Fig. 256.—Mackerel-shark or Salmon-shark, Lamna cornubica (Gmelin). Santa Barbara, Cal.
In the second group the vertebræ have their calcareous matter arranged in rings, one or more about the notochordal center. In all these the anal fin is absent, and in the process of specialization the shark gradually gives place to the flattened body and broad fins of the ray. This group is called Tectospondyli. Those sharks of this group with one ring of calcareous matter in each vertebra constitute the most primitive extreme of a group representing continuous evolution.
From Cladoselache and Chlamydoselachus through the sharks to the rays we have an almost continuous series which reaches its highest development in the devil rays or mantas of the tropical seas, Manta and Mobula being the most specialized genera and among the very largest of the fishes. However different the rays and skates may appear in form and habit, they are structurally similar to the sharks and have sprung from the main shark stem.
Fig. 257.—Star-spined Ray, Raja stellulata Jordan & Gilbert. Monterey, Cal.
The Chimæras.—The most ancient offshoot from the shark stem, perhaps dating from Silurian times and possibly separated at a period earlier than the date of any known shark, is the group of Holocephali or Chimæras, shark-like in essentials, but differing widely in details. Of these there are but few living forms and the fossil types are known only from dental plates and fin-spines. The living forms are found in the deeper seas the world over, one of the simplest in structure being the newly discovered Rhinochimæra of Japan. The fusion of the teeth into overlapping plates, the covering of the gills by a dermal flap, the complete union of the palato-quadrate apparatus or upper jaw with the skull and the development of a peculiar clasping spine on the forehead of the male are characteristic of the Chimæras. The group is one of the most ancient, but it ends with itself, none of the modern fishes being derived from Chimæras.
Fig. 258.—A Deep-sea Chimæra, Harriotta raleighiana Goode & Bean. Gulf Stream.
Fig. 259.—An extinct Dipnoan, Dipterus valenciennesi Agassiz. Devonian. (After Pander.)
The Dipnoans.—The most important offshoot of the primitive sharks is not the Chimæras, nor even the shark series itself, but the groups of Crossopterygians and Dipnoans, or lung-fishes, with the long chain of their descendants. With the Dipnoan appears the lung or air-bladder, at first an outgrowth from the ventral side of the œsophagus, as it still is in all higher animals, but later turning over, among fishes, and springing from the dorsal side. At first an arrangement for breathing air, a sort of accessory gill, it becomes the sole organs of respiration in the higher forms, while in the bony fishes its respiratory function is lost altogether. The air-bladder is a degenerate lung. In the Dipnoans the shoulder-girdle moves forward to the skull, and the pectoral limb, a jointed and fringed archipterygium, is its characteristic appendage. The shark-like structure of the mouth remains.
The few living lung-fishes resemble the salamanders in many regards, and some writers have ranged the class as midway between the primitive sharks and the amphibians. These forms show their intermediate characters in the development of lungs and in the primitive character of the pectoral and ventral limbs. Those now extant give but little idea of the great variety of extinct Dipnoans. The living genera are three in number—Neoceratodus in Australian rivers, Lepidosiren in the Amazon, and Protopterus in the Nile. These are all mudfishes, some of them living through most of the dry season encased in a cocoon of dried mud. Of these forms Neoceratodus is certainly the nearest to the ancient forms, but its embryology, owing to the shortening of its growth stages due to its environment, has thrown little light on the question of its ancestry.
From some ally of the Dipnoans the ancestry of the amphibians, and through them that of the reptiles, birds, and mammals may be traced, although a good deal of evidence has been produced in favor of regarding the primitive crossopterygian or fringe fin as the point of divergence. It is not unlikely that the Crossopterygian gave rise to Amphibian and Dipnoan alike.
In the process of development we next reach the characteristic fish mouth in which the upper jaw is formed of maxillary and premaxillary elements distinct from the skull. The upper jaw of the shark is part of the palate, the palate being fused with the quadrate bone which supports the lower jaw. That of the Dipnoan is much the same. The development of a typical fish mouth is the next step in evolution, and with its appearance we note the decline of the air-bladder in size and function.
The Crossopterygians.—The fish-like mouth appears with the group of Crossopterygians, fishes which still retain the old-fashioned type of pectoral and ventral fin, the archipterygium. In the archaic tail, enameled scales, and cartilaginous skeleton the Crossopterygian shows its affinity with its Dipnoan ancestry. Thus these fishes unite in themselves traits of the shark, lung-fish, and Ganoid. The few living Crossopterygians, Polypterus and Erpetoichthys, are not very different from those which prevailed in Devonian times. The larvæ possess external gills with firm base and fringe-like rays, suggesting a resemblance to the pectoral fin itself, which develops from the shoulder-girdle just below it and would seem to give some force to Kerr's contention that the archipterygium is only a modified external gill. In Polypterus the archipterygium has become short and fan-shaped, its axis made of two diverging bones with flat cartilage between. From this type it is thought that the arm of the higher forms has been developed. The bony basis may be the humerus, from which diverge radius and ulna, the carpal bones being formed of the intervening cartilage.
Fig. 260.—An extinct Crossopterygian, Holoptychius giganteus Agassiz (1835). (After Agassiz, per Zittel.)
The Actinopteri.—From the Crossopterygians springs the main branch of the true fishes, known collectively as Actinopteri, or ray-fins, those with ordinary rays on the paired fins instead of the jointed archipterygium. The transitional series of primitive Actinopteri are usually known as Ganoids. The Ganoid differs from the Crossopterygian in having the basal elements of the paired fins small and concealed within the flesh. But other associated characters of the Crossopterygii and Dipnoans are preserved in most of the species. Among these are the mailed head and body, the heterocercal tail, the cellular air-bladder, the presence of valves in the arterial bulb, the presence of a spiral valve in the intestine and of a chiasma in the optic nerves. All these characters are found in the earlier types so far as is known, and all are more or less completely lost or altered in the teleosts or bony fishes. Among these early types is every variety of form, some of them being almost as long as deep, others arrow-shaped, and every intermediate form being represented. An offshoot from this line is the bowfin (Amia calva), among the Ganoids the closest living ally of the bony fishes, showing distinct affinities with the great group to which the herring and salmon belong. Near relatives of the bowfin flourished in the Mesozoic, among them some with a forked tail, and some with a very long one. From Ganoids of this type the vast majority of recent fishes may be descended.
Fig. 261.—An ancient Ganoid fish, Platysomus gibbosus Blainville. Family Platysomidæ. (After Woodward.)
Fig. 262.—A living Ganoid fish, the Short-nosed Gar, Lepisosteus platystomus Rafinesque. Lake Erie.
Another branch of Ganoids, divergent from both garfish and bowfin and not recently from the same primitive stock, included the sturgeons (Acipenser, Scaphirhynchus, Kessleria) and the paddle-fishes (Polyodon and Psephurus). All these are regarded by Woodward as degenerate descendants of the earliest Ganoids, Palæoniscidæ, of Devonian and Carboniferous time.
Fig. 263.—A primitive Ganoid fish, Palæoniscum macropomum (Agassiz), restored. Permian. Family Potaconiscidæ. (After Traquair.)
Fig. 264.—A fossil Herring, Diplomystus humilis Leidy. (From a specimen obtained at Green River, Wyo.) The scutes along the back lost in the specimen. Family Clupeidæ.
The Bony Fishes.—All the remaining fishes have ossified instead of cartilaginous skeletons. The dipnoan and ganoid traits one by one are more or less completely lost. Through these the main line of fish development continues and the various groups are known collectively as bony fishes or teleosts.
Fig. 265.—A primitive Herring-like fish, Holcolepis lewesiensis Mantell, restored. Family Elopidæ. English Chalk. (After Woodward.)
Fig. 266.—Ten-pounder, Elops saurus L. An ally of the earliest bony fishes. Virginia.
The earliest of the true bony fishes or teleosts appear in Mesozoic times, the most primitive forms being soft-rayed fishes with the vertebræ all similar in form, allied more or less remotely to the herring of to-day, but connected in an almost unbroken series with the earliest ganoid forms. In these and other soft-rayed fishes the pelvis still retains its posterior insertion, the ventral fins being said to be abdominal. The next great stage in evolution brings the pelvis forward, attaching it to the shoulder-girdle so that the ventral fins are now thoracic as in the perch and bass. If brought to a point in front of the pectoral fins, a feature of specialized degradation, they become jugular as in the codfish. In the abdominal fishes the air-bladder still retains its rudimentary duct joining it to the œsophagus.
From the abdominal forms allied to the herring, the huge array of modern fishes, typified by the perch, the bass, the mackerel, the wrasse, the globefish, the sculpin, the sea-horse, and the cod descended in many diverging lines. The earliest of the spine-rayed fishes with thoracic fins belong to the type of Berycidæ, a group characterized by rough scales, the retention of a primitive bone between the eyes, and the retention of the primitive larger number of ventral rays. These appear in the Cretaceous or chalk deposits, and show various attributes of transition from the abdominal to the thoracic type of ventrals.
Fig. 267.—Cardinal-fish, a perch-like fish, Apogon semilineatus Schlegel. Misaki, Japan.
Fig. 268.—Summer Herring, Pomolobus æstivalis (Mitchill). Potomac River. Family Clupeidæ.
Another line of descent apparently distinct from that of the herring and salmon extends through the characins to the loach, carps, catfishes, and electric eel. The fishes of this series have the anterior vertebræ coossified and modified in connection with the hearing organ, a structure not appearing elsewhere among fishes. This group includes the majority of fresh-water fishes. Still another great group, the eels, have lost the ventral fins and the bones of the head have suffered much degradation.
Fig. 269.—Fish with jugular ventral fins, Bassozetus catena Goode & Bean. Family Brotulidæ. Gulf Stream.
Fig. 270.—A specialized bony fish, Trachicephalus uranoscopus. Family Scorpænidæ. From Swatow, China.
The most highly developed fishes, all things considered, are doubtless the allies of the perch, bass, and sculpin. These fishes have lost the air-duct and on the whole they show the greatest development of the greatest number of structures. In these groups their traits one after another are carried to an extreme and these stages of extreme specialization give way one after another to phases of degeneration. The specialization of one organ usually involves degeneration of some other. Extreme specialization of any organ tends to render it useless under other conditions and may be one step toward its final degradation.
Fig. 271.—An African Catfish, Chlarias breviceps Boulenger. Congo River. Family Chlariidæ. (After Boulenger.)
Fig. 272.—Silverfin, Notropis whipplii (Girard). White River, Indiana. Family Cyprinidæ.
We have thus seen, in hasty review, that the fish-like vertebrates spring from an unknown and possibly worm-like stock, that from this stock, before it became vertebrate, degenerate branches have fallen off, represented to-day by the Tunicates and Enteropneustans. We have seen that the primitive vertebrate was headless and limbless and without hard parts. The lancelet remains as a possible direct offshoot from it; the cyclostome with brain and skull is a possible derivative from archaic lancelets. The earliest fishes leaving traces in the rocks were mailed ostracophores. From an unknown but possibly lamprey-like stock sprang the sharks and chimæras. The sharks developed into rays in one right line and into the highest sharks along another, while by a side branch through lost stages the primitive sharks passed into Crossopterygians, into Dipnoans, or lung-fishes, and perhaps into Ostracophores. All these types and others abound in the Devonian Age and the early records were lost in the Silurian. From the Crossopterygians or their ancestors or descendants by the specialization of the lung and limbs, the land animals, at first amphibians, after these reptiles, birds, and mammals, arose.
Fig. 273.—Moray, Gymnothorax moringa Bloch. Family Murænidæ Tortugas.
Fig. 274.—Amber-fish, Seriola lalandi (Cuv. & Val.). Family Carangidæ. Woods Hole.
In the sea, by a line still more direct, through the gradual emphasis of fish-like characters, we find developed the Crossopterygians with archaic limbs and after these the Ganoids with fish-like limbs but otherwise archaic; then the soft-rayed and finally the spiny-rayed bony fishes, herring, mackerel, perch, which culminate in specialized and often degraded types, as the anglers, globefishes, parrot-fishes, and flying gurnards; and from each of the ultimate lines of descent radiate infinite branches till the sea and rivers are filled, and almost every body of water has fishes fitted to its environment.
Geological Distribution of the Families of Elasmobranchs.
| Pliocene | Miocene | Eocene | Cretaceous | Jurassic | Triassic | Permian | Coal Measures | Sub-Carbonifero | Devonian | Silurian | Ordovician | |
|---|---|---|---|---|---|---|---|---|---|---|---|---|
| Cladoselachidæ | █ | █ | ||||||||||
| Acanthodii | █ | █ | █ | █ | ||||||||
| Pleuracanthidæ | █ | █ | █ | █ | ||||||||
| Cladodontidæ | █ | █ | █ | █ | ||||||||
| Petalodontidæ | █ | █ | █ | |||||||||
| Psammodontidæ | █ | █ | █ | |||||||||
| Cochliodontidæ | █ | █ | █ | |||||||||
| Orodontidæ | █ | █ | ||||||||||
| Heterodontidæ | █ | █ | █ | █ | █ | █ | █ | |||||
| Tamiobatidæ | █ | |||||||||||
| Hexanchidæ | █ | █ | █ | █ | █ | |||||||
| Lamnidæ | █ | █ | █ | █ | █ | |||||||
| Mitsukurinidæ | █ | █ | █ | █ | ||||||||
| Odontaspidæ | █ | █ | █ | █ | ||||||||
| Scyliorhinidæ | █ | █ | █ | █ | █ | |||||||
| Carchariidæ | █ | █ | █ | █ | ||||||||
| Squalidæ | █ | █ | █ | █ | ||||||||
| Dalatiidæ | █ | █ | █ | |||||||||
| Squatinidæ | █ | █ | █ | █ | █ | |||||||
| Rhinobatidæ | █ | █ | █ | █ | █ | |||||||
| Pristididæ | █ | █ | █ | █ | ||||||||
| Rajidæ | █ | █ | █ | █ | ||||||||
| Narcobatidæ | █ | █ | █ | |||||||||
| Dasyatidæ | █ | █ | █ | █ | ||||||||
| Myliobatidæ | █ | █ | █ | █ | █ | |||||||
| Ptychodontidæ | █ | |||||||||||
| Chimæridæ | █ | █ | █ | █ | █ |
The Chordate Animals.—Referring to our metaphor of the tree with its twigs as used in the chapter on classification we find the fishes with the higher vertebrates as parts of a great branch from which the lower twigs have mostly perished. This great branch, phylum, or line of descent is known in zoology as Chordata, and the organisms associated with it or composing it are chordate animals.
The chordate animals are those which at some stage of life possess a notochord or primitive dorsal cartilage which divides the interior of the body into two cavities. The dorsal cavity contains the great nerve centers or spinal cord; the ventral cavity contains the heart and alimentary canal. In all other animals which possess a body cavity, there is no division by a notochord, and the ganglia of the nervous system if existing are placed on the ventral side or in a ring about the mouth.
The Protochordates.—Modern researches have shown that besides the ordinary back-boned animals certain other creatures easily to be mistaken for mollusks or worms and being chordate in structure must be regarded as offshoots from the vertebrate branch. These are degenerate allies, as is shown by the fact that their vertebrate traits are shown in their early or larval development and scarcely at all in their adult condition. As Dr. John Sterling Kingsley has well said: "Many of the species start in life with the promise of reaching a point high in the scale, but after a while they turn around and, as one might say, pursue a downward course, which results in an adult which displays but few resemblances to the other vertebrates." In the Tunicates or Ascidians (sea-squirts, sea-pears, and salpas), which constitute the class known as Tunicata or Urochordata, there is no brain, the notochord is confined to the tail and is usually present only in the larval stage of the animal when it has the form of a tadpole. In later life the animal usually becomes quiescent, attached to some hard object, fixed or floating. It loses its form and has the appearance of a hollow, leathery sac, the body organs being developed in a tough tunic. There are numerous families of Tunicates and the species are found in nearly all seas. They suggest no resemblance to fishes and look like tough clams without shells. The internal cavity being usually filled with water it is squirted out through the two apertures when the animal is handled. The class Enteropneusta (Adelochorda, or Hemichordata), includes the rather rare worm-like forms related to Balanoglossus. Bateson has shown that these animals possess a notochord which is developed in the anterior part of the body. They have no fins and before the mouth is a long proboscis. Gill-slits are found in the larval tunicate. In Balanoglossus these persist through life as in the fishes.
The remaining chordate forms constitute the vertebrates proper, not worm-like nor mollusk-like, the notochord not disappearing with age, except as it gives way, by specialized segmentation to the complex structures of the vertebral column. These vertebrates, which are permanently aquatic, are known in a popular sense as fishes. The fish, in the broad sense, is a back-boned animal which retains the homologue of the back-bone throughout life, which does not develop jointed limbs, its locomotive members, if present, being developed as fins, and which breathes through life the air contained in water by means of gills. This definition excludes the Tunicates and Enteropneusta on the one hand and the Amphibia or Batrachia with the reptiles, birds, and mammals on the other. The Amphibia are much more closely related to certain fishes than the classes of fishes are to each other. Still for purposes of systematic study, the frogs and salamanders are left out of the domain of ichthyology, while the Tunicata and the Enteropneusta might well be included in it.
The known branchiferous or gill-bearing chordates living and extinct may be first divided into eight classes—the Enteropneusta, the Tunicata, the Leptocardii, or lancelets, the Cyclostomi, or lampreys, the Elasmobranchii, or sharks, the Ostracophori the Arthrodira, and the Teleostomi, or true fishes. The first two groups, being very primitive and in no respect fish-like in appearance, are sometimes grouped together as Protochordata, the others with the higher Chordates constituting the Vertebrata.
Other Terms used in Classification.—The Leptocardii are sometimes called Acraniata (without skull), as distinguished from the higher groups, Craniota, in which the skull is developed. The Leptocardii, Cyclostomi, and Ostracophori are sometimes called Agnatha (without jaws) in contradistinction to the Gnathostomi (jaw mouths), which include the sharks and true fishes with the higher vertebrates. The sharks and Teleostomes are sometimes brought together as Pisces, or fishes, as distinguished from other groups not true fishes. To the sharks and true fishes the collective name of Lyrifera has been given, these fishes having the harp-shaped shoulder-girdle, its parts united below. The Ostracophores and Arthrodires agreeing in the bony coat of mail, and both groups now extinct and both of uncertain relationship, have been often united under the name of Placoderms, and these and many other fishes have been again erroneously confounded with the Ganoids. Again, the Teleostomi have been frequently divided into three classes—Crossopterygii, Dipneusti or Dipnoi, and Actinopterygii. The latter may be again divided into Ganoidei and Teleostei and all sorts of ranks have been assigned to each of these groups. For our purposes a division into eight classes is most convenient, and lowest among these we may place the Enteropneusta.
The Enteropneusta.—Most simple, most worm-like, and perhaps most primitive of all the Chordates is the group of worm-shaped forms, forming the class of Enteropneusta. The class of Enteropneusta, also called Adelochorda or Hemichordata, as here recognized, consists of a group of small marine animals allied to the genus Balanoglossus, or acorn-tongues (βάλανος, acorn; γλώσσα, tongue). These are worm-like creatures with fragile bodies buried in the sand or mud, or living under rocks of the seashore and in shallow waters, where they lie coiled in a spiral, with little or no motion. From the surface of the body a mucous substance is secreted, holding together particles by which are formed tubes of sand. The animal has a peculiar odor like that of iodoform. At the front is a long muscular proboscis, very sensitive, capable of great extension and contraction, largely used in burrowing in the ground, and of a brilliant orange color in life. Behind this is a collar which overlaps the small neck and conceals the small mouth at the base of the proboscis. The gill-slits behind the collar are also more or less concealed by it.
The body, which is worm-like, extends often to the length of two or three feet. The gill-slits in the adult are arranged in regular pairs, there being upwards of fifty in number much like the gill-slits of the lancelet. As the animal grows older the slits become less conspicuous, their openings being reduced to small slit-like pores.
In the interior of the proboscis is a rod-like structure which arises as an outgrowth of the alimentary canal above the mouth. In development and structure this rod so resembles the notochord of the lancelet that it is regarded as a true notochord, though found in the anterior region only. From the presence of gill-slits and notochord and from the development and structure of the central nervous system Balanoglossus was recognized by William Bateson, who studied an American species, Dolichoglossus kowalevskii, at Hampton Roads in Virginia in 1885, and at Beaufort in North Carolina, as a member of the Chordate series. Unlike the Tunicates it represents a primitively simple, not a degenerate, type. It seems to possess real affinities with the worms, or possibly, as some have thought, with the sea-urchins.
Fig. 275.—"Tornaria" Larva of Glossobalanus minutus. (After Minot.)
A peculiar little creature, known as Tornaria, was once considered to be the larva of a starfish. It is minute and transparent, floating on the surface of the sea. It has no visible resemblance to the adult Balanoglossus, but it has been reared in aquaria and shown to pass into the latter or into the related genus Glossobalanus. No such metamorphosis was found by Bateson in the more primitive genus Dolichoglossus, studied by him. This adult animal may be, indeed, a worm as it appears, but the presence of gill-slits, the existence of a rudimentary notochord, and the character of the central nervous system are distinctly fish-like and therefore vertebrate characters. With the Chordates, and not with the worms, this class, Enteropneusta (ἔντερον, intestine; πνεῖν, to breathe), must be placed if its characters have been rightly interpreted. It is possibly a descendant of the primitive creatures which marked the transition from the archaic worms, or possibly archaic Echinoderms, to the archaic Chordate type.
Fig. 276.—Glossobalanus minutus, one of the higher Enteropneustans. (After Minot.)
It is perhaps not absolutely certain that the notochord of Balanoglossus and its allies is a true homologue of the notochord of the lancelet. There may be doubt even of the homologies of the gill-slits themselves. But the balance of evidence seems to throw Balanoglossus on the fish side of the dividing line which separates the lower Chordates from the worms.
It may be noticed that Hubrecht regards the proboscis of various marine Nemertine worms as a real homologue of the notochord, and other writers have traced with more or less success other apparent or possible homologies between the Chordate and the Annelid series.
Classification of Enteropneusta.—Until recently the Enteropneusta have been usually placed in a single family or even in a single genus. The recent researches of Professor J. W. Spengel of Giessen and of Professor William Emerson Ritter of the University of California, have shown clearly that the group is much larger than had been generally supposed, with numerous species in all the warm seas. In Spengel's recent paper, "Die Benennung der Enteropneusten-Gattungen," three families are recognized with nine genera and numerous species. At least seven species are now known from the Pacific Coast of North America.
Family Harrimaniidæ.—In Harrimania maculosa, lately described by Dr. Ritter from Alaska, the eggs are large, with much food yolk, and the process of development is probably, without Tornaria stage. A second species of Harrimania (H. kupferi) is now recognized from Norway and Greenland. This genus is the simplest in structure among all the Enteropneustans and may be regarded as the lowest of known Chordates, the most worm-like of back-boned animals.
Fig. 277.—Harrimania maculosa (Ritter), the lowest of chordate animals. An Enteropneustan from Alaska. (After Ritter.)
In Dolichoglossus kowalevskii the species studied by Bateson on the Virginia coast, the same simplicity of development occurs. This genus, with a third, Stereobalanus (canadensis), constitutes in Spengel's system the family of Harrimaniidæ.
Balanoglossidæ.—The family Glandicepitidæ contains the genera Glandiceps, Spengelia, and Schizocardium. In the Balanoglossidæ (Ptychoderidæ of Spengel) the eggs are very small and numerous, with little food yolk. The species in this family pass through the Tornaria stage above described, a condition strikingly like that of the larval starfish. This fact has given rise to the suggestion that the Enteropneusta have a real affinity with the Echinoderms.
The Balanoglossidæ include the genera Glossobalanus, Balanoglossus, and Ptychodera, the latter the oldest known member of the group, its type, Ptychodera flava, having been described by Eschscholtz from the Pacific Coast in 1825, while Balanoglossus clavigerus was found by Della Chiaje in 1829.
Low Organization of Harrimaniidæ.—Apparently the Harrimaniidæ, with simpler structure, more extensive notochord, and direct development, should be placed at the bottom as the most primitive of the Enteropneustan series. Dr. Willey, however, regards its characters as due to degeneration, and considers the more elaborate Balanoglossidæ as nearest the primitive type. The case in this view would have something in common with that of the Larvacea, which seems to be the primitive Tunicates, but which may have been produced by the degeneration of more complex forms.
Structure of Tunicates.—One of the most singular groups of animals is that known as Ascidians, or Tunicates. It is one of the most clearly marked yet most heterogeneous of all the classes of animals, and in no other are the phenomena of degeneration so clearly shown.
Among them is a great variety of form and habit. Some lie buried in sand; some fasten themselves to rocks; some are imbedded in great colonies in a gelatinous matrix produced from their own bodies, and some float freely in long chains in the open sea. All agree in changing very early in their development from a free-swimming or fish-like condition to one of quiescence, remaining at rest or drifting with the current.
Says Dr. John Sterling Kingsley: "Many of the species start in life with the promise of reaching a point high in the scale, but after a while they turn around and, as one might say, pursue a downward course which results in an adult which displays but few resemblances to the other vertebrates. Indeed, so different do they seem that the fact that they belong here was not suspected until about thirty-five years ago. Before that time, ever since the days of Cuvier, they were almost universally regarded as mollusks, and many facts were adduced to show that they belonged near the acephals (clams, oysters, etc.). In the later years when the facts of development began to be known, this association was looked on with suspicion, and by some they were placed for a short time among the worms. Any one who has watched the phases of their development cannot help believing that they belong here, the lowest of the vertebrate series."
The following account of the structure and development of the Tunicate is taken, with considerable modification and condensation, from Professor Kingsley's chapter on the group in the Riverside Natural History. For the changes suggested I am indebted to the kindness of Professor William Emerson Ritter:
The Tunicates derive their name from the fact that the whole body is invested with a tough envelope or "tunic." This tunic or test may be either gelatinous, cartilaginous, or leathery. In some forms it is perfectly transparent, in others it is translucent, allowing enough light to pass to show the colors of the viscera, while in still others it is opaque and variously colored. The tunic is everywhere only loosely attached to the body proper, except in the region of the two openings now to be mentioned. One of these openings occupies a more or less central position, while the other is usually at one side, or it may even be placed at the opposite end of the body. On placing one of the Ascidians in a glass dish and sprinkling a little carmine or indigo in the water, we can study some of the functions of the animal. As soon as the disturbance is over, the animals will open the two apertures referred to, when it will be seen that each is surrounded with blunt lobes, the number of which varies with the species. As soon as they are opened a stream of water will be seen to rush into the central opening, carrying with it the carmine, and a moment later a reddish cloud will be ejected from the other aperture. From this we learn that the water passes through the body. Why it does so is to be our next inquiry. On cutting the animal open we find that the water, after passing through the first-mentioned opening (which may be called the mouth) enters a spacious chamber, the walls of which are made up of fine meshes, the whole appearing like lattice-work. Taking out a bit of this network and examining it under the microscope, we find that the edges of the meshes are armed with strong cilia, which are in constant motion, forcing the water through the holes. Of course, the supply has to be made good, and hence more water flows in through the mouth. This large cavity is known as the branchial or pharyngeal chamber. It is, according to Professor Ritter, "as we know from the embryology of the animal, the greatly enlarged anterior end of the digestive tract; and as the holes, or stigmata, as they are technically called, are perforations of the wall for the passage of water for purposes of respiration, they are both morphologically and physiologically comparable with the gill openings of fishes." There can be no doubt, therefore, that the pharyngeal sac of Ascidians is homologous with the pharynx of fishes.
Surrounding the mouth, or branchial orifice, just at its entrance into the branchial chamber is a circle of tentacles. These are simple in some genera, but elaborately branched in others.
In close connection with the cerebral ganglion, which is situated between the two siphons, there is a large gland with a short trumpet-shaped duct opening into the branchial sac a little distance behind the mouth. The orifice of the duct is just within a ring consisting of a ciliated groove that extends around the mouth outside the circle of branchial tentacles. On the opposite side of the mouth from the gland the ciliated groove joins another groove which is both ciliated and glandular, and which runs backward along the upper floor of the pharyngeal sac to its posterior extremity. This organ, called the endostyle, is concerned in the transportation of the animal's food through the pharyngeal sac to the opening of the œsophagus. Comparative embryology makes it almost certain that the subneural gland with its duct, described above, is homologous with the hypophesis cerebri of true vertebrates, and that the endostyle is homologous with the thyroid glands of vertebrates.
The water after passing through the branchial network is received into narrow passages and conducted to a larger cavity—the cloacal or atrial chamber. The general relations can he seen from our diagram, illustrating a vertical and horizontal section. From the atrial chamber the water flows out into the external world.
Now we can readily see how in the older works naturalists were misled as to the affinities of the Tunicates. They regarded the tunic as the equivalent of the mantle of the mollusks, while the incurrent and excurrent openings corresponded to the siphons. In one genus, Rhodosoma, the resemblance was even stronger, for there the tunic is in two parts, united by a hinge line, and closed by an adductor muscle. How and why these views were totally erroneous will be seen when we come to consider the development of these animals.
At the bottom of the pharnygeal sac is the narrow œsophagus surrounded with cilia, which force a current down into the digestive tract. The branchial meshes serve as a strainer for the water, and the larger particles which it contains fall down until they are within reach of the current going down the œsophagus. After passing through the throat, they come to the stomach, where digestion takes place, and then the ejectamenta are carried out through the intestine and poured into the bottom of the atrial cavity.
The heart lies on the ventral side of the stomach and is surrounded by a well-developed pericardium. The most remarkable fact connected with the circulation is that the heart, after beating a short time, forcing the blood through the vessels, will suddenly stop for a moment and then resume its beats; but, strange to say, after the stoppage the direction of the circulation is reversed, the blood taking an exactly opposite course from that formerly pursued. This most exceptional condition was first seen in the transparent Salpa, but it may be witnessed in the young of most genera. We have already referred to the branchial chamber. The walls of this chamber, besides acting as a strainer, are also respiratory organs. The meshes of which they are composed are in reality tubes through which the blood circulates and thus is brought in contact with a constantly renewed supply of fresh water.
The central nervous system in the adults of all except the Larvacea is reduced to a single ganglion placed near the mouth thus indicating the dorsal side. In forms like Cynthia it holds the same relative position with regard to the mouth, but by the doubling of the body (to be explained further on) it is also brought near the atrial aperture, where it is shown in our first diagram.
Development of Tunicates.—The sexes are combined in the same individual, though usually the products ripen at different times. As a rule, the earlier stages of the embryo are passed inside the cloacal chamber, though in some the development occurs outside the body. As a type of the development we will consider that of one of the solitary forms, leaving the many curious modifications to be noticed in connection with the species in which they occur. This will be best, since these forms show the relationship to the other vertebrates in the clearest manner.