THE STORY OF THE DEVIL-FISH^By THEODORE GILL
One of the most remarkable of animals is the great Ray, mostwidely known as Devil-fish, but which bears also several othernames.Devil-fish is a name by no means restricted to any one of the Rays,for it is well known in connection with the gigantic Cuttlefishes andis also used locally in England for the Angler {Lophius piscatorius),and in California for the Gray whale {Rhachianectes glaucus).xA.mong the Rays the name is applied not only to all of the samefamily as the great fish, but also, in some places (for instance. NorthCarolina and the Gulf of Mexico), to species of Eagle-rays. Sea-devil may be considered to be a natural variant of the same name,but it has also been used for the same animals as Devil-fish and evenfor those of another family, the species of the Sharks known asSquatina.Vampire originated in the form "Oceanic A'ampyrc" as a selectivename and was given by Dr. Samuel L. Mitchill, in 1823, as thepopular name for his Ccphaloptenis vampyrus. He claimed that
"this fish being perhaps the largest of the Rays, as the vampyre is ofthe bats, or vespertilios, the name vampyrus may be attached." Thename has somehow been taken up and found limited currency in cer-tain localities where the fish abounds. Thus C. F. Holder' has re-corded that it is in use in southern Florida. When, during a nighton the water about Garden Key, he heard "a rushing, swishing sound ;then a clap as of thunder," a negro boatman exclaimed "Vampa fish,sah," and later alluded to it as "Sea Vampa" or collectively as
"Vampas."
^ Every well-known fish student is more or less frequently asked some ques-tion or questions about the Devil-fish. Not infrequently the student is at a lossfor an answer. The requisite information may have been published, but toobtain it perhaps hundreds of articles may have to be examined. After asearch through such articles the present paper has been compiled and willfurnish answers to many of the questions that may be propounded. It will atleast serve as a basis for investigation and a repertory of what has been ascer-tained or thought to be facts.
' Big Game at Sea, 1908, pp. 2, 3, 4. 155
156 SMITHSONIAN MISCE;lLANEOUS C0LLE:CTI0NS VOL. 52Sea-bat was found by Holder to be in use in the same locality asVampire. When the negro Paublo exclaimed "Sea Vampa, sure,"the Seminole chief in the same boat corroborated his identificationrather than contradicted by exclaiming-, "Sea-bat. . . . They
The Devil-fish. After a photograph/
^ The iconography of the Devil-fish is very defective and the figures herewithgiven are merely provisional. The plate first given by Jordan and Evermann(1900), later reproduced by Fowler (1906), Hugh Smith (1907) and others, isquite inaccurate so far as the tail is concerned. Instead of the tail being muchlonger than the body, as therein represented, it is only about 6/10 as long.Elliott (p. loi) especially criticized De Kay's "characteristic, viz., tail longerthan the body," and affirmed "that the length of the tail is, to that of the body,as six to ten." He had examined "almost twenty individuals." The illustra-tion cited was drawn in Dec, 1894, but the present writer was long unable tolearn what was the basis of the figure. He finally traced it to De Kay, whopublished a composite figure based on Mitchill's and Lesueur's plates. Thereis no specimen of the Devil-fish in the National Museum. The figures herepresented are (i) the old one with the tail modified to suit photographs andElliott's description ; (2) one drawn after the former outline with the undersurface represented from a photographic illustration in Holder's work, and(3) a reproduction of a photograph of a fish caught in 1869 or 1870, during acruise in the Pacific of a revenue cutter (Captain Freeman commanding). Thelast was taken while the fish was suspended from a tripod and the droopingfins may have been partly at least due to the suspension. That fish was about13 feet wide. The photograph is very obscure behind and the reproductionconsequently is unreliable, as are the other figures. Seven photographs orreproductions are at hand, but all are too obscure behind for guidance. Agood one is extremely desirable as are also exact data as to relative propor-tions and weight. All published are deficient. A special article on the subjectwill follow.
NO. 1816 STORY OP THF, DEVIL-FISH—GILL 157jump five—yes, eight—feet high." Bat-fish and Black-bat aresometimes used variants.Another name for the monster Ray has been borrowed from theSpanish. Among the fishermen, and especially the pearl divers ofCentral America and western Mexico, it is known as the Manta;this is a Spanish term, meaning originally blanket, and was given bythe fishermen of parts of Spain and the island of Mallorca to a spe-cies of the Mediterranean^ and extended thence to similar fishes ofother regions. It has been explained that the name was given by theSpaniards of America to the Devil-fish because it was alleged rohover over and cover a fisherman at the bottom as a blanket prepar-atory to killing him for good. The belief, indeed, that the Devil-fish may so attack a man is not only widely spread, but of an ancientorigin.Such an idea, however, is contrary to our knowledge of the fish.Like several other of the gigantic selachians,- its diet is in almostinverse ratio to its size.Inasmuch as Devil-fish is the best known of all these names andhas been long current in story as well as in works on natural history,it will be retained here and will be used for the great fish best knownas such, as well as for its congeners of smaller size. The speciesespecially called Devil-fish is one of a number having the same essen-tial characters and all designated in a general way as Devil-fishes.
IIThe form of the Devil-fishes is extraordinary ; the body, exclusiveof the tail, is about twice as wide as long ; the tail, however, corre-sponds to the hind part of the body in distant relations of the Devil-fish. Different as the animal is from Sharks generally, there is orhas been every gradation from an ordinary Shark to the Devil-fish.
^The Manta of Mallorca, or Majorca, is the Mobiila giorna, and is the Vaccaor Vaca (Cow) with various qualifications of some other localities in theMediterranean. It is also the Bous of Aristotle. The names Vacca and Bousallude to the horn-like caropteres or head-fins. The species is said sometimesto reach a width of 28 feet. Cams, in his Prodromus Faunae Mediterraneae(11, 1893, p. 520), specifies "Longit. 1.5-3 m" Pellegrin in 1901 (Bull. Mus.Hist. Nat., VII, 327) noticed one 5m. 20 wide, and 4m. 15 long. There is recordof one 28 feet wide and 21 feet long and "estimated to weigh a ton" (Z06I.,1899, p. 146). The data are insufficient and a fish of the dimensions notedmust have weighed very much more than a ton."The gigantic Basking Shark {Cctorhinns maximus) and the still largerRhinodon {Rhineodon typus) of the Indian Ocean subsist mainly on theminute crustaceans and other animals living near the surface of the ocean.
158 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 52
Fig. 41.
Fig. 42.
Fig. 43-
Fig. 44-
Fio. 45-
I'u.s. 41 .\.\D 42.^Squaliis acantliias. I'lcs. .\,} TO 45.—A'/(u;(;ta/».v k'niiginosiis.
NO. 1816 STORY OF TIIK DEVIL-FISH—GII.I, 159
.^^
^>/ ::-;' ;; Fig. 46.
Fig. 47
^^ > M
Fig. 48.
Fig. 46.-Raja erinacea. Figs. 47 and 4^.~Dasyhatis sabina.
l60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 52A few forms still living exemplify the manner in which the extrememodification of the last has been attained ; these forms, it is true,are not in the direct line of descent, but they are not very far off.The common Dog-fish of the New England coast (Squalus acan-thias) has a slender tail, but there is a regular gradation from thepreanal region, or trunk, into the postanal, or tail, and the pectoralshave the slender bases characteristic of the Sharks generally. TheGuitar-fishes (Rhinobatidce) still have the regular gradation of thetrunk into the tail, but the pectorals have a broad basis of union withthe body and head, and a narrow disk is thus formed. In the ordi-nary rays {Raiidce) the tail has become disproportionately slenderand the disk wider and more sharply differentiated ; in the Sting-rays (Dasybatidce) the tail has almost entirely lost its muscular de-velopment, but the disk is much like that of an ordinary ray. Thetail of the Sting ray is essentially like that of the Devil-fish, but in theDevil-fish the disk has become extended sideways into acutely angu-lated and wing-like fins. The homologies of the respective parts arethus evident. In the course of evolution, more and more resort hasbeen had to the pectoral fins for progression and the tail correspond-ingly disused; the culmination has been reached in the Devil-fishes,which progress by wing-like flapping of their pectorals and the tailis carried inert behind.The tranformation of shark-like forms into the ray-like type musthave commenced early in Mesozoic times, for well-developed repre-sentatives of the Dasybatids and Myliobatids were living in the Cre-taceous epoch and were abundant in the Eocene. It has been be-lieved that no fossil remains of Devil-fishes have been found, orrather identified. If this had been a fact, it might have been partlyexplained by the pelagic habitat of the species and partly by thereduction of teeth and spines, the parts most likely to be pre-served. There is, indeed, one record of an extinct form which,however, only takes us one stage back in the geological series.The record is of a supracaudal tubercle from the "phosphate beds"of South Carolina, which are supposed to be of post-Pliocene age
;
the tubercle has been considered by Joseph Leidy to represent anextinct species closely related to the living Devil-fish of the sameState and has received from him the name Ccratoptera unios; it wasdescribed and figured in 1877 in the Journal of the Academy of Nat-ural Sciences of Philadelphia (2nd ser., viii, 248-9, pi. 34, figs, i, 2).The individual development of the fishes is to a large extent par-allel with the evolution of the type from the shark-like form to theray-like one.The Devil-fishes form a familv of rav-like Selachians to which the
NO. 1816 STORY OF TIIK DEVIL-FISH—GILL 161names Ccplialoptcn'dcc, PtcroccpJialidcr, MohuUdcv, and Maiitidcchave been given. Mantidce is that used for it by most recent Anieri-
•5\iS >v.
w
Fig. 50.Figs. 49 and 50.—The Devil-fish. After Jordan and Evermann.(With reduced tail.)
•can ichthyologists, as Jordan and Evermann, but it had been previ-ously taken for a family of insects. MohiiUdcc may be used here.
"The essential external characters of the family follow
:
l62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 52
MOBULID.^The jMobnlids or Devil-fishes inckide the largest as well as thewidest of rays. Behind the anus the tail is abruptly attenuated anddeveloped as a whip-like appendage without efficient spines. Themouth, instead of being inferior, as in other types, is in front, andthe jaws have weak teeth or are partially toothless. The pectoralfins are extended outward in a w'ing-like manner, and long, flexible,horn-like processes or fins are developed on each side of the headand bound a preoral space. These processes (caropteres, head-fins,or horns) can be used for grasping, and a number of cases have been
Fig. 51.—Tail of the Devil-lish. After Holmes. (Proceedings Elliott Societyof Natural History, I, pi. 3.) About half natural size. i. Knob andbase of tail. 2. Bone with the small spine as extracted from theknob. 3. Upper view of the same with the posterior spinelet (inwhite).
recorded of a Devil-fish seizing the anchor of a vessel and runningaway with both anchor and vessel for some distance, to the wonderand fear of the sailors. The spines about the base of the whip-liketail, characteristic of the nearest relations of the Devil-fishes, theSting-rays and Eagle-rays, are reduced in size and sometimes to aminimum in the Devil-fishes. In the typical species the spine isquite rudimentary and concealed in a subosscous swelling at the baseof the tail behind the small dorsal fin.Further, the Devil-fishes are peculiar in the possession of pre-branchial organs, to be noticed later.
NO. 1816 STORY OF THE DEVIL-FISH—GILL 163
IIIThe Devil-fishes are inhabitants of warm-water seas. They areto some extent pelagic, though, as a rule, they appear not to extendfar out into the high seas. They belong to the category of tropico-politan forms, some one or other species occurring in every tropicaland every subtropical sea. Besides, some may venture far beyondthe limits of the Tropic of Cancer or of Capricorn, one wanderingoccasionally as far as New York and another into the MediterraneanSea.If we may also believe Turner-Turner, "a characteristic pose isthat of lying motionless, or at most with its disk slightly undulatingwith respiration, in the sand just under the water. Sometimes, in-deed, they are found a yard or so above low-water mark, in pits oftheir own making." This observation needs confirmation for Devil-fishes, although applicable to Sting-rays. But certainly they requireto rest on the ground, and sometimes, wdien harpooned, they descendand (to use a term of the angler) sulk on the bottom. Elliott re-marks that at times one "plunges desperately for the bottom, to whichhe sometimes clings for hours." But they are best known as active
—
and very active—frequenters of the surface waters.Another characteristic of a Devil-fish's action is a tendency toturn somersaults. According to Elliott,^ "It is a very curious exhibi-tion. You first see the feelers thrown out of the water ; then thewhite stomach, marked with five gills, or branchial apertures, oneach side (for the fish is on his back) ; then his tail emerges. Aftera disappearance for a few seconds, the revolution is repeated, some-times as often as six times. It happens occasionally that in makingthese somersets the fish does not rise quite to the surface, but isseveral feet below ; so that his revolutions are detected by the ap-pearance and disappearance of the white or under part of his body,dimly seen through the turbid water in which he delights. Some-times, indeed, he is unseen ; but his presence is shown to the observantsportsman by the boiling of the water from below, as from a greatcaldron. With no better guide than this, the harpoon has beendarted down, and reached him when twelve feet below the surface."These somersaults (or somersets as Elliott- calls them) are oftenmade by the fishes when leaping out of the water. Elliott especiallynoticed a number in 1846 (July ist) at four o'clock in the afternoonnear Hilton Head (S. C.) : "Thev did not show themselves somer-
' Op. cit.. p. 75.
= 0p. cit.,p.85.
l64 SMITHSONIAN MISCKLLANEOUS COLLECTIONS VOL. 52
setting for some time, but after a while began to sport and throwsomersets under the water, but so near to the surface as to show theirbelHes in the evolution. We saw, I do not doubt, as many as twentyfish. We counted eleven that leaped entirely out of the water.They were in the channel, and were further from shore than wherewe had usually met with them ; and, on approaching near to them inour boat, we remarked that those which leaped entirely out of thewater did not again show themselves on the surface until they hadsilently gone a mile or so toward the sea, when they reappeared,gambolled awhile, threw new somersets, and again disappeared fora new seaward movement. The fish which were behind came alongsporting until they had reached the spot where the first had throwntheir somersets. They, too, then threw their somersets, and disap-peared like the first. Usually they leaped twice—leaping from theirbacks, and falling likewise on their backs ; leaping, I should say, atleast ten feet above the water."The appearance and evolutions of the Devil-fish are indeed im-pressive and startling. Holder^ thought that "no more diabolicalcreature could be imagined. They resembled enormous bats, andin following one another around the circle raised the outer tip of thelong wing-like fin high out of the water in a graceful curve, theother being deeply submerged." They might be seen, "now glidingdown with flying motion of the wings; sweeping, gyrating upwardwith a twisting vertical motion marvelous in its perfect grace ; nowthey flashed white, again black, so that one would say they wererolling over and over, turning somersaults, were it possible for solarge a fish to accomplish the feat." Such evolutions, Holderlearned, were "really a common practice of the big rays." But it isthe great leaps out of the water that are most striking, especiallyduring the stillness of the night. Holder,^ on such an occasion onthe outer Florida reef, first encountered the fish. "There came outof the darkness, near at hand, a rushing, swishing noise ; then a clapas of thunder, which seemed to go roaring and reverberating awayover the reef, like the discharge of a cannon. So startling was thesound, so peculiar, that the negroes stopped rowing, and one or twodropped their oars in consternation."
•Op. cit.,p.8.
' Op. cit., p. 2.
NO. 1816 STORY OF the; devil-fish—GILI. 16 =IVIn some warm sea a fortunate observer may find perhaps a Devil-fish or a couple swimming on or near the surface ; not rarely aschool, or "shoal," of them. (Shoal is the word used by the Hon.William Elliott in his earliest full treatise on them as subjects ofsport. ^) Frequently they project themselves in the air to a -consid-erable height and for some distance. Their progression indeed israther of the nature of flight than swimming, and has been likenedto "the flight of a bird of prey"; it is by flaps of the wing-like pec-toral fins and not at all by the tail, as in Sharks and fishes generally.
Fig. 52.—Eagle-rays in motion. After Mangelsdorff. (Natiir und Haus,8, 1900, p. 255.)In fact, the progression of the Devil-fishes is quite similar to thatof their near relatives, the Eagle-rays, which have been portrayedfrom life by Mangelsdorff. Meanwhile, according to Holder, theircaropteres, or head-fins, otherwise called arms, feelers, claspers, orhorns, are "in constant motion, being whirled about like the tentaclesof a squid."Mr. Hector von Beyer, of the U. S. Bureau of Fisheries, informedDr. Hugh Smith^ that he had "observed the animal in the Gulf of
^ Carolina Sports by land and water, including incidents of Devil-fishing,[etc.]. Charleston, 1846. (2d edition, N. Y., 1850; 3d edition, N. Y., 1859.)
" The Fishes of North Carolina, 1907, p. 48.
l66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 52California" and noticed that "each of these appendages may becurved on itself like an elephant's trunk, and can firmly grasp ob-jects within reach." According to Elliott/ "It is the habit of thisfish to ply these arms rapidly before its mouth while it swims, and toclasp with the utmost closeness and obstinacy whatever body it hasonce inclosed. In this way, the boats of fishermen have often beendragged from their moorings and overset by the Devil-fish havinglaid hold of the grapnel."That these "arms" are muscular and powerful has been demon-strated on many occasions. The natural movement of the head-finsor caropteres is inward, and wdien any object strikes between them itis instinctively held, a proceeding which explains the undoubted factthat these fishes can run away with quite large vessels. Many suchcases of towing vessels have been recorded.One of the characteristics for which the Devil-fishes are celebratedis the capture of vessels and carrying them off far from their moor-ings. In one of the earliest notices of the Devil-fish, by John Law-son in "The History of Carolina" (1714), this peculiarity is de-scribed. "The Devil-fish," he says, "has been known to weigh aship's anchor, and run with the vessel a league or two, and bringher back, against tide, to almost the same place." Later notices donot give the animal credit for the same accommodating treatment!A number of accounts, however, corroborate the tendency indicated.William Elliott noticed several instances, and, in later times. Holder(p. 18) records that "at least instances of this were heard of on thereef occurring from Tampa Bay to Garden Key." He adds : "Inevery case the vessels, always at anchor, suddenly moved oft' in amysterious manner and were towed greater or less distances. TheRay had collided with the chain, and, true to its instincts, threw itstwo tentacular feelers or claspers around it and rushed ahead, thuslifting the anchor."In accordance, too, with this proclivity to seize upon objects whichbar their progress, Devil-fishes have been charged with damage anddestruction to wharves which extend into the water. "It was inobeying this peculiarity of their nature that a shoal of these fish, asthey swept by in front of 'Elliott's' grandfather's residence, wouldsometimes, at floodtide, approach so near to the shore as to come incontact with the water fence, the firm posts of which they wouldclasp and struggle to uptear, till they lashed the water into a foamwith their powerful wings.'"- Any such action, however, would beentirely exceptional and the statement requires authentication.
' Op. cit., p. 16.Op. cil., p. if).
NO. 1816 STORY OF THE DEVIL-FISH
—
Clhh 167VThe food of the Devil-fishes, so far from being large animals andoccasionally a man or so, as has been alleged, appears to be chieflythe small crustaceans and young or small fishes which swarm in cer-tain places near the surface of the w^ater. Rarely does one prey onlarge fishes. Once only did the man who had the most experiencewith the fish (Hon. William Elliott) see evidence of disposition toresort to scalv fish ; he gives this testimony:^ "I have frequently ex-amined the contents of their stomachs, and found little else in themthan portions of shell-fish, highly triturated, resembling the shellsof shrimps. Once a small crab was found entire ; but I sought invain for the scales of small fish, which I supposed to be their food,partlv because the Devil-fish make their appearance in our waters inMay, before the shrimps are found on our shores, and would thusbe anticipating their food—a mistake which fish are not apt tomake—and partly because I witnessed a performance on the part ofa Devil-fish which could scarcely be referred to anything else but toan occasional indulgence in a fish diet.
"I w^as w^atching a Devil-fish, who was playing close to the shore.But in shallow water he is often alarmed by the noise of the oars,and he would not suffer my approach within striking distance.While thus engaged, I observed a shoal of small mullets swimmingnear the surface, and showing signs of extraordinary agitation, whensuddenly the open mouth of the Devil-fish was protruded frombelow, and the small fry disappeared from view\ and w^ere receivedinto it, as into the mouth of an enormous funnel. I do not think itwas mere wantonness on the part of the fish, but that he was, onthat occasion, indulging a caprice of appetite, and substituting a chetof scale-fish for his ordinary mess of shrimps."We have, in this observation, a hint as to the function of the
"horns" or head fins; these may not only serve by their extensionto partly confine the prey, but they may be actively used to drive orscoop them in. The stories of their grasping intentionally may bereceived with some skepticism, although they do so accidentally.It is, indeed, largely by means of the head fins, or caropteres, thatthe Devil-fishes secure their food. That consists at least in part ofcrustaceans and other organisms which live about the surface of theseas they frequent. In the Gulf of California, where the Devil-fishesare most numerous, such animalcules are said by one observer to soabound that a thick sheet (nappe epaisse) of the organisms is
' Op. cit., pp. 84, 85.
l68 SAIITHSONIAN MISCELLANIiOUS COLI.ECTIONS VOL. 52"formed at the surface of the water. The fishermen in such localitiesaffirm that they never find any large animals in the stomachs of theDevil-fishes.But, if Richard HilP is to be credited, some Devil-fishes may bealso "ground feeders." They are, he thought, "formed for shoving^through the fields of turtle grass, testudinaria, but, unlike the Rays,which are likewise ground feeders," one of the Devil-fishes "doesnot seize its prey on the ground, but, pushing on through the marineherbage, it takes into its wide-open mouth the congregated livingthings that are in the way—it may be the fish that nestle in the vege-tation or the naked mollusca that depasture there—at once swallow-ing them, or rather cramming them in with its cranial arms into itsmouth and stomach, without deglutition, having no CEsophagus. Asthe animal in this gathering in of food can not see forward, it mu'itdepend on casualties in the course it steers through the marinemeadows for prey. The rolled-up head-fins between the crescentedhead sufficiently direct the food to the mouth."In the Gulf of Mexico and elsewhere, the Devil-fish has beencharged with feeding on shell-fish and complaint has been made thatit does considerable damage to oyster beds. This charge is duesimply to the fact that the animal has been confounded with theEagle-rays, whose large molar teeth eminently fit them for crushingshells. The general resemblance as well as real relationship of theDevil-fish to the Eagle-rays is indeed such as to leave no room towonder that the same name is applied to species of both families, butthe singular head-fins of the Devil-fish distinguish it from all itsrelations of dififerent families.Probably connected with the food and feeding of the Devil-f^:hesare peculiar organs within the mouth, called by Panceri- and Dume-
ril,^ who first described them, "prebranchial appendages.""On examining at the bottom of the mouth the pharyngeal aper-tures of the branchial chambers, or separating the walls of theirexternal apertures, we see, in front of each of the respiratory sur-faces, a very regular series of organs which do not occur in anyother fish, whether bony or cartilaginous.
"These organs are elongated lamellae, the aspect of which some-what reminds us of that of the stems of ferns, but with the leaflets
' Tlie Devil-fish of Jamaica. Intellectual Observer, 2, 1862, p. 167-176.
"Panceri (P.) e Leone de Sanctis. Sopra alcuni organi delle CephalopteraGiorna, M. H. Atti Accad. Pontoniana, Napoli, vol. 9, 1871, pp. 335-3/0, 2 pis.
^ Dumeril (A.). On the presence of peculiar organs belonging to theBranchial Apparatus in the Rays of tlie Genus Ccplialoptcra. Ann. Mag. Nat.Hist. (4), 5, 1870, pp. 38s, 386.
NO. 1816 STORY OF THE DEVIL-FISH—GILL 109turned back toward the branchise. Each being formed of a fold ofmucous membrane supported by a cartilage, these lamellae areattached to the anterior surface of the branchial arches, in front ofthe membranous and vascular folds of the respiratory organs; andit is their position that has suggested the name of prebranchial ap-pendages, by which they are designated by the Italian anatomist.
"Thev do not serve for respiration. By means of injections, M.Panceri has ascertained that they receive arterial vessels, like theother organs, and not branches of the branchial artery."These organs are thought by Panceri (and Dumeril did not dis-sent) to be "destined, on account of the remarkable size of the aper-tures of the branchial chambers, the orifices of which are much
J / ii' ( /
^'^
. im''''''''.
.
r .* f
Fig. 53.—Anterior Hemibranch of the Fourth Left Pouch.
a. Fourth branchial arch.b. Section of the special muscle of the branchial arch or adductor of the twoceratobranchial and epibranchial portions.c. Branchial lamellae.d. Prebranchial appendages.X. Fold of the mucosa which partly covers the branchial lamellae.
smaller in the other Rays, to retain the water and prevent it fromtraversing these cavities with a rapidity which would be injuriousto the perfect accomplishment of the act of hsematosis."A more probable use for these organs would be as strainers, sub-serving thus the same function, or rather an analogous one, as thatof the gill-rakers of the giant Sharks. They would retain the smallorganisms contained in the ingesta taken into the mouth, while the12
lyo SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 52water itself would find exit as usual, relieved of a large part of itslife.
Fig. 54.—A Branchial Arch with Annexed Organs; transverse section next tothe articulation of the arch (semischematic).0. Branchial arch with fossa of adductor muscle.b. One of the cartilaginous rays of the branchial diaphragm adherent to theanterior branchial lamellae.
c. Accessory stem which connects with the arch.d. Muscle of the branchial diaphragm or interbranchial muscle to whichposterior branchial lamellae adhere.e. Adductor muscle of the ceratobranchial and epibranchial parts of the arch.ff. Branchial lamellae whose external surface as usual is folded.gg. Cartilaginous stems of the bases of the branchial lamellae.hh. Muscles which unite the latter to the arch.it. Hydrophorous canals.jj. Prebranchial appendages in profile./. Branch of the branchial artery.mm. Branchial veins with efferent lamellar branches, from which proceed thebranches for the prebranchial appendages.n. Principal nervous trunk.X. Fold of mucosa covering partly the branchial lamellae.VIThe Devil-fishes, of course, like other Selachians, come togetherin sexual intercourse.^ The details of their union as well as the
'According to Risso (Hist. Nat. Europe Men, 1826, p. 165) the female of theM. giorna is always (toujours) much larger than the males. No observationshave been made on the American Devil-fish ; it is to be hoped that somemay be.
NO. 1816 STORY OF THE DEVIL-FISH—GILL I7Ilength of gestation are unknown. Even the exact date of the oneobservation that has been pubHshed has not been given, although itappears to have been some time in July. It is, indeed, quite possiblethat the appearance of the animals close to the coasts of the SouthernStates may be for the purpose of finding a suitable place for thebirth of the young. By Elliott,^ it was found, in the first years ofhis experience with them, that they appeared "only in August"; in1843, "for the first time, in July," and in 1844 thev were "taken inJune."Care seems to be extended even to the place of parturition by theSting-rays, so that the young shall encounter the least danger fromthe tide as well as from living enemies. Alcock tells that all thesmall Sting-rays (Dasybatis zvalga) with embryos he observed "w^erefound in shallow little tidal pools lying behind natural breakwatersof sand," and he urges, "it seemed as if this comparatively safe sit-uation had been deliberately chosen by the mother as a nursery forher expected family, as, in the opinion of Professor Mcintosh, is thecase with the viviparous Blenny (Zoarccs) of northern seas." Anal-ogous care may therefore be exercised by the Devil-fishes, the rela-tions of the Sting-rays.A pair of these huge animals, male and female, were seen in unionby Mr. Elliott and described by him.*VIIWhatever be the size or other characters of the Devil-fishes, so faras observed, they agree among themselves and differ from most otherfishes* by having, normally, only a single young one at a birth. Thegiant mothers noticed by Duhamel, Risso, Mitchill, and Lamont
' Op. cit., p. 67.
^ Subito, laeva—sed longiore spatio, quam, si jaculatus essem, speraverimtransfigere ictu—duos pisces cephalopteras aspexi, amplexu conjunctos. Ven-tribus juxtapositis—capitibus erectis, et supra undam oblatis—antennis lasciveintersertis—coitum salacem, ut solet genus squalus, ipso contactu corporis,tunc sine dubio exercuere. Ferire, ob distantiam non licitum, aut duoscephalopteras, solo ictu transfixisse, gloria inopinata mihi contegisset. Cym-bam appropinquantem, hastamque minantem, circumspecte evitant—et, inprofundo paulisper latentes, iterum, dextra emergunt, ludosque lascivosrepetunt. Tunc, quasi deliciis satiati, saltatione in aere, utrinque facta
—
aper-tum mare petivere. Hoc concursu tarn raro notato—antennis albis, cum nigrisadmixis utsi lacertis—imago foedi et immundi coitus, nudi Africani cumCaucasiana, plane praefigurabatur. (Elliott, Carolina Sports, 3d edition, pp.93, 94).*The Stingrays (Dasybatids) of some species at least have only a singleyoung.
1/2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 52
each had only one (one or two, according to Risso^). In case of thesmall species named Ceratobatis robertsii or massenoidea, the motherlikewise had a single foetus (a foetus sixteen inches wide).Although only one young is formed, that one is worthy of thegiant mother and larger than any of the full-grown common Rays ofordinary size. It is practically immune from danger from the cus-tomary enemies of fishes and well able to take care of itself.Nature is economical in her methods and there is some adjustmentof ways and means. In the case of egg-laying fishes of inferior sizeand when no care is taken of the eggs, many thousands—even mil-lions—may be laid by a single fish, and yet the number of adultsremains practically the same, generation after generation. In thecase of viviparous fishes like the Devil-fishes, a single young one ata birth is enough to keep up the species.The fishermen of Jamaica, according to Hill, "say that the motherfish makes the violent leaps she is seen to take out of the water toeject the foetus from the matrix ; that the young fish is then observedto fall from her ; and that for a time it swims upon the parent's back,and possibly enters the wide mouth-sack when necessary to seekshelter from apprehended danger." All this is improbable. It ap-pears to be certain that the "leaps" are habitual to males and femalesalike, and it is probable that they are the extension of their peculiarmode of progression or "flight."A pregnant female, 15 feet wide and which with difficulty fortymen with two lines attached to it could drag along the ground, waslanded, after a five hours' fight, at Port Royal, Jamaica, in 1824."On opening it a young, about 20 pounds weight, was taken out,perfectly formed" ; it was five feet broad. An account of the cap-ture was given by Lieutenant Lamont in the Edinburgh Philosophi-cal Journal (xi, T13-118).Two observations respecting the procreation of Devil-fishes re-quire attention.That the Devil-fishes have only one young each, and consequentlyare viviparous, is the statement made by all observers. This vivi-parity is in analogy with the gestation in the relatives of the Devil-fishes, all the Sting-rays and Eagle-rays. Nevertheless a gentleman
* Risso, in his "Remarques" on the "Cephalopteres" gives the following data
:
L'epoqne de leurs amours est I'hiver ; les femelles mettent has en Septembre una deux petits, qui originairement sont renfermes dans un oeuf oblong jaunatre.Les males paraissent quelquefois n'abandonner leur compagne qu'apres qu'ellea depose ses foetus; et si I'un des deux se jette dans un filet, I'autre ne tardejamais a le suivre. Risso Hist. Nat. Europe Mer., 3, 1826, p. 165.
NO. 1816 STORY OF THE DHVIL-FlSIi—GILL 173with considerable knowledge of ichthyology, Swinburne Ward, oncethe Civil Commissioner of the Seychelles Islands, after an accountof the capture of a Devil-fish which "ten men could not haul" up onthe beach, concluded with the affirmation that "she was full of eggs."The idea might be (and has been) derived that this may have beena case of oviparity or multiparity, but the eggs (if they were such)were possibly the reserve stock left perhaps after the birth of ayoung one. The statement is in great need of confirmation.Mitchill, in 1823, tells that a "female that was struggling afterhaving been w^ounded brought forth in her agony a living youngone, as Captain Potter related, and Mr. Patchen, while he showed[Mitchill] the orifices through which sucking is probably performed,declared that on dissection mammary organs were found, which dis-charged as much as a pailful of milk." This at first incomprehen-sible and incredible statement may be reconciled with facts when werecall the mode of nutrition of the embryo among the Sting-rays,described by Alcock. It was the honest statement of an inex-perienced observer who misinterpreted facts.A remarkable provision among the Sting-rays for the nutrition ofthe embryo within the body of the mother has been made known byA. Alcock, on whose description, published in 1902, we may draw.^It is by means of a secretion which is regarded as "analogous tomilk" that the embryo is for some time fed. The mucous membraneof the oviduct is "shaggy, with vascular filaments [named trophone-inata\ dripping with milk" or rather a milk-like fluid, and on micro-scopic examination it was found that "each filament was providedwith superficial muscles whose contraction must serve to squeeze themilk out. Some such mechanism is undoubtedly necessary, seeingthat the young one has no power of extracting the secretion foritself. On examination of the young one, the mother's milk wasfound inside the modified first pair of gill-clefts or spiracles (theother gill-clefts being tightly closed), and also in large clots withinthe spiral valve of the intestine, so that there can be no doubt thatin these viviparous Rays the unborn young ones may be said to
^Alcock (A.). A Naturalist in Indian Seas [etc.], London, 1902, pp. 210,71, 159. See, also, Observations on the Gestation of some Indian Sharks andRays. Journ. Asiat. Soc. Bengal, 59, pt. 2, 1890, pp. 51-56, pi. i ; On the UterineVilliform Papillae of Pteroplatea micrura, [etc.] Proc. Roy. Soc., 49, 1891,pp. 359-367, pis. 7, 8; Further observations on the Gestation of Indian Rays;[etc.] ; Proc. Roy. Soc, 50, 1891, pp. 202-209. On Utero-gestation in Trygonhleekcri. Ann. Mag. Nat. Hist., (6), 9, pp. 417-427, pi. 19, 1892; Some Obser-vations on the Embryonic History of Pteroplataa micrura. Ann. Mag. Nat.Hist, (6), 10, pp. 1-8, pi. 4, 1892.
1/4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 52
'drink its mother's milk' like a mammal, even though the milk-likesecretion does not go in at the mouth, but by channels homologouswith the ear-drum of air-breathing vertebrates."
Fig. 55.
—
Ptcrophitcca viicnira. After Alcock.
NO. 1816 STORY OF THE DEVIL-FISH—GILL 175EXPLANATION OF FIG. 54.
1. Embryo of Pteroplata:a micrura, from dorso-lateral aspect; nat. size, butwith only a few of the gill-filaments represented, for the sake of clear-ness, s, spiracle.2. End of a gill-filament, showing marginal capillary filled in places withblood-clot. X 42.3. Transverse section of a gill-filament, showing the marginal capillary insection and the single fold of epithelium. X 188. For the sake of clear-ness the blood-clot is represented in one limb of the capillary only, andthe spaces between the nuclei of the surface epithelium are a littleexaggerated.4. End of a trophonema, or nursing-filament, seen as a transparent object inglycerine, showing the marginal artery and the superficial capillaryplexus. X 42. The median vein is not seen so near the end.5. Obliquely transverse section through a nursing-filament, showing the glandsstill in the form of solid bulbs lying beneath a still unbroken surface ofepithelium. X no. aa, arteries; v, vein; cc, superficial capillaries.Doubtless an analogous provision for the nutrition of the embryois developed in the Devil-fishes, and thus we have a satisfactory ex-planation of the statements of Patchen and Mitchill. Somethinglike milk is secreted by the mother fish and is ingested by the young,but it is chemically different from milk, and instead of being suckedin by the mouth is absorbed through the postocular spiracles. Thestatements which have been much ridiculed have therefore a soundfoundation in fact and are susceptible of a natural explanation.Nothing is known respecting the development of the embryo ofany Devil-fish, but undoubtedly it is similar to that of the Sting-rays.The very young embryo of the Sting-rays, as of all other Rays,contrasts remarkably with the mother, especially in the case of thevery wide forms, such as the Pteroplateines. The embryo at anearly stage has a form very like that of a Shark, but with pectoralsprovided with basilar extensions free from the head, and extendingforward parallel with it in advance of the eyes. These extensionslater unite with the sides of the head, and the regular Ray has thenbecome developed. Essentially, the form of the mother has beenattained by the young when ready for extrusion from the mother'swomb. This much at least is known of the new born of Devil-fishes, VIIIThe various species of Devil-fishes are representatives apparentlyof three different generic types, distinguished by differences of deri-vation. Mobula (also called Aodon, Cephaloptera, or Diccrobatis'.
1/6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 52has teeth in both jaws; Manta (or Ceratoptera) has teeth confinedto the lower jaw, and Ceratobatis has teeth only in the upper jaw.The species also differ in size and the character of the dorsal spine.While a width of twenty feet or more may be attained by some,others become sexually mature when four feet wide. In most ofthem the tail is short and the dorsal spine characteristic of Sting-rays is obsolete, but it is asserted to be well developed in the Mobulagiorna.The number of species of Devil-fishes is uncertain. In 1870 sevenspecies were recognized, five of the genus Dicerobatis (Mobula)^ andtwo of Ceratoptera (Manta). One representing a new generic type(Ceratobatis) was added in 1897. One of gigantic size, generallysupposed to be Manta vampyrus, has been observed at many places.Whether there are more than one species is uncertain.- There is adiscrepancy in the length of the tail assigned to some. Most of thegiants have a tail nearly as long as the body, but one referred to byHill, about fifteen feet wide, had a tail only two feet long.^ The spe-cies of Mobula dififer. The M. giorna of the Mediterranean is saidto have a tail about three times longer than the width of the disk;the M. japonica one "nearly thrice as long as the body," and theM. olfersii of Brazil and the Caribbean Sea one about as long as thedisk and much less than its width. The Ceratobatis robertsii has thetail not much less than twice the length of the disk (620: 350), butconsiderably less than its width (620:780).*One species—the true Devil-fish of the United States, Mantavampyrus—is not uncommon in the warm American waters and ap-pears on the South Carolina coast in summer in "shoals."The Manta vampyrus has a body or disk nearly twice as wide aslong, and a tail about 6/10 as long as the body ; the body and tail arerough from the development of small tubercles which extend almosteverywhere; the band of teeth (confined to the lower jaw) extendsover almost the whole width of the jaw and is composed of about a
* Three nominal species were described later
—
Dicerobatis draco. Giinther,1872; D. moiis/niin Khinzinger, 1871, and Cephaloptcra tarapacana, Philippi,1894.
'This subject will be considered in a future article.
' The tail may have been decurtated in youth.
* The figure in Day's Fishes of India (1878, p. 745), which he "surmises"may represent "Ceratoptera chrenbergii" is nothing but an illustration of a notuncommon monstrosity of an ordinary Ray (Rata) with free anterior exten-sions of the pectorals, resulting from arrest of development. (See Proc. U. S.Nat. Museum, 1895, PP- 195-198.)
NO. 1816 STORY OF THE DEVIL-FISH—GILL 177hundred transverse rows; the rows are separated from each otherby well-marked interspaces. It is said to attain a width of 30 feet.'This or a very closely related species has been found not only :nthe West Indian and Carolinian seas, but along the west coast ofAmerica, along the African coast, and in the Indian Ocean. ADevil-fish fourteen feet six inches wide, caught near Durban, Natal,also presented the same proportions as the American species. Aplate representing it from before and behind was published in theZoologist for April, 1899.Like most other large Selachians, the Devil-fish is beset by Echen-eidids, commonly known as Sucking-fish or Suckers and oftenconfounded with the Pilot-fish. Elliott- noted that "he is attendedby a band of parasites," which "followed him into shoal water" and"adhered so closely after he was aground that several suffered them-selves to be taken by the hand."^IXThe Devil-fish from time to time has been the object of sport. Hewho indulged most in it and captured almost twenty has givenanimated pictures of some of his adventures. One of the most con-densed and entertaining accounts may be welcome here.One day in late June (24th), sailing toward "Hilton Head"(South Carolina), Mr. Elliott with his crew went after Devil-fish.Soon he saw "a shoal" of them "sweeping along the beach, travelingrapidly downward with the tide" and freely showing themselves atthe surface. After an inefifectual cast with a harpoon, "three showedthemselves below and one above."
'The records of size are very defective. The largest actually measured byElliott was 17 feet wide (p. 64), another 16 feet (p. 80), and another 15 feet(p. 43). Another lost after being dragged "into three feet water" was esti-mated to be larger; "there he lay, extending twenty feet by the wings" (p.51). One taken in the Gulf of California in 1846 was 19 feet wide, 3 feet 6inches thick, and had a mouth 3 feet 5 inches wide (Z06I., 1849, p. 2358).Another noticed by Gosse (The Ocean, p. 193-194, Amer. Edit., p. 189) takenat La Guayra, was 20 feet wide, with a "length from end of tail to end oftusks [caropteres] 18 feet," a "mouth 4 feet wide," and "its weight 3.502pounds."
* Op. cit., p. 44.^Le Vaillant, near the African coast, met three Devil-fishes ("diable"), oneof which was accompanied by a sucking-fish ("pilote du diable") attached toeach horn ("corne") of the Devil-fish. His account is unreliable. The para-site is the Remora remora according to Street (Bull. U. S. Nat. Mus., 7, p. 54),and Pellegrin (Bull. Mus. Hist. Nat, Paris, vii, 327).
178 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 52Now he shall speak for himself:^
"I pushed at one that showed his back fairly above water, as heswam; but he sank just before I reached him, and I drove down theharpoon at a venture. He had a narrow escape, for the staff struckhim. At this moment, three showed themselves below and oneabove. I pushed for the latter, and when I approached the spot, Isaw the water boiling up like a caldron—from which sign I knew thatthe fish was throwing his somersets below the surface (in the waywhich is so very peculiar to them). Making the oarsmen check theheadway with their oars, I looked anxiously for a view, when, unex-pectedly, I saw the white of his belly far beneath the water, andquite away toward the stern. He was thus behind me, but wheelingsuddenly to the right, I pitched the harpoon at him, across the oars,and felt a sensation of surprise, as well as pleasure, in finding thatI had struck him. The fish dashed out violently for the channel,and we payed him out thirty fathoms of rope until, headway beinggiven to the boat, we brought him to a dead pull ; and now his mo-tions were very erratic ; unlike some that I had before struck, he didnot take a direct course for the sea, but sometimes drew the boatagainst the tide, then suddenly turned and ran directly toward us, soas to give slack line. I inferred from these signs that he was mor-tally hurt. As often as he approached the Middle Bank and shoaledthe water, he drew off in alarm, and would not cross it until he hadgot to its tail ; his course was then for Paris Bank, which, suitingwell with our intention to land him, if we could, at Bay Point, we didnot interrupt. About this time he came to the surface without beingpulled, and showed great distress—and we resolved, then, to drawupon him and get a second harpoon planted. It was after variousfruitless efforts, and by shortening the rope as far as we prudentlycould, that we at length drew him so far up that the dark shadowof his body was indistinctly seen beneath. The second harpoon wasnow driven, and the gush of blood to the surface showed that it haddone its work. We now drew mainly on this second, leaving only amoderate strain upon the first—and after a few convulsive runs,brought him up helplessly to the surface, and with a spear dispatchedhim outright. With a hatchet we now cut a hole in one of hisfeelers, and inserting a rope, passed it to the stern, drawing solelyon this, so that the resistance of the fish through the water shouldbe as small as practicable. The wind was now due east and moder-ately fresh ; we raised both sails, and, helped at the same time by theoars, made some way in our tedious progress on towing our prize toland. At this time, espied a boat beating down from Beaufort, andon signalizing her, she proved to be that of Col, De Treville, thenon his way to Bay Point. His offer of assistance was accepted, anda tow-line being passed to his boat, we landed our fish at the Pointexactly at sunset. This fish measured sixteen feet across, which Isuppose to be the medium size of those that visit our waters. Thefirst harpoon had struck it near the center of the belly—had pierced
' Op. cit., pp. 68-72. The punctuation of the original is preserved.
NO. 1816 STORY Q-e- THE DEVIL-flSH—GILL 1 79the liver, and passed nearly through to the back. The second hadpassed from the back into his lungs or gills—so that the full powerof so large a fish was never fairly exerted against us. Had the samefish been struck in the wings, or other parts not vital, his capturewould have been uncertain—and would at any rate have cost us thework of many hours.
"I suppose the shoal of Devil-fish was a large one; the third whichappeared we struck at—the fourth we harpooned—and as we wererapidly drawing off from the shore, a fifth was seen. How manywere still behind, we had not leisui-e to observe; but conjecture thiswas but the advance guard of the column."Later adventurers after sport with the Devil-fish have hunted italong the Florida coast as well as in the Gulf of Mexico and theCaribbean Sea. C. J. Holder has told of his experience in "Trailingthe Sea-bat" in "Outing" for 1900, and J. Turner-Turner has de-voted two chapters of his book entitled "The Giant Fish of Florida"(1902) to the "Enormous Rays, or Devil-fish," which he pursued.The article by Holder has been republished i;i that author's workentitled "Big Game at Sea," published in 1908 (pp. 1-35).The pursuit of such a giant as the Devil-fish is necessarily attendedwith some danger, but this incident adds to the zest the sportsmanfeels. Elliott records that he had been "carried twenty-five miles inthe course of a few hours by two of these fish (having struck a relaywhen the first sea-horse escaped, and losing both), with three boatsin train."According to Leon Diguet^ (1898), in the Gulf of California,where Devil-fishes are numerous, the pearl-fishers, when caughtduring a calm away from mooring places, always take the precautionof dropping two anchors at night for fear that one should be seizedby a Devil-fish and hauled afar by it. Diguet went in pursuit of aspecimen for the Musee d'Histoire Naturelle of Paris, and, after onehad been harpooned, it turned back on the boat, seized the bow withits headfins, and held it in its clasp till it was lanced a second time.But this clasping is largely automatic, and the Devil-fish only makesfor the boat from which it has been attacked when it experiencesthe stress through the line from that direction. It is not like theattack of some sharks when wounded. The Devil-fish, in fact, hasbeen called a "timid animal" by Diguet.The Devil-fish, nevertheless, is the object of considerable dreadamong the fishermen of the Gulf of California; for, although notaggressive, it is frequently encountered, and Diguet tells that numer-
^Vaillant (L.) et L. Diguet. Snr le Cephaloptere du Golfe de Californie.Bull. Mus. Hist. Nat., Paris, 1898, pp. 127-128.
l8o SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 52ous cases have occurred of death resulting to divers , as well asbathers from encounters with the Devil-fish, or Manta, as the mencall it.^ On the other hand, the carcasses of many that are killed areused for bait for other fishes.
^ An accomplished naturalist of the second quarter of the last century, Col.Hamilton Smith, "once witnessed the destruction of a soldier by one of theseCephalopteri off Trinidad. It was supposed that the soldier, being a goodswimmer, was attempting to desert from the ship, which lay at anchor in theentrance of the Boca del Toro. * * * The Colonel is positive as to thisfish being a Cephalopterus." The full account is given in Griffith's edition ofCuvier's Animal Kingdom ("The Class Pisces," p. 654). The evidence isvery unsatisfactory.