Fig. 278.—Development of the larval Tunicate to the fixed condition. (From Seeliger, per Parker & Haswell.) a, larva; b, intermediate stage; c, adult.

The egg undergoes a total segmentation and a regular gastrulation. Soon a tail appears, and under the microscope the young embryo, which now begins its free life, appears much like the tadpole of the frog. It has a large oval body and a long tail which lashes about, forcing the animal forward with a wriggling motion. Nor is the resemblance superficial; it pervades every part of the structure, as may be seen from the adjacent diagram. The mouth is nearly terminal and communicates with a gill-chamber provided with gill-clefts. At the posterior end of the gill-chamber begins the alimentary tract, which pursues a convoluted course to the vent. In the tail, but not extending to any distance into the body, is an axial cylinder, the notochord, which here, as in all other vertebrates, arises from the hypoblast; and above it is the spinal cord (epiblastic in origin), which extends forward to the brain, above the gill-chamber. Besides, the animal is provided with organs of sight and hearing, which, however, are of peculiar construction and can hardly be homologized with the corresponding organs in vertebrates. So far the correspondence between the two types is very close, and if we knew nothing about the later stages, one would without doubt predict that the adult tunicate would reach a high point in the scale of vertebrates. These high expectations are never fulfilled; the animal, on the contrary, pursues a retrograde course, resulting in an adult whose relationship to the true vertebrates never would have been suspected had its embryology remained unknown.

Fig. 279.—Anatomy of Tunicate. (After Herdman, per Parker & Haswell.)

After the stage described this retrograde movement begins. From various parts of the body lobes grow out, armed on their extremities with sucking-disks. These soon come in contact with some subaquatic object and adhere to it. Then the notochord breaks down, the spinal cord is absorbed, the tail follows suit, the intestine twists around, and the cloaca is formed, the result being much like the diagram near the head of this section. In forms like Appendicularia, little degeneration takes place, so far as is known, the tail, with its notochord and neural chord, persisting through life.

Reproduction of Tunicates.—As to the reproduction of the Tunicates, Dr. Ritter writes: "In addition to the sexual method of reproduction, many tunicates reproduce asexually by budding. The capacity for bud reproduction appears to have been acquired by certain simple Ascidians in connection with, probably as a result of, their having given up the free-swimming life and become attached and consequently degenerate.

"Instructive as the embryonic development of the creatures is from the standpoint of evolution, the bud method of development is scarcely less so from the same point of view. The development of the adult zooid from the simple bud has been conclusively shown to be by a process in many respects fundamentally unlike that by which the individual is developed from the egg. We have then in these animals a case in which practically the same results are reached by developmental processes that are, according to prevailing conceptions of animal organizations, fundamentally different. This fact has hardly a parallel in the animal kingdom."

Habits of Tunicates.—The Tunicates are all marine, some floating or swimming freely, some attached to rocks or wharves, others buried in the sand. They feed on minute organisms, plants, or animals, occasional rare forms being found in their stomachs. Some of them possess a single median eye or eye-like structure which may not do more than recognize the presence of light. No fossil Tunicates are known, as they possess no hard parts, although certain Ostracoderms have been suspected, though on very uncertain grounds, to be mailed Tunicates, rather than mailed lampreys. It is not likely that this hypothesis has any sound foundation. The group is divided by Herdman and most other recent authorities into three orders, viz., the Larvacea, the Ascidiacea, and the Thaliacea.

Larvacea.—In the most primitive order the animals are minute and free-swimming, never passing beyond the tadpole stage. The notochord and the nervous chord persist through life, the latter with ganglionic segmentations at regular intervals. The species mostly float in the open sea, and some of them form from their own secretions a transparent gelatinous envelope called a "house." This has two apertures and a long chamber "in which the tail has room to vibrate."

The order consists of a single small family, Appendiculariidæ. The lowest type is known as Kowalevskia, a minute creature without heart or intestine found floating in the Mediterranean. It is in many respects the simplest in structure among Chordate animals. Oikopleura (Fig. 288) is another genus of this group.

Ascidiacea.—In the Ascidiacea the adult is usually attached to some object, and the two apertures are placed near each other by the obliteration of the caudal area. The form has been compared to a "leathern bottle with two spouts."

Fig. 280.Ascidia adhærens Ritter. Glacier Bay, Alaska. (After Ritter.)

The suborder Ascidiæ simplices includes the solitary Ascidians or "sea-squirts," common on our shores, as well as the social forms in which an individual is surrounded by its buds. The common name arises from the fact that when touched they contract, squirting water from both apertures. The Ascidiidæ comprise the most familiar solitary forms, some of them the largest of the Tunicates and represented on most coasts. In the Molgulidæ and most Ascidiæ compositæ the young hatch out in the cloaca, from which "these tadpoles swim out as yellow atoms," while in a new genus, Euherdmania, described by Ritter, from the coast of California, the embryos are retained through their whole larval stage in the oviduct of the parent. They form, according to Kingsley, adhesive processes on the body, but those of Molgula cannot use them in becoming attached to rocks, since they are entirely inclosed in a peculiar envelope. This envelope is after a while very adhesive, and if the little tadpole happens to touch any part of himself to a stone or shell he is fastened for life. Thus "I have frequently seen them adhere by the tail, while the anterior part was making the most violent struggles to escape. Soon, however, they settle down contentedly, absorb the tail, and in a few weeks assume the adult structure."

In the family Cynthiidæ the brightly-colored red and yellow species of Cynthia are known as sea-peaches by the fishermen. The sea-pears, Boltenia, are fastened to long stalks. These have a leathery and wrinkled tunic, to which algæ and hydroids freely attach themselves. Into the gill-cavity of these forms small fishes, blennies, gobies, and pearl-fishes often retreat for protection.

Fig. 281.Styela yacutatensis (Ritter), a simple Ascidian. Family Molgulidæ. Yakutat Bay, Alaska. (After Ritter.)

The social Ascidians constitute the Clavellinidæ. They are similar to the Ascidiidæ in form, but each individual sends out a bud which forms a stern bearing another individual at the end. By this means large colonies may be formed.

The suborder, Ascidiæ compositæ, contains the compound Ascidians or colonies enveloped in a common gelatinous "test." These colonies are usually attached to rock or seaweed, and the individuals are frequently regularly and symmetrically arranged. The bodies are sometimes complex in form.

Fig. 282.Styela greeleyi Ritter. Family Molgulidæ. Lukanin, Pribilof Islands. (After Ritter.)

Fig. 283.Cynthia superba Ritter. A Tunicate from Puget Sound. Family Cynthiidæ. (After Ritter.)

In the Botryllidæ and Polystyelidæ the individuals are not segmented and in the former family are arranged in star-shaped groups about a common cloaca, into which the atrial siphons of the different individuals open. The group springs by budding from the tadpole, or larva, which has attached itself to some object. These forms are often brightly colored. Botryllus gouldi is a species very common along our North Atlantic coast, forming gray star-shaped masses sometimes an inch across on eel-grass (Zostera) and on flat-leaved seaweeds. Goodsiria dura, a representative of the Polystyelidæ, is one of the most common Ascidians on the California coast southward, where the brick-red masses incrusting on seaweeds of various kinds, and on other Ascidians, are frequently thrown ashore in great quantities during heavy storms.

Fig. 284.Botryllus magnus Ritter. A compound Ascidian. Shumagin Islands, Alaska. (After Ritter.)

In Didemnidæ the body is more complex, of two parts, called the "thorax" and "abdomen." In Amarœcium, the "sea pork" of the fishermen, the body is in three parts and the individuals are very long. These sometimes form great masses a foot or more long, "colored like boiled salt pork, but more translucent." Other families of this type are the Distomidæ and the Polyclinidæ.

In the suborder Luciæ, including the family Pyrosomidæ, the colonies are thimble-shaped and hollow, the incurrent openings being on the outer surface of the thimble, the outgoing stream opening within. Pyrosoma is highly phosphorescent. In the tropical seas some colonies reach a length of two or three feet. It is said that a description of a colony was once written by a naturalist on a page illumined by the colony's own light. "Each of the individuals has a number of cells near the mouth the function of which is to produce the light."

Thaliacea.—In the order Thaliacea the Tunicates have the two orifices at opposite ends of the body. All are free-swimming and perfectly transparent. The principal family is that of Salpidæ. The gill-cavity in Salpa is much altered, the gills projecting into it dividing it into two chambers.

In these forms we have the phenomena of alternation of generations. A sexual female produces eggs, and from each hatches a tadpole larva which is without sex. This gives rise to buds, some at least of the individuals arising which in turn produce eggs.

Fig. 285.Botryllus magnus Ritter. Part of colony. (After Ritter.)

In the family Salpidæ two kinds of individuals occur, the solitary salpa, or female, and the chain salpa, or bisexual males. The latter are united together in long bands, each individual forming a link in the chain held together by spurs extending from one to the next. From each solitary individual a long process or cord grows out, this dividing to form the chain. Each chain salpa produces male reproductive organs and each develops as well a single egg. The egg is developed within the body attached by a sort of placenta, while the spermatozoa are cast into the sea to fertilize other eggs. From each egg develops the solitary salpa and from her buds the chain of bisexual creatures. Dr. W. K. Brooks regards these as nursing males, the real source of the egg being perhaps the solitary female. Of this extraordinary arrangement the naturalist-poet Chamisso, who first described it, said: "A salpa mother is not like its daughter or its own mother, but resembles its sister, its granddaughter, and its grandmother." But it is misleading to apply such terms taken from the individualized human relationship to the singular communal system developed by these ultra-degenerate and strangely specialized Chordates.

Fig. 286.Botryllus magnus Ritter, a single Zooid. Shumagin Islands, Alaska. (After Ritter.)

Fig. 287.Aplidiopsis jordani Ritter, a compound Ascidian. Lukanin Beach, Pribilof Islands. (After Ritter.)

The Salpas abound in the warm seas, the chains often covering the water for miles. They are perfectly transparent, and the chains are often more than a foot in length. In Doliolum the body is barrel-shaped and the gills are less modified than in Salpa. The alternation of generations in this genus is still more complicated than in Salpa, for here we have not only a sexual and a non-sexual generation, the individuals of which differ from each other, but there is further a differentiation among the asexually produced individuals themselves; so that we have in all three instead of two sorts of animals in the complete life cycle. Besides the proliferating stolon situated on the ventral side, the bud-producing individual possesses a dorsal process larger than the stolon proper. The buds become completely severed from the true stolon at an early stage and actually crawl along the side of the parent up to the dorsal process, upon which they arrange themselves in three rows, two lateral and one median. The buds of the lateral rows become nutritive and respiratory zooids, while those of the median row, ultimately at least, give rise in turn to the egg-producing individuals.

Origin of Tunicates.—There can be little doubt that the Tunicata form an offshoot from the primitive Chordate stock, and the structure of their larva in connection with that of the lancelet throws a large light on the nature of their common parents. "We may conclude," says Dr. Arthur Willey, "that the proximate ancestor of the Vertebrates was a free-swimming animal intermediate in organization between an Ascidian tadpole and Amphioxus, possessing the dorsal mouth, hypophysis, and restricted notochord of the former and the myotomes, cœlomic epithelium, and straight alimentary canal of the latter. The ultimate or primordial ancestor of the Vertebrates would, on the contrary, be a worm-like animal whose organization was approximately on a level with that of the bilateral ancestors of the Echinoderms."

Fig. 288.—Adult Tunicate of the group Larvacea, Oikopleura. Family Appendiculariidæ. (After Fol, per Parker & Haswell.)

Degeneration of Tunicates.—There is no question, furthermore, Professor Ritter observes, "that most of the group has undergone great degeneration in its evolutionary course. Just what the starting-point was, however, is a matter on which there is considerable difference of opinion among authorities. According to one view, particularly championed by Professor W. K. Brooks, Appendicularia is very near the ancestral form. The ancestor was consequently a small, marine, free-swimming creature. From this ancestor the Ascidiacea were evolved largely through the influence of the attached habit of life, and the tadpole stage in their development is a recapitulation of the ancestral form, just as the tadpole stage in the frog's life is a repetition of the fish ancestry of the frog.

"According to the most common view Appendicularia is not an ancestral form at all, but is the tadpole stage of the Ascidiacea that has failed to undergo metamorphosis and has become sexually mature in the larval condition, as the larva of certain Amphibians and insects are known to never pass into the adult state but reproduce their kind sexually in the larval condition. By this view the tadpole of such Ascidian as Ciona, for example, represents more closely the common ancestor of the group than does any other form we know. This view is especially defended by Professor K. Heider and Dr. Arthur Willey."


CHAPTER XXVII
THE LEPTOCARDII, OR LANCELETS

The Lancelet.—The lancelet is a vertebrate reduced to its very lowest terms. The essential organs of vertebrate life are there, but each one in its simplest form unspecialized and with structure and function feebly differentiated. The skeleton consists of a cartilaginous notochord inclosed in a membranous sheath. There is no skull. No limbs, no conspicuous processes, and no vertebræ are present. The heart is simply a long contractile tube, hence the name Leptocardii (from λεπτός, slender; καρδία, heart). The blood is colorless. There is a hepatic portal circulation. There is no brain, the spinal cord tapering in front as behind. The water for respiration passes through very many gill-slits from the pharynx into the atrium, from which it is excluded through the atripore in front of the vent. A large chamber, called the atrium, extends almost the length of the body along the ventral and lateral regions. It communicates with the pharynx through the gill-slits and with the exterior through a small opening in front of the vent, the atripore. The atrium is not found in forms above the lancelets.

The reproductive organs consist of a series of pairs of segmentally arranged gonads. The excretory organs consist of a series of tubules in the region of the pharynx, connecting the body-cavity with the atrium. The mouth is a lengthwise slit without jaws, and on either side is a row of fringes. From this feature comes the name Cirrostomi, from cirrus, a fringe of hair, and στόμα, mouth. The body is lanceolate in form, sharp at either end. From this fact arises a third name, Amphioxus, from ἀμφί, both; ὀξύς, sharp. Dorsal and anal fins are developed as folds of the skin supported by very slender rays. There are no other fins. The alimentary canal is straight, and is differentiated into pharynx and intestine; the liver is a blind sac arising from the anterior end of the intestine. A pigment spot in the wall of the spinal cord has been interpreted as an eye. Above the snout is a supposed olfactory pit which some have thought to be connected with the pineal structure. The muscular impressions along the sides are very distinct and it is chiefly by means of the variation in numbers of these that the species can be distinguished. Thus in the common lancelet of Europe, Branchiostoma lanceolatum, the muscular bands are 35+14+12=61. In the common species of the Eastern coasts of America, Branchiostoma caribæum, these are 35+14+9=58, while in the California lancelet, Branchiostoma californiense, these are 44+16+9=69.

Habits of Lancelets.—Lancelets are slender translucent worm-like creatures, varying from half an inch (Asymmetron lucayanum) to four inches (Branchiostoma californiense) in length. They live buried in sand in shallow waters along the coasts of warm seas. One species, Amphioxides pelagicus, has been taken at the depth of 1000 fathoms, but whether at the bottom or floating near the surface is not known. The species are very tenacious of life and will endure considerable mutilation. Some of them are found on almost every coast in semi-tropical and tropical regions.

Species of Lancelets.—The Mediterranean species ranges northward to the south of England. Others are found as far north as Chesapeake Bay, San Diego, and Misaki in Japan, where is found a species called Branchiostoma belcheri. The sands at the mouth of San Diego Bay are noted as producing the largest of the species of lancelets, Branchiostoma californiense. From the Bahamas comes the smallest, the type of a distinct genus, Asymmetron lucayanum, distinguished among other things by a projecting tail. Other supposed genera are Amphioxides (pelagicus), dredged in the deep sea off Hawaii and supposed to be pelagic, the mouth without cirri; Epigonichthys (cultellus), from the East Indies, and Heteropleuron (bassanum), from Bass Straits, Australia. These little animals are of great interest to anatomists as giving the clue to the primitive structure of vertebrates. While possibly these have diverged widely from their actual common ancestry with the fishes, they must approach near to these in many ways. Their simplicity is largely primitive, not, as in the Tunicates, the result of subsequent degradation.

Fig. 289.—California Lancelet, Branchiostoma californiense Gill. (From San Diego.)

The lancelets, less than a dozen species in all, constitute a single family, Branchiostomidæ. The principal genus, Branchiostoma, is usually called Amphioxus by anatomists. But while the name Amphioxus, like lancelet, is convenient in vernacular use, it has no standing in systematic nomenclature. The name Branchiostoma was given to lancelets from Naples in 1834, by Costa, while that of Amphioxus, given to specimens from Cornwall, dates from Yarrell's work on the British fishes in 1836. The name Amphioxus may be pleasanter or shorter or more familiar or more correctly descriptive than Branchiostoma, but if so the fact cannot be considered in science as affecting the duty of priority.

The name Acraniata (without skull) is often used for the lower Chordates taken collectively, and it is sometimes applied to the lancelets alone. It refers to those chordate forms which have no skull nor brain, as distinguished from the Craniota, or forms with a distinct brain having a bony or cartilaginous capsule for its protection.

Origin of Lancelets.—It is doubtless true, as Dr. Willey suggests, that the Vertebrates became separated from their worm-like ancestry through "the concentration of the central nervous system along the dorsal side of the body and its conversion into a hollow tube." Besides this trait two others are common to all of them, the presence of the gill-slits and that of the notochord. The gill-slits may have served primarily to relieve the stomach of water, as in the lowest forms they enter directly into the body-cavity. The primitive function of the notochord is still far from clear, but its ultimate use of its structures in affording protection and in furnishing a fulcrum for the muscles and limbs is of the greatest importance in the processes of life.

Fig. 289a.—Gill-basket of Lamprey.


CHAPTER XXVIII
THE CYCLOSTOMES, OR LAMPREYS

The Lampreys.—Passing upward from the lancelets and setting aside the descending series of Tunicates, we have a long step indeed to the next class of fish-like vertebrates. During the period this great gap represents in time we have the development of brain, skull, heart, and other differentiated organs replacing the simple structures found in the lancelet.

The presence of brain without limbs and without coat-of-mail distinguishes the class of Cyclostomes, or lampreys (κυκλός, round; στόμα, mouth). This group is also known as Marsipobranchi (μαρσιπίον, pouch; βράγχος, gill); Dermopteri (δέρμα, skin; πτερόν, fin); and Myzontes (μυζάω, to suck). It includes the forms known as lampreys, slime-eels, and hagfishes.

Structure of the Lamprey.—Comparing a Cyclostome with a lancelet we may see many evidences of specialization in structure. The Cyclostome has a distinct head with a cranium formed of a continuous body of cartilage modified to contain a fish-like brain, a cartilaginous skeleton of which the cranium is evidently a differentiated part. The vertebræ are undeveloped, the notochord being surrounded by its membranes, without bony or cartilaginous segments. The gills have the form of fixed sacs, six to fourteen in number, on each side, arranged in a cartilaginous structure known as "branchial basket" (fig. 289a), the elements of which are not clearly homologous with the gill-arches of the true fishes. Fish-like eyes are developed on the sides of the head. There is a median nostril associated with a pituitary pouch, which pierces the skull floor. An ear-capsule is developed. The brain is composed of paired ganglia in general appearance resembling the brain of the true fish, but the detailed homology of its different parts offers considerable uncertainty. The heart is modified to form two pulsating cavities, auricle and ventricle. The folds of the dorsal and anal fins are distinct, supported by slender rays.

The mouth is a roundish disk, with rasping teeth over its surface and with sharper and stronger teeth on the tongue. The intestine is straight and simple. The kidney is represented by a highly primitive pronephros and no trace exists of an air-bladder or lung. The skin is smooth and naked, sometimes secreting an excessive quantity of slime.

From the true fishes the Cyclostomes differ in the total absence of limbs and of shoulder and pelvic girdles, as well as of jaws. It has been thought by some writers that the limbs were ancestrally present and lost through degeneration, as in the eels. Dr. Ayers, following Huxley, finds evidence of the ancestral existence of a lower jaw. The majority of observers, however, regard the absence of limbs and jaws in Cyclostomes as a primitive character, although numerous other features of the modern hagfish and lamprey may have resulted from degeneration. There is no clear evidence that the class of Cyclostomes, as now known to us, has any great antiquity, and its members may be all degenerate offshoots from types of greater complexity of structure.

Supposed Extinct Cyclostomes.—No species belonging to the class of Cyclostomes has been found fossil. We may reason theoretically that the earliest fish-like forms were acraniate or lancelet-like, and that lamprey-like forms would naturally follow these, but this view cannot be substantiated from the fossils. Lancelets have no hard parts whatever, and could probably leave no trace in any sedimentary deposit. The lampreys stand between lancelets and sharks. Their teeth and fins at least might occasionally be preserved in the rocks, but no structures certainly known to be such have yet been recognized. It is however reasonably certain that the modern lamprey and hagfish are descendants, doubtless degraded and otherwise modified from species which filled the gap between the earliest chordate animals and the jaw-bearing sharks.

Conodontes.—Certain structures found as fossils have been from time to time regarded as Cyclostomes, but in all such cases there is doubt as to the real nature of the fossil relic in question or as to the proper interpretation of its relationship.

Thus the Conodontes of the Cambrian, Silurian, and Devonian have been regarded as lingual teeth of extinct Cyclostomes. The Cycliæ of the Devonian have been considered as minute lampreys, although the vertebral segments are highly specialized, to a degree far beyond the condition seen in the lampreys of to-day. The Ostracophores have been regarded as monstrous lampreys in coat of mail, and the possibility of a lamprey origin even for Arthrodires has been suggested. The Cycliæ and Ostracophori were apparently without jaws or limbs, being in this regard like the Cyclostomes, but their ancestry and relationships are wholly problematical.

Fig. 290.Polygnathus dubium Hinde. A Conodont from the New York Devonian. (After Hinde.)

The nature of the Conodontes is still uncertain. In form they resemble teeth, but their structure is different from that of the teeth of any fishes, agreeing with that of the teeth of annelid worms. Some have compared them to the armature of Trilobites. Some fifteen nominal genera are described by Pander in Russia, and by Hinde about Lake Erie and Lake Ontario. Some of these, as Drepaniodus, are simple, straight or curved grooved teeth or tooth-like structures; others, as Prioniodus, have numerous smaller teeth or denticles at the base of the larger one.

Orders of Cyclostomes.—The known Cyclostomes are naturally divided into two orders, the Hyperotreta, or hagfishes, and the Hyperoartia, or lampreys. These two orders are very distinct from each other. While the two groups agree in the general form of the body, they differ in almost every detail, and there is much pertinence in Lankester's suggestions that each should stand as a separate class. The ancestral forms of each, as well as the intervening types if such ever existed, are left unrecorded in the rocks.

The Hyperotreta, or Hagfishes.—The Hyperotreta (ὑπερῴα, palate; τρετός, perforate), or hagfishes, have the nostril highly developed, a tube-like cylinder with cartilaginous rings penetrating the palate. In these the eyes are little developed and the species are parasitic on other fishes. In Polistotrema stouti, the hagfish of the coast of California, is parasitic on large fishes, rockfishes, or flounders. It usually fastens itself at the throat or isthmus of its host and sometimes at the eyes. Thence it works very rapidly to the inside of the body. It there devours all the muscular part of the fish without breaking the skin or the peritoneum, leaving the fish a living hulk of head, skin, and bones. It is especially destructive to fishes taken in gill-nets. The voracity of the Chilean species Polistotrema dombeyi is equally remarkable. Dr. Federico T. Delfin finds that in seven hours a hagfish of this species will devour eighteen times its own weight of fish-flesh. The intestinal canal is a simple tube, through which most of the food passes undigested. The eggs are large, each in a yellowish horny case, at one end of which are barbed threads by which they cling together and to kelp or other objects. In the California hagfish, Polistotrema stouti, great numbers of these eggs have been found in the stomachs of the males.

Fig. 291.—California Hagfish, Polistotrema stouti Lockington.

Similar habits are possessed by all the species in the two families, Myxinidæ and Eptatretidæ. In the Myxinidæ the gill-openings are apparently single on each side, the six gills being internal and leading by six separate ducts to each of the six branchial sacs. The skin is excessively slimy, the extensible tongue is armed with two cone-like series of strong teeth. About the mouth are eight barbels.

Of Myxine, numerous species are known—Myxine glutinosa, in the north of Europe; Myxine limosa, of the West Atlantic; Myxine australis, and several others about Cape Horn, and Myxine garmani in Japan. All live in deep waters and none have been fully studied. It has been claimed that the hagfish is male when young, many individuals gradually changing to female, but this conclusion lacks verification and is doubtless without foundation.

In the Eptatretidæ the gill-openings, six to fourteen in number, are externally separate, each with its own branchial sac as in the lampreys.

The species of the genus Eptatretus (Bdellostoma, Heptatrema, and Homea, all later names for the same group) are found only in the Pacific, in California, Chile, Patagonia, South Africa, and Japan. In general appearance and habits these agree with the species of Myxine. The species with ten to fourteen gill-openings (dombeyi: stouti) are sometimes set off as a distinct genus (Polistotrema), but in other regards the species differ little, and frequent individual variations occur. Eptatretus burgeri is found in Japan and Eptatretus forsteri in Australia.

The Hyperoartia, or Lampreys.—In the order Hyperoartia, or lampreys, the single nostril is a blind sac which does not penetrate the palate. The seven gill-openings lead each to a separate sac, the skin is not especially covered with mucus, the eyes are well developed in the adult, and the mouth is a round disk armed with rasp-like teeth, the comb-like teeth on the tongue being less developed than in the hagfishes. The intestine in the lampreys has a spiral valve. The eggs are small and are usually laid in brooks away from the sea, and in most cases the adult lamprey dies after spawning. According to Thoreau, "it is thought by fishermen that they never return, but waste away and die, clinging to rocks and stumps of trees for an indefinite period, a tragic feature in the scenery of the river-bottoms worthy to be remembered with Shakespeare's description of the sea-floor." This account is not far from the truth, as recent studies have shown.

The lampreys of the northern regions constitute the family of Petromyzonidæ. The larger species (Petromyzon, Entosphenus) live in the sea, ascending rivers to spawn, and often becoming land-locked and reduced in size by living in rivers only. Such land-locked marine lampreys (Petromyzon marinus unicolor) breed in Cayuga Lake and other lakes in New York. The marine forms reach a length of three feet. Smaller lampreys of other genera six inches to eighteen inches in length remain all their lives in the rivers, ascending the little brooks in the spring, clinging to stones and clods of earth till their eggs are deposited. These are found throughout northern Europe, northern Asia, and the colder parts of North America, belonging to the genera Lampetra and Ichthyomyzon. Other and more aberrant genera from Chile and Australia are Geotria and Mordacia, the latter forming a distinct family, Mordaciidæ. In Geotria, a large and peculiar gular pouch is developed at the throat. In Macrophthalmia chilensis from Chile the eyes are large and conspicuous.

Food of Lampreys.—The lampreys feed on the blood and flesh of fishes. They attach themselves to the sides of the various species, rasp off the flesh with their teeth, sucking the blood till the fish weakens and dies. Preparations made by students of Professor Jacob Reighard in the University of Michigan show clearly that the lamprey stomach contains muscular tissue as well as the blood of fishes. The river species do a great deal of mischief, a fact which has been the subject of a valuable investigation by Professor H. A. Surface, who has also considered the methods available for their destruction. The flesh of the lamprey is wholesome, and the larger species, especially the great sea lamprey of the Atlantic, Petromyzon marinus, are valued as food. The small species, according to Prof. Gage, never feed on fishes.

Fig. 292.—Lamprey, Petromyzon marinus L. Woods Hole, Mass.

Metamorphosis of Lampreys.—All lampreys, so far as known, pass through a distinct metamorphosis. The young, known as the Ammocœtes form, are slender, eyeless, and with the mouth narrow and toothless. From Professor Surface's paper on "The Removal of Lampreys from the Interior Waters of New York" we have the following extracts (slightly condensed):

Fig. 293.Petromyzon marinus unicolor (De Kay). Mouth of Lake Lamprey, Cayuga Lake. (After Gage.)

Fig. 294.Lampetra wilderi Jordan & Evermann. Larval Brook Lamprey in its burrow in a glass filled with sand. (After Gage.)

Fig. 295.Lampetra wilderi Jordan & Evermann. Mouth of Brook Lamprey. Cayuga Lake. (After Gage.)

"In the latter part of the fall the young lampreys, Petromyzon marinus unicolor, the variety land-locked in the lakes of Central New York, metamorphose and assume the form of the adult. They are now about six or eight inches long. The externally segmented condition of the body disappears. The eyes appear to grow out through the skin and become plainly visible and functional. The mouth is no longer filled with vertical membranous sheets to act as a sieve, but it contains nearly one hundred and fifty sharp and chitinous teeth, arranged in rows that are more or less concentric and at the same time presenting the appearance of circular radiation. These teeth are very strong, with sharp points, and in structure each has the appearance of a hollow cone of chitin placed over another cone or papilla. A little below the center of the mouth is the oral opening, which is circular and contains a flattened tongue which bears finer teeth of chitin set closely together and arranged in two interrupted (appearing as four) curved rows extending up and down from the ventral toward the dorsal side of the mouth. Around the mouth is a circle of soft membrane finally surrounded by a margin of fimbriæ or small fringe. This completes the apparatus with which the lamprey attaches itself to its victims, takes its food, carries stones, builds and tears down its nest, seizes its mate, holds itself in position in a strong current, and climbs over falls."

Mischief Done by Lampreys.—"The most common economic feature in the entire life history of these animals is their feeding habits in this (spawning) stage, their food now consisting wholly of the blood (and flesh) of fishes. A lamprey is able to strike its suctorial mouth against a fish, and in an instant becomes so firmly attached that it is very rarely indeed that the efforts of the fish will avail to rid itself of its persecutor. When a lamprey attaches itself to a person's hand in the aquarium, it can only be freed by lifting it from the water. As a rule it will drop the instant it is exposed to the open air, although often it will remain attached for some time even in the open air, or may attach itself to an object while out of water.

"Nearly all lampreys that are attached to fish when they are caught in nets will escape through the meshes of the nets, but some are occasionally brought ashore and may hang on to their victim with bulldog pertinacity.

"The fishes that are mostly attacked are of the soft-rayed species, having cycloid scales, the spiny-rayed species with ctenoid scales being most nearly immune from their attacks. We think there may be three reasons for this: 1st, the fishes of the latter group are generally more alert and more active than those of the former, and may be able more readily to dart away from such enemies; 2d, their scales are thicker and stronger and appear to be more firmly imbedded in the skin, consequently it is more difficult for the lampreys to hold on and cut through the heavier coat-of-mail to obtain the blood of the victim; 3d, since the fishes of the second group are wholly carnivorous and in fact almost exclusively fish-eating when adult, in every body of water they are more rare than those of the first group, which are more nearly omnivorous. According to the laws and requirements of nature the fishes of the first group must be more abundant, as they become the food for those of the second, and it is on account of their greater abundance that the lampreys' attacks on them are more observed.

"There is no doubt that the bullhead, or horned pout (Ameiurus nebulosus), is by far the greatest sufferer from lamprey attacks in Cayuga Lake. This may be due in part to the sluggish habits of the fish, which render it an easy victim, but it is more likely due to the fact that this fish has no scales and the lamprey has nothing to do but to pierce the thick skin and find its feast of blood ready for it. There is no doubt of the excellency of the bullhead as a food-fish and of its increasing favor with mankind. It is at present the most important food- and market-fish of the State (New York), being caught by bushels in the early part of June when preparing to spawn. As we have observed at times more than ninety per cent. of the catch attacked by lampreys, it can readily be seen how very serious are the attacks of this terrible parasite which is surely devastating our lakes and streams."

Migration or "Running" of Lampreys.—"After thus feeding to an unusual extent, their reproductive elements (gonads) become mature and their alimentary canals commence to atrophy. This duct finally becomes so occluded that from formerly being large enough to admit a lead-pencil of average size when forced through it, later not even liquids can pass through, and it becomes nearly a thread closely surrounded by the crowding reproductive organs. When these changes commence to ensue, the lampreys turn their heads against the current and set out on their long journeys to the sites that are favorable for spawning, which here may be from two to eight miles from the lake. In this migration they are true to their instincts and habits of laziness in being carried about, as they make use of any available object, such as a fish, boat, etc., that is going in their direction, fastening to it with their suctorial mouths and being borne along at their ease. During this season it is not infrequent that as the Cornell crews come in from practice and lift their shells from the water, they find lampreys clinging to the bottoms of the boats, sometimes as many as fifty at one time. They are likely to crowd up all streams flowing into the lake, inspecting the bed of the stream as they go. They do not stop until they reach favorable spawning sites, and if they find unsurmountable obstacles in their way, such as vertical falls or dams, they turn around and go down-stream until they find another, up which they go. This is proved every spring by the number of adult lampreys which are seen temporarily in Pall Creek and Cascadilla Creek. In each of these streams, about a mile from its mouth, there is a vertical fall over thirty feet in height which the lampreys cannot surmount, and in fact they have never been seen attempting to do so. After clinging with their mouths to the stones at the foot of the falls for a few days, they work their way down-stream, carefully inspecting all the bottom for suitable spawning sites. They do not spawn in these streams because there are too many rocks and no sand, but finally enter the only stream (the Cayuga Lake inlet) in which they find suitable and accessible spawning sites.