, W. HAROLD.

1920. The comparative morphology of the secondarysexual characters of elasmobranch fj.shes. Memoir1. Journal of Morphology, vol. 34, No.2, pp.245-265.

1921. Idem, Memoir II. Ibid., vol. 35, No.2, pp.359-380.

1924. Idem, Memoir VI. Ibid., vol. 39, No.2, pp.553-566.

MEEK, SETH E., ·and SAMUEL F. HILDEBRAND.1923. The marine fishes of Panama. Field Museum

Natural History Publication No. 215, ZoologicalSeries, vol. 15, pt. I, pp. 1-330.

MULLER, JOHANNES, and F. G. J. HENLE.1841. Systematische beschreibung del' plagiostomen.

Berlin, pp. 3-200, pIs. 1-60.MURPHY, ROBERT CUSHMAN, and JOHN TREADWELL

NICHOLS.1916. The shark situation in the watNs about New

York. Brooklyn Museum Science Quarterly, vol. 3,No.4, pp. 145-160.

NARDO, G. D.1827. Prodromus ObservaUonum et disquisitionum

Adriaticae Ichthy010giae. Giornale fisica, chimica,storia naturale, medicina ed arti, vol. 10, pp.22-40. Pavia (1827).

NICHOLS, JOHN TREADWELL.1918. Ground sharks of Florida and elsewhere.

[Abstract.] Copeia, No. 53, p. 13.1929. The fishes of Puerto Rico and the Virgin

Islands. New York Academy of Science Survey,vol. 10, Pt. 2, pp. 161-295.

NICHOLS, JOHN TREADWELL, and CHARLES M. BREDER, JR.1927. The marine fishes of New York and southern

New England. Zoologica, vol. 9, No.1, pp. 1-192.OLSEN, A. M.

1954. The biology, migration, and growth rate ofthe school shark. Galeorhinu8 aU8trali8 (Macleay)(Carcharhanidae) in south-eastern Australian waters.Australian Journal of Marine and FreshwaterResearch, vol. 5, No.3, pp. 353-410.

PARKER, G. H., and R. E. SHELDON.1913. The sense of smell in fishes. Bulletin of the

U.S. Bureau of Fisheries, vol. 32, Doc. No. 775.pp.35-46.

PUGSLEY, L. I.1939. Factors influencing the vitamin A and D po­

tency of grayfish liver oil, Sqlla.l1l8 8ucklii (Girard).Journal of the Fisheries Research Board of Canada.vol. 4, No.5, pp. 405-408.

RADCLIFFE, LEWIS.1916. The sharks and rays of Beaufort, North

Carolina. Bulletin of the U.S. Bureau of Fisheries,vol. 34, Doc. No. 882, pp. 241-284, pIs. 38-49.

RIBIERO, ALIPIO DE MIRANDA.1923. Fauna Braziliense, Peixes, vol. 2.

RIPLEY, WILLIAM ELLIS.1946. The soupfin shark and the fishery. California

Division of Fish and Game, Fish Bulletin 64,pp.6--37.

RIPLEY, WILLIAM ELLIS, and RENE A. BOLOMEY.1946. The relation of the biology of the soupfin

shark to the liver yield of vitamin A. CaliforniaDivision of Fish and Game, Fish Bulletin 64,pp.39-72.

ROCKWELL, ROBIlRT H.1916. Long Island sharks for the Museum. Brooklyn

Museum Quart.erly, vol. 3, No.4, pp. 160-167.ROSENBI,ATT, RICHARD H., and WAYNE J. BALDWIN.

1958. A review of the eastern Pacific sharks of thegenus CarcharhiJl.1t8, with a redescription of C.malapoen8i8 (Fowler) and California records ofC. remotus (Dumeril). California Fish and Game,vol. 44, No.2, pp. 137-159.

SCHWARTZ, :F. J.1958. The cub shark, Carcharhinl/8 lencas, taken at

Flag Pond, Chesapeake Bay, Maryland. Maryland. Tidewater News, vol. 14, No.4, pp. 14 and 16.

38 FISHERY BULLETIN OF THE FISH AND WILDLIFE SERVICE

SMITH, HOMER W.1936. The retent.ion and physiological role of urea.in

Ehlsmobranchii. Biological Reviews, vol. 11,pp. 49-81.

SPRINGER, STEWART.1938. Notes on the sharks of Florida. Proceedings

of t.he Florida Academy of Science, vol. 3, pp. 9-41.1940. The sex ratio and seasonal distribution of

some Florida sharks. Copeia, No.3, pp. 188-194.1946. A collection of fishes from the stomachs of

sharks taken off Salerno, Florida. Copeia, No.3,pp.174-175.

1949. An outline for a Trinidad shark fishery. Pro­ceedings of the Gulf and Caribbean Fisheries Insti­tute, 2d annual session, pp. 17-26.

1950. A revision of Nort.ll American sharks allied tothe genus Carcha.rhin1ts. American Museum Novi­tat.es, No. 1451, pp. 1-13.

1951. The effect of the availability of sharks on ashark fishery. Proceedings of the Gulf and Carib­bean Fisheries Institute, 4th annual session, pp.140-145.

1954. Laboratory experiments with shark repel­lents. Proceedings of the Gulf and CaribbeanFisheries Institute, 7th annual session, pp. 159-163.

1957.. Some observations on the behavior of schoolsof fishes in the Gulf of Mexico and adjacent waters.Ecology, vol. 38, No. I, pp. 166-171.

SPRINGER, STEWART, and PRICE M. FRENCH.1944. Vitamin A in shark liver oils. Industrial and

Engineering Chemistry, vol. 36, p. 190.

o

SPRISSLER, G. PAUL.1942. An investigation of the proteinase of the gas­

tric mucosa of shark. Catholic University of Amer­ica, Contributions from Biological Laboratory, No.42, pp. 1-52.

STRASBURG, DONALD W.1958. Distribution, abundance, and habits of pelagic

sharks in the Central Pacific Ocean. U.S. Fish andWildlife Service, Fishery Bulletin 138, vol. 58, pp.335-361.

TEMPLEMAN, WILFRED.1944. The life-history of the spiny dogfish (Squallls

acanthias) and the vitamin A values of dogfish liveroil. Newfoundland Department of Natural Re­sources Research BulleUn No. 15 (Fisheries), pp.1-102.

THORNE, EDWIN.1916. Occurrence of ground sharks, Carcharhinus, in

Great South Bay. Copeia, No. 35, pp. 71-96.TORTONESE, ENRICO.

1938. Revisione degli Squali del Museo Civico diMilano. Atti della Societa Italiana di ScienzeN aturali, vol. 77, pp. 283-318.

1951. Studi sui Plagiostomi V, U1teriori considerazi­oni sulle specie Mediterranee dei generi Sphyrna eCarcharhin1ts. Doriana, Supplemento agli Annalidel Museo Civico di Storia Naturale "G. Doria",vol. I, No. 20, pp. 1-8.

1956. Fauna d'Italia, vol. 2, Leptocardia, Ciclosto­mata, Selachii. Edizioni Calderini, Bologna, pp.1-332.

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