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Proceedings of

the United States
National Museu m
SMITHSONIAN INSTITUTION

.

WASHINGTON,

D.C.

A REVISION OF THE HAMMERHEAD SHARKS
(FAMILY SPHYRNIDAE)
By Carter

R. Gilbert

Introduction

The hammerhead sharks, so named because of the unusual lateral
expansion of the head, comprise a moderately large family of sharks
occurring in tropical and temperate waters throughout the world.
They are a rather old group; fossil remains have been found in Eocene
deposits from Vu-ginia (Shelton P. Applegate, pers. comm.).
The
family belongs to the relatively speciose and poorly known
suborder
Galeoidea, in which are found most of the large and all of the

dangerous

kinds of sharks.

Since the

hammerheads

are well defined, and, with

one exception, adequate study material exists for
form an ideal taxon for systematic study.

all

the species, they

The most recent works of a revisionary nature that have appeared
on the hammerheads (Tortonese, 1950,
pp. 1-39; Fraser-Brunner,
1950, pp. 213-219) list eleven and ten species, respectively; however,
the above authors were unable to examme examples of all
the species
treated or

all

the extant

clusions were reached that
'

Florida State

types.

Consequently, a number of conto be erroneous.
The following

now appear

Museum, University

of Florida, Gainesville.

PROCEEDINGS OF THE NATIONAL

MUSEUM

a comparison of the systematic arrangements of the family Sphyrby Tortonese and Fraser-Brunner, with a classifi-

is

nidae, as presented

cation as determined from the present study:
Fraser-Branaer, 1950

Tortonese, 1950

Proposed

Subgenus Eusphyra Gill, 1861
Eusphyra
1. Sphyrna blochii (Cuvier,

blochii (Cuvier

Sphyrna

blochii

(Cuvier, 1817)

1817)

1817)

Subgenus Sphyrna Rafinesque, 1810
2.

Sphyrna mokarran

Sphyrna mokarran

(Ruppell, 1835)

(Ruppell, 1835)

Sphyrna tudes
(Valenciennes, 1822)
3.

Sphyrna zygaena

Sphyrna zygaena

(Linnaeus, 1758)

(Linnaeus, 1758)
4.

Sphyrna lewini (Griffith
and Smith, 1834)

Sphyrna lewini
(Griffith, 1834)

Sphyrna oceanica
(Garman, 1913)
Sphyrna diplana

Sphyrna tudes
(Valenciennes, 1822)

Sphyrna

ligo,

new

species

Sphyrna zygaena
(Linnaeus, 1758)

Sphyrna lewini
(Griffith, 1834)

Sphyrna oceanica
(Garman, 1913)

Springer, 1941
5.

Sphyrna couardi
Cadenat, 1951
Subgenus Platysqualus

6.

Sphyrna

Swainson, 1839

Sphyrna

tiburo tiburo

Sphyrna

tiburo vespertina

Springer, 1940

Springer, 1940

Sphyrna corona

9.

Sphyrna tudes

Sphyrna corona

Springer, 1940

(Valenciennes, 1822)

Another nominal
described

tiburo

Sphyrna media

Sphyrna media

8.

Sphyrna

(Linnaeus, 1758)

Springer, 1940
7.

tiburo

(Linnaeus, 1758)

(Linnaeus, 1758)

tiburo

(Linnaeus, 1758)

Sphyrna

tiburo

(Linnaeus, 1758)

Sphyrna media
Springer, 1940

Sphyrna corona

Springer, 1940

Springer, 1940

Sphyrna bigelowi

Sphyrna bigelowi

Springer, 1944

Springer, 1944

species,

by Sadowsky

Sphyrna

(1965),

Sphyrna nana, which was recently
is

regarded as a synonym of S. media,

for reasons discussed elsewhere in this paper.

above species, with
have not seen an entire specimen
of this species, although I have examined a head; however, the morphology of the head and chondrocranium, together with the patterns
of the Ampullae of Lorenzini, indicate that S. couardi is a valid form.
A number of structures have been studied that have proved to be

Adequate study material

exists for all of the

the exception of Sphyrna couardi.

I

All species can
usefid in the delineation of the species of Sphyrna.
be distinguished on the basis of head shape alone. Head shape ranges
from evenly rounded and spadelike (Sphyrna tiburo tiburo) to extremely wide and narrow {Sphyrna blochii). Some forms have an

HAMMERHEAD SHARKS

NO. 3539

GILBERT

3

indentation on the median anterior margin of the head, while in
others this indentation

is

lacking.

The presence

or absence of an

outer narial and/or inner narial groove and the relative length of the
latter help to distinguish

even the smallest individuals of a particular

species.

The various chondrocrania

also are distinctive.
Shape of both
and posterior parts of the preorbital process, degree of
development of the "wmgs" on the olfactory cartilage, development

the anterior

"wings," shape of the rostral cartilage, presence or
absence of accessory rostral cartilages, presence or absence of a
rostral fenestra, and shape of the anterior fontanelle all differ from
of the rostral

species to species.

The arrangement
surface of the head

arrangement

of the
is

Ampullae

of Lorenzini

on the ventral

relatively constant for each species, with the

of those in the anterior-median area being particularly

These ampullae, which are long canals

filled with mucus,
comprise a highly modified part of the lateral-line system, and it has
been suggested that they may be involved in the detection of temperature changes, hydrodynamic pressure changes, tactile stimulation,
and/or electric gradients (Young, 1962, p. 172). For brevity, these
structures are referred to as mucal pores throughout this paper.
Teeth vary according to species, although in several instances the

diagnostic.

differences are rather slight.
as well as the presence or

Shape and arrangement

of the teeth,

absence of serrations, are distinctive.

There are differences between the species in size and shape of the
and in the positions of these structures relative to each other.
The positions of the foiu"th and fifth gill slits in relation to the insertion
of the pectoral fin and the relative lengths of the gill slits are important.
fins

Finally, the various species differ in the presence or absence of a

lower precaudal pit and the shape of the upper precaudal pit.
Recent studies by Drs. Victor G. Springer and J. A. F. Garrick
(1964) on carcharhinid sharks indicate that the natiu"e and nimiber of
the vertebrae can be an extremely useful taxonomic tool, often showing
differences not evident

fortunately,

from the study

of external

morphology.

Un-

the value of vertebral characters was not recognized

while my study was in progress, and consequently only a few counts
were made for the species of Sphyrna.
Other characters that are probably of taxonomic importance in

hammerheads

morphology of the pelvic claspers and the
serum proteins. The latter were not
studied because of the unavailability of fresh specimens and lack of
facilities.
Because of the relatively small size and immaturity of
the available specimens, claspers of only three species, Sphyrna tudes,
S. media, and S. tiburo, were examined.
are the

electrophoretic patterns of the

and Mrs. WilKam C. Robert H. Weitzman. University of California. Drs. such data probably woidd not alter the present taxonomic conclusions. S. U. that are not evident in the embryos and young. Stanford. Bohlke. who supervised the project. In the case of S. The decision reached concerning the status of Sphyrna mokarran. L.C. Ofiice of Naval Research (ONR no. Dr. National Museum. Hubbs. Harvard University. the late Mrs. Robert Wisner. Washington. Calif. Daniel M. therefore. Drs. Applegate. 1354(09)). D. Giles W. does not rest on as firm a basis as for S.. Carl L. Washington. Walker and Mr. Field Museum of Natural History. lU.S. with the exception of the pelvic claspers. U. The aid and cooperation of the following people have been invaluable during the coiu-se of this work: Dr. Garrick. U. Ernest A. Fish and Wildlife Service. couardi was represented only by a head. France. and Miss Pearl Sonoda. Stanley H.C. Academy of Natm-al Sciences of Philadelphia. Museum National d'Histoire Naturelle. Springer. . Mead and Henry B. Marion Grey. A. Bauchot. Drs. D. tudes. Dr. Paris. Dr.4 PROCEEDINGS OF THE NATIONAL IVIUSEUM vol. Collette. Dr. Woods. Loren P.S. however. zygaena because far fewer specimens of mokarran were seen and the geographic coverage of the Only one specimen of Sphyrna last two species was much better.. for most of the information on dental variation in Sphyrna lewini and for other information on shark classification. James E. Schroeder. know of no taxonomically significant characters. National Museum. and was carried out while the author was a research associate of the U. Stewart Springer. I Atlantic area is Imown by me to be present in collections. Mr. and Richard H. Los Angeles County Museum. Shelton P. and. and Mr.S. Bigelow. U. . WiUiam Ralph Taylor. Acknowledgments This study was supported by contracts between the Smithsonian Institution. tudes from the Mediterranean was examined. Although it would be desirable to know the full extent of the various morphometric changes occiu-ring during growth. Victor G. Boyd W. Rosenblatt. Myvanwy Dick. Mr. Schidtz. F. as stated previously. Kanazawa. Chicago. Leonard P. The degree of confidence in the taxonomic conclusions reached in this paper is dependent largely upon the number of specimens available and on the extent of geographic coverage. for informing me of a specimen of Sphyrna couardi he had collected. J. 119 Another problem encountered was the scarcity of large specimens. U.. Fish and Wildlife Service. Mr. and the Atomic Energy Commission.S. Bruce B.S.S. Dr. Madame M. Cohen. Fish and Wddlife Service. Museum of Comparative Zoology. for loan of the type specimens of Zygaena tudes. National Museum. lewini and S. Scripps Institution of Oceanography. Lachner. only one other specimen of this species outside of the western however.

and Mr. Dr. Mr. M. Schultz. William I. the late Miss Margaret H. University of Michigan. Craig Phillips. David K. Frank Williams. Harvard University museum (ANSP) Academy . respectively. Genoa. Dr. University of Michigan. 1957. Various external morphological characters are illustrated and labelled in figures 1 and 2.. University of California at Los Angeles. Head-pore and chondrocranial terminologies (figs. University of Florida. Benazzi and Mr. equally to the hammerhead sharks. 29-40). University of Pisa (Italy) (UP). University of Florida (UF).. Guinean Trawling Survey. Dorothea B. University of Miami. 562-568). Proportional measm-ements for aU specimens were made according to the methods described by Springer (1964. Materials and Methods Specimens examined or recorded are from the following collections: British Academy of Natiu-al Sciences of Philadelphia Museiun (Natural History) (BMNH) (CAS). John C. Field Museum Chicago Natiual History Museum) of Sciences of California . University of Washington. Dr. Frankfurt-on-Main. 2. FoUett and Mrs. Storey. 59) and Tortonese (1950a. pp. Prof. Welander and Kelshaw Bonham. Nigeria. Measurements are expressed in thousandths of total length (TL). Prof. HAMMERHEAD SHARKS NO. Director. West Germany. Museo Civicio de Storia Natm-ale. Bailey and Robert Rush Miller. Institute of Marine Science (UMML). Museum (MCZ) Museum parative Zoology. California Academy of Sciences. Drs. Senckenberg Museum. Earl S. Dr. Paul Laessle. University of South Florida. Herald and Mr. University of California. pp. of Com- National d'Histou-e Naturelle (Paris) (MNHN) Scripps Institution of Oceanography (SIO). Natural History (formerly (FMNH). M . Lagos. Stanford University (SU). most of the characters apply . 3) are adapted from Daniel (1922. Baldwin. who are responsible staff for the illustrations and graphs. Museum of Zoology (UMMZ). 9). Wayne artist of the Department of Zoology. Los Angeles County Museum. Enrico Tortonese. Steinhart Aquarium. p. Wolfgang Klausewitz. While the specimen on w^hich figure 1 is based is not a member of the family Sphyrnidae. Arthur D. Briggs. San Francisco. Mrs. I woidd like to express my special appreciation to Mr. Calif. Los Angeles (UCLA). Italy. Caldwell. 3539 —GILBERT 5 J. Lillian Dempster. Italy. the authorities of the British Museum of Natural History. Robert Dempster. United States National Museum (USNM). All radiographs of the chondrocrania were made on type Kodak Industrial film with a "hard ray" machine (Miller. and to Finally. Mr. Reeve M. Drs. p. Florida State Museum. Pisa. Stanford University. University of Pisa Museum. Alberto Lanfranchi.

however. NEARLy SYMMETRICAL CAUDAL OF HEAD —External features of a shark with indications body. pp. 119 Additional proportional measurements. 17). This difference has been expressed as a ratio by dividing the rostral cartilage length by the distance betw^een the tips of the preorbital cartilages (fig. ventral view of head and comparison of caudal from Garrick and Schultz. media is rostral cartilage length. Phillips. since they give no information not already expressed by the measure- ments in the tables. These have not been included.PROCEEDINGS OF THE NATIONAL 6 MUSEUM vol. 5. 1963.) indicated in plates 7c. lateral view (Drawings by Craig fins. other than those found in the tables. were also taken. one of the main chondrocranial features separating Sphyrna corona and S. As SIDE b. 6. DISTANCE BETWEEN NOSTRILS NOSTRILS OR NARES VENTRAL Figure of 1. for FIN measurement: a. Values were obtained by measuring the distances . d.

1950a. showing important morphological features and mucal pore patterns (terminology of mucal pores adapted from Tortonese. morphometric changes re- sulting from growth. no diagnosis or description is given under the account of the species. media may also be distinguished by The values have been expressed as a difference in snout length. if the nomenclatm'al history of a species is of sufficient is complexity. p. in most of the the head examined this species tables. This is subject to variation. additional discussion is presented elsewhere in the species account. however. in no discussion of morphological variation. ANTERIOR-LATERAL PORES NARIAL OPENING INNER NARIAL GROOVE SUBNASAL PORES SUBOCULAR PORES OUTER NARIAL GROOVE ORBIT Figure 2. In addition. 3539 GILBERT 7 X (rostral cartilage length) and Y between the points marked by (distance between tips of preorbital cartilage) in figure 3. S. Also. these are compared in figure 16. 9. new combinations of names. More complete synonymies. drawing by Paul Laessle). Since a whole specimen of Sphyrna couardi was not available and was seen after most of the figures were prepared. Information regarding life histories. and zoogeography are usually included arate sections rather than in the individual species accounts. based on references for the I . corona and S.HAMMERHEAD SHARKS NO. it may be assumed that no noteworthy variation was discovered. or in figures 4 and 22. —Ventral side of head. determination of the patterns of the Ampullae of Lorenzini on the ventral surface of the head becomes progressively easier with an increase in size of the individual. New names. and information pertaining to original descriptions and type material have been included in the synonymies. there in sepIf. even within the same series of specimens. ratio by dividing snout length by head width. has not been included in the species key. In general. •MEDIAN INDENTATION LATERAL INDENTATION ANTERIOR-MEDIAN PORES. a species account.

POP OW (postorbital process). been accorded family status by most of the investigators responsible for the major works on . Localities the distribution IOC POP Figure 3. OC (posterior mediorbital connection). may MUSEUM be found in Bigelow and Schroeder (1948). as much as practicable. In cases where paucity of specimens does not permit an adequate comparison. RF (rostral (Drawing by Paul Laessle). (anterior mediorbital connection). (rostral cartilages). Classification The hammerhead sharks have. fenestra). (accessory rostral cartilage). PRC PMC RN (terminology adapted from Daniel. (olfactory wing). In the tabidation of proportions values have been separated. (preorbital process). RW values have been combined. —Diagram of hammerhead chondrocranium 1922. and a comparison with material from the western Atlantic was desirable. IOC AMC PRP (rostral node). from which specimens were examined are represented on maps by solid symbols. (preorbital cartilage). (rostral wing). and if the specimens do not appear to be different from those from other areas. This specimen is the holotype. 59): AF (anterior fontanelle). exception to this was made in the case of the single specimen of Sphyrna tudes from the Mediterranean. p. in the past. by geographic area. An RC ARC (olfactory cartilage). (innerorbital cartilage).PROCEEDINGS OF THE NATIONAL 8 western Atlantic region. literatm-e records are represented by open symbols.

(6) an asterospondylic type of vertebrae. differ from carcharhinid sharks in the following ways: (1) The vertebrae. . 1923. their value in determining interspecific relationships has been subject to question. and (2) in the teeth the lower blade has a smooth. and (9) an ovo viviparous type of reproduction. Accordmg to White (1937. 403-412.. G. While the pelvic claspers obviously constitute a valuable taxonomic character for the separation of various action apparently species. in cross section. Shelton P. Springer and J. also occurs in the triakids (V. which has the appearance of a "maltese cross" when viewed in cross section. p. the nictimembrane.) have regarded this group as a subfamily of the Carcharhinidae. Jordan. Berg's was based primarily on the studies by White (1937). ms. pp." Final judgment should be reserved until a more intensive study is made of claspers] most or all the species. Recent unpublished studies by Dr. pp. Little was said about the relationships of the sphyrnids and carcharhinids. some of which are not intimately related to these two groups.. Norman.). Although White (1937) retained the Sphyrnidae as a distinct family. show a convexity of the uncalcified canal of the neural and haemal arches that is "butterfly-like" in shape. Both have (1) the first dorsal fin anterior to the pelvic fins. as it is in the carcharhinids tating Garrick. The sphyrnids and carcharhinids share a large number of morphological characters. In addition. symmetrical arc. Applegate indicate that the hammerheads. I regard these differences as sufficient justification for continued recognition of the hammerheads as a distinct family. in Htt. F. A. she gave no convincing reasons supporting this decision. in addition to the pronounced lateral expansion of the head. Berg. while others (Regan. (5) teeth few in number. however. rather than straight. (8) three rows of valves on the conus arteriosus. (2) no naso-oral groove. 94). pattern of the vertebrae is as well developed in the family Triakidae and sphyrnids. 1861. so inconsistently that they are of little value in establishing relationships. 1906. (3) the keels of the dermal and dentate on the posterior edge and situated on a denticles complete flat surface.HAMMERHEAD SHARKS NO. 3539 GILBERT 9 fish classification (Gill. The and carcharhinids have so many basic most of these featm-es are foimd also in other famihes. never multicuspid ate. which has been thought to be unique to the latter two families. The "maltese cross" fact that the sphja-nids characters in common suggests a close relationship. (4) a nictitating membrane. 1940). "They [the tend to vary within the order [Galea] . iinpubl. (7) a "scroll type" spiral valve. 722-758. and the only places where they were compared were in the key and in the discussion of the pelvic claspers. whereas in carcharhinid genera there is a definite break in the arc on the side away from the crown.

1810. Bull. 125. 1828. Ann.) Zygoena Risso. (Substituted for Sphyrna Rafinesque. 16. 119 Genus Sphyrna Rafinesque Walbaum. of Cornell University. (Apparently a typographical error. 403. by subsequent designation of Gill. 1861. U. for Lepidoptera. Cestrorhinus Blainville. p. 1815. dermal denticles not closely overlapping. 403. Squalus tiburo Linnaeus.) (Type species. by original designation. in the past. p. p. in addition to the ^^'idely expanded head. 318. Geol. p. and thus should be viewed with caution.) Reniceps Gill. (Type species. Nat. 1861. 1861. labial furrow either small or absent. 1883. pp. Linnaeus. 26. 1839. substituted for Sphyrna Rafinesque. figs. Zygaena blochii Cuvier. 93. 1817. p. 1826.) Platysqualus Swainson.) Sphyra van der Hoeven.) Sphyrnias Rafinesque. (Type species. (Type species. 77. 408. (Type species. (Emended spelling for Sphyrna Rafinesque. pp. m which serum proteins were analyzed using electrophoretic techniques. (Type species.) — Phylogeny. Bull. Squalus tiburo Linnaeus. body deeply compressed pectoral fin short. p. Squalus zygaena 1792. Lyceum Nat.) Sphyrnichthys Thienemann.) Eusphyra Gill. Surv. vol 4.PROCEEDINGS OF THE NATIONAL 10 MUSEUM vol. New Jersey. Squalus zygaena Linnaeus. The most important of these are: No middorsal ridge on the posterior part of the back. 1 and 2). 1817. (Name p. Of the above characters. 1775. NewYork. on birds) has shown that the . p. Squalus zygaena Linnaeus. Sibley.) Zygana Swainson. as result of International Commission rulings of 1907 and 1910. by original designation. by subsequent designation of Fowler. Cestracion Klein in p. (Emended spelling for Zygaena Cuvier. the last two may have evolved in direct response to the lateral widening of the head. 318. basic morphometric characters. 412. Mus. Squalus zygaena Linnaeus. 580. (Type species. 1816. 60.) Sphyrna Rafinesque. 1839. by absolute tautonymy. name inadmissable. p. 1855. Work done by various investigators (the most important and extensive being that of C. which indicate derivation from a single common ancestor. 1911. by subsequent designation of Jordan and Gilbert. 68. 412. 403. teeth moderately to strongly oblique. pp. and are thus regarded as fimdamental evidence of a close relationship among the species of Sphyrna.S. G. pp. by original designation. 46. 1810. vol 7. 127. p. should be spelled Zygaena. genus name preoccupied by Zygaena Fabricius. There has. p. 292-295. Hist. The remaining characters are not likely to have been affected by such a change. Perhaps the most convincing study supporting a monophyletic derivation has resulted from the work by Starr and Fosberg (1957.) Zygaena Cuvier. been some question as to whether the genus Sphyrna has had a nionophyletic or polyphyletic Members of the genus are characterized by certain apparently origin. . 121. 1810. 1810.

and two of Aprionodon isodon. first. analyses Avere made of the serum proteins from one specimen of Sphyrna levnni. 7 a. On the other hand. The fifth species. The respective lengths of the anal fin and anal fin base. the patterns in Sphyrna levnni and S. tihuro showed a marked similarity. which is the most divergent species of hammerhead and the one most likely to have evolved independently. an outer narial groove. AU possess a number of common closer hammerheads morphological features. the presence of total number of vertebrae. the various characters most closely approach those found in the abcrve species is S. The extreme expansion some of . In aU likelihood. and several chondrocranial characters all are similar in these fom* species. blochii are similar to those of other hammer- heads. although he has not yet studied the vertebrae of this species. In addition. of the head. the relative positions of the orbits and anterior part of the mouth.). the family Sphyrnidae has had a monophyletic origin. the greatly decreased and the marked enlargement of the verte- . Sphyrna tudes. the appearance of the upper precaudal pit. 4). Applegate (in litt. these four apparently closely related species are grouped together in the subgenus Platysqualus. and pis. 7 c. all apparently reaching a maximum size of about 5 feet. the three species figs. This assumption is based. and Sphyrna media show a phylogenetic relationship than any other species of (fig. is strikingly different from all other species of Sphyrna in several characters. corona. 4. d and 8). 14-22. The patterns of the latter two species. the species are comparable in size. share several other characters which. Within the subgenus Platysqnalus. 3539 H GILBERT patterns resulting from tliese analyses may be valuable indicators of higher relationships in the various groups of vertebrate animals. which are not found in other members of the genus (table 1. differed notably from each other and from the patterns in the two species of Sphyrna. two specimens of Rhizoprionodon terraenovae. figs. Sphyrna media seems to show a slightly closer relationship to S. the diameter of the orbit in relation to the distance from the anterior edge of the orbit to the outer margin of the narial opening. the presence of a lower precaudal pit. 7 a-d). however. Sphyrna blochii. as well as similarities in appearance of the anterior-median pore patch (figs. It woidd be desirable to rim similar tests on Sphyrna blochii. Sphyrna corona. fiu-ther demonstrate a close relationship. tihuro (table 1. thus suggesting a close phylogenetic relationship. on the greater overall similarity of the chondrocrania (pis.HAMMERHEAD SHARKS NO.21. In the paper by Starr and Fosberg. both of which are members of the family Carcharhinidae. informs Sphyrna me tihuro. tihuro than does S. two of that the teeth in S. the relative lengths of the gill slits. and pis. 22e-h). and 22. b) as a result. although not confined to them The form in which alone. 11. Finally.

nothing was said regarding head width. In the preceding discussion. S. Eusphyra. a deeply falcate anal fin. tudes. blochii have. and S. blochii. media. the only lewini Despite the difficulties encountered in interpreting the relationships above foiu. This suggests that they probably are more closely related to each other than to S. S. S. mokarran is miique. S. S. such featui-es in as a a short anal fin greater than the pectoral base. base. the length of which is never more than slightly S. a long slender anterior part of the preorbital process of the chondrocranium. mokarran are the only members of the genus that ordinarily have a rostral fenestra. lewini. couardi) and S'. among the hammerheads. corona. couardi. they appear to be related more closely to one another than to any other group of species of Sphyrna and thus are of the included in the subgenus Sphyrna. and S. S. mokarran: The overall shape of the rostral node. although this Tortonese (1950a. a rather tall second dorsal fin. mokarran and S. Although I have followed both of these allocations (in modified form). indicating S. table 1. zygaena. S. lewini (including S. For example. with a definite ridge on the anterior margin. and S'. a blunt. pp. 119 brae situated over the posterior part of the coelom are all unique to this species. more than twice that attained by any other members of the genus. S. mokarran are more difficult. as do S. tudes. mokarran has. and a deep. in having a deeply falcate pelvic fin. tiburo. zygaena and S. 4.species. The following examples. feet. blochii is so different from the other hammerheads that recognition of Eusphyra as a genus would not be unwarranted. zygaena and probably S. The main featiu-e a fairly close interrelationship among these species is All reach a total length of at least 12 similarity in maximum size. 3-6) placed Sphyrna tiburo in the subgenus Platysqualus and placed Sphyrna blochii in a separate genus. S. however. and S.PROCEEDINGS OF THE NATIONAL 12 MUSEUM vol. most mokarran are hammerheads lacking a wing on the olfactory cartilage. and S. Contrary to the relative ease with which the relationships of the preceding species are discerned. those of Sphyrna zygaena. Eusphyra. and figs. lewini (and common. is almost the only basis for previous separations of the genus Sphyrna into separate genera and subgenera. and it is consequently placed in a separate subgenus. widely V-shaped upper precaudal pit. . S. couardi) have certain low second dorsal fin with a long lobe. as do S. illustrate the difficulties one encounters in attempting to determine the relationships of *S'. tiburo. 6-10). clublike knob at the end of the anterior part of the preorbital process of the chondrocranium. S. and anterior part of the preorbital process in the chondrocraniiun of Sphyrna zygaena are similar to those found in S. mokarran (fig. corona. anterior-median pore patch. media. S.

couardi. an outer narial groove. becoming gradually straighter with increase in size and age until sometimes. Those head characters of greatest importance in distinguishing species of sphyrnids. It has long been thought that such changes in head shape would make hammerhead taxonomy a difficult problem. corona. —-Certain modifications in shape of the head of sphyi-nids occur with increase in size and age. likely evolved from a single common ancestor. S. media. a reduced number of vertebrae. this margin is nearly straight (figs. Sphyrna blochii possesses several characters that are markedly different from those found in other hammerheads. Sphyrna tihuro. and S. 3. Such is not the case. lewini. although elevation of this taxon to the generic level might be justified. on the basis of morphological evidence and electrophoretic analyses of serum proteins. particularly embryos. and S. 3539 the differences in head width have my —GILBERT had comparatively 13 httle to do with decision. such as structure of the chondrocranium. the anterior margin of the head is more or less broadly rounded. and Sphyrna media. With an increase in size of the individual. coelom. Sphyrna corona. S. for which there is nothing comparable in other carcharhinid sharks. 56).HAMMERHEAD SHARKS NO. In young hammerheads. which has the most widely expanded head of any member of the genus. mokarran are more difficult to determine. The most important feature shared by the four species is their large maximum size. 4. with numerous identical morphological characters. lewini. S. Eusphyra. these structures become situated nearly perpendicular to the body (fig. are regarded as the most closely related members of the genus Sphyrna. because of their close relationship. the lateral extensions of the head in embryonic and early postembryonic specimens are nearly parallel to the body axis (fig. 2. and pattern of the mucal . nature of the intranarial and extranarial grooves. Sphyrna tucles. 7a and 20c). In Sphyrna blochii. They are referred to the sub- genus Sphyrna. such as an extremely expanded head. the four species are placed in the subgenus Platysqualus. The interrelationships of Sphyrna zygaena. and couardi seem to show a closer relationship to each other than to mokarran. The phylogenetic conclusions may be summarized as follows: 1. as in Sphyrna mokarran and Sjphyma tudes. The nine species of Sphyrna. tucles. some of which are unique. Growth changes. with a nmnber of morphological characters similar to those in S. although zygaena. 5c). It is 5. Therefore. appears to occupy a phylogenetic position intermediate between these three species and other members of the genus Sphyrna. and enlarged vertebrae over the posterior part of the placed in a separate subgenus.

in figures 21/. measuring 356 and 900 mm. with an mcrease in total body length.000 from specimens of Sjjhyrna media likely reaches its final is illustrated. Negative allometric growth occurs in the head of sphyrnids since. g. closely related to S. pp. (S. appear to change very Httle throughout life. Since most size. and this was not included. —Hypothetical phylogeny of the genus Sphyrna couardi. Other morphometric changes are indicated with an increase in size (tables 2-9) although more large specimens must be examined before the validity of these changes can be established. however. changing with an increase in undergoes the greatest modification during size of the individual. tip of snout to orbit. 546-548). and intranarial width. diameter of orbit. is (figure by Paul Laessle). whereas the shape of the fins may be modified with an increase in body size. The shape of the upper precaudal pit.PROCEEDINGS OF THE NATIONAL MUSEUM 14 pores on the ventral surface of the head. long. occur in squaloid sharks (Garrick. lezvini. Other postcephalic features seem to remain fairly constant. 1960. there is a proportional decrease in the following characters: Width of head. which early growth mm. considerable changes in body proportions can occur with a relatively small increase in total squaloids do not reach a very large . particularly in length of the trunk between the axil of the pectoral fin and the insertion of the pelvic fin. tip of snout to symphysis of jaws. TL. have been shown to . Pronounced modifications. HYPOTHETICAL PHYLOGENY OF THE GENUS SPHYRNA GARGHARHINID ANCESTOR Figure 4.) form before the shark is 1. tip of snout to nares.

showing an increased number of "teeth" on the posterior margin. this does not result from changes in the original denticle. No detailed discussion of hammerhead ecology is presented here. couardi) apparently occur both in inshore and and have even been known to move into brackish seems to prefer shallow embayments and rarely ventures far from land. since most of the available information has appeared in works by Springer (1939. tiburo. pp. although the reason for (particularly the first dorsal) this is not clear. and. although I have not yet these extremities are black. a straightening of the fins and fin lobes body length. in litt. of the remaining three species. Bigelow and Schroeder (1948. In this process the fins become increasingly more erect and pointed. since there verified captures is doubtful if have been no below a hundred fathoms (Stewart Springer. pp. It species descends to very great depths. 407-449). 3539 GILBERT 15 The galeoid sharks. crabs. lesser degree for S.). In Sphyrna lewini there is a progressive darkening of the tips of the pectoral fins with increasing size and age. zygaena. is a summary of the in- The larger species of Sphyrna (S. on the other hand. mokarran. Hammerheads swim near any the surface of the water. and squid and also may take such items as shrimp. whereas the anal fin tends to become somewhat more falcate. 188194). occurs as the shark approaches maturity. 96-152). in contrast. but instead occurs as the result of one or more denticle replacements. In contrast to these true morphometric changes are tlie unfolding of the fins after bu-th. S. and sharks. pp. S. The following formation appearing in their papers. The fish larger hammerheads are regarded as danger- . As indicated previously. As indicated by Bigelow and Schroeder (1957. 1940b. Little is known about the habitat offshore waters areas. S. 24). much larger size. S. 9-41.HAMMERHEAD SHARKS NO. usually reach a and thus proportional changes are not likely to be so pronounced within a comparable size range. and Baughman and Springer (1950. presumably. The dermal denticles undergo a noticeable morphological change with an increase in size and age. a phenomenon that might be compared with the situation in a newly emerged butterfly. stingrays. among specimens of the same species is discussed under the accounts of the individual species. mokarran. p. so that in large specimens This also may be true to a greater or zygaena and S. substantitated Variation it. lewini. — Life history and ecology. however. Curling of the fins is most pronounced in embryos of Sphyrna mokarran. pp. including individuals of Sphyrnids feed largely on their own species.

tiburo. media. Presumably similar migrations occur in other Oils. tiburo one another. S. to the south. the largest (38) in S. partial Insufficient evidence exists to explain the present distribution of S. — The present distribution of the hammerhead sharks indicates. and *S'. from Miami. The smallest recorded brood (8) was found in S. tiburo is to it possible to estimate Only how long it has taken in the case for evolution have occurred. although it is impossible tades. S. of S. tiburo tiburo. couardi. The fact that the only specimens of Sphyrna zygaena seen by me from south of the Carolinas (UMML 5639 and 10444. 418). S. S.) were taken in winter seems to confirm this. 417. and thus pre- S. The number of young in a brood appears to be directly proportional both to size of the individual and to maximum size of the species. Whether this is due to a partial ecological separation or simply to lack of material is presently not known. UMML parts of the world. often traveling in schools in both inshore and offshore waters (Bigelow and Schroeder. mokarran. media and S. Although S. corona have been taken in the same areas from Panama Bay north to southern Mexico (map 5). Possibly the initial separation goes farther back and the intermediate examples have resulted from a reamalgamation of the stocks. corona. pp. . 442-444). Hammerheads along the eastern North American coast are known to migrate northward in summer and southward in winter. lewini. a slow rate of evolution. 119 and there are substantiated records of fatal attacks on humans (Gudger. pp. lewini is the predominant species. S. 1937. S. corona. the presence of individuals of this species with a head shape more or less intermediate between typical tiburo and vespertina indicates that the above explanation may not be as simple as first appears. and S. Mexico) were the two actually taken together. zygaena. because the two subspecies probably differentiated following the most recent (Pliocene) emergence of the Central Ameri- can isthmus. and S. To the north. on the whole. The members of the subgenus Sphyrna (S. mokarran) apparently have undergone no intraspeciiic differentiation. However. in only one of the collections examined (from Chiapas. Zoogeography. tildes. Evolution of these three species is almost certainly correlated Avith the various submergences and emergences of the Central American land mass. whereas. media. vespertina) are morphologically similar to sumably have undergone a more recent evolution. despite worldwide distributions (for three of the four species) and long periods of separation of the various segments of their respective populations. 1948. Fla. The five forms included within the subgenus Platysqualus (S.PROCEEDINGS OF THE NATIONAL 16 MUSEUM vol. Sphyrna zygaena is the only species involved in such migrations.

outer narial groove present. Sphyrna zygaena (Ekman. 1953. later followed by two species. Greatest maximum width of head not more than 32 percent of total body length. anterior.8 to 13.5 to 9. and body length. upper precaudal pit as in figure 21o. nares present near eyes. caudal pit not as in figure 21a. tudes evolved first.—GILBERT HAMMERHEAD SHARKS NO. It is impossible to determine the center of origin of the subgenus Sphyrna. chondrocranium as in plate 5. when equatorial waters were cooler than at present. . or to a long period of isolation during which the two populations rence have. . nares not present near eyes. differentiation of the other in few species display such a distribution pattern. a prehistoric body of water that formerly extended from the Eastern Pacific-West Indian region to what is now southeast Asia (Ekman. is restricted to that region. Key to Species of Sphyrna mm.5 percent. one or more species of the subgenus would likely still be found there.9 percent of total first gill slit. distribution of antitropical distribution a classic example of an is 250). chondrocranium not as in plate 5 2 Anal fin and base longer. p. pp. fifth gill slit about as both somewhat shorter than three middle slits. One may conclude from present distributions that the subgenus Platysqualus evolved in the western Tethys Sea. As the glaciers retreated and seas became warmer. or smaller) Greatest maximum width of head almost 50 percent of total body length.median notch. Sphyrna blochii (Cuvier) Indo-Pacific region. fifth gill slit situated above insertion of pectoral fin. zygaena gradually disappeared from the tropical regions. its horizontal diameter less than shortest distance from anterior edge of 219-937—67 2 latter from long as 6. The presence of S. S. for The some reason. Should it have evolved in the Indo-Pacific region. anterior-median pores on ventral surface of head as in figure 22a. failed to differentiate to the species level. lb. since environmental conditions in this area have remained stable over a long period of time. from Persian Gulf to northern Australia and southern Philippines. perhaps remaining for a time in deeper waters where temperatures were cooler (equatorial submergence). . orbit smaller. 1953. as the only included species.000 la. TL (Based primarily on specimens 1. 63-67). situated more than half distance from eyes to anterior. since three of the species in this group have a worldwide distribution.median pores on ventral surface of head not as in figure 22a. The subgenus Eusphyra apparently evolved in the Indo-Pacific. This common occurmay be due either to recent migration in one direction or the other. former measuring from 9. upper pre- 2a. . Presumably the now-disjunct north-south distribution of this species (map 2) was continuous during one or more of the Pleistocene glacial periods. Sphyrna hlochii. however. S. 3539 17 Judging from morpho- to determine the exact sequence of events. tudes both the Mediterranean and western Atlantic is unusual in that logical evidence. outer narial groove absent.

usually with a more or less distinct median notch (sometimes evenly rounded in aS. usually less than 40 percent of head width (table 7. 146. lewini) posterior lobe of first dorsal fin extending beyond insertion of pelvic fin (in S. media only) maximum size smaller. also recorded from western Mediterranean Sea. chondrocranium not as in plates 7a. usually coming to a point at tip of rostrum. greatest width of head measuring slightly more as shown in table 6. 156) anterior-median pores on ventral surface of head as in figure 22/. b anterior margin shown slightly less as 4 (fig. with a more pointed apex (figs. line drawn between posterior margins of orbits passing well anterior to symphysis of lower jaw. origin of first dorsal fin situated well posterior mouth. . greatest in table 6. posterior lobe of first dorsal fin terminating anterior to . . anal fin more deeply falcate. to axil of pectoral fin. 12a). 4a.. the largest specimens probably not over 4 or 5 feet TL . maximum size possibly larger. 126) . b. greatest width about 21 percent of total body length (usually less than 20 percent) anterior margin of head evenly rounded or slightly pointed in median line. eyes not situated distinctly anterior to line drawn between posterior margins of orbits either intersecting or passing just anterior to symphysis of lower jaw. 3 Head not widely expanded.. Distance from tip of snout to symphysis of upper jaw slightly shorter. no well-defined median notch present on anterior margin of head (poorly defined notch may be present in some individuals) anterior-median pores on ventral surface of head 6 not as in figure 22/i. 119 orbit to outer margin of narial opening (at least in smaller specimens). the least width 22 percent of total body length. posterior lobe of first dorsal fin terminating posterior to above or slightly posterior to axil of pectoral fin. 15a) mouth slightly broader (figs. well-defined median notch present on anterior margin of head. . fig. tudes. anterior-median pores on ventral surface of head as in figure 22h. origin of first dorsal fin situated . upper precaudal pit as in figure 21e. PROCEEDINGS OF THE NATIONAL MUSEUM 18 vol. Inner narial groove present. Head with less evenly rounded anterior margin (fig. anterior-median pores on ventral surface of head as in figure 22e. corona. and S. S. b 5 5a. . upper precaudal pit not as in figure 21e. fig. insertion of pelvic fin. Head widely expanded. teeth relatively weak. eyes situated anterior to mouth. chondrocranium as Head with more evenly rounded width of head measuring in plates 7a. Sphyrna tudes (Valenciennes) Gulf of Mexico south to Uruguay. . median ones often more or less needle-like in appearance. 3a. . chondrocranium not as in plates 8a. anterior-median pores on ventral surface of head not as in figure 22e. 14a. never with median notch. the largest specimen insertion of pelvic fin. media) . 4b. Sphyrna tiburo vespertina Springer Eastern Pacific. from New England to southern Brazil. anterior margin of head not always evenly rounded. . 3b. 13.. from southern California to Equador. chondrocranium as in plate 7c. b 6a. lower precaudal pit present (may also be present in S. Inner narial groove absent. 16). 5b. fig 13 Sphyrna tiburo tiburo (Linnaeus) Western Atlantic. chondrocranium as in plates 8a.

from Panama to Gulf of California. and. rostral fenestra ordinarily absent. greater in length than margin . maximum size larger. also in southern Caribbean and southwestern Atlantic.. teeth smooth in small individuals. from Panama to Brazil. the largest specimen . but apparently absent from the more chondrocranium as in plate . its horizontal diameter greater than shortest distance from anterior edge of orbit to outer margin of narial opening. not extending nearly to caudal pit. Inner narial groove absent. when extended vertically upward.o. Circumtropical in isolated oceanic islands. lower precaudal pit absent. inner narial groove extending at least halfway from narial opening to median part of anterior margin of head. 7b. pelvic fin falcate. maximum size possibly smaller. lobe of second dorsal fin shorter. fig. 16). . teeth broad. when extended vertically upward. posterior lobe of first dorsal fin not extending to . from Colombia to southern Mexico. even in small individuals. but absent from intervening tropical waters. above insertion of pelvic fin. with a more rounded apex (fig. 6a . and. Inner narial groove present. base of anal fin shorter. extending nearly to caudal pit. chondrocranium as in plate 6c. 18b) anterior-median pores on ventral surface of head as in figure 22(j.. pelvic fin not falcate. base of anal fin longer. anterior-median pores on ventral surface of head as in figure 22c. 18a). examined a little over 2 feet long Sphyrna corona Springer Eastern Pacific. orbit larger. Median notch absent from anterior margin of head. mouth slightly narrower (fig. fifth gill slit shorter than first gill slit (fourth gill slit about as long as first gill slit) fifth gill slit situated behind insertion of pectoral fin. chondrocranium as in plate 7d. definitely exceeding greatest height of fin. HAMMERHEAD SHARKS 3539 GILBERT 19 examined about dy> feet long Sphyrna media Springer Eastern Pacific. Sphyrna zygaena (Linnaeus) Antitropical in distribution. some individuals reaching 20 feet TL 7 7a.median pores on ventral surface of head not as in figure 226. about equal in length to base of second dorsal fin and slightly less than base of pectoral fin. teeth strongly serrate. length of its anterior margin greater than length of anterior margin of anal fin. Anal fin and base shorter. occurring in cooler waters of northern and southern hemispheres. . anal fin less deeply falcate. anterior. chondrocranium not as in plate 6a 8 8a. often weakly serrate in large individuals. anterior. usually more than 40 percent of head width (table 8. former measuring from 8.2 to 10 percent. second dorsal fin higher. 6b. and latter from 3.8 to 6. 8b. not needle-like in appearance.7 percent of total body length. 2b. lobe of second dorsal fin longer.median pores on ventral surface of head as in figure 226. about equal to greatest height of fin. length of its anterior margin less than length of anterior margin of anal fin. . inner narial groove not extending halfway from narial opening to median part of anterior of head. Median notch present on anterior margin of head. Sphyrna mokarran (Riippell) distribution. Distance from tip of snout to symphysis of upper jaw slightly longer. second dorsal fin lower.

119 of either second dorsal or pectoral fins. 10). and (according to the original description) in having the tips of the pectoral fins white. rostral fenestra present. pi. vol. vertebrae over distinguished posterior part of coelomic cavity expanded in size. Dignaosis of Family and Genus Distinguished primarily by the shape of the head. 117.) Zygaena blochii Cuvier. 1817. 117. and only about half as wide. j . and various chondrocranial characters (pi. 5). in having a deeper and more robust head (pi. orbit separated from nares by a distance many times diameter of orbit. common warm in all seas. 22a. pointed angle at outer-posterior corner and a more or less straight posterior edge (fig. which is expanded and which is accompanied by a corresponding modification laterally. lewini. 127. Eyes situated at tips of lateral expansions.2 percent of total body length. 5 1. however. (Original description [in footnote]: "Ajoutez I'espfece representee par Bl. 21a. Sphyrna (Eusphyra) blochii (Cuvier) Figures 5. Nob. Plates Squalus zygaena (misidentification) Bloch. reconnaissable k ses narines Sa deuxifeme dorsale est placees bien plus prfes du milieu [Z.0 to 49. and nares situated at varying distances along anterior margin of head. Blochii]. | . Olfactory capsules and orbital region widely expanded. dorsal fin always much smaller than Body compressed. there being from 15 to 16 fewer body than caudal vertebrae. 9) in minor differences in the chondrocranium (pi. anterior-median pores on ventral surface of head as in figure 22d. (Figure. body vertebrae 51 to 54. when pit present in some present. not examined. Subgenus Eusphyra Gill The subgenus Eusphyra. absent in others. 1. the patch characterized by a sharp. Caudal peduncle about two-thirds as wide as deep. but was not included here since a whole specimen was It diSers from S. first dorsal. upper precaudal pit merely a narrow longitudinal groove (fig. 22a). of the chondrocranium. 21a). Second of back posterior to first dorsal fin without a middorsal ridge. *Sphyrna covardi Cadenat keys out to this point.20 PROCEEDINGS OF THE NATIONAL MUSEUM vol. not as prominent as upper pit species. which includes only Sphyrna blochii. the depth of the trunk (measured at origin of first dorsal) from 4 to 5K times in distance from tip of snout to upper precaudal pit. 1785. p. is by the following unique characters: Head extremely expanded. total number of vertebrae few (117 and 124 in two specimens examined). a unique anteriormedian pore patch on underside of head. outer narial groove present. in having the anteriomedian pore patch completely instead of partially divided. Upper precaudal pit Lower precaudal strongly developed. and the three Midline rostal cartilages broadly expanded where joined anteriorly.* Sphyrna lewini (Griffith and Smith) Circum tropical in distribution. the greatest width from 42.

USNM 195846 69397 (1). Nomenclature. Krusadai Island. p. Malaya: BMNH 1902." specimen.19. new generic name. 1838. 315. Gulf of Mannar. blochi. For reasons that are discussed elsewhere in this paper. 1851. Sumatra. Burma: SU 14495 (1). (Originally written Latycephala. (Name. but in an even greater number of has been considered a subgenus of Sphyrna. (2). (Synonymy. 237. range. 1860. coast of Perak. —Although Sphyrna blochii. (Figure. p. Diagnosis. Eusphyra is the only genus which has been erected (Gill. based on adult specimen of S. pp. Singapore. In some of the most important recent works (Bigelow and Schroeder. Sandakan. MCZ 1389 mutilated head). (Original description. There has been some difference of opinion among recent investigators as to whether Eusphyra should be regarded as a good genus. (Tale Sap. 2. Cestracion blochii (Singapore. Characters mentioned in the diagnosis of the subgenus Eusphyra are not repeated here. 1948. 1924. 1950a) this has been followed. 412) GiU considered that two unique morphological features of blochii (the position of the nares.) Zygaena latycephala van Hasselt. p. (Reference. 1861. and the extreme elongation of the lateral extensions of the head) constituted sufficient reason for for the sole reception of this species.) Eusphyra blochii Gill. no type locality. p. p. blochii with other members of the genus is presented in table 1 classification cases it — — . p. sous le Description based on figure in Bloch.) Zygana laticeps Swainson.) Sphyrna blochi Pellegrin.) Sphyrna blochii Mliller and Henle. (Locality unknown. Description. South Island. 1841. Pacific Ocean: British North Borneo: SU 13815 (1).890 (1 (1). placing the species in a distinct genus. (Benkulen. Zygaena laticeps Cantor. 54.3. Java: FMNH 15655 (1). Bombay. (1). 403. p. p. UMMZ 177111 (6). p.) Zygaena blochi Volz. 134. BMNH ANSP (2).) Sphyrna {Eusphyra) blochii McCuUoch and Whitley. 1925. (Range in size: 333-880 mm.) Cestracion (Zygaena) blochii Dumeril. 1904.5. (Illustration of head. 315. 1823.) (Indian Ocean. 50. 1907. HAMMERHEAD SHARKS NO. 1912. 464. 450. I follow the latter alternative. description. Southern Moscos gi'oup. 1861. Singapore. Sphyrna zygaena (misidentification) Bridge. Siam. type locality. donnde. 1837.37 (1). no type de pantouflier. 383. Tortonese.) Cestracion blochi Hora. 129. FMNH 21836 (1). p. aussi bien plus prfes de la caudale. See diagnosis of subgenus Eusphyra. Calcutta. p. TL). FMNH 15656 Batavia. in common with the other hammerheads. generic name misspelling of Zygaena. Pumbau District.) — Specimens examined.) Sphyrnias blochii Gray. Meristic data appear in table 2.28. has been placed in several different genera. Madras. (India. 156-157.) Garman. Java. 1865.. A comparison of *S^. which are nearer the tip of the rostrum than the eyes. no further data. 412. 3539 —GILBERT 21 L'espfece k large tete. 1785. 1913. nom type locality. FMNH 40914 India: SU 41989 (1). pp. p.

extending about half of distance from narial opening to Sphyrna hlochii: a. outer posterior corner of lateral expansions of head situated posterior to corner of mouth in specimens of all sizes. 0. narial flap broad. head of juvenile. 622 mm. Figure 5. greatest transverse distance between corners of mouth about one-seventh of greatest width of head. length of snout about one-eighth of head width. blunt at tip. 333 mm. c. anterior margin of head broadly curved medially.2 X natural size (FMNH 21836). h. symphysis of jaws . dermal denticles (FMNH (Drawings 21836). with the outer edge curving gradually inward.) scallop. Schultz.6 X natural size (UMMZ 177111) d. 0. 21836). by Dorothea median B.— PROCEEDINGS OF THE NATIONAL 22 MUSEUM Intranarial distance from one-seventh to one-sixth of TL. from Batavia. inner narial groove present and distinct. Java. TL. North Borneo (FMNH head of adult. teeth series from left side of upper and lower jaws (FMNH 21836). e. the lateral expansions of the head pointing posteriorly in embryonic individuals and extending nearly perpendicularly from body in larger specimens. female. TL. from Sandakan. distinct median scallop present on anterior margin of head. narial depression present.

fifth gUl above slit above or slightly anterior to axil of pectoral drawn ventrally from origin usually inter- situated distinctly behind. absent. Origin of fin. pelvic fin about five-sLxths length of anal fin. accessory rostral cartilages absent. anterior fontanelle (viewed ventrally) . terminal part of caudal about twoninths of total length of fin. second dorsal fin base about fovir-sevenths length of anal base. its length about twice greatest height of fin. and fourth gill very slightly posterior to) insertion of pectoral fin. not perfectly erect. pectoral fin about threefifths as broad as long. first dorsal fin relatively slender. height of anal fin about two-sevenths again as much as height of second dorsal fin. labial furrows absent. and about equal to greatest height of first dorsal. posterior margin of lower caudal lobe nearly straight. Anterior margin of rostral node of chondrocranium straight. fu'st through fourth gill slits nearly equal in length. anterior margin of pectoral slightly convex. and all longer than fifth gill slit.HAMMERHEAD SHARKS NO. sloping posteriorly at about a 35° angle from a perpendicular to the body axis. upper margin of caudal slightly convex. 3639 situated posterior to a transverse line GILBERT drawn between 23 posterior margin of orbits. anal fin deeply falcate toward apex and nearly straight toward tip of lobe. first dorsal jEin a perpendicular line secting axil. the lower half falcate. pectoral fin base four-fifths to seven-eighths length of anal fin base. anterior margin of lower caudal lobe convex. lower caudal lobe not erect. lobe of second dorsal fin long. lower lobe of caudal moderately narrow and short. apex of pectoral pointed. terminal apex of caudal broadly pointed. distal margin of pectoral slightly falcate. posterior lobe of first dorsal fin terminating well anterior to insertion of pelvic fin. length of caudal fin about one-thu'd of TL. anterior margin of pectoral fin three-fourths to four-fifths length of head. base of first dorsal fin about three-fifths head length and almost twice length of pectoral fin base anterior margin of first dorsal fin nearly straight. lobe of first dorsal fin (free posterior margin) slightly less than one-half length of base. inner corner of pectoral narrowly rounded. and extending ahnost to upper precaudal pit. its greatest height five-eighths to five-sevenths as long as base. and about five-sevenths greatest lieight of anal fin. upper half of distal margin of first dorsal fin straight. posterior margin of terminal part of caudal fin shallowly falcate. origm of second dorsal fin situated over posterior third of anal base. slightly curved near apex. lower precaudal pit slit (or . second dorsal fin moderately tall. a line drawn perpendicularly from apex of fin intersecting free rear margin of fin (posterior lobe) about halfway along its length. length of lower caudal lobe one-third to two-fifths as long as upper lobe. anal fin base slightly longer than pectoral and pelvic bases and more than 1}^ times as long as second dorsal fin base.

119 appearing extremely broad and shallowly U-shaped. cusps visible on all teeth. feet. is of restricted to the hammerheads not found . as uddth. Maximum total length not recorded.) Sphyrna blochii specimens examined. hollow denticles (examined from upper part of back beneath first dorsal fin) not closely overlapping. with none (except the median tooth) becoming progressively more oblique toward corners of mouth. Individuals gray or grayish brown dorsally. denticle (measured to tip of median marginal "tooth") as great. pointed wings present on sides of rostral node. p. the blades steeply raised length of . fig. pomted. and those nearest the corners having cusps pointing almost straight back. teeth 6 through 10 in upper jaw notably larger than others. becoming more pale below. two species Hemisphere. and as many bluntly pointed teeth usually present on posterior margin of and teeth sometimes rudimentary or absent. pedicel short and broad. teeth 2 through 10 in lower jaw gradually in perfectly erect. or nearly as great. 12) eastward throughout the East Indies. It is L0ppenthin. no markings on fins. anterior wing of olfactory cartUage prominent. Range. and projecting medially. tooth 1 notably smaller than tooth 2 in lower jaw. A (Solid Dermal = O Sphyrna mokarran = confirmed literature references. teeth not serrated. posterior part of preorbital process terminating in an angular knob. five strong ridges usually present. median notch. mmM'^m Map 1. with a shallow. Teeth — (one specimen examined). broad. the outer ridges particularly in small individuals. although those near corner of mouth in lower jaw poorly developed. possibly not more than several denticle. and is one —Sphyrna blochii of the Indo-West Pacific area in the Western known from the Persian Gulf (Blegvad and 41. teeth both jaws notably oblique. median tooth on denticle ordinarily slightly longer than more distal teeth. the teeth increasing in size. 1944. small. — Distribution of species.PROCEEDINGS OF THE NATIONAL 24 MUSEUM vol. anterior part of preorbital process long and slender.

(General. identification probable because teeth described as "gezalmelte. p. individuals reaching a size of from 12 couardi. as evidenced by its absence from outlying oceanic islands (map 1). (Red Sea. . fifth gill slit situated distinctly behind. while perhaps not as closely restricted to continental waters as Sphyrna tlhvro. type specimen. 7. is distinguished by only two unique characters. teeth. 303. (Original description figures of head. 1841. 1908. (Identification probable because teeth serrate. Sphyrna (Sphyrna) motcarran Figures 6. These are: A large orbit. S. S. and a large maximiun size. mokarran. illustration of head. . 3539 —GILBERT 25 China and the southern Philippines (Herre.5 percent. (Identification by description of head and reference to Lacepede. Australia (Ogilby.) Sphyrna mokarran Miiller and Henle. Plates 2. and S. 235. the greatest horizontal diameter greater than distance from anterior margin of orbit to narial opening. of TL. 66-67. Senckenberg Museum no.) Zygaena tudes Valenciennes. types from Mediterranean Sea. 225-226. The species also has been recorded . a male 2510 mm. lewini. and body.000 mm. (In part. 3590. absence of an outer narial gi-oove. type locality. very rare.2 percent of total body length (in specimens less than 1. 226.0 percent. 6a ?Squalus tiburo (misidentification) Risso. (Riippell) 216. Agassiz. specific name tudes subsequently restricted to a species other than one here called Sphi/rna mokarran. 1835. and fourth gill slit above (or very slightly anterior to) insertion of pectoral fin. p.HAMMERHEAD SHARKS NO. zygaena. total number of vertebrae ranging from 174 to 206 (usually 190 or more). and all somewhat longer than fifth gill slit. pp. original description. Massaua. nevertheless appears to be incapable of traversing broad stretches of ocean.9 to 30. p. anal fin and anal fin base relatively short. and Coromandel. the former not over 11. Red Sea. 1838. 35. 408. 1822. and north to southern 1930. 142). type specimens apparently represent more than one species. pp. 1810. presence of a deep narial depression. p. 1828. pp.") Zygaena mokarran Riippell. TL. with those over posterior part of coelomic cavity not expanded in size. to 20 feet in total length. Sphyrna blochii. and the latter not over 7. lobe of first dorsal fin terminating w^ell anterior to insertion of pelvic fin. Subgenus Sphyrna Rafinesque The subgenus Sphyrna. the greatest width from 22. which includes S. first through fourth gill slits nearly equal in length. from Queensland.) . p. TL). The subgenus is also distinguished by: A moderately expanded head. Cayenne. 4) this area apparently represents both the eastern and southern limits of its range. "Squale pantouflier" Nice. 54.) Sphyrna malleus (misidentification) L.) Sphyrnichthys zygaena (misidentification) Thienemann.

23. an embryo. 81°47' W. New South Wales. 1851. 41. p. figure. 1890.. 44. pp. 435-675 USNM tograph seen). 1847. 383. 25°56' N. is readily identified by the shape of In addition. 1865. Beaufort. fig. pi. female. MCZ 35247 (1). type locality. radiograph of head seen.. USNM —Zygaena mokarran was described by Riippell The (1835. 1916. Baja California. p. (Original description. 499. p. (In part. New South Wales. 1887. Englewood. particularly fins. (Not seen. 119 Sphyrna tudcs Mtiller and Henle.C. Mexico: 29645 (1). Cuba. Cape Lookout. 215. 293). Pacific Ocean: Australia: 40026 (1). p. p. p. Clarence River. Goree. (Original description. 108454 (1). in illustration as serrate. 111.) BMNH Specimens examined. p. p. identification p. the conformation of the the head. 1822. (In part. at Massaua. (Figure of S. UCLA 52-252 (1). N. 1822.9. p. 1872. MCZ 969 (1). Captiva Pass. (In part. 1890. 2).) Sphyrna lewini (misidentification in part) Fowler. Bahia Almejas. 40014 (1).C. 3590) has been illustrated recently by Klausewitz (1960. 115. (In part. identification by reference to Valen- ciennes. TL. 1896. identification from reference to Valencennes. (Identification specimens 12 ft. 6 in.) Sphyrnias tudes Gray. (Identification by shape of head in photo. — (Range mm. (Name.) Zygaena dissimilis Mm-ray. p. Kurrachee. 231 (x-ray of head of holotype of Sphyrna ligo). heads Atlantic Ocean: Senegal: (pho- in size: of larger specimens also examined). 17. Isla Santa Margarita. SU 14330 (1). 103.) Cestracion (Zygaena) mokarran Dum^ril.) Platysqualus tudes Howell-Rivero. 217. 66-67.) Cestracion mokarran Bleeker. Australia. Clarence River. p. 1868.) Squalus zygaena (misidentification) de la Blanchere. long. N. (Identification probable because of large size. 37162 (1). by reference to Valenciennes.. 10. the type specimen (Senckenberg Museum no. Richmond River.) ISphijrna chiereghini Nardo.) Sphyrna ligo Fraser-Brunner.") Cestracion tudes Ninni. p. 63092 (1). mokarran incorrectly labelled S.) Cestracion zygaena by description (misidentification) of teeth Radcliffe. p.) Sphyrna zygaena (misidentification in part) Coles. 12 to 14 ft. 1822. Mazatlan.) Sphyrnias mokarran Gray. . pp. India: 12622 (1). Madras.) Cestracion (Zygaena) tudes Dumeril. 3) from the Red Sea. 1865. 263. 50.23. (Teeth de- scribed as "dentelees. 50. (Name only. 384. 10 in. West Indies and Mediterranean. p. type locality.PROCEEDINGS OF THE NATIONAL MUSEUM 26 vol. illustration (pi. 53. holotype. 186087 (2). of 13 ft. references to specimens from Gulf of California and Indian Ocean perhaps do not pertain to this species. 530. 213-219.) Sphyrna (Platysquabis) tudes Jordan and Evermann. 1936. 1919. lewini. (Chinese drawing. 1873. 1851. USNM FMNH FMNH USNM USNM USNM USNM BMNH USNM USNM Nomenclature.9. 1950. though inaccurate in certain details. p. 1941.231. 108453 (1). (Compiled. 1841.. 1822. New South Wales. United States (Florida): 106543 (1). description from Valenciennes.

S. known Figure Sphyrna mokarran: 6. Fla. one can assume that the Coromandel individual was probably Sphyrna mokarran. mokarran. TL. juvenile male. dermal denticles. one of which may be S. neither of which However. a tall second dorsal fin. the length of the lobe equal to height of fin and extending only about one-half of distance from posterior part of fin base to upper sizes. from Englewood. b. about 49 X p. c. this structure represented by a thin line. mokarran. French Guiana). head. since the missing specimen is from an is S. pp. India) where the species represented by the two existing types does not occur. and since there is a fairly close superficial resemblance between the two species in question. (USNM (USNM 106543).) — Diagnosis. (From Bigelow and Schroeder. Tortonese (1950b. 673 X (USNM 106543). (scallop) . Characters mentioned in the diagnosis of the subgenus Sphyrna are not repeated here. absence of an inner narial groove. a short second dorsal lobe. the height of the fin slightly greater than height of anal fin.2 a. area (Coromandel. 1030-1033) was the first to point out that the three type specimens of Zygaena tudes actually comprise two species. the second from Cayenne. Sphyrna mokarran is unique among members of the genus Sphyrna in having a deeply falcate pelvic fin and strongly serrated teeth at all mokarran is also characterized by a deep median indentation on anterior margin of head. mm.— HAMMERHEAD SHARKS — GILBERT 27 For many years the species herein called Sphyrna mokarran was as Sphyrna tudes. 430. Only two of the types are apparently now extant (one from Nice. 0. France. 106543). 1948.

head of small adult. mm. b./. from Englewood. 1948. 1.. teeth series from left side of upper and lower 108453). long). p. shallow scal- lops present on anterior margin of head about halfway between nares and tip of snout. 216). about 1. Sphyrna mokarran: 0. narial flap broad. eleventh lower tooth. presence of a rostral fenestra. A comparison of S. outer posterior corner of lateral posterior to corner of mouth expansion of head in smaller specimens. mokarran are not repeated here. .9 to 27. X. — Meristic data appear in table Characters men- 3. a shallow upper precaudal pit. broadly pointed at tip. Fla.000 mm. TL. tioned in the diagnoses of the subgenus Sphyrna and S.) Head moderately expanded. Figure 7.5 d. 6a). c. 226). 1. the horizontal diameter almost equal to length of shortest (fifth) gill slit.— PROCEEDINGS OF THE NATIONAL MUSEUM 28 precaudal pit. anterior to corner of mouth in larger individuals. lower tooth. percent of TL (in specimens the greatest width from 22. about natural size. fifth natural size (USNM a.09 X jaws (USNM e. mokarran wdth other members of the genus is presented in table Description. fifth (c-f.) twelfth upper tooth.3 less than 1. an anterior-median pore patch in which the outer-posterior corners are sharply angled and the posterior edge is straight (fig.660 108453). upper tooth. (From Bigelow and Schroeder. with the outer edge ciu"ving abruptly inward. 429. orbit large. with an acutely pointed posterior ridge to upper precaudal pit (fig. and several other chondrocranial characters (pi.

pelvic fin base slightly less than length of anal base. 3539 GILBERT 29 length of snout from one-third to one-fourth of head width. sloping posteriorly at about a 45° angle from a perpendicular to body axis. and second dorsal fin bases. where it is broadly curved. posterior margin of terminal part of caudal fin notably falcate. origin of second dorsal fin situated slightly behind origin of anal fin. apex of pectoral broadly pointed. first dorsal fin tall. accessory rostral ventrally) shallow cartilages absent. terminal part of caudal about onefourth of total length of fin. greatest transverse distance between corners of mouth about one-foiu-th of gi-eatest width verse Une of head. anal fin deeply falcate toward apex. symphysis drawn between of jaws situated posterior to a trans- posterior margins of orbits. lower caudal lobe not erect. anterior margin of pectoral longer than one-half length of head and a vertical height of first dorsal fin. posterior margin of lower caudal lobe nearly straight or slightly concave. a line of fin lobe. anterior margin of pelvic nearly straight. the lower one-third distinctly falcate. second dorsal fin base slightly greater than foiu-of first dorsal fin . length of anal fin slightly greater than length of pelvic fin. base of first dorsal fin slightly less than one-half head length and almost twice length of pectoral fin base. anterior margin broadly cm'ved (excessively cm-ved in embryonic and early post-embryonic individuals) upper two-thirds of distal margin of first dorsal fin straight or slightly falcate toward tip. anal fin base slightly (up to one-sixth) longer than pectoral. or slightly behind. and pointing strongly posteridrawn perpendicularly from apex of fin passing well back this point. length of lower caudal lobe one-third to two-fifths as long as upper lobe. . orly. Anterior margin of rostral node of chondrocranium straight. inner corner of pectoral broadly rounded. a perpendicular origin intersecting at. anterior fontanelle (viewed and nearly straight on posterior margin. anterior . fifths length of anal fin base. length of caudal fin nearly a third of TL. labial fuiTows absent. prominent. pectoral fin base about four-fifths length of anal fin base. lobe of fu'st dorsal fin about onethird length of base. pectoral fin fin slightly than greatest two-thirds as broad as little less long. margin of pectoral broadly rounded (strongly curved in embryos and early post-embryos) distal margin of pectoral broadly falcate. Origin of line first dorsal fin above axil of pectoral drawn ventrally from fin. slender. upper margin of caudal slightly convex. terminal apex of caudal broadly pointed. pelvic. except near apex.HARIMERHEAD SHARKS NO. nearly straight toward tip of lobe. anterior margin of lower caudal lobe convex. lower lobe of caudal narrow and long. length of pelvic fin almost equal to length of anal fin. the distance from angle included by the upper and lower caudal lobes to tip of latter going about 23^ times in anterior margin of lower caudal lobe.

no markings on fins. Sphyrna mokarran is circum tropical in distribution (map However. and this. a species of occupying a similar range. Individuals with gray or grayish brown dorsally. Maximum total length reputedly about 20 feet. — lewini. 16 or 17-1 to 3-16 or 17 upper teeth triangidar. on expanded f & ^^ > but the subsequent upper teeth increasingly oblique toward corners of mouth. as well as the fact that neither of the other heads of worldwide distribution {S. lewini) show any differentiation. small specimens with from three to five. together with the paucity of specimens. but with skin often visible in between. the blades thin and little arched length of denticle (measured to tip of median marginal "tooth") about same as width. larger specimens with five or even six. it does not usually appear to be as common as Sphyrna 1). Range. In addition. it seems best to recognize Sphyrna mokarran. orbital process terminating in Teeth . the first tooth erect. For bases. outermost teeth in both jaws with distinct cusps. becoming more pale below. symmetrical. hammer- zygaena and S. and of these a large percentage (including all individuals from the Indo-Pacific region) were very small embryos.— Comparatively few specimens of Sphyrna mokarran were available for study. between specimens from the Atlantic and Pacific regions (table 3). S. Total number of vertebrae 205 to 206 (in two specimens examined) body vertebrae 95 to 98. radiographs of the chondrocrania show no apparent differences. and as many short. no anterior wing on olfactory cartilage. these reasons. there being from 10 to 11 fewer body than caudal vertebrae. bluntly rounded wings present on sides of rostral node. including the Gulf Mexico and Caribbean Sea. posterior part of pre- an evenly rounded knob. smooth-topped ridges.. u9 broad. PROCEEDINGS OF THE NATIONAL 30 MUSEUM vol. Variation. median tooth on denticle slightly longer than more distal teeth pedicel short and thick. such as orbital diameter and length of second dorsal fin base. tip of anterior part of preorbital process very short. noticeable intraspecific but one species. . and rather thick. broadly pointed marginal teeth on dermal denticle. one or two series of teeth functional in alternating rows along sides of upper jaw and usually two rows along sides of lower jaw. . tends to negate apparent differences. blunt. second to tenth or eleventh teeth the largest in both upper and lower jaws. Specimens of this species are characterized by strongly curled bodies at this stage of development. mokarran is found in the western Atlantic. from North Carolina south to Brazil. . Dermal denticles (examined from upper part of back beneath first dorsal fin) evenly and closely spaced.

1925. as certain its from other species. p. lewini.) ISphyrna {Zygaena) zygaena Imms. 21c.) p. skull and . 1861. pers. identification by references. Provincetown. in the northern Indian Ocean. 121. 403. Sphyrna (Sphyrna) zygaena (Linnaeus) Figures 8. 1810. type presumably Cape Cod. p. 510. 1817. 129. 3539 It is —GILBERT 31 present in the eastern Atlantic. specimens from Mediterranean may apply in part to S. 1862.) malleus Valenciennes. ISphyrna (Sphyrna) zygaena McCulloch and Whitley. p. p. 1758. 1788. 36). 59. 1686. Springer. (Reference. ("French" coasts. Mediterranean. p. "Europe. (South Africa. lewini. identification by references. 1855. 70. Australia.) ICestracyon zygaena Poey. (At least in part. p. p. 377. only . 9. 234. identification by reference to Linnaeus. 1822. evidently based on Zygaena (Refers primarily to Sphyrna lewini. 46. no type specimen. type locality. 1950a.) Sphyrna (Cestracion) zygaena von Bonde. 621.) Sphyrna malleus van der Hoeven. (Identification by included references. 1848. p.") Squalus zygena Bonnaterre.) Cestracion zygaena Gill. 1891. such as the carcharhinid shark Rhizoprionodon acutus (Victor G. (Name only. p.) (Identified by description of 2 ft. p. 121.) ICestrorhinus pidus Blainville.) Zygaena Zygaena Zygaena Zygaena zygaena Cuvier. in MCZ Sphyrnias zygaena Gray. (May refer entirely or in part to Sphyrna lewini. but also includes lewini [misspelled "lewisii"]. (In part. 1951. 127. p. 1758. 9. may name refer entirely or in part to S.) subarcuata Storer. speci- [not located]. identionly.) vulgaris Cloquet. refer in part to S. Brazil. 96-97) presence in the last two areas. Sicily. may p.) Sphyrna zygaena Rafinesque.) ICestrorhinus caroliniensis Blainville. p. only. 1905.) Cestracion subarcuata GiU. there are no substantiated records the intervening Mediterranean (Tortonese. 43. 1848.) p. (General. p. out the East Indies and westward. 1816. 262. 1816. comm. nerves. 22c. mokarran is found throughthe more eastern parts of Oceania. (Name only. (Name subarcuata Storer." "America. 1933. 348. at least off the northwest coast and in the Red Sea. (Name (General. Plate 6c Squalus zygaena Linnaeus. zygaena.) IZygoena malleus Sauvage. 48. although it east Africa. fication by reference to Linnaeus.). identification by photo of head. p. general. (Pharyngeal denticles. to It occurs in America. 223. p. 1851. 1830. (By reference to illustration by Willughby. type locality. show a similar hiatus in distribution. Sphyrna mokarran of California south is present in the eastern Pacific from the Gulf to Panama and probably northwestern South the Pacific from southern Japan south to appears to be absent from New Zealand and S. pp.HAMMERHEAD SHARKS NO. Despite of Africa (Cadenat. This is probably not due to lack of adequate collecting. p. 1881. men. (Atlantic and Mediterranean.

(New Jersey): 93845 (3). Long Island. b. of head.7 third lower tooth. Monroe Co. (in Gulf Stream). {c-f. Alligator Reef. 39992 (1).^:i£:i£S^^3. c. about 3. ninth lower tooth. (Massachusetts): USNM 10418 (1).) — Specimens examined. (2). (MCZ X. (1). 687 upper and lower teeth.8 X. second upper tooth. d. Rio de 604 (1). Seaside Park. USNM 33268 (1). juvenile female. UMML iAi^dM^^y:h^^^i. 115903 (1). (Range in size: 457-2. /. Japan: 23383 (stuffed skin). 4°13' S. MCZ 36425 (1). (no 5639 (head). MUSEUM (Portugal. MCZ 35364-35365 Livorno. Italy: AICZ 451 USNM 28452 (1). AtOcean (including Mediterranean Sea): Azores: USNM 94502 lantic (1). no further data. 437. Holly Beach. Bigelow and Schroeder. MCZ 1159 (1). Norfolk. from Nahant.. TL). land): UMMZ USNM USNM USNM USNM USNM USNM USNM | . MCZ 383 (1). 425. Barnegat Bay. Buzzards Bay. left side. 1948. (Virginia): USNM 119699 6090 (1). off North Miami. Woods Hole. Pacific Ocean: Australia: 29020 (head). 71779 (1). Sandy Hook.) (From p. MCZ 1416 (1).^ Figure Sphyrna zygaena: 8. Sydney (Port Jackson). Mass. about mm. MCZ 136 (2). 1159).— PROCEEDINGS OF THE NATIONAL 32 Zygaena zigaena Nobre. Galapagos Islands. Janeiro. 1935. Graves End Bay. Cape Cod. ANSP 522 (1). Dade Co. (1). off Virginia Beach. USNM 83980 (1). 81°13' W. TL. Brazil: Terceira. Nahant.) USNM USNM USNM 76681-76683 (3). (New York): CAS 19189 (1). 120495 (11). further data). a. Stuart's Pond. off ANSP UMML United States (Florida): 10444 (1). Gulf of Guayaquil. identification by description p. upper tooth.. Buz(Maryzards Bay. Woods Hole.663 mm. e. ninth 1.. Tokyo market. 116447 (5). Ocean City. USNM 31527 (1). Ecuador: 127758 (head).

and several other chondrocranial characters (pi. Diagnosis. a slightly more deeply pit. Nagasaki. Peru: USNM 77711 (head). Yenoshima. being separated by a distance equal to about half of greatest horizontal diameter of eye.. percent of TL (in 219-937—67—3 specimens the greatest width from 26. Suruga Bay. Misaki. I. MCZ 421 (1). HAMMERHEAD SHARKS NO. vicinity of Choshi. which extends at least 50 percent of distance from inner margin of nares to tip of snout. lewini. Magdalena Bay. SU 13003 (1). the length of which is slightly less than length of pectoral base. Wakanoura. the deepest part of the notch on the distal margin drawn perpendicularly from the posterior axil of the fin. off La Jolla. SIO 58-382 (jaws). Payta. San Diego County.1 to 29. 24125 (1). long). Head moderately expanded. a shorter anal base. LACM uncataloged (jaws and chondrocranium). northern Kyushu.000 mm. at most. the patch characterized by a broad. the length of the former about twice greatest height of fin. pointed posterior extension in the FAINH UMMZ UMMZ UMMZ USNM UMMZ UMML — median -posterior area S. 51291 (head). 179080 (1). orbit closer to nares. 190592 (10). zygaena — are not repeated here. Characters mentioned in the diagnoses of the subgenus Sphyrna and of S. United States (CaHfornia): SIO 58-224. Meristic data appear in table 4. shallow . about two-thirds of distance from second dorsal base to pit. upper precaudal tip of second dorsal lobe not reachmg as near to the lobe extending. Lobos de Tierra. Hakata Bay. "off Peru". no fiu-ther data. oft' San Diego. North Island. Cedros New Zealand: imcataloged (2). Gulf of California. 179079 (1). AICZ 1042 (1). Panama: MCZ 515 (1). falcate anal fin. Sphyrna zygaena is characterized (as is S. lewini is (scallop) on anterior margin by lack of a median of head. no further data. its closest relative) by a long second dorsal fin lobe and a low second dorsal fin. 21c). absence of a rostral fenestra. SIO 58-29 (1). 57423 (1). MCZ 441 (3). The anteriormedian pore patch on the imderside of the head is unique. Bay of Islands. Characters mentioned in the diagnosis of the subgenus Sphyrna are not repeated here. A comparison of Sphyrna zygaena with other members of the genus Sphyrna is presented in table 1 Description. 3539 GILBERT 33 USNM USNM 51289 (head). smoothly rounded angle at the outerposterior corner and a broad. a longer inner narial groove. 61032 (1).0 less than 1. Mexico: southern end of Angel de la Guarda Island. Island. Osaka market. distinguished from S. MCZ 382 (2). 6c). 22c). a more acutely pointed situated slightly posterior to a line posterior ridge to the upper precaudal pit (fig. SU 179078 (1). absence of a lower precaudal pit. zygaena indentation (fig.

e.) b. illustrated in figure 8 second dorsal and anal fifth lower tooth. 1948. Sphyrna zygaena: ventral view. p. fins.— PROCEEDINGS OF THE NATIONAL MUSEUM 34 ^^f-^^^B b Figure 9. head of specimen natural size. ^.45 X a. about 0. about 9 X.) (From . (MCZ 1159). Bigelow and Schroeder. about 45 X. 438. fourth upper tooth. c. {d-e. dermal denticles.

height of anal fin about a third again as much as height of second dorsal. . the horizontal diameter about equal to length of longest (second) gill slit. second dorsal fin base from three-fifths to four-fifths length of anal fin base. a line drawn perpendicularly from apex of fin passing close behind base. posterior margin of terminal part of caudal fin moderately falcate. . terminal part of caudal about one-fifth of total length of fin. distal margin of first dorsal fin strongly falcate. with the outer edge curving evenly inward. about halfway between nares and tip of snout. length of lower caudal lobe one-third to two-fifths as long as upper lobe. origin of second dorsal fin situated about halfway back above base of anal fin. greatest transverse distance be- tween corners of mouth from one-fifth to one-foui'th of greatest width symphysis of jaws situated posterior to a transverse line drawn between posterior margins of orbits in specimens of all sizes. upper margin of caudal fin nearly straight. lower caudal fairly erect. Origin of fii'st dorsal fin above posterior axil of pectoral fin. lobe of first dorsal fin about one-thii'd length of base. with the part nearest the apex very strongly curved. anterior margin of pectoral moderately convex. length of anal fin equal to or slightly greater than length of pelvic fin. the distal margin nearly straight. pectoral fin about two-thirds as broad as long. and from three-fourths again to nearly twice length of pectoral fin base. orbit large. . apex and inner corner of pectoral rounded length of pelvic fin about foiu--fif ths length of anal fin. length of snout from onefifth to one-fourth of head width.. first dorsal fin broad. moderately pointed at tip. length of caudal fin nearly a third of TL. HAMMERHEAD SHARKS NO. rather erect (not as erect as in large examples of Sphyrna tudes). distal margin of pectoral slightly falcate. sloping posteriorly at about a 35° angle from a of head. the distance from angle included by upper and lower caudal lobes to tip of latter going about twice in anterior margin of lower caudal lobe. a perpendicular line drawn ventrally from origin intersecting axil. anterior margin of first dorsal fin evenly cm-ved. about one-fifth as long as greatest horizontal diameter of eye. labial fiu-row present at corners of lower jaw only (concealed when mouth is closed). terminal apex of caudal narrowly rounded lower lobe of caudal relatively narrow and long. length of pelvic fin base approximately equal to length of anal fin base. anterior margin of pectoral fin from one-half to two-thirds length of head (higher values usually apply to larger individuals) and about equal to or slightly greater than greatest vertical height of first dorsal fin. probably in specimens of all sizes narial flap broad. base of first dorsal fin from two-fifths to ninetwentieths of length of head. 3539 scallops present GILBERT 35 on anterior margin of bead. anterior margin of pelvic fin weakly convex. outer posterior corner of lateral expansion of head posterior to corner of mouth.

Total number of vertebrae 196 (in one specimen examined). there being six more body than caudal vertebrae. as width. sharp-toothed ridges. represented only of preorbital process long and by a small "bump". though well-developed cusps. median tooth on denticle slightly longer than more distal teeth. posterior margin of lower caudal lobe nearly straight. m • thi-ee. comm. PROCEEDINGS OF THE NATIONAL MUSEUM 36 vol. median indentation. and from Argentina and Uruguay to (probably) northern Brazil (exact northern limits of range in South American waters uncertain). body vertebrae 101. Dermal denticles (examined from upper part of back beneath first dorsal fin) closely overlapping. 13 to 15-0 i — large serrate teeth weakly Teeth . an antitropical distribution.^ ^ m-^ 12 to 14-1-12 to 14 dividuals. though smaller. shading to white ventrally. their outer margins deeply incised and with the outermost teeth with very small. anterior fontanelle (viewed ventrally) rather deep and evenly rounded. length of denticle (measured to tip of median marginal "tooth") as great. small specimens usually with — . very shallow cartilages absent. It is present in the western Atlantic from Nova Scotia and northern New England south to (occasionally) southern Florida (UMML 5639 and 10444). anterior part and without a sharp angle preorbital process terminating in an slender. Maximum probably between 12 and 13 feet.. one small symmetrical tooth at symphysis of lower jaw. their inner margins straight or evenly convex. ^ . . anterior margin of lower caudal lobe weakly convex. or 1-13 to 15. first four teeth in lower jaw rather erect and with narrow cusps.. lower teeth similar to upper. and from three to five pointed marginal teeth present on denticle.— . with no noticeable median notch. occurring in cooler waters of both the Northern and Southern Hemispheres but apparently absent from the intervening warmer areas (map 2). 119 perpendicular to the body axis. . first upper tooth nearly symmetrical and erect. . accessory rostral or with a broad. rounded wings present on sides of rostral node. Stewart Springer (pers. and two to three series functional toward center. two to three series functional in lower jaw. pedicel very short.) states that he Range. short. posterior part of evenly rounded knob. one to two series of teeth functional along sides of upper jaw. tip of ventral sm'face of pectoral fin sometimes faintly gray black in life. none or one in upper jaw. . . . Individuals deep olive to brownish gray dorsally. large specimens with up to seven. total length —Sphyrna zygaena has UMML . on anterior edge. . or nearly as great. Anterior margin of rostral node of chondrocranium nearly straight. subsequent upper teeth strongly oblique. anterior wing of olfactory cartilage absent. the blades thin and moderately arched.

south coast of New Holland. it is likely that the species does occur there. (In part. 22d.) Zygaena malleus Valenciennes. 1816. 47) may not have been based on this however. and Smith) 6b. 89. (Name only. 1947. species. 46. p. but is present off New Zealand and southern Australia. however. (Name only. and the Galapagos Islands) and (one record) Panama. (Griffith Plates 3. 48. zygaena farther south along the coast of Africa (Irvine. 1823. (Type locality: Java. 90) and from South Africa (Smith 1953. It seems to be absent from the Hawaiian Islands and the East Indian area. I ffiTF ^Htaffi ^SfeaSy Map (Solid 2. malleus included in synonymy. 121. pp.) ICestrorhinus pictus Blainville. p. 10b caroliniensis Blainville. Van Hasselt. and from southern Cahfornia and Mexico (south to the Gulf of Cahfornia). illustration of 1 Cestrorhinus head.) "^Zygaena indica Zygaena lewini dom. Mediterranean Sea and Atlantic Ocean. Peru. specimen. 1951. in Cuvier. 640. p. 121.) Sphyrnias zygaena (misidentification in part) Gray. 1851. 50. zygaena is common around southern Japan. 1822. 1834. — Distribution of ^^ Sphyrna zygaena. 9b. figure.HAMMERHEAD SHARKS knows of no authentic record for this species or Caribbean. 46). and Smith. zygaena.) no type (In part. pi. which literature records applying to there. . given. In the northwestern Pacific S. teeth described as denticulate. type locality. p.) and Smith. pp.) In the eastern Pacific Sphyrna zygaena is known from western South America (Chile. 315. in northwest Africa (Cadenat. Sphyrna {Sphyrna) lewini Figures 10. hollow = confirmed literature references. description. In the eastern Atlantic Sphyrna zygaena is known from Scotland south to Senegal. 223. is GILBERT common all 37 from the Gulf of Mexico Sphyrna lewini. Records of S. p. able for S. 21d. Animal King- Griffith 10. (Original description. p. but mtensive collecting has failed to reveal its presence between the above two areas. and also from the Mediterranean Sea. since in neither case were adequate descriptions and the hammerheads pictured represent species other than Considering that the cool waters off South Africa are suitzygaena. = specimens examined. although the exact northern Umits of its range there are uncertain. vol. It probably is also present in the southern parts of the Indian Ocean as well. 1816. p. Griffith.

lantic UMML Sui-inam: (2 heads). (1). 666. 383.) (Description. p. pp. no fm-ther data. Society Islands.) zygoena (probable misidentification) Chevey. (Possibly in part. "Europe": MCZ 673 (1). Messurado River beach. 46-52. 54°47' to 54°51' W. (chondrocranium). 1913. type locality. (Possibly in part. (May refer ISphyrna {Zygaena) zygaena (possible misidentification) Imms. may refer entirely or in part to Sphyrna zygaena. Sphyrna mokarran. Mexico: MCZ 39646 (1).) Cestracion oceanica Carman. Specimens examined. MCZ 470 (1). 94. East Indies. SU 52717-52719 (3). p. (Pharyngeal denticles. (Moreton Bay. off west end Grand Bahamas: further data. 271. Englewood. Rio de Janeu-o. AtOcean: "Atlantic": MCZ 393 (1). name only. 1932. Italy: UP uncataloged (1). 129. type locality. 06°20' N. Mediterranean area. SU 52874 Cuba: USNM 111350 (1). Fla.) Cestracion {Zygaena) leeuwenii Dumeril. zygaena. "West Indies": SU 11830 (1). p. . 261. (Reference. SU 11696 (1). (Moreton p. Bay. head and fins). Campeche. MCZ 314 (1). Senegambia.) p. pp. no 10867 (1). 1865. Malabar. name only. no further data. p. 263. Queensland. 1925. Jamaica: USNM 29999 (1). 43.) Cestracion lewini Ogilby.) p. 158-159. p. 129. entirely or in part to Sphyrna zygaena.) Cestracion Sphyrna diplana Springer. 1871. 1941. off Ciudad Carmen. Florianopolis. SU 14046 (1). 348.) ISphyrna tudes (possible misidentification) Ogilby. West Africa. 1891. Bahamas Island. 1916. only. USNM 79285 (1). 1858. 6. 1881. Gorr6e. pp. 49937 USNM USNM 159197 159196 (1). 81. 1865. USNM 179705-179706 (2).PROCEEDINGS OF THE NATIONAL 38 MUSEUM vol. 23..) IZygoena malleus (possible misidentification) Sauvage.) Sphyrna {Sphyrna) lewini McCuUoch and Whitley. from C zygaena. refers primarily to Sphyrna lewini. Santos. specimens from Mediterranean may apply in part to S. 119 Sphyrna lewini Dumeiil.) tSphyrna zygaena (possible misidentification) Studer. Sphyrna oceanica Fowler. (Cochin (Original description. China. Port Jackson. p.) Zygaena leuwinii Ramsay..755 mm. 576. — (Range in size: 386-1. p.) (Sphyrna) (Possibly in part. 510.) 1925. TL). Brazil: MCZ 462 (4). Australia. zygaena (misidentification) Steindachner. Guiana. p. Rio Grande do Sul. 1908.) ISphyrna (Sphyrna) zygaena (possible misidentification) McCuUoch and Whitley. Leeuwen" ICestracion ("Cotes de la terre de [Australia]. 9° 34' N. no fiu-ther data.) (Porto Rico. 1881. 1870. p. p. no further Panama: USNM 79282 (1). no further data. Guinea: USNM uncataloged (1. (Copied. p. misidentification probable because of locality. (Name in synonymy. Livorno.. distinguished Cestracion leeuwenii Day. 1889. 1905. USNM 06°23' to 159199 (1). p. 1928. (May refer in part to Sphyrna lewini.) name Zygaena erythraea (Ehrenberg) Klunzinger. ANSP 49936data. Colon. Cestracyon zygaena (misidentification) Poey. Liberia: 16° 18' W. (Original (May refer to description. 96. 4.

South Moscos Island group. is 106544 : (1). 1°29' N. mouth of St. — . (South Carolina): 37798 fl). Freeport. Englewood. 640. 1834. UMMZ USNM USNM : USNM FMNH MCZ USNM USNM USNM MCZ : BMNH MCZ USNM USNM MCZ USNM USNM USNM USNM MCZ MCZ no further data. USNM 29289 (1)." as it was then called. 52642 (1). SU 35828 (3). between Panama City and Punta Gorda.32. 50) from the south coast of Australia. Gulf of California. Society Islands: MCZ 460 (3). at Massaua. Manila market. UCLA 53-275 (3). New Caledonia: USNM 111201 (head). Biu-ma: SU 14496 (1).) 155050 (1). 3639 GILBERT 39 FMNH 06°27' to 06°25' N. or "New Holland. pi. Englewood. Trinidad: 3165 (2). Simons 25180 (1). (Texas): 35826Aransas. (This locality.. Foochow. p. Mexico: SU 11696 (1). 108451 (holotype 133580 (1). Sphyrna lewini was originally described (as Zygaena leivini Griffith and Smith. Eritrea: 49330 (1). India: SU 14398 (1). the accompanying picture (pi. Although the description lists none of the important diagnostic features. MCZ 1059-1061 (3). 151539 (1). China: SU 14150 (1). Makassar market. SU 19193 (1). 130604 (1). San Miguel Island. Honolulu. no further data. USNM United States (Florida) not close to Trinidad. 170557 (1). Red Sea. no further data. Batavia. Honolulu. USNM 56337 (1). Charleston. no further data. Panama: ANSP 565 (1). USNM 170555-170556 (2). Panama Bay." no fiu-ther data. 79 (1). 3) is identifiable. USNM 111243-111244 (2 dried skins). near Panama. Panama Bay. a circumstance resulting from the inclusion of his (1).: UF 455 (1). USNM 153587 (1) (syntypes of Cestracion oceanica). Pacific Ocean (including Indian Ocean and Red Sea) Aden 1925. USNM 120376 (head). Celebes: 479 (1).. Panama Bay. Costa Rica: SIO 52-94 (1). SU 14088 (1). Brevard Co. no fiu-ther data. Tinghai. UF 4208 (1). "Pacific side.5). and can be recognized by the distinct median indentation on the anterior part of the head and the low second dorsal fin with a long lobe.. 52644 (1).37 (6). Philippines: USNM 170554 (1). Japan: UAIMZ 177119 (1). 18390 (1). SU 12765 (1). Garachine Pt. Gulf of Aden. St. Glynn Co. Port USNM. 10°39' W. BB^Od' to 55°10' W. Nomenclature. Iloilo market. Nassau Co. Sound. MCZ 1088-1092 (. Cavite.. of Sphyrna diplana). Calicut. (Georgia) UF 5463 (1). 456 listed. 28°31'00" N.. Hawaii: 471 (1). Galveston. no further data. 32°42' N. Fernandina.20. as no further data. USNM 111414 (stuffed skin). ANSP 82186 (2). 118660 (1). 141°47'00" W. MCZ 1063 (1).. Matanzas Inlet.7. 158561 (1). Chu Shan Island. Java: USNM 72476 (1). Johns Co. no further data. 79°02' W. Puntarenas. off San Marcos Island. In the past Griffith has been cited as the original describer of Sphyrna lewini.. San Miguel Bay. Merritt Island. 35223 (1) 116449 (3). Southern Chosen (South Korea?).: HAMMERHEAD SHARKS NO. MCZ 517 (2).

however. he was unaware that the Atlantic and Pacific populations of lewini are morphologically indistinguishable. 46-52) v/as the first to demonstrate conclusively that S. diplana.]. in his monograph of the sharks (1913. and therefore. 28). [and] similar in most respects. At the time. past confusion has resulted because only Griffith's page (entitled "The Animal Kingdom") as author of the additions. For a long time Sjphyrna lewini was confused with the superficially similar S. a suspicion that is confirmed by examination of the four syntypes (MCZ 460 [3 spec. He did not synonymize the two species. however. remains constant at all sizes. 119 name on the title page as author of the supplementary sections in the English (1834) edition of Cuvier's "The Animal Kmgdom. is the fact that there are actually two consecutive title pages to each volume. he described a new hammerhead. It is clear from the above that Cuvier had no part in the preparation of these supplements. however. The length of the inner narial groove in oceanica. zygaena and *S'. since. although the words "and others" are written in small type underneath. zygaena. Although Garman distinguished the young of the above species. diplana and S. pp. due to a lack of material from critical areas. 21-28)." In the comparison of this species with zygaena. as he correctly suspected. lewini. Tortonese doubted the validity of Sphyrna oceanica (1950a. Springer (1941. p. which. and. strongly questioned their taxonomic separation.PROCEEDINGS OF THE NATIONAL 40 MUSEUM vol. 153587 [1 spec." the names of Edward Griffith and Charles Hamilton Smith appear. and he consequently described the former population as a new species. which he noted was "closely allied to C. reference was made to the long inner narial groove of the latter (extending more than halfway from the nares to the middle of the snout). there is no evidence that he recognized the adults. which he recognized in the yoimg. Griffith and Smith should be regarded as describers of Sphyrna lewini and other new forms appearing in this volume. Garman seems to have been the first to recognize the distinctness of the two species. pp. Tortonese (1950a. although conthuiing to recognize both S. is identical with lewini. together with other characters mentioned in the original description. In addition. Cestracion oceanica. thereby eliminating him as a codescriber of the species named therein.]). a situation that is difficult to understand in view of the fact that the difference in length of the inner narial groove. S. 158-159). thus indicating their responsibility for the new the first title The name appears on sections on the orders of fishes. under the heading "The Class Pisces. lewini are specifically distinct. zygaena." Apparently overlooked. would lead one to suspect that this species is identical with Sphyrna lewini. USNM . pp. on the second page of volume 10.

c. the patch characterized by a rather sharp. size of second dorsal fin. a shorter inner narial groove. 213-219) synonymized S. Figure 10. 639 mm. The anterior-median pore patch on the underside of the head is unique.20. 415. {d-g. eleventh lower tooth. diplana with although he continued to recognize S. from Rio de Janeiro. orbit farther from nares. in 22d). citing supposed diiferences in size of eye. pp.37). (From Bigelow and Schroeder. b. which is devoid of pores. — Sphyrna are not repeated here. smoothly rounded angle at the outer-posterior comer and by a narrow triangular the median-posterior region (fig. h. being separated by a distance slightly less than greatest horizontal diameter of eye. and shape of head. Characters mentioned in the diagnosis of the subgenus S.) g. about 42 X . lewini. its closest relative) by a long second dorsal fin lobe and a low second dorsal fin.7. about natural size. lewini is distinguished (scallop) . about 0. Examination of specimens from the Gulf of Aden (BMNH 1925. dermal denticles. juvenile female. Brazil upper and lower teeth. the lobe extending from three-fourths to four-fifths of distance from second dorsal base S.55 X. twelfth upper tooth. (MCZ Sphyrna 462). 32. 1948. p. the length of the former about twice greatest height of fin. zygaena. d. relative position of first dorsal fin. zygaena by a median indentation on anterior margin of head. fourth upper tooth. fails to substantiate his conclusion that this nominal species is a valid form. area.) Sphyrna lewini is characterized (as is S. oceanica as valid. ventral view of head. which extends 40 to 45 percent of distance from inner margin of nares to tip of snout. which were identified by Fraser-Brunner as S. e. lewini: a. length of snout. oceanica. third lower tooth. TL. from S. left side. tip of second dorsal lobe reaching closer to upper precaudal pit.— HAMMERHEAD SHARKS —GILBERT 41 Fraser-Brunner (1950./. Diagnosis.

—Meristic data appear Characters men- in table 5. Head moderately expanded.0 to 30. first dorsal about one-half length of head and from half again to almost twice length of pectoral fin base. about one-fifth to one-fourth as long as greatest horizontal diameter of eye. labial furrow present at corner of lower jaw only (concealed when mouth is closed). distal margin of pectoral fin . A comparison of Sphyrna lewini with other members of the genus is presented in table Description. a more obtusely pointed posterior ridge to the upper precaudal pit (fig. with the outer edge curving evenly inward. . pectoral fin about two-thirds as broad as long. tioned in the diagnoses of the subgenus Sphyrna and of lewini are aS. long). a line drawn perpenfin. .42 PROCEEDINGS OF THE NATIONAL MUSEUM vol. length of snout about one-thu'd of head width in smaller specimens. narial flap broad. dicularly from apex of fin passing close behind base base of . 119 to pit. the greatest width from 24. orbit large. anterior margin of finest dorsal fin evenly curved. a slightly less deeply falcate anal fin. to about one-fifth of head width corners of in larger specimens. outer posterior corner of lateral expansion of head posterior to corner of mouth. probably in specunens of all sizes. shallow scallops present on anterior margin of head. the deepest part of the notch on the distal margin situated slightly anterior to a Ime drawn perpendicularly from the posterior axil of the fin. a longer anal base.2 percent of TL (in specimens less than 1. second dorsal fin base from three-fifths to fourfifths length of anal fin base. 2ld). and several chondrocranial characters (pi. not repeated here. presence of a rostral fenestra. distal margin of first dorsal fin strongly falcate lobe of first dorsal fin about one-third length of base origin of second dorsal fin situated about halfway back above base of anal fin.000 mm. the horizontal diameter almost equal to length of shortest (fifth) gill slit. moderately pointed at tip. with part nearest apex very strongly curved. greatest transverse distance mouth from one-fifth to one-fourth of greatest between width of head. 1. first dorsal fin broad. Origin of first dorsal fin above or slightly posterior to axU of pectoral a perpendicidar line drawn ventrally from origin intersecting anterior thu'd of pectoral fin. anterior margin of pectoral fin moderately convex. rather erect (not as erect as in large specimens of Sj^hyrna tudes). anterior margin of pectoral fin from one-half to two-thirds length of head (higher values usually apply to larger individuals) and about equal to greatest vertical height of first dorsal fin. symphysis of jaws situated posterior to a transverse line drawn between posterior margins of orbits in specimens of all sizes. occasional presence of a lower precaudal pit. about halfway between nares and tip of snout. 6b). the length of which is about equal to length of pectoral base.

lower margin nearly straight. caudal fin nearly a thii'd of TL. sloping posteriorly at about a 35° angle from a perpendicular to body axis. upper margin of caudal fin nearly straight. the relative number of body to caudal vertebrae varying from six more to twelve less. posterior margin of lower lobe nearly straight. prominent. similarly oblique. or nearly as great. only one individual with less than 192). the distance from angle included 1% by upper and lower caudal to 2 times in anterior margin lobes to tip of latter going from of lower caudal lobe. . pointed. the 15th Dermal dorsal fin) length of from upper part of back beneath first and moderately arched denticle (measured to tip of median marginal "tooth") as denticles (examined closely overlapping. large individuals. anterior wing of olfactory cartilage prominent. teeth 1 through 3 in upper jaw nearly symmetrical and erect. lower caudal lobe fairly erect. anal fin pelvic fin. lower caudal lobe one-third to two-fifths as long as upper lobe. and projecting medially. apex rounded. terminal part of caudal about one-fifth of total length of fin. Anterior margin of rostral node of chondrocranium nearly straight. body vertebrae 86 to 100. median indentation. bluntly rounded wings present on sides of rostral node. posterior part of preorbital process terminating in an evenly rounded knob. teeth sometimes weakly serrate in 15 or 16-1 or 2-15 or 16 and 16th very small. cusps present on all teeth. 3539 GILBERT 43 and inner corner of pectoral fin equal in length or slightly less (about four-fifths as straight or slightly falcate. pelvic long) than anal anal fin fin fin. with no noticeable median notch. as width. height of anal fin about a third again as lobe of caudal relatively narrow and long. anterior margin of pelvic fin weakly convex. first tooth smaller than second. and slender. anterior margin of lower caudal lobe weakly convex. anterior fontanelle (viewed ventrally) appearing broad and shallowly U-shaped. posterior margin of terminal part of caudal fin moderately falcate. accessory rostral cartilages or with a very shallow absent. but with successive series tending to become more erect and their cusps relatively more narrow with growth. with a sharp angle on anterior edge. HAMMERHEAD SHARKS NO. lower teeth with somewhat narrower cusps than uppers. anterior part of preorbital process long Total number of vertebrae 174 to 204 (in nine specimens examined. small specimens usually with thi'ee. pelvic fin base approximately equal in length to base. with the subsequent teeth becoming increasingly oblique toward corners of mouth. terminal apex of caudal narrowly rounded. Teeth .. the distal equal in length or slightly greater than much as height of second dorsal fin. the blades thin great.

and. sharp-toothed ridges. 419) . With the possible exception of the teeth. Sphyrna lewini shows little noticeable geographic variation in external morphological characters. Individuals deep olive to brownish-gray dorsally. 1948. This can be seen from table 5. I choose to regard S. S. Shelton P. the length and development of the intranarial groove. lewini. median tooth on denticle slightly longer than more distal teeth. slender. p. remain quite constant. tip of ventral siu"face of pectoral fin faintly gray-black to dark black in Maximum life. 1965. However. series of nine vertebral Range — Sphyrna lewini In the western Atlantic it is circumtropical in distribution (map 3). probably between 12 and 13 feet. The reduced count in the other specimen (USNM 25180—from Charleston. 1948. in some cases they may be very weakly serrate. It is diflBcult to say at this time what the significance of this unusually low count is. 119 and as manypointed marginal teeth present on denticle.PROCEEDINGS OF THE NATIONAL MUSEUM 44 vol. In the eastern Atlantic its range is not so well defined. Applegate has recently informed me that he has seen specimens of this species from the western (in litt. S. such as the structure of the chondrocranium (pi.) Atlantic with serrated teeth. Variation. and the shape and position of the fins. in fact. Those characters that are considered to be of basic taxonomic importance. The northern der. shading to white ventrally. variation in eight of the nine individuals is only from 192 to 204. was one of the reasons for Springer's naming the Atlantic popidation as a new species. 22d). 6b). moderately large specimens with four or five. 1960. p. This apparently occurs total length — only in larger specimens. diplana. in which morphometric data have been recorded from specimens from throughout the range. the position and length of the gill slits. pedicel very short. Although the teeth in Sphyrna lewini are ordinarily smooth. Backus. p. Dr.) is due both to a decreased number of body and caudal vertebrae. involved. 419). diplana as a A and because there seem more than one species to suggest that synonym of S. p. 244). occurs from southern Brazil (and possibly Uruguay) north to New Jersey (Bigelow and Schroeder. the form of the anterior-median pore patch on the underside of the head (fig. to be is In view of no concomitant characters this. counts is available for specimens from widely scattered parts of the range. p. The serrated condition has long been thought to be characteristic only of the Pacific population (Sadowsky.C. 2). 1948. Although the total number of vertebrae ranges from 174 to 204. 419. It occurs in the Mediterranean Sea (Bigelow and Schroe- and along the African coast south at least to Liberia and Ghana (Bigelow and Schroeder. at least 5 or 6 feet in length.

although the form was said to resemble S. although it has never been recorded from the British Isles. which also logically might be expected to occur there. (The most westerly record from Africa mens examined. 31. — . in which the temperature does not rise above the low Although its range overlaps that of S. 1961. pp. together (Herald. bigelowi. has been compared with the head of a typical specimen of S. p. zygaena to some extent.—GILBERT HAMMERHEAD SHARKS 45 limits of its range in the eastern Atlantic are uncertain. This head. which is 425 mm. diplana ( = S. 98-99." The description was very brief and there was no illustration. wide (USNM mm. compared with tips of pectoral fins described as wliite. couardi.).) S. -Guinea: USNM uncataloged (1 head. diplana locality: — (Original description. Bruce B.e. "off West Africa". WhUe I have not seen a whole specimen of S. wide. solid = :speci- literature references. differing from that species (and from all other members of the family Sphyrnidae) in having the inferior part of the pectoral fin white and a longer and smaller head.. 12°06' N. and S. The fact that it is the only species of hammerhead definitely recorded from the Hawaiian Islands (Gosline and Brock. 1960. 1951. 17°22' W. 425 in width). lewini). hollo is now known w= confirmed to represent S. and in the eastern Pacific from northwestern South America to the Gulf of California. p. 70's F. CoUette from off West Africa. I have examined a large head (USNM uncatalogued) collected by Dr. Sphyrna lewini is found in the western Pacific from southern Australia north to southern Japan and Korea. pp.) there are relatively few records of the two species having been taken waters . Sphyrna (Sphyrna) couardi Cadenat Plates 9a. 98-99) from "off West Africa. lewini.) Sphyrna lewini apparently does not extensively penetrate cool (i. 10a Sphyrna couardi Cadenat. type Specimens examined. 91) may indicate that it is a more pelagic species than Sphyrna mokarran. r I mS\ mii^-iriAxAiyjzj Map 3. 445 mm. Discussion Sphyrna couardi was described by Cadenat (1951. fig. couardi. — Distribution of Sphyrna lewini.

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PROCEEDINGS OF THE NATIONAL MUSEUM

46

vol.

no

from the same geographic area. Photographs of these
appear in plate 10 and radiographs in plate 9.
The most striking difference between the two heads is the deeper,
uncataloged)

,

more robust appearance
10).

This

is

measurements in table
Cadenat described S. couardi as having a

of S. couardi (also see

confusing, since

"longer and smaller" head than S. lewini; however, this apparent
The overall
discrepancy may simply be a matter of semantics.
appearance of the head, in fact, seems closer to S. mokarran than S.
lewini; however, couardi and mokarran may readUy be distinguished
by the presence of a distinct intranarial groove and nonserrated teeth
in the former species, and by chondrocranial structure and differences
in pore patterns on the ventral surface of the head.

The mucal pore patches on the ventral surface of the head are
basicaDy similar in S. couardi and S. lewini, although the anteriormedian patch in the former is completely separated medially instead
The
of being divided about halfway as in the latter (fig. 22d).
chondrocrania of the two forms also are similar (pis. 9 a, b), the main
difference being in the appearance of the preorbital process. In (S.
couardi this structure is shorter and has a distinct knob at the angle,
whereas in S. lewini it is longer and lacks a distinct knob. In addition,
the olfactory wings in S. couardi are blunt and do not come to a sharp
point as in S. lewini. In this case, however, the wing tips are rather
ragged in appearance, suggesting that this possibly may be an aberrant
condition.

In conclusion, the above evidence seems to indicate that Sphyrna
is a valid species of restricted range and that it is most closely

couardi

related to

Sphyrna

lewini.

Subgenus Platysqualus Swainson
The subgenus Platysqualus, which is here considered to include
Sphyrna tudes, S. media, S. corona, and S. tiburo, is distinguished by
the following unique characters: First and fifth gill slits nearly equal
in length and both somewhat shorter than the three middle slits anal
fin and anal fin base longer than in any other group of sphyrnids, the
former 11.0 to 13.5 percent, and the latter 7.5 to 9.0 percent, of TL;
anal base a fourth to half again as long as pectoral base; upper pre;

caudal pit broadly semicircular or slightly pointed in shape, the ends
pointing anteriorly (figs. 21e-'i) lower precaudai pit present (this
featm-e also sometimes present in S. lewini); fifth gill slit above in;

sertion of pectoral fin (slightly posterior in S. tiburo); lobe of first
dorsal fin extending past insertion of pelvic fin (in all but S. tiburo)

teeth weak, nonserrated, with narrow pointed cusps; accessory rostral
cartilages often present (none observed in S. tiburo).

;

HAMMERHEAD SHARKS

NO. 3539

GILBERT

47

In addition, the subgenus is characterized by: A fairly high second
fin, its greatest height about equal to greatest height of anal fin
a shallowly falcate anal fin; e^^es situated anterior to mouth, a line
drawn between the posterior margins of orbits passing anterior to symphysis of jaws (intersecting symphysis in larger specimens of S. tiburo)
a small orbit, separated from nares by a distance greater than horizontal diameter of orbit; absence of an outer narial groove; a variable total
number of vertebrae, ranging in number from 142 to 202, mth those
over posterior part of coelomic cavity not enlarged; a small maximum
Perhaps the maximum size
size, probably not exceeding 4 or 5 feet.
dorsal

;

in

some

cases

is

even

less (e.q., S. corona).

Sphyrna (Platysqualus) tiburo (Linnaeus)

Diagnosis.

—-Characters mentioned in the diagnosis of the subgenus

Platysqualus are not repeated here.

Sphyrna tiburo is unique among members of the genus Sphyrna in
having the head only slightly expanded, either spade or shovel shaped,
and measuring from 14.4 to 24.5 percent of TL in specimens up to
1,000 mm. long (higher values usually apply to smaller individuals
and/or to individuals of the subspecies vespertina). The anteriormedian pore patch on the underside of the head is also unique, the
patch characterized by a rather sharp, but smoothly rounded angle at
the posterior corner and by a long, narrow median posterior extension
(fig.

226).

Sphyrna tiburo is also distinguished by the anterior margin of the
head being more or less evenly rounded between the ej^es, with neither
a median scallop nor narial depressions present; no inner narial groove;
origin of first dorsal fin distinctly posterior to axil of pectoral fin, a per-

pendicular line drawn ventrally from origin of dorsal intersecting posterior part of pectoral fin; teeth not serrated, with short, weak, rather

blunt cusps.

A

comparison of Sphyrna tiburo with other members

presented in table

of the

genus

is

1.

Description.— Meristic data appear

in table 6.

Characters men-

tioned in the diagnoses of the subgenus Platysqualus and of S. tiburo
are not repeated here.

Intranarial distance from one-tenth to one-sixth of

values usually apply to larger individuals)

TL

(lower

outer posterior corners
of lateral expansions of liead broadly rounded and situated posterior
to corners of mouth; narial flap broad, blunt at tip, with the outer
;

edge curving gradually inward; length of snout about two-fifths
of head width; greatest transverse distance between corners of mouth
from one-third to a little more than two-fifths of greatest width
of head; sjTnphysis of jaws situated, in smaller individuals, slightly

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PROCEEDINGS OF THE NATIONAL

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vol. 119

drawn between posterior margins of
drawn between these points
passes through the symphysis; a weakly marked labial furrow present
on lower jaw only, the furrow extending only a short distance (concealed when mouth is closed)
Fourth gill slit situated anterior to, and fifth gill slit slightly posanterior to a transverse line

orbits, while in larger individuals a line

terior to, insertion of pectoral fin; posterior lobe of first dorsal fin
terminating just anterior to insertion of pelvic fin (the dorsal and
pelvic fins being in closer proximity only in S. tudes, S. corona, and
first dorsal fin relatively slender, not perfectly erect, a
S. media)
;

line

drawn perpendicularly from apex

of fin; base of first dorsal fin

of fin barely intersecting lobe

about two-fifths length

least half again length of pectoral fin base; anterior

head and at
margin of first

of

dorsal fin evenly curved, with the part nearest the apex

curved; distal margin of
first

first

more strongly

dorsal fin moderately falcate; lobe of

dorsal fin from four-ninths to almost one-half length

of first

dorsal base; origin of second dorsal fin situated about a third of

way

back above base of anal fin second dorsal fin rather tall, its greatest
height about two-thirds length of base, and about equal to greatest
height of anal fin; second dorsal fin base about five-eighths of anal
fin base; lobe of second dorsal fin rather long, its length about a third
again greatest height of same fin, and extending about two-thirds
of distance from posterior part of second dorsal base to upper precaudal pit; anterior margin of pectoral fin from one-half to nearly
two-thirds length of head (higher values usually apply to larger
individuals); pectoral fin four-fifths as broad as long; pectoral fin
;

base five-eighths to three-fourths length of anal fin base; anterior
margin of pectoral moderately convex; distal margin of pectoral
nearly straight; apex and inner corner of pectoral rounded; pelvic
fin about three-fourths length of anal fin; pelvic fin base about threefourths length of anal fin base; anterior margin of pelvic fin weakly
convex; distal margin of pelvic fin very shallowly falcate; anal fin
about one-fourth again as long as pelvic fin; anal fin base from onefourth to over one-third again as long as pectoral and pelvic fin
bases, and from three-fifths again to nearly twice as long as second
dorsal fin base; height of anal fin about equal to greatest height of
second dorsal fin; anal fin weakly falcate toward apex and nearly
straight toward tip of lobe; caudal fin from one-fourth to nearly

body length; upper margin of caudal nearly straight;
terminal part of caudal about one-fourth of total length of fin; posterior
margin of terminal part of caudal fin nearly straight or shghtly
falcate; both terminal and lower apices of caudal narrowly rounded

one-third of total

somewhat more pointed in larger individuals;
lower caudal lobe appearing relatively broad and short, the distance

in smaller specimens,

accessory rostral cartilages absent. anterior wing of olfactory cartilage prominent. the eighth and subsequent teeth without cusps or definite cutting edges. posterior margin of lower caudal lobe nearly straight. and three series functional functional toward corner of Dermal mouth denticles (examined of lower jaw. the anterior evenly rounded and projects forward. the relative number of body to caudal vertebrae varying from 9 more to 2 less. but subsequent upper teeth becoming increasingly oblique toward corners of mouth. denticle. body vertebrae 71 to 88. times blunted or broken) usually present at posterior margin of and teeth sometimes rudimentary or absent. the fourth through seventh or eighth slightly oblique. . rostral fenestra absent. 423). . lower teeth shorter than upper teeth. usually two functional along side. lower caudal lobe sloping posteriorly at about a 45° angle from a perpendicular to the body axis. first upper ^^ tooth 12 to 14-1-12 to 14 erect and symmetrical.u Teeth teeth not serrate. p. or nearly as great.HAMMERHEAD SHARKS NO. with a distinct. with terior part of preorbital process terminating in a corner of which is . rounded. five strong ridges usually present. lower lobe of caudal one-third to two-fifths as long as upper lobe. ^ n . length of denticle (measured to tip of median marginal "tooth") as great. and the posterior corner of which is somewhat more angulate (pis. denticle ordi- pedicel short and slender. 3539 —GILBERT 49 from the angle included by the upper and lower caudal lobes to tip of latter going from 2 to 2^ times in anterior margin of lower caudal lobe. . pos- knob. . _ 12 to 14-0 or 1-12 to 14 ^ . anterior fontanelle (viewed ventraUy) appearing broad and shallowly U-shaped. ' ' narrower cusps and slightly broader bases. . the outer ridges particularly in smaller individuals. and projecting medially. from upper part of back beneath first the blades rather steeply raised. . as width. anterior margin of lower caudal lobe weakly convex. pointed. Anterior margin of rostral node of chondrocranium either straight or with a poorly defined indentation. anterior part of preorbital process short and rather blunt. 1948. three series of teeth functional in front one or two and along side of upper jaw. teeth 3 through 10 or 11 largest in both jaws. cusps evident only on teeth 1 through 9 in specimens examined (through 11 or 12 according to Bigelow and Schroeder. prominent pointed wings present on sides of rostral node. the first through third teeth erect and symmetrical. and as many long. narily slightly longer than 219-937—67 4 more median tooth on distal teeth . 7 a. four to six series functional in front. b) Total number of vertebrae 142 to 172 (seven specimens examined). sharp-pointed teeth (these teeth are somedorsal fin) closely overlapping. median notch. with a moderately sharp angle on anterior edge.

b) Individuals gray or grayish sively — This difference is also reflected in proportional measurements. Sphyrna vespertina. 173. F. 1948. Garrick (1964) indicates the existence of a number of closely This suggests that those specimens of S. Herre. some individuals are intermediate in head shape (fig. p. 161. . In addition. A. For several reasons the two populations are regarded here as having First. 119 brown dorsally. 27. ulations of the Range.. same subspecies only by number of vertebrae. pp. 12a. the width of the head and transverse distance between the nares averaging slightly greater for specimens from the Pacific (table 6 and fig. tiburo tiburo and three of S. Springer and vertebrae. while those with the higher coimts (159 and 170) are from Rhode Island and Brazil. G. differences are not attributable solely to variation in number of caudal Although it is possible that larger series of counts will show this apparent difference to be clinal in nature. — Despite statements to the contraiy (Bigelow and Schroe- 424-425. recently completed by Drs. although Bigelow and Schroeder (1948. The two specimens of S. V. and 1961. Variation. tiburo tiburo and the three specimens of S. are interesting both because of the wide range in variation found in the typical subspecies and also because of the close agreement in vertebral number between two of the individuals of S. These counts. and often with small dark spots on side of body. (It should be mentioned here that these differentiated only to the subspecies level. As a result. p. p. 162) described the Pacific population as a new species. 12c). no differences comparable to those found for head shape were discovered for other morphometric characters (table 6). there appear to be no substantiated records of Sphyrna tiburo outside of the Western Hemisphere. becoming progresmore pale below. pp. inshore waters of the ranges from southern Brazil northward. tiburo tiburo with the lowest counts (142 and 143) are from the eastern Gulf of Mexico. tiburo tiburo from the eastern Gulf of Mexico represent a sibling form that may be distinguished from other poprelated species differing only in vertebral counts. a study of vertebral variation in sharks. As mentioned previously. 13). such specimens possibly may have resulted from a partial reamalgamation of the populations following initial differentiation of the two forms. Springer (1940a. which range from 163 to 172 for the latter form and from 142 to 170 for the former. p. 1960. where common it in shallow. Briggs. Vertebral counts were made on four specimens of S.) J. 428) later placed it in the synonymy relative of S. tiburo vespertina. PROCEEDINGS OF THE NATIONAL MUSEUM 50 vol. tiburo. tiburo vespertina. 552). occasional spots on caudal fin. Sphyrna tiburo tiburo is western Atlantic. 1953. der. Specimens of Sphyrna tiburo from the Pacific usually have more pointed heads than specimens from the Atlantic (figs.

1839. Ponta de Mucuripe. Vitoria market. Although the be well established on Bimini. pp.) — Specimens examined.) Jordan and Evermann. Brazil: MCZ 464 (3). 1822. 1758. than 100 miles from the mainland. 121. MCZ 162 (5). to southern New —GILBERT North Carolina and. Mexico: MCZ 39645 (1). 1836.) Cestrorhinus tiburo Blainville. ANSP 49938-49940 (3 heads). Panama: USNM 79315 (1). 59. in the western Bahamas. 13. (Photo of jaws. 06°20' to 06°19' N. yolk yolk. MCZ 439 (2). 06°27' to 06°25' N. type locality. Bahia. SU 14047 (1). FMNH .) Platy squalus tiburo Swainson. (Range in size: 350-910 mm. 412. USNM 104318 (2). pi. MCZ 394 (4). illustration of head. UMMZ .HAMMERHEAD SHARKS NO. Punta Gorda..) Zxjgaena tiburo Valenciennes. off Ciudad Carmen. 12/>. MCZ 461 (5). and it is possible that records from the latter area have resulted from market specimens that were caught on the other side of the Florida Straits. there is no evidence of a permanent population on Cuba. misspelled "liburo.) Gill. (Description. new generic name list. 226-227. USNM 43359 (head and tail). fig. MCZ 416 (4). near Puerto Colombia (and Barranquilla). Recife (Pernambuco) 52714 (1). p. (Illustration. MCZ 27 (1).) (Name. 46997 (1). p. (Description. 50. 54°33. p. SU 52715 (1). 403. pp.. despite availability of Platy squalus. evidently mislabeled by accident. (Range. Brazil. these exceptions are from Bimini. SU 52873 SU 52713 (1). Fortaleza. Cuba: USNM 2710 (1). USNM 159201 United States (1). 1896. no type specimen.5' W. p. 51 occasionally. (Original description. new generic name proposed (In North American catalogue. 234.) Sphyrna tiburo Rafinesque. Colon. 22e. SU 52872 (1). 217. 1916.) Sphyrnias tiburo Gray.) Cestracion tiburo Gill 1862. 22. 318. Bahamas: 174384 (1). MCZ 153 (5). 1810. MCZ 522 (2).. early embryo. proposed without comment. species is less known Sphyrna to tiburo vespertina is conomon in similar habitats in the eastern Pacific from southern California to Equador. 3539 regularly." obviously a typographical error. 1758. and England. 55°05' to 55°10' W. Colombia: USNM 94754 (1). to southern All but three records are from continental waters. Sphyrna (Platysqualus) tiburo tiburo (Linnaeus) Figures 11. ANSP 4984149847 (7). c.) Zygaena (Squalus) tiburo Leuckart. p. Nicaragua: USNM 78180 (1). (New generic name for Squalus tiburo Linnaeus. no further data. Sphyrna (Reniceps) tiburo Cestracion zygaena (misidentification) Kadcliffe. "America". Plate 7a Squalus tiburo Linnaeus. 21e. Locality records for Sphyrna tiburo are plotted on map 4. p. Reniceps tiburo (In 1861. Campeche. 2. 1851. 47. 1816. (Name only. 43. Sm-inam: USNM 159198 (1). stalk and p. MCZ 90 (2). SU (1). TL). MCZ 469 (1). Santos. MCZ 450 (2). p. Bimini. Caribbean Sea. Rio de Janeiro.5' to 54°38. and from Cuba.

Englewood. USNM USNM USNM USNM no further data. USNM 51894 (3).. 116888 (1). Tortugas. Beaufort. about 3 X . 91°00' W. Nassau Co. g. Sable. about 14 X (MCZ 90). UF 8948 (1). (MCZ 848). 812 mm. off Cape 184870 (1). Bayou Fifi. Gulf of Mexico. 421. lateral and apical views of dermal denticle. 9270 (1). Johns Co. UF 4208 (1). 125763 (1). (i-A. about 26 X... . Charlesport. off 127114-127115 (2). 1407 (4). 35847 (1). tenth upper tooth.. (1). St. Pine Island Sound. 90). e. UF 5649 (1). south of New Orleans. 108455 (1). 62773 (1). Fernandina Beach. USNM 26582 (1).. UF near Cedar Pt. New(South Carolina): USNM 25171 (1). Alligator Light. TL. seventh lower tooth. UF 1416 (1). 118995 (1). TL. Barataria Bay channel. (Louisiana): 155648 127116 (1). from Rio de Janeiro. Fla. St.. (New 147793 (1).) Figure (MCZ 11. Key West. (Georgia): UF 5463 (1). Simons Island. Spanish Harbor Key (west end-Atlantic side). 755 (1). Daytona Beach pier. from Tortugas. Delaware Bay. Volusia Co.5 X d. juvenile female. p. Tampa Bay. UF 4211 (1). tenth lower tooth. c. . 30687 (1). between Great Egg and Peck Bays.. Mississippi Sound. near West Pass. SU 14329 (1).) (From Bigelow and Schroeder. head of same specimen. Pensacola. 57242 (1). Brazil b. 1948./. Marys River. FMNH 21590 (1). I. fourth upper tooth. Apalachicola Bay. (Alabama): (Florida): USNM USNM USNM USNM USNM USNM USNM USNM FMNH MCZ USNM MCZ USNM USNM USNM Sphyrna tihuro: a. St. 7038 (1).. UF 7061 (1). of male. 367 mm. 34733-34734 (2). left side. St. 28°36' N. Cedar Key. 47334 (1). Jersey): USNM . second lower tooth. h. MCZ 78 (skin). Glynn Co. Levy Co. upper and lower teeth. dermal denticles. UF 7061 (1). (Mississippi): Horn Island. (Rhode Island): ANSP 581 (1). (North CaroUna): Beesleys Pt. 39882 (1). 92594 (1). Augustine.— PROCEEDINGS OF THE NATIONAL MUSEUM 52 USNM 119827 (1). ventral view. CAS 19306 (1). Monroe Co. about 1.

USNM 155964 (18). (Original description. USNM 127104 (1). male. Sonora. Schultz. Sphyrna vespertina Springer. vespertina. specimen 636 h. 15 uador: (1). range. TL (USNM specimen c. a 769 City. from Guayaquil. TL (UCLA tiburo.) — in Diagnosis. Ocean View. SU 11881. 13 mi. same locality as holotype. from Texas. UCLA 58-46 (1). MCZ (5). Bocochibampo Bay. tiburo vespertina having a slightly narrower and more evenly rounded head. SU 11584. FMNH 59350 277-880 mm. showing pointed head. from Gulf of California. Sinaloa. EcGuayaquil. Acajutla. MCZ 36240-36245 (9). 637 S. north of Mazatlan. 37794-37796 (3). mm. El Salvador: USNM 21627 (1). head of 127106). USNM 116450 35824-35825 (2). Ecuador. showing rounded head. mm. FMNH . Chame Point Pass. com- mm. 1940a. CAS (W 55-167) (1). Sphyrna (Platysqualus) tiburo vespertina Springer Figure 12a. UCLA 51-4 (1). parisons. (Texas): ton. ca. paratypes. Sphyrna tiburo: a. mm. USNM 127106 (1). These differences are illustrated in figures 12 and 13. Port Aransas. from southern Florida. Gulf of California. TL). type locality. Galveston. Camorore Beach. Point Isabel. Sinaloa. Baja California. /. t. Hampton Roads. CAS 3987 (1). (Range in size: USNM 88677 (1). MCZ 138 (skin). t. female. Lynnhaven Koads. head of S. 1622 (1). Mexico: USNM 190595 (1). USNM 127101 (1). Almejas Bay.— —GILBERT HAMMERHEAD SHARKS 53 USNM 82710 (2). showing intermediate (Drawings by Dorothea B. 5675. and formerly Carnegie Museum no. 164-166. off Isla San Ignacio and Isla Macapule. . Aransas. TL (USNM tiburo. Harbor Island. a head of S. Parateria (Grand Pass) USNM 127109 (1). a 57242). Guaymas. a a 938 mm. pp. Plate 7b 161. Corpus Christi. SIC 47-53 (1). south of Bahia Topolobampo. Laguna Madre. just west of Cape Henry light. Gulf of California. Gulf of California. (Virginia): USNM 125760 (1). UCLA 52-250 (4). specimen 880 condition of head. a 58-46). ca. Panama 520 mm. holotype.) — Specimens examined. off Bay Chalaud. MCZ 36157 (1). I I I FMNH FMNH Figure 12. south end of Angel de la Guarda Island. female. 11195-11198 (4). Gulf of CaUfornia. Sphyrna tiburo tiburo differs from S. at anchorage off Santa Margarita Island. 3 mi.

vespertina) Panama City. . MCZ — MCZ are illustrated in figures 12 250 ?40 9^ < < <n 111 X3 and 13. from FMNH . SU 11881 (one paratype of S. Sphyrna tiburo vespertina differs from S. Baja California. Panama City. Diagnosis. from Rio Pacora. Panama: 190591 (2). 80 (1). Panama Bay. Panama Bay. no further data. 516 (1).PROCEEDINGS OF THE NATIONAL IMUSEUM 54 USNM mouth of Rio Colorado. 62438 (2). Panama Bay. Panama Bay. between Panama City and Punta Gorda. 1-3 mi. SU 11584 (holotype of Sphyrna vespertina). Punta Chame and Punta Anton. offshore between Rio Chico and Punta de la Plata. mi. tiburo tiburo These differences in having a slightly broader and more pointed head. SE of Isla Taboga and 2-4 mi. CJCLA 53-275 (3). UCLA 58-305 (1).

(all originally UCLA 58-304). which were at first thought to be due to individual variation. first dorsal fin of larger specimens (600 mm.HAMMERHEAD SHARKS GILBERT 55 USNM Pacific Ocean: Costa Rica: 196140 (6). but noneWith a firm basis for separation thus theless constant. Initially Sphyrna media and Sphyrna corona were regarded by me as synonymous. Panama Bay. off Isla San Ignacio and Isla Macapule. offshore between Punta de Hicacal and Rio Pasiga. or more) less erect and less broadly triangular. tudes in the following characters: Anterior margin of head more rounded (particularly in large specimens). The various distinguishing features are discussed in the diagnoses of the respective species. UCLA 58-304 (4.. corona. 50376 (1). does not remain constant. Sinaloa. Sphyrna media differs from the closely related S. In addition. available. Mazatlan. 9 A (Solid Sphyrna tiburo tiburo Sphyrna tiburo vespertina = specimens examined. a line drawn perpendicularly from apex of fin not intersecting base of fin. SU 11583 (holotype of Sphyrna media partial skin). it was found that other morphological differences. tudes and with a pointed median-posterior extension (fig. which was the primary basis for Springer's origmal separation of the two forms. —Characters mentioned in the diagnosis of the subgenus Platysqualus are not repeated here. differences. 22/) narial depression very shallow. SIO 60-88 (3). the chondrocrania of the two forms are very similar. the head shape grading from the more evenly rounded condition of S. 21°55' N. Closer examination of the chondrocrania. includmg three heads). This conclusion was based mainly on the fact that difference in anterior contour of the head. 28160 (1 paratype of Sjihynia media). however. including one bead). Gulf of California. anterior-median pore patch on underside of head with the posterior margin more broadly rounded than in ^S. Gulf of California. media to the more lobate situation found in S. as are FMNH — USNM USNM USNM Map 4. also could be used to separate the two species. Panama: 190593 (4). Sinaloa. south of Bahia Topolobampo. . Diagnosis. hollow: confirmed literature references. . —Distribution of subspecies. Gulf of Nicoya. Mazatlan.) the various external morphological features. no further data. 63093 (2). revealed several relatively smaD. Sinaloa. Mexico: UCLA 58-47 (2. above San Benito.. Chiapas. 105°37' W. with no pronounced median notch. inner narial groove absent.

5 percent. corona in having a narrower postorchondrocranium. 16.— . rather than anterior. 900 mm. these have been plotted and may be seen in figure 17. TL. media. from Panama Bay (UCLA 58-304).5 X. corona from 25. corona in having a slightly shorter snout. d. The last difference has been objectively shown by dividing the transverse distance between the tips of the preorbital cartilages into the 14. two species. dermal denticles of same specimen. with the anterior edge of this structure posterior. and 186. PROCEEDINGS OF THE NATIONAL MUSEUM 56 from S. c.1 to 24. A comparison of snout lengths may be seen in figures Although there is some overlap in values for the 156. and shorter rostral cartilages. (Drawings by Dorothea B. upper and lower teeth.0 percent. Figure b. Sphyrna media also differs from S. when specimens of a comparable size are compared lower length of the rostral cartilage. values are obtained for S. about 0. about 1. Schultz. obtained is from 18. .9 to 30. of same specimen.12 X. to the angle formed by the Sphyrna media differs bital process to the juncture of the innerorbital and anterior mediorbital cartilages (pi. Sphyrna viedia: a. adult male. head of same specimen.) For S. media the range of values thus and for S. 7c) a slightly deeper anterior fontanelle. left side.

from Chiapas.3 percent of head somewhat variable. corona in having: The anal fin and with a more pointed apex (figs. corona (the holotype) measured only 672 mm. must be seen A to definitely confirm this. Characters mentioned in the diagnoses of the subgenus Platysqualus and of S. bluntly pointed at tip. When the transverse distance between the corners of the mouth is divided by the head width. Sphyrna media may be a larger species. and with a broad. a. comparison of Sphyrna media with other members of the genus is presented in table 1. anterior-median pore patch on underside of head with the posterior margin more broadly rounded and with a pointed median-posterior extension (fig. narial flap broad. 14a.^ -Meristic data appear in table 7.— HAMMERHEAD SHARKS —GILBERT 57 from S. very shallow median depression in others. 21 g). media The largest specimen Finally. outer posterior corner of lateral expansion of head situated anterior to corner of mouth. character of anterior-median margin of 22. evenly rounded in some individuals. with the broadly curved outer edge curving sharply inward. with no median indentation. the largest More specimens of S. Sphyrna media: 63093). 15a). Mexico (FMNH (Drawings by Paul Laessle. a .3 to 27. 22/) a more pointed upper precaudal pit in smaller specimens (fig. of Sphyrna media seen (a head) was calculated to be 1025 mm.) Head moderately expanded. the values obtained are somewhat greater for S. the greatest width measuring from TL. — Figure IS. TL. Description. Sphyrna media more deeply also differs falcate . TL. b. media are not repeated here. however. mm. juvenile female. 356 head of same specimen. while of the 17 specimens of Sphyrna corona examined. and a slightly wider mouth.

second dorsal fin tall. PROCEEDINGS OF THE NATIONAL MUSEUM 58 vol. outer third slightly convex. anterior margin of pelvic fin straight or slightly convex. inner twothirds of anterior margin of pectoral fin straight. its length about a third again greatest height of fin and extending from two-thirds to three-fourths of distance from posterior part of second dorsal base to upper precaudal pit. sloping posteriorly at about a 25 to 30° angle from a . length of pectoral fin base from five-eighths to thi'ee-fourths length of anal fin base. its greatest height about threefourths length of base and from three-fourths to four-fifths of greatest height of anal fin. particularly in larger individuals. lobe of second dorsal fin rather long. lower caudal lobe about one-third as long as upper lobe. particularly fin from two-fifths margin and evenly curved. the distance from the angle included by the upper and lower caudal lobes to tip of latter going about 1% times in anterior margin of lower caudal lobe. distal margin of pectoral fin slightly falcate. lobe of first dorsal to one-half length of first dorsal base. not noticeable in smaller specimens. length of anal fin base onefourth to four-ninths again as long as pectoral and pelvic fin bases and from three-fifths again to nearly twice as long as second dorsal fin base. particularly of first dorsal fin rather deeply on lower two-thirds of fin. distal falcate. terminal part of caudal slightly less than one-fourth of total length of fin.. Origin of first dorsal fin slightly posterior to axil of pectoral drawn ventrally from a perpendicular line fin. ii9 very short labial furrow present at corner of lower jaw in larger specimens. origin intersecting anterior third of pectoral fin.third of total body length. length of caudal fin from one-fourth to nearly one. anterior margin of first dorsal fin broadly in larger individuals. upper margin of caudal fin evenly convex. pectoral fin from two-fifths to two-thirds length of head (higher values usually apply to larger individuals) pectoral fin two-thirds as broad as long. second dorsal fin base from one-half to five-eighths of anal fin base. slightly more acute in larger individuals. distal margin of pelvic very slightly falcate. both terminal and lower apices of caudal broadly pointed. apex of pectoral broadly pointed in small specimens. length of pelvic fin from two-thirds to four-fifths length of anal fin. lower posterior margin of terminal part of caudal fin nearly straight or slightly falcate. origin of second dorsal fin situated about two-fifths of way back above base of anal fin. lower caudal lobe rather erect. inner corner of pectoral narrowly rounded. anal fin from one-tenth to one-third again as long as pelvic fin. height of anal fin one-fifth to one-fourth greater than height of second dorsal fin. base of first dorsal fin one-third to one-half length of head and about twice length of pectoral fin base. length of pelvic fin base from five-ninths to three-fourths length of anal fin base. lower lobe of caudal appearing relatively narrow and long.

more strongly convex toward apex. . obtusely pointed wings present . Anterior margin of rostral node of chondrocranium varying sUghtly. posterior margin of lower caudal lobe nearly straight. rostral fenestra absent. a distinct median notch present on anterior fontanelle. from broadly convex to broadly and shallowly concave.HAMMERHEAD SHARKS —GILBERT 59 perpendicular to the body axis* anterior margin of lower caudal lobe weakly convex toward base. tudes) broad. anterior fontanelle (viewed dorsally) deep and broadly U-shaped. fairly prominent (though not as prominent as in S. with the sides of the U slanting slightly inward.

Dermal denticles (examined from upper part of back beneath first dorsal fin) evenly and closely spaced. the outer teeth very low. for those characters that undergo Httle or no proportional change with an increase in total body length. media in having a shorter snout. overlapping but little. If one compares these measurements with those listed for S. nana: (169). snout to second dorsal origin (637). fortunately. length of anal lobe (33). a Ungreater interdorsal distance. no markings on fins. [1940a. For those characters that undergo pronounced . The following measurements were given Head width (190). This form was said to differ from S. length of denticle (measured to tip of median marginal "tooth") less than width. becoming increasingly so toward corners vol. a larger orbit. larger specimens usually with up to five. becoming progressively more pale below. from southern Brazil. media. nana generally fall within the range of values for S. pedicel short and broad. no mention was made either of the morphology of the chondro cranium or of the arrangement of the mucal pores on the (first oblique. median tooth on denticle slightly longer than more distal teeth. snout to fu-st gill sht for the holotype of S. a shorter upper caudal lobe. and without a definite cusp. 119 upper teeth in upper jaw. p. round. Individuals gray or grayish brown dorsally. length of pelvic base (34). teeth 2 through 8 longest (tooth 1 not markedly smaller however). length of pectoral base (54). 163]). Sphyrna nana. in lower jaw. the blades thick and rather strongly arched. based on a single adult female.330 milUmeters long. the values for S. snout to fu'st dorsal origin (308). height of second dorsal fin (37) length of anal base (80). Status of Sphyrna nana. one or two series of teeth functional in alternating rows along sides of upper jaw and two to three rows along sides of lower j aw. Sadowsky (1965) recently described a new species of hammerhead. teeth 6 to 9 or 10 longest. height of anal fin (36). the outermost two or three teeth very short.PROCEEDINGS OF THE NATIONAL MUSEUM 60 two counts listed by Springer. horizontal diameter of orbit (15) length of first dorsal base (95). media in table 7. a shorter distance between the eye and nares. distance between first and second dorsal bases (233). length of caudal fin (251). a shorter distance between the nares. length of second dorsal base (43) length of second dorsal lobe (45). snout to pectoral insertion (204) snout to pelvic insertion (466). but with a definite cusp. the anterior margin of the head not regularly oval and with prenarial convexities. and a longer first dorsal fin base. 1. sharp-topped ridges. small specimens mth three. it can be seen that. of mouth — ventral siu-face of the head. and as many sharp-pointed marginal teeth (especially deep and pointed in small individuals) on denticle.

146). 156). media are very similar. corona. while the validity of S. Presence of the species in this area is further substantiated by Stewart . from the Gulf of CaHfornia south at least as far as Panama Bay. was at first identified as on the basis of head shape. In conclusion. and length of caudal fin) seem to fall outside the range of variation one would normally expect to find in a large individual of S. only three (distance from snout to pelvic insertion. in the southern Caribbean. The occurrence of Sphyrna media in the Caribbean is confirmed by two specimens (USNM 79284. media. The contour of the anterior margin of the head.3539 —GILBERT 61 allometric change with an increase in length (head width. from media.300 milhmeters long. distance between first and second dorsal bases. distance from snout to symphysis. Sphyrna media occurs in the eastern Pacific. I so far can find no evidence to indicate that Variation. while in others the head is faintly lobate. with a slight median indentation or scallop (fig. —Sphyrna media shows moredistinct than the usual amount it is specifically S. Those individuals with more lobate heads may be distinguished only with difficulty from specimens of aS'. a 900 mm. nana cannot be categorically denied until the natures of the chondrocranium and the pore patches on the ventral siu-face of the head are known. Thus. of Some specimens have an evenly rounded head (fig. media (as discussed and elsewhere in this paper). MCZ 500) from the Colon (Panama) market. nana S. the values given for S. corona — Range.HAMMERHEAD SHARKS NO. and those in which there is slight disagreement are not sufficiently different to warrant recognition of a second species. with few exceptions the proportional measurements in S. and in the southwestern Atlantic (map 5). S. another character that supposedly differentiates S. media. which were collected around the tm-n of the century. falls within the normal range of variation for S. Variation in the median scallop on the anterior margin of the head was found to result from a comparable variation in the tip of the rostral cartilage. media examined. That this condition is not strictly a function of size is shown by the fact that one of the largest specimens of S. and snout to pectoral insertion). nana. Considerable variation was found also in the degree of development and/or presence of the accessory rostral cartilages. nana and S. male (UCLA 58-304) from Panama Bay. a single series of specimens sometimes showing all degrees of development. Of the various measurements listed for S. nana are about what one would expect in individuals of media around 1. head length. horizontal diameter of orbit. variation in configuration of the anterior margin of the head.

17-26). and further exploration will likely show the species to be more widely There men of is. Trinidad. the next closest place from which S. the this species distributed in the western Atlantic than presently appears. from the Gulf of Paria. containing eight embryos. head of a speci(SU 52716) from Vitoria. 1. the intervening area has not been weU collected. who lists (under the name "scoophead shark") a female. media has been recorded. pp. Since this locality is so far removed from Trinidad. in the Stanford University collection. suspicion. one might regard the record with However. long. Brazil. .125 mm. which was collected in April 1949.PROCEEDINGS OF THE NATIONAL 62 MUSEUM Springer (1949.

d) a and longer rostral cartilages. S. 21h). anterior-median pore patch on underside of head with the posterior margin less broadly rounded and without a pointed median-posterior extension (fig. corona and S. — Sphyrna corona differs from S. media in having a broader postorbital process to the chondrocranium. (Fig. . b. Panama Bay. TL. from Chiapas.6 Dorothea B. Characters mentioned in the diagnosis of the subgenus Platysqualus are not repeated here. between Panama City and Punta Gorda. a less deeply falcate anal fin with a more rounded apex (fig. tudes (figs. media.) of the innerorbital X natural size. 22g) a more broadly rounded . the shape of the anterior-median pore patch on the underside of the head is very similar in aS*. 18a). however. ex 63093).HAMMERHEAD SHARKS GILBERT 63 53-275 (9). Finally. Unlike S. and a slightly narrower mouth. about 0. to the angle formed by the juncture 18— Sphyrna corona: a. 7c. upper precaudal pit in smaller specimens (fig. 371 mm. Schultz. 402 mm. a drawn by Paul Laessle. fig. Mexico (FMNH 72521. rather than posterior. corona may not reach as large a max- .) Sphyrna corona also differs from S. media). media in having: A slightly longer snout (fig. 186). 22g. Sphyrna corona differs from the closely related S. with the anterior edge of this structure anterior. juvenile female. (The last character is discussed at greater length in the diagnosis of S. Diagnosis. slightly shallower anterior fontanelle. media differs from that species (see diagnosis of S. head of juvenile male. SU 11882 (holotype of Sphyrna corona). by b and anterior mediorbital cartilages (pis. tudes in much the same way that S. media. TL. Panama City. from Panama Bay Figure (UCLA 53-275).h).

media A comparison of Sphyrna corona with other members of the genus presented in table 1. a 395 mm. a 385 mm. male [USNM 87682]. p. type locality. p. of the species of Sphyrna (Platysqualus) tudes (Valenciennes) Figures 19. MUSEUM half as large as the largest specimen of S. 1822. 1841. — Distribution of species. West- ern Atlantic Ocean (including Caribbean Sea and Gulf of Mexico): . 1944. or the description of *S'. — (Range in size: 213-900 mm. the following Only those description has been condensed so as to avoid repetition.7 to 29.1 percent of by Sprmger. 7nokarran]. the greatest width measuring from 13— ?— 13 (count given for holotype 1^—1 — lo teeth ' ' and ^^_~_. original description [apparently based on two species. while type from Coromandel probably is S. tudes. 53. 1822. identification by reference to Valenciennes. 274-276. 22h.) characters in which actual counts or measurements are cited are For other characters see either the listed in the following account. (Illustration of head. 20. S. occurring only in the eastern Pacific from southern Mexico south at least to the coast of Colombia (map 5). 1940a. 225-226. TL. Description. pp. TL).) Sphyrna bigelowi Springer.^^ Range. media. media. as here recognized. (Original description. male from Uruguay [USNM 120751]. Uruguay holotype. Since most of the morphological characters in Sphyrna corona are similar to those found in S. mokarran. sphyrna tudes Sphyrna media sphyrna corona A O V ^ wmm Sphyrna media and Sphyrna corona (Solid = specimens examined.) Sphyrna tudes Mi'iUer and Henle.) . Brazil [MCZ 463].PROCEEDINGS OF THE NATIONAL 64 imum vol. is — ]Map 5. hollow = confirmed literature references. types from Nice and Cayenne are S. mm. 163) 8. tudes and *S. and a 900 mm. Specimens examined. paratypes. 23. diagnoses of S. — Sphyrna corona has one j^z^' most restricted ranges of any hammerhead. (In part. illustration. Plates 4. media. corona or the subgenus Platysgualus. Meristic data appear in table Head moderately expanded. 8 Zygaena tudes Valenciennes. 21z. female from Rio de Janeiro. pp. 119 because the largest specimen examined (672 size as S.) was little over that was seen.

1019). and Coromandel. media margin of head straighter (particularly in large specimens). Sphyrna tudes from the closely related S. p. Venezuela: 123217 (head). 1049) on which the illustration in the original description was based fin (pl. 55°04' to 55°08'W. India (specimen lost). Nomenclature. ANSP 516 (1). with a pronounced median notch. 54°54'W. 2110). Gulf of Venezuela. 06°22'N. which is characterized by the extending beyond the insertion of the pelvic sUts nearly equal in length. 480). French 1019 (lectoparatype of Zygaena tudes). 12.. (no further data). 55°05' to 55°10'W. (Oregon Sta. . United States (Mississippi): 195957 (8). Diagnosis. Gulf of Mexico. Guiana: Surinam: ANSP 377 (1). 120751 (paratype of S. and a large maximum size (up to 20 feet). first dorsal fin of larger specimens (600 mm. a Une drawn perpendicularly from differs in the following characters: Anterior apex of fin intersecting 219-937—67 5 base of fin. to 06°19. figs.. with the latter situated above the insertion of the pectoral fin.. 159197 159199 (1). —The original description of Zygaena tudes (Va- lenciennes. the first gill sht longer than the fifth gill slit. (1). Cayenne. 29°54'N. 159227 (2). 88°06'W. As has previously been shown. mokarran (Gilbert. Uruguay: 87682 (holotype of Sphyrna bigelowi). 1961). Anway Bay. 3539 65 MCZ 463 (paratype of S. France (cat. Nice. Cayenne. Mediterranean Sea: France: 1049 (lectotype of Zygaena Brazil: MNHN USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM MNHN tudes). characterized by the tip of the first dorsal not reaching the insertion of the pelvic fins. female (no.5'N. and a small maximum size (perhaps not over 4 or 5 feet) and the one presently called Sphyrna tip of the first dorsal fin fins. to 54°49'W. bigelowi). 225-226. 06°27' to 06°25'N. la and 6) was based on specimens. 1 mi. with the latter situated posterior to the insertion of the pectoral fin. 1822. no 1049). inner narial groove present. southeast of Paramaraibo Ught ship. 159229 (4). Kio de Janeiro. pi. 4).. pp. no.. 159228 (4). bigelowi). no further data. 226). 1961.5'N. corona and S. or more) more erect and more broadly triangular. 06°20. wide.—GILBERT HAMMERHEAD SHARKS NO. French Guiana (cat. these specimens comprise two species: The one currently called Sphyrna tudes (Gilbert. no further data. median indentation (fig.. I here restrict the name tudes to those specimens from Nice and Cayenne. anterior-median pore patch on underside of head divided into two broadly rounded lobes and separated by a deep. from Nice. 156721 (1). narial depression sUghtly deeper. the first and fifth gill . and designate as lectotype of Zygaena tudes the 346 mm.— Characters mentioned in the diagnosis of the subgenus Platysqualus are not repeated here. in the Museum National d'Histoire Naturelle.

with the broadly curved outer edge curving sharply inward. not noticeable in smaller specimens. Brazil h. dermal denticles of MCZ 463. Characters men- tioned in the diagnoses of the subgenus Platysqualus and of S. —Meristic data appear in table 9. c. pointed at tip.) Figure (MCZ 19. apical view.Ar^ 'Jl^S^5==^ Sphyrna tudes: a. i. . p. narial flap rather mouth in narrow (particularly in smaller specimens). 119 of the genus is 1. MUSEUM Sphyrna tudes with other members vol. outer posterior corner of lateral expansion of head situated posterior to UMJJJU'. anterior to corner of larger individuals. 410.5 percent of percent of distance from narial openings to median notch. g.5 to 4. tudes are not repeated here. about 17 X A. 886 mm.5 to 31. about 34 X . 463). Head moderately expanded. fifth lower tooth. upper and lower teeth. Origin of first dorsal fin slightly posterior to axil of pectoral a perpendicular line drawn ventrally from fin. greatest transverse distance between corners of mouth from one-fifth to one-fourth of greatest width of head.third of head width in larger specimens. d. from Rio de Janeiro. the greatest width measuring from TL.0 27. left side.— PROCEEDINGS OF THE NATIONAL 66 A comparison of presented in table Description./. denticle. fifth upper tooth. about 34 X. twelfth upper tooth. {c-f. length of snout about two-fifths of head width in smaller specimens to about one. dermal denticle. side view. a very short labial furrow present at corner of lower jaw in larger specimens. of same specimen. TL. origin intersecting anterior . about L5 X. dermal (From Bigelow and Schroeder.) corner of mouth in smaller specimens. inner narial gTOOve extending 3. 1948. e. about 3 X. subadult male. twelfth lower tooth.

about 0. its greatest height from two-thirds to four-fifths length of base and from two-thirds to four. second dorsal fin base slightly greater than one-half of anal fin base. anterior margin of first dorsal fin broadly and evenly curved. Sphyrna tudes: a. Schultz. head of small adult. . fifths of greatest height of anal fin. apex of pectoral broadly pointed.18 X. origin of second dorsal fin situated about two-fifths of way back above base of anal fin. length of pectoral fin base from two-thirds to three-fourths length of anal fin base. from Gulf of Venezuela (USNM 123217). pectoral fin two-thirds as broad as long. TL. lower margin broadly but distinctly falcate. length from a third to half again greatest height of fin. second dorsal fin tail. with the part nearest the apex a little more strongly curved upper distal margin of first dorsal broadly convex. c.3 X. and extending from two-thirds to three-fourths of distance from posterior its part of second dorsal base to upper precaudal pit. base of first dorsal fin three-eighths to four-ninths length of head and from a third again to not quite twice length of pectoral fin base. juvenile female. h. distal margin of pectoral fin nearly straight or very slightly falcate. 402 mm. outer third distinctly convex. from Mississippi (USNM head of same specimen. lobe of second dorsal fin long. pectoral fin from one-half to two-thirds length of head (higher values usually apply to larger individuals). (Drawings by Dorothea B. particularly in larger mdividuals. about 0. inner two-thirds of anterior margin of pectoral fin straight.) FiGURE 20. 195957). lobe of first dorsal fin a little less than one-half length of first dorsal base.— HAMMERHEAD SHARKS —GILBERT 67 third of pectoral fin.

upper margin of caudal fin slightly convex. mokarran. h by Paul Laessle.PROCEEDINGS OF THE NATIONAL 68 MUSEUM vol. media. 119 inner corner of pectoral fin narrowly rounded. anal fin deeply falcate toward apex. lower posterior margin of terminal part of caudal fin nearly straight or slightly falcate. height of > S d f — Figure 21. 356 mm. a-f. from Florida (USNM 106543). 675 mm. 900 mm. 880 drawn by Dorothea B. S. female. both terminal and lower apices of caudal broadly pointed. length of caudal fin nearly a third of total body length. S. S. d. t. anal fin from oneeighth to one-third again as long as pelvic fin. /. g. viedia. S. anal fin from one-fifth to one-third greater than height of second dorsal fin. S. mm. 825 mm. lower lobe of .) e. b. length of pelvic fin from two-thirds to nine-tenths length of anal fin. length of anal fin base from one-third to one-half again as long as pectoral and pelvic fin bases and about twice as long as second dorsal fin base. S. male. male. h. lewini. zy gaena. (Figs. 371 mm. 580 mm. Schultz. from Chiapas (FMNH 63093). female. tiburo. 615 mm. from Texas (USNM 127106). length of pelvic fin base from five-ninths to seven-eighths length of anal fin base. female. from Chiapas (FMNH 72521. ex. from Virginia (USNM 119699). 495 mm. anterior and distal margins of pelvic fin nearly straight. nearly straight toward tip of lobe. male. i female. from North Borneo (FMNH 21836). figs. from Panama Bay (UCLA 58-304). S. c. S. Appearance of upper precaudal pit: a. from Texas. g. corona. tudes. terminal part of caudal about one-fourth of total length of fin. male. Sphyrna blochii. i. from Surinam (USNM 156721). 63093). female.

but with a definite cusp. lower caudal lobe rather erect. overlapping but little. lower caudal lobe about one-third as long as upper lobe.^ ^ J 1 ^^ latter count counted. without a definite cusp.. Teeth . rostral fenestra usually absent (present in one individual examined) anterior fontanelle (viewed dorsally) deep and broadly V-shaped. median indentation. posterior margin of lower lobe nearly straight. broad. mouth — . the blades thick and rather strongly arched. and as many sharp-pointed marginal teeth (especially deep and pointed in small individuals) on denticle. Variation. Individuals gray or grayish brown dorsally. Anterior margin of rostral node of chondrocranium with a broad. obtusely pointed wings present on sides of rostral node. anterior wing of olfactory cartilage prominent. pedicel long and rather slender. sharp-topped ridges. pointed. and ^ 16-1-15 15-1-15 postorbital process terminating in corner of which may . Dermal denticles (examined from upper part of back beneath first dorsal fin) evenly and closely spaced. Total number of vertebrae numbering from 195 to 202 (iBve specimens examined). upper teeth oblique. HAMMERHEAD SHARKS NO. female from Nice. the anterior be either broadly pointed or narrowly rounded. prominent. and two to three rows along sides of lower jaw. becoming progressively more pale below. the distance from the angle included by the upper and lower caudal lobes to tip of latter going about IK times in anterior margin of lower caudal lobe. . France. and projecting medially. rounded. indicates that the Mediterranean and western Atlantic populations of Sphyrna tudes are morphologically indistinguishable. an angular knob. the outermost two very short. a 346 mm. in lower jaw teeth 2 to 7 or 8 longest. accessory rostral cartilages usually present. the outer teeth very low.^ (two specunens . rather thick. sloping posteriorly at about a 25 to 30° angle from a perpendicular to the body axis. no markings on fins. The only unfor lectotype) corners of . ^ J 16-0-16 .. length of denticle (measured to tip of median marginal "tooth") less than width. body vertebrae numbering from 107 to 110. 3539 —GILBERT 69 caudal appearing relatively narrow and long. Examination of the lectotype (designated elsewhere in this paper) of Zygaena tudes. mokarran). and the posterior corner of which forms a broadly rounded angle. with an indistinct median notch on the posterior margin. deep. teeth 4 or 5 to 11 longest. tip of preorbital process short (though not as short as in S. small specimens with three. median tooth on denticle slightly longer than more distal teeth. with a sharp angle on the anterior edge. anterior margin of lower caudal lobe weakly convex.. there being from 12 to 25 more body than caudal vertebrae. larger specimens usually with up to five. 15-0-15 . one or two series of teeth functional in alternating rows along sides of lower jaw. becoming increasingly so toward in upper jaw.

480). the occasional presence of such a fenestra in <S. S. lewini. 1950b. f. media. (Drawings Laessle. H g ' side of head: a. S. In the western Atlantic it occurs from Uruguay (Springer. S. d. Sphyrna blochii. 1961. S. by Paul e. 1950a. zygaena and S. 1031). p. France (the type locality) and Livorno. tudes. corona. 32. Figure c. Italy (Tortonese. p. 8b). g.) — Range.PROCEEDINGS OF THE NATIONAL 70 usual feature of the lectotype is MUSEUM the presence of a small rostral fen- However. 274) to the northern Gulf of Mexico (Gilbert. S. tiburo. 1944. b. h. pp. . 22. S. —Distribution of pores on ventral S. zygaena. p. blochii (in which species it is ordinarily absent) suggests that the type specimen is merely a variant with regard to this estra (pi. mokarran. 31. Sphyrna tudes is restricted to the western Atlantic Ocean and the western Mediterranean Sea. character. In the western Mediterranean it is recorded from Nice.

3.. bridge Natural History. The East Pacific barrier Evolution. 545-554. pp. and atlas P. 113-154. 2. vol. Thomas William 1904. pt. 389 pp.. 1960. 5. 4 pis. 171-180. 71 . 3. Jean 1951. vol. and Springer. Res. Nat. Sharks. Switzerland. Tableau encyclop^dique et m^thodique des Ichthyologie. 3. distribution syst^matique Philomatique Paris. both recent and Classification of fishes Acad. 1948. Naturgeschichte der auslandischen Fische. Soc. Inst. Fishes. no. identification. Leopold 1785. 243-245. 139-537. Richard H. Baughman. Mem. Mich. no. Ann Arbor. Notes on western North Atlantic sharks. pp. Copeia. 215 pp. vol. In Fishes of the western North Atlantic. Literature Cited Aqassiz.. 8. 4. no. 9 pts. faune ichthyologique de Chine. Louis Recherches sur les poissons fossiles. Bonnaterre. study of the sharks of the suborder Squaloidea. pp. 3. 324 col. 1950. Copeia. Midi.. 87-517.] by and Schroeder. 1. 1. . A Zool. distribution of marine shore fishes. B. [not 1950]. 2. Imprimerie de Petit-pierre. Marie Ducrotay de Prodrome d'une nouvelle Bull. Henry fossil. no. pp. Zool.. 1-150.. pp. 241 figs. 3 text figs. Mar. Sci. L. pp. and the 4. vol. Blainville.. 1960. pis. no. 1. 105-124. Bridge. Pieter 1873. Inc. trois regnes de la nature . exclusive of the systematic account of Teleostei. pt. in (3 vols. vol. 190 text figs. J. volume on fishes.. Franc. 1. Fishes of worldwide (circumtropical) distribution. text Berg. Joseph 1788. 345 pp . 102 pis. 44. Backus. [ReEdwards Brothers. 1944. 3. 59-546. Dierk. 117. BiGELOw. Neder. 1961. and L0ppenthin. Bull. Cadenat. 247 pp. Stewart Biological and economic notes on the sharks of the Gulf of Mexico. M6moire sur la du regne animal. Trav. pp. B. 106 text figs. 16 text figs. Blegvad. Briggs. Bloch. H. In Danish Scientific Investigations Fishes of the Iranian Gulf. Neuchatel. 1957. no. pp. vol. 96-152... figs. 2. Poissons de Mer du S^n^gal. Leo 1940. John C. URSS. 1960.). Inst. vol. In Cam- pp. Comp. 19 S. Sears Found. Tijdschr.. Amer. Henri 1816. 1947. vol.. Initiations Africaines 3. William C. Bleeker. with special reference to those of Texas and with a key for their 1838. 135 text in figs. Mus. 15. no. printed in English. pp. Iran.. 11 pis. d'Afrique Noire. 1960.

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102 pis. 574 pp. James A. 77 pis. pp. Ogilby. Samuel 1923. 10th McCuLLOCH. Zool. pp. Augusto Vertebrados. and 824 pp. opera intitolata: Descrizione de' Crostacei.. MiJLLER. vol. 9. Pacific coast of Mexico. rappresentati in figure . 215 Hist.). Soc. . Murray. Brisbane. Syst. 8. Synopsis der Fische des Rothen Meeres. vol. vol. (Zool. families and genera of recent and Unpublished manuscript. and Hildebrand. vol. 1957. 103-105. J. draft synopsis of the orders. and Fishes of Queensland.. Y. pp. 1916. Selachians. 6. no vol.. The marine fishes of Panama. Brisbane. 1. Friedrich Sigismund 1836. Porto. Henri db 1868. 75 pp. P. 44 pp. ed.. Zool.. registrate nell' pi. vol. KuMADA. . Meek.. Mem. A. Nat. La p^che et les p^ches. vol. Leitckart.. In Fauna marinha de Portugal. 441-668.. 2. Mus.. Untersuchungen uber die ausseren Kiemen der Embryonen von Rochen und Hayen.. pt. xi pp. land Mus. G. Systema naturae. Giovanni Domenico 1847. ... 1. deir Abate St. NiNNi. 1935. (Conte) Alessandro Pericle Revista critica delle specie dei pesci adriatici descritti 1872. Field Publ. In Nouveau dictionnaire general des poissons. Hist. Whitley. A 1925. Ser. land Mus. Theil. 1841. 1. Johannes. dall' le Abate Chiereghini Ven. Queens- F. list of the fishes recorded from Queensland waters. A new species Journ... 2. Ges. de' Testacei e de' Pesci che abitano Lagune S. 5. 1. Wien.- T. Nardo. 1908. Chiereghini di Chioggia. figs. 24 pis. Bot. 125-182. 1. Carl 1758. [Photo by British Museum of Natural History. R. 21. Nobre. vol. F. 29-40. no. J. [printed in double column]. pp. 5 pis. dissimili^) Nat. 1. Mem. 128 cols. Ixxxiv + Norman. R. and Hiyama. Queensfishes in the Queensland museum. pp. from the Kurrachee harbour.] vertebrates. pt. 1-41. Brisbane. Linnaeus.. G. Miller. fishlike offset copies distributed Douglas New or little known Ann. pt. applicata per commissione govern- ativa. 1937. Check-list of the Cephalochordates. 15. Verh. Robert Rush Utilization of x-rays as a tool in systematic zoology. 60 pis. Queensland Mus. vol. Sinonimia moderna delle specie xxii + 200 pp.. 1-330. ein Beitrag zur Entwicklungsgeschichte der Abtheilung der Knorpelfische angehorenden Plagiostomen. A J. La BLANCHfcRE. pp.PROCEEDINGS OF THE NATIONAL 74 Klitnzinger. Systematische Bcschreibung der Plagiostomen. Marine fishes of the MUSEUM 2. Clodiense. 1887. e Golfo Veneto. 859 pp. no. 70-98. Carl Benjamin 1871. pis. and Henle. nell' opera MS. pp. Seth E. of Zygaena Bombay {Z.

Nat. Zool. 1 pi. 1815.HAMMERHEAD SHARKS NO. no. An 274-276. Proc. Sphyrna 1949. 16 pis. Bull. Mar.. 5. Ann. (1938). 38-49.. 1912. with a list of other sharks Proc. 1941. 1810. 10 Risso. with the description of a new species. 1 text fig. pp. 11 pis. vol. Henri Emilb 1891. 564 pp. + 1826. U. C. bigelowi. (1940). hammerhead shark from off the Atlantic Florida Acad. pp. Inst.. 722-758.. Proc.. 46-52. naturelle de Madagascar. 3. 1. pis. 34. 1940a. L. pp. Felipe In Gundlach. of some Florida sharks. Florida Acad. Histoire physique. 70 pp. Notes on the sharks of Florida. vol. 1944. 34. 10. 27.. outline for a Trinidad shark fishery. vol. Poissons. Three new sharks of the genus Sphyrna from the Pacific coast of Stanford Ich. Sadowsky.. 1. Stewart 1939. Jacques Poissons du Mus6e de Naples provenant des expeditions du "Vettor Pisani" et du "Dogali" et de la mer Rouge. 161-169. tropical America. 61 pis. Soc. WiLHELM PeTER EdUARD SiMON Neue Wirbelthiere zu der Fauna von Abyssinien gehorig. Tate A 1906. Fish.. Lewis The sharks and rays 1916. new ser. la fauna puerto-riquena. 48 pp. vol. Journ. North Carolina. 1835. 33 pis. classification of the Selachian fishes. Victor 1965. The sea fishes of southern Africa. Brazil. 1940.. 317-350. Madrid. Poet. Ramsay. 3539 —GILBERT 75 Pellegrin.. Anal. Sci. Sci. vol. taken in Port vol. 1953. Paris. 17-26. et politique J. Histoire naturelle des principales productions de I'Europe M^ridionale et particuUSrement de celles des environs de Nice et des Alpes Maritimes. 15. 2 pis. of Beaufort. 239-284.. ou histoire naturelle des Alpes Maritimes. Mus. Rafinesque. pp.S. Edward Pierson Notes on Galeocerdo 1881. vol. hammerhead shark Proc. . B. 3. no. 3rd ed. Linn. 224 pp. Regan. 5. 188-194. pp. 1-12. vol. Constantine Samuel Indice d'ittiologia Siciliana. Soc. pp. Antoinb Ichthyologie de Nice. 6 figs. Napoli. figs. Springer. xvi 480 pp. In Grandidier. sex ratio and seasonal distribution Copeia. pis. with notes on Sphyrna mokarran from New South Wales. Sauvage. 102 pis. vol. Gulf Caribbean Fish. Bur. 95-97. ou tableau de I'univers et de corps organises. + RtJPPELL. pp. pp. A new species of 3. pp. session. a new coast of South America. New South Wales. 16. Jackson.. 5. rayneri.. The hammerhead sharks of the Littoral Zone of Sao Paulo. Apuntes para Peces. London. Sci. The figs. Analyse de la nature. Soc. pp. Zool. of the genus Sphyrna. (1914). Radcliffe. no.. 7 text 1940b. xxii 388 pp. pp. vol. 1-11. Proc. 1810. no. Washington Acad. vol. pp. 3. Smith. Bull. Sci. Bull. 2nd ann. 1881. 543 pp. 9-41. Espanola Hist.

377-399.] Studer. Nat. Petri Artedi . 11 text figs. no. Algemeene Konst. Handboek der dierkunde. Mass.. Mus. und Geologie. 3. Indie. pp. A survey of vertebral numbers in sharks. Humphreys [Occurrence of the Chatham. 1964. 12. v-vl figs. Walter 1907. C. VAN Hasselt. Tortonese. 73-96. Springer. 115. vol. 20. Franz 1870..M. 8 bijvoegsels pis. 2. after a study Ann. 3 pis. Inst. Storer. 1. Boston Soc. Wien. 1839. or history and classification of fishes. 61. 1950a. . Victor G. and Fosberg. 35-250. A revision of the carcharhinid shark genera Scoliodon. 1839. 4. no. Enrico vol. J. 1823..S. 2.. Mus. Mem. and 99 reptiles. vol. Lehrbuch der Zoologie. + 368 pp. 559-632. Valenciennes. . . pp. vol. 2. U. famiglia Sphymidae (Pesci martello). pp. 1030-1033. Joum. vi + 322 pp. Johann Julius 1792.. 8 pis. no. 3. vol. Theophil. Hist. Cecil 1933.. 70-71. 1: The craniology and neurology of a hammerhead shark. Zool. Zur Fischfauna des Senegal. 119 Springer. figs. Nat. 1-9. Genera Piscium Ichthyologica. no. pp. Sitzber. Rhizoprionodon.. Proc. VON BoNDE. jaar 1823. VAN DER HOEVEN. Artedi renovati .en Letter-bode voor het 188-419. Natuurk. and U.. Hist. Mus. pp... Sphyrna (Cestracion) zygaena (Linnaeus). 3496. 1-4. 314-317. Nat. VoLZ. 1889. Victor G. vol. tweede verbeterde uitgave. 33 pis. amphibians. Walbaum. David 1848. "Ga- 1889. Comp. pp. no. 66. vi + 452 pp. of the types. monocardian animals. 1957. Die Forschungsreise S... Loxodon. . vol. A. 1838 figs. Copeia. Pt. 1950b.S. Zoologie zelle" hammerhead shark at Provincetown and at Proc. no. ser.. 1874 bis 1876 . Akad. J. pp. Univ. 533-583.. . 3 in P. Wiss. corologia e sistematica della Boll. 292-295. Sur le sons-genre marteau. vol. Starr. William The natural 1838. Hist. Theodore 1957. and Garrick. met en aanmerkingen door Leuckart. 135 vol. Nederl. A note on the hammerhead shark. pis. Torino. (8 col. Pt. Proc. vol. 2. JaNUS 1855. Thienemann. 2.. 2: Evoluzione. J.) Swainson. 723 pp. 3 in . 9. Achille 1822.S. vol. 1. Tijdschr. Nat. no. MUSEUM PROCEEDINGS OF THE NATIONAL 76 vol.. 2 pis. Neurol. Mag. Nat. Sphyrna tudes Val. Mus. Filter Steindachner. 39 pp. 116.. Catalogue of the fishes of Sumatra. F. text figs. Studi sui Plagiostomi. vol. 3493. pp. 3.. Walter paper electrophoresis of serum proteins from sharks.. Zygaena. Contributions to the morphology of the Elasmobranchii. vol. 222-228. pp. 58. Friedrich August Ludwig 1828. ed. pp. 1964. pp.

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78 PROCEEDINGS OF THE NATIONAL II MUSEUM .

— HAMMERHEAD SHARKS Table 2. —GILBERT 79 Measurements of Sphyrna blochii (proportions expressed in thousandths of total length) .

— PROCEEDINGS OF THE NATIONAL MUSEUM 80 Table 3. Measurements of Sphyrua .

— HAMMERHEAD SHARKS Table 4. —GILBERT Measurements of Sphyrna zj'gaena {proportions expressed 81 ^ .

— PROCEEDINGS OF THE NATIONAL 82 Table 4. Measurements of Sphyrna zygaena sandths of total length) MUSEUM (proportions — Continued expressed in thou- .

Measurements —GILBERT of Sphyrna lewini (proportions expressed Continued of total length) — 83 in thousandths .— HAMMERHEAD SHARKS Table 5.

— PROCEEDINGS OF THE NATIONAL 84 Table 5. Measurements of MUSEUM Sphyrna lewini (proportions expressed in thousandths Continued of total length) — .

— Table 6. Measurements of subspecies of Sphyrno tiburo {proportions expressed in thousandths of total length) Continued — .

- Measurements of Sphyma media (proportions expressed in thousandths of total length) .Table 7.

Measurements of Sphyma —GILBERT 87 tudes (proportions expressed in thousandths of total length) .— HAMMERHEAD SHARKS Table 9.

Measurements of Sphyrna tudes {proportions expressed in thousandths Continued of total length) — .— PROCEEDINGS OF THE NATIONAL MUSEUM 88 Table 9.

NAT K MUS VOL 119 GILBERT -PLATE 1 .PROC. US.

GILBERT— PLATE 119 I ^ ^v<^ c 2 . NAT. VOL.PROC U. MUS.S.

GILBERT— PLATE 119 i 3 . NAT. MUS. VOL.PROC.S. U.

la.S. MUS.PROC. 12. b). NAT. pi. GILBERT — PLATE 119 t '"M ^ ^ Original illustration of Zygaena tudes. U. from Valenciennes (1822. . VOL. figs.

U.S. MUS. NAT.PROC. 119 GILBERT-PLATE 5 . VOL.

PROC.

U.S.

NAT. MUS. VOL.

GILBERT— PLATE

119

Radiographs of chondrocrania of several species of Sphyrna: A,

from Florida

(USNM

C, S. zygaena, 590

186087); B, S. lewini, 447

mm. TL, from

Italy

(USNM

S.

mm. TL, from
28452).

mokarran, 560
Liberia

6

mm. TL,

(USNM

179706);

PROC.

U.S.

NAT. MUS. VOL.

GILBERT^PLATE

119

Radiographs of chondrocrania of several species of Sphyrna: A,

TL, from Nicaragua

(USNM

from Mexico
ex. 63093).

388 mm.
mm. TL, from Mexico

S. liburo tiburo,

78180; B, S. tiburo vespertma, 545

(UCLA 51^); C, S. media, 337 mm. TL,
mm. TL, from Mexico (FMNH 72521,

371

7

(FMNH

63093); D,

S. corona,

PROC.

U.S.

GILBERT— PLATE

NAT. MUS. VOL

Radiographs of chondrocrania of Sphyrna tudes: A,
S.

bigelowi)

S. tudes)

from Uruguay

from France

(USNM

(MNHN

a

specimen 390

87682); B, a specimen 346

mm. TL (holotype
mm. TL (lectotype

8

of

of

1049).

I

S. NAT.PROC. wide. from Guinea (USNM uncat. head 445 mm.). from Guinea wide. B.). lewini. VOL. GILBERT -PLATE 119 Radiographs of chondrocrania: A. U. head 425 mm. 5. couardi. (USNM 9 . MUS. uncat. S.

from Guinea wide. NAT. head 445 head 425 mm. U. i .). GILBERT— PLATE 119 S. mm. 10 B uncat. lezuini. MUS. VOL. S. from Guinea (USNM (USNM uncat. wide. Photographs of heads: A.S.). coMarJi.PROC.