The gonads are among the oldest organs, the few other organs that we find in the Platodes between the gut-wall and body-wall being later evolutionary products. One of the oldest and most important of these are the kidneys or nephridia, which remove unusable matter from the body (Figure 2.240 nc). These urinary or excretory organs were originally enlarged skin-glands—a couple of canals that run the length of the body, and have a separate or common external aperture (nm). They often have a number of branches. These special excretory organs are not found in the other Coelenteria (Gastraeads, Sponges, Cnidaria) or the Cryptocoela. They are first met in the Turbellaria, and have been transmitted direct from these to the Vermalia, and from these to the higher stems.

Finally, there is a very important new organ in the Turbellaria, which we do not find in the Cryptocoela (Figure 2.239) and their gastraead ancestors—the rudimentary nervous system. It consists of a couple of simple cerebral ganglia (Figure 2.241 g) and fine nervous fibres that radiate from them; these are partly voluntary nerves (or motor fibres) that go to the thin muscular layer developing under the skin; and partly sensory nerves that proceed to the sense-cells of the ciliated epiderm (f). Many of the Turbellaria have also special sense-organs; a couple of ciliated smell pits (na), rudimentary eyes (au), and, less frequently, auditory vesicles.

On these principles I assume that the oldest and simplest Turbellaria arose from Platodaria, and these directly from bilateral Gastraeads. The chief advances were the formation of gonads and nephridia, and of the rudimentary brain. On this hypothesis, which I advanced in 1872 in the first sketch of the gastraea-theory (Monograph on the Sponges), there is no direct affinity between the Platodes and the Cnidaria.

(FIGURE 2.240. A simple turbellarian (Rhabdocoelum). m mouth, sd gullet epithelium, sm gullet muscles, d gastric gut, nc renal canals, nm renal aperture, au eye, na olfactory pit. (Diagram.)

FIGURE 2.241. The same, showing the other organs. g brain, au eye, na olfactory pit, n nerves, h testicles, male symbol male aperture, female symbol female aperture, e ovary, f ciliated epiderm. (Diagram.)

(FIGURES 242 AND 243. Chaetonotus, a rudimentary vermalian, of the group of Gastrotricha. m mouth, s gullet, d gut, a anus, g brain, n nerves, ss sensory hairs, au eye, ms muscular cells, h skin, f ciliated bands of the ventral surface, nc nephridia, nm their aperture, e ovaries.))

Next to the ancient stem-group of the Turbellaria come a number of more recent chordonia ancestors, which we class with the Vermalia or Helminthes, the unarticulated worms. These true worms (Vermes, lately also called Scolecida) are the difficulty or the lumber-room of the zoological classifier, because the various classes have very complicated relations to the lower Platodes on the one hand and the more advanced animals on the other. But if we exclude the Platodes and the Annelids from this stem, we find a fairly satisfactory unity of organisation in the remaining classes. Among these worms we find some important forms that show considerable advance in organisation from the platode to the chordonia stage. Three of these phenomena are particularly instructive: (1) The formation of a true (secondary) body-cavity (coeloma); (2) the formation of a second aperture of the gut, the anus; and (3) the formation of a vascular system. The great majority of the Vermalia have these three features, and they are all wanting in the Platodes; in the rest of the worms at least one or two of them are developed.

Next and very close to the Platodes we have the Ichthydina (Gastrotricha), little marine and fresh-water worms, about 1/250 to 1/1000 inch long. Zoologists differ as to their position in classification. In my opinion, they approach very close to the Rhabdocoela (Figures 2.240 and 2.241), and differ from them chiefly in the possession of an anus at the posterior end (Figure 2.242 a). Further, the cilia that cover the whole surface of the Turbellaria are confined in the Gastrotricha to two ciliated bands (f) on the ventral surface of the oval body, the dorsal surface having bristles. Otherwise the organisation of the two classes is the same. In both the gut consists of a muscular gullet (s) and a glandular primitive gut (d). Over the gullet is a double brain (acroganglion, g). At the side of the gut are two serpentine prorenal canals (water-vessels or pronephridia, nc), which open on the ventral side (nm). Behind are a pair of simple sexual glands or gonads (Figure 2.243 e).

While the Ichthydina are thus closely related to the Platodes, we have to go farther away for the two classes of Vermalia which we unite in the group of the "snout-worms" (Frontonia). These are the Nemertina and the Enteropneusta. Both classes have a complete ciliary coat on the epidermis, a heritage from the Turbellaria and the Gastraeads; also, both have two openings of the gut, the mouth and anus, like the Gastrotricha. But we find also an important organ that is wanting in the preceding forms—the vascular system. In their more advanced mesoderm we find a few contractile longitudinal canals which force the blood through the body by their contractions; these are the first blood-vessels.

(FIGURE 2.244. A simple Nemertine. m mouth, d gut, a anus, g brain, n nerves, h ciliary coat, ss sensory pits (head-clefts), au eyes, r dorsal vessel, l lateral vessels. (Diagram.)

FIGURE 2.245. A young Enteropneust (Balanaglossus). (From Alexander Agassiz.) r acorn-shaped snout, h neck, k gill-clefts and gill-arches of the fore-gut, in long rows on each side, d digestive hind-gut, filling the greater part of the body-cavity, v intestinal vein or ventral vessel, lying between the parallel folds of the skin, a anus.

Figure 2.246. Transverse section of the branchial gut. A of Balanoglossus, B of Ascidia. r branchial gut, n pharyngeal groove, asterisk ventral folds between the two. Diagrammatic illustration from Gegenbaur, to show the relation of the dorsal branchial-gut cavity (r) to the pharyngeal or hypobranchial groove (n).)

The Nemertina were formerly classed with the much less advanced Turbellaria. But they differ essentially from them in having an anus and blood-vessels, and several other marks of higher organisation. They have generally long and narrow bodies, like a more or less flattened cord; there are, besides several small species, giant-forms with a width of 1/5 to 2/5 inch and a length of several yards (even ten to fifteen). Most of them live in the sea, but some in fresh water and moist earth. In their internal structure they approach the Turbellaria on the one hand and the higher Vermalia (especially the Enteropneusta) on the other. They have a good deal of interest as the lowest and oldest of all animals with blood. In them we find blood-vessels for the first time, distributing real blood through the body. The blood is red, and the red colouring-matter is haemoglobin, connected with elliptic discoid blood-cells, as in the Vertebrates. Most of them have two or three parallel blood-canals, which run the whole length of the body, and are connected in front and behind by loops, and often by a number of ring-shaped pieces. The chief of these primitive blood-vessels is the one that lies above the gut in the middle line of the back (Figure 2.244 r); it may be compared to either the dorsal vessel of the Articulates or the aorta of the Vertebrates. To the right and left are the two serpentine lateral vessels (Figure 2.244 l).

After the Nemertina, I take (as distant relatives) the Enteropneusta; they may be classed together with them as Frontonia or Rhyncocoela (snout-worms). There is now only one genus of this class, with several species (Balanoglossus); but it is very remarkable, and may be regarded as the last survivor of an ancient and long-extinct class of Vermalia. They are related, on the one hand, to the Nemertina and their immediate ancestors, the Platodes, and to the lowest and oldest forms of the Chordonia on the other.

The Enteropneusta (Figure 2.245) live in the sea sand, and are long worms of very simple shape, like the Nemertina. From the latter they have inherited: (1) The bilateral type, with incomplete segmentation; (2) the ciliary coat of the soft epidermis; (3) the double rows of gastric pouches, alternating with a single or double row of gonads; (4) separation of the sexes (the Platode ancestors were hermaphroditic); (5) the ventral mouth, underneath a protruding snout; (6) the anus terminating the simple gut-tube; and (7) several parallel blood-canals, running the length of the body, a dorsal and a ventral principal stem.

On the other hand, the Enteropneusta differ from their Nemertine ancestors in several features, some of which are important, that we may attribute to adaptation. The chief of these is the branchial gut (Figure 2.245 k). The anterior section of the gut is converted into a respiratory organ, and pierced by two rows of gill-clefts; between these there is a branchial (gill) skeleton, formed of rods and plates of chitine. The water that enters at the mouth makes its exit by these clefts. They lie in the dorsal half of the fore-gut, and this is completely separated from the ventral half by two longitudinal folds (Figure 2.246 A*). This ventral half, the glandular walls of which are clothed with ciliary epithelium and secrete mucus, corresponds to the pharyngeal or hypo-branchial groove of the Chordonia (Bn), the important organ from which the later thyroid gland is developed in the Craniota (cf. Chapter 2.16). The agreement in the structure of the branchial gut of the Enteropneusts, Tunicates, and Vertebrates was first recognised by Gegenbaur (1878); it is the more significant as at first we find only a couple of gill-clefts in the young animals of all three groups; the number gradually increases. We can infer from this the common descent of the three groups with all the more confidence when we find the Balanoglossus approaching the Chordonia in other respects. Thus, for instance, the chief part of the central nervous system is a long dorsal neural string that runs above the gut and corresponds to the medullary tube of the Chordonia. Bateson believes he has detected a rudimentary chorda between the two.

Of all extant invertebrate animals the Enteropneusts come nearest to the Chordonia in virtue of these peculiar characters; hence we may regard them as the survivors of the ancient gut-breathing Vermalia from which the Chordonia also have descended. Again, of all the chorda-animals the Copelata (Figure 2.225) and the tailed larvae of the ascidia approach nearest to the young Balanoglossus. Both are, on the other hand, very closely related to the Amphioxus, the Primitive Vertebrate of which we have considered the importance (Chapters 2.16 and 2.17). As we saw there, the unarticulated Tunicates and the articulated Vertebrates must be regarded as two independent stems, that have developed in divergent directions. But the common root of the two stems, the extinct group of the Prochordonia, must be sought in the vermalia stem; and of all the living Vermalia those we have considered give us the safest clue to their origin. It is true that the actual representatives of the important groups of the Copelata, Balanoglossi, Nemertina, Icthydina, etc., have more or less departed from the primitive model owing to adaptation to special environment. But we may just as confidently affirm that the main features of their organisation have been preserved by heredity.

We must grant, however, that in the whole stem-history of the Vertebrates the long stretch from the Gastraeads and Platodes up to the oldest Chordonia remains by far the most obscure section. We might frame another hypothesis to raise the difficulty—namely, that there was a long series of very different and totally extinct forms between the Gastraea and the Chordaea. Even in this modified chordaea-theory the six fundamental organs of the chordula would retain their great value. The medullary tube would be originally a chemical sensory organ, a dorsal olfactory tube, taking in respiratory-water and food by the neuroporus in front and conveying them by the neurenteric canal into the primitive gut. This olfactory tube would afterwards become the nervous centre, while the expanding gonads (lying to right and left of the primitive mouth) would form the coeloma. The chorda may have been originally a digestive glandular groove in the dorsal middle line of the primitive gut. The two secondary gut-openings, mouth and anus, may have arisen in various ways by change of functions. In any case, we should ascribe the same high value to the chordula as we did before to the gastrula.

In order to explain more fully the chief stages in the advance of our race, I add the hypothetical sketch of man's ancestry that I published in my Last Link [a translation by Dr. Gadow of the paper read at the International Zoological Congress at Cambridge in 1898]:—

A. MAN'S GENEALOGICAL TREE, FIRST HALF: EARLIER SERIES OF ANCESTORS, WITHOUT FOSSIL EVIDENCE.

COLUMN 1 : CHIEF STAGES. COLUMN 2 : ANCESTRAL STEM-GROUPS. COLUMN 3 : LIVING RELATIVES OF ANCESTORS.

STAGES 1 TO 5. PROTIST ANCESTORS. UNICELLULAR ORGANISMS.

1 to 2. Protophytes. : 1. Monera. Without nucleus. : Chromacea. (Chroococcus.) Phycochromacea.

1 to 2. Protophytes. : 2. Algaria. Unicellular algae. : 2. Paulotomea. Palmellacea. Eremosphaera.

3 to 5. Protozoa. : 3. Lobosa. Unicellular (amoebina) rhizopods. : 3. Amoebina. Amoeba Leucocyta.

3 to 5. Protozoa. : 4. Infusoria. Unicellular. : 4. Flagellata. Euflagellata. Zoomonades.

3 to 5. Protozoa. : 5. Blastaeades. Multicellular hollow spheres. : 5. Catallacta. Magosphaera, Volvocina, Blastula.

STAGES 6 TO 11. INVERTEBRATE METAZOA ANCESTORS.

6 to 8. Coelenteria, without anus and body-cavity. : 6. Gastraeades. With two germ-layers. : 6. Gastrula. Hydra, Olynthus, Gastremaria.

6 to 8. Coelenteria, without anus and body-cavity. : 7. Platodes I. Platodaria (without nephridia). : 7. Cryptocoela. Convoluta, Proporus.

6 to 8. Coelenteria, without anus and body-cavity. : 8. Platodes II. Platodinia (with nephridia). : 8. Rhabdocoela. Vortex, Monotus.

9 to 11. Vermalia, with anus and body-cavity. : 9. Provermalia. (Primitive Worms.) Rotatoria. : 9. Gastrotricha. Trochozoa, Trochophora.

9 to 11. Vermalia, with anus and body-cavity. : 10. Frontonia. (Rhynchelminthes.) Snout-worms. : 10. Enteropneusta. Balanoglossus, Cephalodiscus.

9 to 11. Vermalia, with anus and body-cavity. : 11. Prochordonia. Chorda-worms. : 11. Copelata. Appendicaria. Chordula-larvae.

STAGES 12 TO 15. MONORHINA ANCESTORS.

Oldest vertebrates without jaws or pairs of limbs, single nose. : 12. Acrania I. (Prospondylia.) : 12. Amphioxus larva.

Oldest vertebrates without jaws or pairs of limbs, single nose. : 13. Acrania II. More recent. : 13. Leptocardia. Amphioxus.

Oldest vertebrates without jaws or pairs of limbs, single nose. : 14. Cyclostoma I. (Archicrania.) : 14. Petromyzonta larvae.

Oldest vertebrates without jaws or pairs of limbs, single nose. : 15. Cyclostoma II. More recent. : 15. Marsipobranchia. Petromyzonta.

B. MAN'S GENEALOGICAL TREE, SECOND HALF: LATER ANCESTORS, WITH FOSSIL EVIDENCE.

COLUMN 1 : GEOLOGICAL PERIODS. COLUMN 2 : ANCESTRAL STEM-GROUPS. COLUMN 3 : LIVING RELATIVES OF ANCESTORS.

Silurian. : 16. Selachii. Primitive fishes. Proselachii. : 16. Natidanides. Chlamydoselachius. Heptanchus.

Silurian. 17. Ganoides. Plated-fishes. Proganoides. : 17. Accipenserides. (Sturgeons.) Polypterus.

Devonian. : 18. Dipneusta. Paladipneusta. : 18. Neodipneusta. Ceratodus. Protopterus.

Carboniferous. : 19. Amphibia. Stegocephala. : 19. Phanerobranchia. Salamandrina. (Proteus, triton.)

Permian. : 20. Reptilia. Proreptilia. : 20. Rhynchocephalia. Primitive lizards. Hatteria.

Triassic. : 21. Monotrema. Promammalia. : 21. Ornithodelphia. Echidna. Ornithorhyncus.

Jurassic. : 22. Marsupalia. Prodidelphia. : 22. Didelphia. Didelphys. Perameles.

Cretaceous. : 23. Mallotheria. Prochoriata. : 23. Insectivora. Erinaceida. (Ictopsida +.)

Older Eocene. : 24. Lemuravida. Older lemurs. Dentition. 3. 1. 4. 3. : 24. Pachylemures. (Hyopsodus +), (Adapis +).

Neo-Eocene. : 25. Lemurogona. Later lemurs. Dentition. 2. 1. 4. 3. : 25. Autolemures. Eulemur. Stenops.

Oligocene. : 26. Dysmopitheca. Western apes. Dentition. 2. 1. 3. 3. : 26. Platyrrhinae. (Anthropops +), (Homunculus +).

Older Miocene. : 27. Cynopitheca. Dog-faced apes (tailed). : 27. Papiomorpha. Cynocephalus.

Neo-Miocene. : 28. Anthropoides. Man-like apes (tail-less). : 28. Hylobatida. Hylobates. Satyrus.

Pliocene. : 29. Pithecanthropi. Ape-men (alali, speechless). : 29. Anthropitheca. Chimpanzee. Gorilla.

Pleistocene. : 30. Homines. Men, with speech. : 30. Weddahs. Australian negroes.

CHAPTER 2.21. OUR FISH-LIKE ANCESTORS.

Our task of detecting the extinct ancestors of our race among the vast numbers of animals known to us encounters very different difficulties in the various sections of man's stem-history. These were very great in the series of our invertebrate ancestors; they are much slighter in the subsequent series of our vertebrate ancestors. Within the vertebrate stem there is, as we have already seen, so complete an agreement in structure and embryology that it is impossible to doubt their phylogenetic unity. In this case the evidence is much clearer and more abundant.

The characteristics that distinguish the Vertebrates as a whole from the Invertebrates have already been discussed in our description of the hypothetical Primitive Vertebrate (Chapter 1.11, Figure 1.98 to 1.102). The chief of these are: (1) The evolution of the primitive brain into a dorsal medullary tube; (2) the formation of the chorda between the medullary tube and the gut; (3) the division of the gut into branchial (gill) and hepatic (liver) gut; and (4) the internal articulation or metamerism. The first three features are shared by the Vertebrates with the ascidia-larvae and the Prochordonia; the fourth is peculiar to them. Thus the chief advantage in organisation by which the earliest Vertebrates took precedence of the unsegmented Chordonia consisted in the development of internal segmentation.

The whole vertebrate stem divides first into the two chief sections of Acrania and Craniota. The Amphioxus is the only surviving representative of the older and lower section, the Acrania ("skull-less"). All the other vertebrates belong to the second division, the Craniota ("skull-animals"). The Craniota descend directly from the Acrania, and these from the primitive Chordonia. The exhaustive study that we made of the comparative anatomy and ontogeny of the Ascidia and the Amphioxus has proved these relations for us. (See Chapters 2.16 and 2.17.) The Amphioxus, the lowest Vertebrate, and the Ascidia, the nearest related Invertebrate, descend from a common extinct stem-form, the Chordaea; and this must have had, substantially, the organisation of the chordula.

However, the Amphioxus is important not merely because it fills the deep gulf between the Invertebrates and Vertebrates, but also because it shows us to-day the typical vertebrate in all its simplicity. We owe to it the most important data that we proceed on in reconstructing the gradual historical development of the whole stem. All the Craniota descend from a common stem-form, and this was substantially identical in structure with the Amphioxus. This stem-form, the Primitive Vertebrate (Prospondylus, Figures 1.98 to 1.102), had the characteristics of the vertebrate as such, but not the important features that distinguish the Craniota from the Acrania. Though the Amphioxus has many peculiarities of structure and has much degenerated, and though it cannot be regarded as an unchanged descendant of the Primitive Vertebrate, it must have inherited from it the specific characters we enumerated above. We may not say that "Amphioxus is the ancestor of the Vertebrates"; but we can say: "Amphioxus is the nearest relation to the ancestor of all the animals we know." Both belong to the same small family, or lowest class of the Vertebrates, that we call the Acrania. In our genealogical tree this group forms the twelfth stage, or the first stage among the vertebrate ancestors (Chapter 2.20). From this group of Acrania both the Amphioxus and the Craniota were evolved.

The vast division of the Craniota embraces all the Vertebrates known to us, with the exception of the Amphioxus. All of them have a head clearly differentiated from the trunk, and a skull enclosing a brain. The head has also three pairs of higher sense-organs (nose, eyes, and ears). The brain is very rudimentary at first, a mere bulbous enlargement of the fore end of the medullary tube. But it is soon divided by a number of transverse constrictions into, first three, then five successive cerebral vesicles. In this formation of the head, skull, and brain, with further development of the higher sense-organs, we have the advance that the Craniota made beyond their skull-less ancestors. Other organs also attained a higher development; they acquired a compact centralised heart with valves and a more advanced liver and kidneys, and made progress in other important respects.

We may divide the Craniota generally into Cyclostoma ("round-mouthed") and Gnathostoma ("jaw-mouthed"). There are only a few groups of the former in existence now, but they are very interesting, because in their whole structure they stand midway between the Acrania and the Gnathostoma. They are much more advanced than the Acrania, much less so than the fishes, and thus form a very welcome connecting-link between the two groups. We may therefore consider them a special intermediate group, the fourteenth and fifteenth stages in the series of our ancestors.

(FIGURE 2.247. The large marine lamprey (Petromyzon marinus), much reduced. Behind the eye there is a row of seven gill-clefts visible on the left, in front the round suctorial mouth.)

The few surviving species of the Cyclostoma are divided into two orders—the Myxinoides and the Petromyzontes. The former, the hag-fishes, have a long, cylindrical, worm-like body. They were classed by Linne with the worms, and by later zoologists, with the fishes, or the amphibia, or the molluscs. They live in the sea, usually as parasites of fishes, into the skin of which they bore with their round suctorial mouths and their tongues, armed with horny teeth. They are sometimes found alive in the body cavity of fishes (such as the torsk or sturgeon); in these cases they have passed through the skin into the interior. The second order consists of the Petromyzontes or lampreys; the small river lamprey (Petromyzon fluviatilis) and the large marine lamprey (Petromyzon marinus, Figure 2.247). They also have a round suctorial mouth, with horny teeth inside it; by means of this they attach themselves by sucking to fishes, stones, and other objects (hence the name Petromyzon = stone-sucker). It seems that this habit was very widespread among the earlier Vertebrates; the larvae of many of the Ganoids and frogs have suctorial disks near the mouth.

The class that is formed of the Myxinoides and Petromyzontes is called the Cyclostoma (round-mouthed), because their mouth has a circular or semi-circular aperture. The jaws (upper and lower) that we find in all the higher Vertebrates are completely wanting in the Cyclostoma, as in the Amphioxus. Hence the other Vertebrates are collectively opposed to them as Gnathostoma (jaw-mouthed). The Cyclostoma might also be called Monorhina (single-nosed), because they have only a single nasal passage, while all the Gnathostoma have two nostrils (Amphirhina = double-nosed). But apart from these peculiarities the Cyclostoma differ more widely from the fishes in other special features of their structure than the fishes do from man. Hence they are obviously the last survivors of a very ancient class of Vertebrates, that was far from attaining the advanced organisation of the true fish. To mention only the chief points, the Cyclostoma show no trace of pairs of limbs. Their mucous skin is quite naked and smooth and devoid of scales. There is no bony skeleton. A very rudimentary skull is developed at the foremost end of their chorda. At this point a soft membranous (partly turning into cartilage), small skull-capsule is formed, and encloses the brain.

The brain of the Cyclostoma is merely a very small and comparatively insignificant swelling of the spinal marrow, a simple vesicle at first. It afterwards divides into five successive cerebral vesicles, like the brain of the Gnathostoma. These five primitive cerebral vesicles, that are found in the embryos of all the higher vertebrates from the fishes to man, and grow into very complex structures, remain at a very rudimentary stage in the Cyclostoma. The histological structure of the nerves is also less advanced than in the rest of the vertebrates. In these the auscultory organ always contains three circular canals, but in the lampreys there are only two, and in the hag-fishes only one. In most other respects the organisation of the Cyclostoma is much simpler—for instance, in the structure of the heart, circulation, and kidneys. We must especially note the absence of a very important organ that we find in the fishes, the floating-bladder, from which the lungs of the higher Vertebrates have been developed.

When we consider all these peculiarities in the structure of the Cyclostoma, we may formulate the following thesis: Two divergent lines proceeded from the earliest Craniota, or the primitive Craniota (Archicrania). One of these lines is preserved in a greatly modified condition: these are the Cyclostoma, a very backward and partly degenerate side-line. The other, the chief line of the Vertebrate stem, advanced straight to the fishes, and by fresh adaptations acquired a number of important improvements.

(FIGURE 2.248. Fossil Permian primitive fish (Pleuracanthus Dechenii), from the red sandstone of Saarbrucken. (From Doderlein.) I Skull and branchial skeleton: o eye-region, pq palatoquadratum, nd lower jaw, hm hyomandibular, hy tongue-bone, k gill-radii, kb gill-arches, z jaw-teeth, sz gullet-teeth, st neck-spine. II Vertebral column: ob upper arches, ub lower arches, hc intercentra, r ribs. III Single fins: d dorsal fin, c tail-fin (tail-end wanting), an anus-fin, ft supporter of fin-rays. IV Breast-fin: sg shoulder-zone, ax fin-axis, ss double lines of fin-rays, bs additional rays, sch plates. V Ventral fin: p pelvis, ax fin-axis, ss single row of fin-rays, bs additional rays, sch scales, cop penis.

FIGURE 2.249. Embryo of a shark (Scymnus lichia), seen from the ventral side, v breast-fins (in front five pairs of gill-clefts), h belly-fins, a anus, s tail-fin, k external gill-tuft, d yelk-sac (removed for most part), g eye, n nose, m mouth-cleft.)

The Cyclostoma are almost always classified by zoologists among the fishes; but the incorrectness of this may be judged from the fact that in all the chief and distinctive features of organisation they are further removed from the fishes than the fishes are from the Mammals, and even man. With the fishes we enter upon the vast division of the jaw-mouthed or double-nosed Vertebrates (Gnathostoma or Amphirhina). We have to consider the fishes carefully as the class which, on the evidence of palaeontology, comparative anatomy, and ontogeny, may be regarded with absolute certainty as the stem-class of all the higher Vertebrates or Gnathostomes. Naturally, none of the actual fishes can be considered the direct ancestor of the higher Vertebrates. But it is certain that all the Vertebrates or Gnathostomes, from the fishes to man, descend from a common, extinct, fish-like ancestor. If we had this ancient stem-form before us, we would undoubtedly class it as a true fish. Fortunately the comparative anatomy and classification of the fishes are now so far advanced that we can get a very clear idea of these interesting and instructive features.

In order to understand properly the genealogical tree of our race within the vertebrate stem, it is important to bear in mind the characteristics that separate the whole of the Gnathostomes from the Cyclostomes and Craniota. In these respects the fishes agree entirely with all the other Gnathostomes up to man, and it is on this that we base our claim of relationship to the fishes. The following characteristics of the Gnathostomes are anatomic features of this kind: (1) The internal gill-arch apparatus with the jaw arches; (2) the pair of nostrils; (3) the floating bladder or lungs; and (4) the two pairs of limbs.

The peculiar formation of the frame work of the branchial (gill) arches and the connected maxillary (jaw) apparatus is of importance in the whole group of the Gnathostomes. It is inherited in rudimentary form by all of them, from the earliest fishes to man. It is true that the primitive transformation (which we find even in the Ascidia) of the fore gut into the branchial gut can be traced in all the Vertebrates to the same simple type; in this respect the gill-clefts, which pierce the walls of the branchial gut in all the Vertebrates and in the Ascidia, are very characteristic. But the EXTERNAL, superficial branchial skeleton that supports the gill-crate in the Cyclostoma is replaced in the Gnathostomes by an INTERNAL branchial skeleton. It consists of a number of successive cartilaginous arches, which lie in the wall of the gullet between the gill-clefts, and run round the gullet from both sides. The foremost pair of gill-arches become the maxillary arches, from which we get our upper and lower jaws.

The olfactory organs are at first found in the same form in all the Gnathostomes, as a pair of depressions in the fore part of the skin of the head, above the mouth; hence, they are also called the Amphirhina ("double-nosed"). The Cyclostoma are "one-nosed" (Monorhina); their nose is a single passage in the middle of the frontal surface. But as the olfactory nerve is double in both cases, it is possible that the peculiar form of the nose in the actual Cyclostomes is a secondary acquisition (by adaptation to suctorial habits).

A third essential character of the Gnathostomes, that distinguishes them very conspicuously from the lower vertebrates we have dealt with, is the formation of a blind sac by invagination from the fore part of the gut, which becomes in the fishes the air-filled floating-bladder. This organ acts as a hydrostatic apparatus, increasing or reducing the specific gravity of the fish by compressing or altering the quantity of air in it. The fish can rise or sink in the water by means of it. This is the organ from which the lungs of the higher vertebrates are developed.

(FIGURE 2.250. Fully developed man-eating shark (Carcharias melanopterus), left view. r1 first, r2 second dorsal fin, s tail-fin, a anus-fin, v breast-fins, h belly-fins.)

Finally, the fourth character of the Gnathostomes in their simple embryonic form is the two pairs of extremities or limbs—a pair of fore legs (breast-fins in the fish, Figure 2.250 v) and a pair of hind legs (ventral fins in the fish, Figure 2.250 h). The comparative anatomy of these fins is very interesting, because they contain the rudiments of all the skeletal parts that form the framework of the fore and hind legs in all the higher vertebrates right up to man. There is no trace of these pairs of limbs in the Acrania and Cyclostomes.

Turning, now, to a closer inspection of the fish class, we may first divide it into three groups or sub-classes, the genealogy of which is well known to us. The first and oldest group is the sub-class of the Selachii or primitive fishes; the best-known representatives of which to-day are the orders of the sharks and rays (Figures 2.248 to 2.252). Next to this is the more advanced sub-class of the plated fishes or Ganoids (Figures 2.253 to 2.255). It has been long extinct for the most part, and has very few living representatives, such as the sturgeon and the bony pike; but we can form some idea of the earlier extent of this interesting group from the large numbers of fossils. From these plated fishes the sub-class of the bony fishes or Teleostei was developed, to which the great majority of living fishes belong (especially nearly all our river fishes). Comparative anatomy and ontogeny show clearly that the Ganoids descended from the Selachii, and the Teleostei from the Ganoids. On the other hand, a collateral line, or rather the advancing chief line of the vertebrate stem, was developed from the earlier Ganoids, and this leads us through the group of the Dipneusta to the important division of the Amphibia.

(FIGURE 2.251. Fossil angel-shark (Squatina alifera), from the upper Jurassic at Eichstatt. (From Zittel.) The cartilaginous skull is clearly seen in the broad head, and the gill-arches behind. The wide breast-fin and the narrower belly-fin have a number of radii; between these and the vertebral column are a number of ribs.)

The earliest fossil remains of Vertebrates that we know were found in the Upper Silurian (Chapter 2.18), and belong to two groups—the Selachii and the Ganoids. The most primitive of all known representatives of the earliest fishes are probably the remarkable Pleuracanthida, the genera Pleuracanthus, Xenacanthus, Orthocanthus, etc. (Figure 2.248). These ancient cartilaginous fishes agree in most points of structure with the real sharks (Figures 2.249 and 2.250); but in other respects they seem to be so much simpler in organisation that many palaeontologists separate them altogether, and regard them as Proselachii; they are probably closely related to the extinct ancestors of the Gnathostomes. We find well-preserved remains of them in the Permian period. Well-preserved impressions of other sharks are found in the Jurassic schist, such as of the angel-fish (Squatina, Figure 2.251). Among the extinct earlier sharks of the Tertiary period there were some twice as large as the biggest living fishes; Carcharodon was more than 100 feet long. The sole surviving species of this genus (C. Rondeleti) is eleven yards long, and has teeth two inches long; but among the fossil species we find teeth six inches long (Figure 2.252).

From the primitive fishes or Selachii, the earliest Gnathostomes, was developed the legion of the Ganoids. There are very few genera now of this interesting and varied group—the ancient sturgeons (Accipenser), the eggs of which are eaten as caviare, and the stratified pikes (Polypterus, Figure 2.255) in African rivers, and bony pikes (Lepidosteus) in the rivers of North America. On the other hand, we have a great variety of specimens of this group in the fossil state, from the Upper Silurian onward. Some of these fossil Ganoids approach closely to the Selachii; others are nearer to the Dipneusts; others again represent a transition to the Teleostei. For our genealogical purposes the most interesting are the intermediate forms between the Selachii and the Dipneusts. Huxley, to whom we owe particularly important works on the fossil Ganoids, classed them in the order of the Crossopterygii. Many genera and species of this order are found in the Devonian and Carboniferous strata (Figure 2.253); a single, greatly modified survivor of the group is still found in the large rivers of Africa (Polypterus, Figure 2.255, and the closely related Calamichthys). In many impressions of the Crossopterygii the floating bladder seems to be ossified, and therefore well preserved—for instance, in the Undina (Figure 2.254, immediately behind the head).

Part of these Crossopterygii approach very closely in their chief anatomic features to the Dipneusts, and thus represent phylogenetically the transition from the Devonian Ganoids to the earliest air-breathing vertebrates. This important advance was made in the Devonian period. The numerous fossils that we have from the first two geological sections, the Laurentian and Cambrian periods, belong exclusively to aquatic plants and animals. From this paleontological fact, in conjunction with important geological and biological indications, we may infer with some confidence that there were no terrestrial animals at that time. During the whole of the vast archeozoic period—many millions of years—the living population of our planet consisted almost exclusively of aquatic organisms; this is a very remarkable fact, when we remember that this period embraces the larger half of the whole history of life. The lower animal-stems are wholly (or with very few exceptions) aquatic. But the higher stems also remained in the water during the primordial epoch. It was only towards its close that some of them came to live on land. We find isolated fossil remains of terrestrial animals first in the Upper Silurian, and in larger numbers in the Devonian strata, which were deposited at the beginning of the second chief section of geology (the paleozoic age). The number increases considerably in the Carboniferous and Permian deposits. We find many species both of the articulate and the vertebrate stem that lived on land and breathed the atmosphere; their aquatic ancestors of the Silurian period only breathed water. This important change in respiration is the chief modification that the animal organism underwent in passing from the water to the solid land. The first consequence was the formation of lungs for breathing air; up to that time the gills alone had served for respiration. But there was at the same time a great change in the circulation and its organs; these are always very closely correlated to the respiratory organs. Moreover, the limbs and other organs were also more or less modified, either in consequence of remote correlation to the preceding or owing to new adaptations.

(FIGURE 2.252. Tooth of a gigantic shark (Carcharodon megalodon), from the Pliocene at Malta. Half natural size. (From Zittel.))

In the vertebrate stem it was unquestionably a branch of the fishes—in fact, of the Ganoids—that made the first fortunate experiment during the Devonian period of adapting themselves to terrestrial life and breathing the atmosphere. This led to a modification of the heart and the nose. The true fishes have merely a pair of blind olfactory pits on the surface of the head; but a connection of these with the cavity of the mouth was now formed. A canal made its appearance on each side, and led directly from the nasal depression into the mouth-cavity, thus conveying atmospheric air to the lungs even when the mouth was closed. Further, in all true fishes the heart has only two sections—an atrium that receives the venous blood from the veins, and a ventricle that propels it through a conical artery to the gills; the atrium was now divided into two halves, or right and left auricles, by an incomplete partition. The right auricle alone now received the venous blood from the body, while the left auricle received the venous blood that flowed from the lungs and gills to the heart. Thus the double circulation of the higher vertebrates was evolved from the simple circulation of the true fishes, and, in accordance with the laws of correlation, this advance led to others in the structure of other organs.

(FIGURE 2.253. A Devonian Crossopterygius (Holoptychius nobilissimus, from the Scotch old red sandstone. (From Huxley.)

FIGURE 2.254. A Jurassic Crossopterygius (Undina penicillata), from the upper Jurassic at Eichstatt. (From Zittel.) j jugular plates, b three ribbed scales.

FIGURE 2.255. A living Crossopterygius, from the Upper Nile (Polypterus bichir).

FIGURE 2.256. Fossil Dipneust (Dipterus Valenciennesi), from the old red sandstone (Devon). (From Pander.)

FIGURE 2.257. The Australian Dipneust (Ceratodus Forsteri). B view from the right, A lower side of the skull, C lower jaw. (From Gunther.) Qu quadrate bone, Psph parasphenoid, PtP pterygopalatinum, Vo vomer, d teeth, na nostrils, Br branchial cavity, C first rib. D lower-jaw teeth of the fossil Ceratodus Kaupi (from the Triassic).)

The vertebrate class, that thus adapted itself to breathing the atmosphere, and was developed from a branch of the Ganoids, takes the name of the Dipneusts or Dipnoa ("double-breathers"), because they retained the earlier gill-respiration along with the new pulmonary (lung) respiration, like the lowest amphibia. This class was represented during the paleozoic age (or the Devonian, Carboniferous, and Permian periods) by a number of different genera. There are only three genera of the class living to-day: Protopterus annectens in the rivers of tropical Africa (the White Nile, the Niger, Quelliman, etc.), Lepidosiren paradoxa in tropical South America (in the tributaries of the Amazon), and Ceratodus Forsteri in the rivers of East Australia. This wide distribution of the three isolated survivors proves that they represent a group that was formerly very large. In their whole structure they form a transition from the fishes to the amphibia. The transitional formation between the two classes is so pronounced in the whole organisation of these remarkable animals that zoologists had a lively controversy over the question whether they were really fishes or amphibia. Several distinguished zoologists classed them with the amphibia, though most now associate them with the fishes. As a matter of fact, the characters of the two classes are so far united in the Dipneusts that the answer to the question depends entirely on the definition we give of "fish" and "amphibian." In habits they are true amphibia. During the tropical winter, in the rainy season, they swim in the water like the fishes, and breathe water by gills. During the dry season they bury themselves in the dry mud, and breathe the atmosphere through lungs, like the amphibia and the higher vertebrates. In this double respiration they resemble the lower amphibia, and have the same characteristic formation of the heart; in this they are much superior to the fishes. But in most other features they approach nearer to the fishes, and are inferior to the amphibia. Externally they are entirely fish-like.

(FIGURE 2.258. Young ceratodus, shortly after issuing from the egg, magnified ten times. k gill-cover, l liver. (From Richard Semon.)

FIGURE 2.259. Young ceratodus six weeks after issuing from the egg. s spiral fold of gut, b rudimentary belly-fin. (From Richard Semon.))

In the Dipneusts the head is not marked off from the trunk. The skin is covered with large scales. The skeleton is soft, cartilaginous, and at a low stage of development, as in the lower Selachii and the earliest Ganoids. The chorda is completely retained, and surrounded by an unsegmented sheath. The two pairs of limbs are very simple fins of a primitive type, like those of the lowest Selachii. The formation of the brain, the gut, and the sexual organs is also the same as in the Selachii. Thus the Dipneusts have preserved by heredity many of the less advanced features of our primitive fish-like ancestors, and at the same time have made a great step forward in adaptation to air-breathing by means of lungs and the correlative improvement of the heart.

Ceratodus is particularly interesting on account of the primitive build of its skeleton; the cartilaginous skeleton of its two pairs of fins, for instance, has still the original form of a bi-serial or feathered leaf, and was on that account described by Gegenbaur as a "primitive fin-skeleton." On the other hand, the skeleton of the pairs of fins is greatly reduced in the African dipneust (Protopterus) and the American (Lepidosiren). Further, the lungs are double in these modern dipneusts, as in all the other air-breathing vertebrates; they have on that account been called "double-lunged" (Dipneumones) in contrast to the Ceratodus; the latter has only a single lung (Monopneumones). At the same time the gills also are developed as water-breathing organs in all these lung-fishes. Protopterus has external as well as internal gills.

The paleozoic Dipneusts that are in the direct line of our ancestry, and form the connecting-bridge between the Ganoids and the Amphibia, differ in many respects from their living descendants, but agree with them in the above essential features. This is confirmed by a number of interesting facts that have lately come to our knowledge in connection with the embryonic development of the Ceratodus and Lepidosiren; they give us important information as to the stem-history of the lower Vertebrates, and therefore of our early ancestors of the paleozoic age.

CHAPTER 2.22. OUR FIVE-TOED ANCESTORS.

With the phylogenetic study of the four higher classes of Vertebrates, which must now engage our attention, we reach much firmer ground and more light in the construction of our genealogy than we have, perhaps, enjoyed up to the present. In the first place, we owe a number of very valuable data to the very interesting class of Vertebrates that come next to the Dipneusts and have been developed from them—the Amphibia. To this group belong the salamander, the frog, and the toad. In earlier days all the reptiles were, on the example of Linne, classed with the Amphibia (lizards, serpents, crocodiles, and tortoises). But the reptiles are much more advanced than the Amphibia, and are nearer to the birds in the chief points of their structure. The true Amphibia are nearer to the Dipneusta and the fishes; they are also much older than the reptiles. There were plenty of highly-developed (and sometimes large) Amphibia during the Carboniferous period; but the earliest reptiles are only found in the Permian period. It is probable that the Amphibia were evolved even earlier—during the Devonian period—from the Dipneusta. The extinct Amphibia of which we have fossil remains from that remote period (very numerous especially in the Triassic strata) were distinguished for a graceful scaly coat or a powerful bony armour on the skin (like the crocodile), whereas the living amphibia have usually a smooth and slippery skin.

The earliest of these armoured Amphibia (Phractamphibia) form the order of Stegocephala ("roof-headed") (Figure 2.260). It is among these, and not among the actual Amphibia, that we must look for the forms that are directly related to the genealogy of our race, and are the ancestors of the three higher classes of Vertebrates. But even the existing Amphibia have such important relations to us in their anatomic structure, and especially their embryonic development, that we may say: Between the Dipneusts and the Amniotes there was a series of extinct intermediate forms which we should certainly class with the Amphibia if we had them before us. In their whole organisation even the actual Amphibia seem to be an instructive transitional group. In the important respects of respiration and circulation they approach very closely to the Dipneusta, though in other respects they are far superior to them.

This is particularly true of the development of their limbs or extremities. In them we find these for the first time as five-toed feet. The thorough investigations of Gegenbaur have shown that the fish's fins, of which very erroneous opinions were formerly held, are many-toed feet. The various cartilaginous or bony radii that are found in large numbers in each fin correspond to the fingers or toes of the higher Vertebrates. The several joints of each fin-radius correspond to the various parts of the toe. Even in the Dipneusta the fin is of the same construction as in the fishes; it was afterwards gradually evolved into the five-toed form, which we first encounter in the Amphibia. This reduction of the number of the toes to six, and then to five, probably took place in the second half of the Devonian period—at the latest, in the subsequent Carboniferous period—in those Dipneusta which we regard as the ancestors of the Amphibia. We have several fossil remains of five-toed Amphibia from this period. There are numbers of fossil impressions of them in the Triassic of Thuringia (Chirotherium).

(FIGURE 2.260. Fossil amphibian from the Permian, found in the Plauen terrain near Dresden (Branchiosaurus amblystomus). (From Credner.) A skeleton of a young larva. B larva, restored, with gills. C the adult form, natural size.)

The fact that the toes number five is of great importance, because they have clearly been transmitted from the Amphibia to all the higher Vertebrates. Man entirely resembles his amphibian ancestors in this respect, and indeed in the whole structure of the bony skeleton of his five-toed extremities. A careful comparison of the skeleton of the frog with our own is enough to show this. It is well known that this hereditary number of the toes has assumed a very great practical importance from remote times; on it our whole system of enumeration (the decimal system applied to measurement of time, mass, weight, etc.) is based. There is absolutely no reason why there should be five toes in the fore and hind feet in the lowest Amphibia, the reptiles, and the higher Vertebrates, unless we ascribe it to inheritance from a common stem-form. Heredity alone can explain it. It is true that we find less than five toes in many of the Amphibia and of the higher Vertebrates. But in all these cases we can prove that some of the toes atrophied, and were in time lost altogether.

The causes of this evolution of the five-toed foot from the many-toed fin in the amphibian ancestor must be sought in adaptation to the entire change of function that the limbs experienced in passing from an exclusively aquatic to a partly terrestrial life. The many-toed fin had been used almost solely for motion in the water; it had now also to support the body in creeping on the solid ground. This led to a modification both of the skeleton and the muscles of the limbs. The number of the fin-radii was gradually reduced, and sank finally to five. But these five remaining radii became much stronger. The soft cartilaginous radii became bony rods. The rest of the skeleton was similarly strengthened. Thus from the one-armed lever of the many-toed fish-fin arose the improved many-armed lever system of the five-toed amphibian limbs. The movements of the body gained in variety as well as in strength. The various parts of the skeletal system and correlated muscular system began to differentiate more and more. In view of the close correlation of the muscular and nervous systems, this also made great advance in structure and function. Hence we find, as a matter of fact, that the brain is much more developed in the higher Amphibia than in the fishes, the Dipneusta, and the lower Amphibia.

The first advance in organisation that was occasioned by the adoption of life on land was naturally the construction of an organ for breathing air—a lung. This was formed directly from the floating-bladder inherited from the fishes. At first its function was insignificant beside that of the gills, the older organ for water-respiration. Hence we find in the lowest Amphibia, the gilled Amphibia, that, like the Dipneusta, they pass the greater part of their life in the water, and breathe water through gills. They only come to the surface at brief intervals, or creep on to the land, and then breathe air by their lungs. But some of the tailed Amphibia—the salamanders—remain entirely in the water when they are young, and afterwards spend most of their time on land. In the adult state they only breathe air through lungs. The same applies to the most advanced of the Amphibia, the Batrachia (frogs and toads); some of them have entirely lost the gill-bearing larva form.* (* The tree-frog of Martinique (Hylades martinicensis) loses the gills on the seventh, and the tail and yelk-sac on the eighth, day of foetal life. On the ninth or tenth day after fecundation the frog emerges from the egg.) This is also the case with certain small, serpentine Amphibia, the Caecilia (which live in the ground like earth-worms).

(FIGURE 2.261. Larva of the Spotted Salamander (Salamandra maculata), seen from the ventral side. In the centre a yelk-sac still hangs from the gut. The external gills are gracefully ramified. The two pairs of legs are still very small.)

The great interest of the natural history of the Amphibia consists especially in their intermediate position between the lower and higher Vertebrates. The lower Amphibia approach very closely to the Dipneusta in their whole organisation, live mainly in the water, and breathe by gills; but the higher Amphibia are just as close to the Amniotes, live mainly on land, and breathe by lungs. But in their younger state the latter resemble the former, and only reach the higher stage by a complete metamorphosis. The embryonic development of most of the higher Amphibia still faithfully reproduces the stem-history of the whole class, and the various stages of the advance that was made by the lower Vertebrates in passing from aquatic to terrestrial life during the Devonian or the Carboniferous period are repeated in the spring by every frog that develops from an egg in our ponds.

(FIGURE 2.262. Larva of the common grass-frog (Rana temporaria), or "tadpole." m mouth, n a pair of suckers for fastening on to stones, d skin-fold from which the gill-cover develops; behind it the gill-clefts, from which the branching gills (k) protrude, s tail-muscles, f cutaneous fin-fringe of the tail.)

The common frog leaves the egg in the shape of a larva, like the tailed salamander (Figure 2.261), and this is altogether different from the mature frog (Figure 2.262). The short trunk ends in a long tail, with the form and structure of a fish's tail (s). There are no limbs at first. The respiration is exclusively branchial, first through external (k) and then internal gills. In harmony with this the heart has the same structure as in the fish, and consists of two sections—an atrium that receives the venous blood from the body, and a ventricle that forces it through the arteries into the gills.

We find the larvae of the frog (or tadpoles, Gyrini) in great numbers in our ponds every spring in this fish-form, using their muscular tails in swimming, just like the fishes and young Ascidia. When they have reached a certain size, the remarkable metamorphosis from the fish-form to the frog begins. A blind sac grows out of the gullet, and expands into a couple of spacious sacs: these are the lungs. The simple chamber of the heart is divided into two sections by the development of a partition, and there are at the same time considerable changes in the structure of the chief arteries. Previously all the blood went from the auricle through the aortic arches into the gills, but now only part of it goes to the gills, the other part passing to the lungs through the new-formed pulmonary artery. From this point arterial blood returns to the left auricle of the heart, while the venous blood gathers in the right auricle. As both auricles open into a single ventricle, this contains mixed blood. The dipneust form has now succeeded to the fish-form. In the further course of the metamorphosis the gills and the branchial vessels entirely disappear, and the respiration becomes exclusively pulmonary. Later, the long swimming tail is lost, and the frog now hops to the land with the legs that have grown meantime.

This remarkable metamorphosis of the Amphibia is very instructive in connection with our human genealogy, and is particularly interesting from the fact that the various groups of actual Amphibia have remained at different stages of their stem-history, in harmony with the biogenetic law. We have first of all a very low order of Amphibia—the Sozobranchia ("gilled-amphibia"), which retain their gills throughout life, like the fishes. In a second order of the salamanders the gills are lost in the metamorphosis, and when fully grown they have only pulmonary respiration. Some of the tailed Amphibia still retain the gill-clefts in the side of the neck, though they have lost the gills themselves (Menopoma). If we force the larvae of our salamanders (Figure 2.261) and tritons to remain in the water, and prevent them from reaching the land, we can in favourable circumstances make them retain their gills. In this fish-like condition they reach sexual maturity, and remain throughout life at the lower stage of the gilled Amphibia.

(FIGURE 2.263. Fossil mailed amphibian, from the Bohemian Carboniferous (Seeleya). (From Fritsch.) The scaly coat is retained on the left.)

We have the reverse of this experiment in a Mexican gilled salamander, the fish-like axolotl (Siredon pisciformis). It was formerly regarded as a permanent gilled amphibian persisting throughout life at the fish-stage. But some of the hundreds of these animals that are kept in the Botanical Garden at Paris got on to the land for some reason or other, lost their gills, and changed into a form closely resembling the salamander (Amblystoma). Other species of the genus became sexually mature for the first time in this condition. This has been regarded as an astounding phenomenon, although every common frog and salamander repeats the metamorphosis in the spring. The whole change from the aquatic and gill-breathing animal to the terrestrial lung-breathing form may be followed step by step in this case. But what we see here in the development of the individual has happened to the whole class in the course of its stem-history.

The metamorphosis goes farther in a third order of Amphibia, the Batrachia or Anura, than in the salamander. To this belong the various kinds of toads, ringed snakes, water-frogs, tree-frogs, etc. These lose, not only the gills, but also (sooner or later) the tail, during metamorphosis.

The ontogenetic loss of the gills and the tail in the frog and toad can only be explained on the assumption that they are descended from long-tailed Amphibia of the salamander type. This is also clear from the comparative anatomy of the two groups. This remarkable metamorphosis is, however, also interesting because it throws a certain light on the phylogeny of the tail-less apes and man. Their ancestors also had long tails and gills like the gilled Amphibia, as the tail and the gill-arches of the human embryo clearly show.

For comparative anatomical and ontogenetic reasons, we must not seek these amphibian ancestors of ours—as one would be inclined to do, perhaps—among the tail-less Batrachia, but among the tailed lower Amphibia.

The vertebrate form that comes next to the Amphibia in the series of our ancestors is a lizard-like animal, the earlier existence of which can be confidently deduced from the facts of comparative anatomy and ontogeny. The living Hatteria of New Zealand (Figure 2.264) and the extinct Rhyncocephala of the Permian period (Figure 2.265) are closely related to this important stem-form; we may call them the Protamniotes, or Primitive Amniotes. All the Vertebrates above the Amphibia—or the three classes of reptiles, birds, and mammals—differ so much in their whole organisation from all the lower Vertebrates we have yet considered, and have so great a resemblance to each other, that we put them all together in a single group with the title of Amniotes. In these three classes alone we find the remarkable embryonic membrane, already mentioned, which we called the amnion; a cenogenetic adaptation that we may regard as a result of the sinking of the growing embryo into the yelk-sac.

All the Amniotes known to us—all reptiles, birds, and mammals (including man)—agree in so many important points of internal structure and development that their descent from a common ancestor can be affirmed with tolerable certainty. If the evidence of comparative anatomy and ontogeny is ever entirely beyond suspicion, it is certainly the case here. All the peculiarities that accompany and follow the formation of the amnion, and that we have learned in our consideration of human embryology; all the peculiarities in the development of the organs which we will presently follow in detail; finally, all the principal special features of the internal structure of the full-grown Amniotes—prove so clearly the common origin of all the Amniotes from single extinct stem-form that it is difficult to entertain the idea of their evolution from several independent stems. This unknown common stem-form is our primitive Amniote (Protamnion). In outward appearance it was probably something between the salamander and the lizard.

It is very probable that some part of the Permian period was the age of the origin of the Protamniotes. This follows from the fact that the Amphibia are not fully developed until the Carboniferous period, and that the first fossil reptiles (Palaehatteria, Homoeosaurus, Proterosaurus) are found towards the close of the Permian period. Among the important changes of the vertebrate organisation that marked the rise of the first Amniotes from salamandrine Amphibia during this period the following three are especially noteworthy: the entire disappearance of the water-breathing gills and the conversion of the gill-arches into other organs, the formation of the allantois or primitive urinary sac, and the development of the amnion.

One of the most salient characteristics of the Amniotes is the complete loss of the gills. All Amniotes, even if living in water (such as sea-serpents and whales), breathe air through lungs, never water through gills. All the Amphibia (with very rare exceptions) retain their gills for some time when young, and have for a time (if not permanently) branchial respiration; but after these there is no question of branchial respiration. The Protamniote itself must have entirely abandoned water-breathing. Nevertheless, the gill-arches are preserved by heredity, and develop into totally different (in part rudimentary) organs—various parts of the bone of the tongue, the frame of the jaws, the organ of hearing, etc. But we do not find in the embryos of the Amniotes any trace of gill-leaves, or of real respiratory organs on the gill-arches.

With this complete abandonment of the gills is probably connected the formation of another organ, to which we have already referred in embryology—namely, the allantois or primitive urinary sac (cf. Chapter 1.15). It is very probable that the urinary bladder of the Dipneusts is the first structure of the allantois. We find in these a urinary bladder that proceeds from the lower wall of the hind end of the gut, and serves as receptacle for the renal secretions. This organ has been transmitted to the Amphibia, as we can see in the frog.

The formation of the amnion and the allantois and the complete disappearance of the gills are the chief characteristics that distinguish the Amniotes from the lower Vertebrates we have hitherto considered. To these we may add several subordinate features that are transmitted to all the Amniotes, and are found in these only. One striking embryonic character of the Amniotes is the great curve of the head and neck in the embryo. We also find an advance in the structure of several of the internal organs of the Amniotes which raises them above the highest of the anamnia. In particular, a partition is formed in the simple ventricle of the heart, dividing into right and left chambers. In connection with the complete metamorphosis of the gill-arches we find a further development of the auscultory organs. Also, there is a great advance in the structure of the brain, skeleton, muscular system, and other parts. Finally, one of the most important changes is the reconstruction of the kidneys. In all the earlier Vertebrates we have found the primitive kidneys as excretory organs, and these appear at an early stage in the embryos of all the higher Vertebrates up to man. But in the Amniotes these primitive kidneys cease to act at an early stage of embryonic life, and their function is taken up by the permanent or secondary kidneys, which develop from the terminal section of the prorenal ducts.

(FIGURE 2.264. The lizard (Hatteria punctata = Sphenodon punctatus) of New Zealand. The sole surviving proreptile. (From Brehm.))

Taking all these peculiarities of the Amniotes together, it is impossible to doubt that all the animals of this group—all reptiles, birds, and mammals—have a common origin, and form a single blood-related stem. Our own race belongs to this stem. Man is, in every feature of his organisation and embryonic development, a true Amniote, and has descended from the Protamniote with all the other Amniotes. Though they appeared at the end (possibly even in the middle) of the Paleozoic age, the Amniotes only reached their full development during the Mesozoic age. The birds and mammals made their first appearance during this period. Even the reptiles show their greatest growth at this time, so that it is called "the reptile age." The extinct Protamniote, the ancestor of the whole group, belongs in its whole organisation to the reptile class.

The genealogical tree of the amniote group is clearly indicated in its chief lines by their paleontology, comparative anatomy, and ontogeny. The group succeeding the Protamniote divided into two branches. The branch that will claim our whole interest is the class of the Mammals. The other branch, which developed in a totally different direction, and only comes in contact with the Mammals at its root, is the combined group of the reptiles and birds; these two classes may, with Huxley, be conveniently grouped together as the Sauropsida. Their common stem-form is an extinct lizard-like reptile of the order of the Rhyncocephalia. From this have been developed in various directions the serpents, crocodiles, tortoises, etc.—in a word, all the members of the reptile class. But the remarkable class of the birds has also been evolved directly from a branch of the reptile group, as is now established beyond question. The embryos of the reptiles and birds are identical until a very late stage, and have an astonishing resemblance even later. Their whole structure agrees so much that no anatomist now questions the descent of the birds from the reptiles. On the other hand, the mammal line has descended from the group of the Sauromammalia, a different branch of the Proreptilia. It is connected at its deepest roots with the reptile line, but it then diverges completely from it and follows a distinctive development. Man is the highest outcome of this class, the "crown of creation." The hypothesis that the three higher Vertebrate classes represent a single Amniote-stem, and that the common root of this stem is to be found in the amphibian class, is now generally admitted.

(FIGURE 2.265. Homoeosaurus pulchellus, a Jurassic proreptile from Kehlheim. (From Zittel.))

The instructive group of the Permian Tocosauria, the common root from which the divergent stems of the Sauropsids and mammals have issued, merits our particular attention as the stem-group of all the Amniotes. Fortunately a living representative of this extinct ancestral group has been preserved to our day; this is the remarkable lizard of New Zealand, Hatteria punctata (Figure 2.264). Externally it differs little from the ordinary lizard; but in many important points of internal structure, especially in the primitive construction of the vertebral column, the skull, and the limbs, it occupies a much lower position, and approaches its amphibian ancestors, the Stegocephala. Hence Hatteria is the phylogenetically oldest of all living reptiles, an isolated survivor from the Permian period, closely resembling the common ancestor of the Amniotes. It must differ so little from this extinct form, our hypothetical Protamniote, that we put it next to the Proreptilia. The remarkable Permian Palaehatteria, that Credner discovered in the Plauen terrain at Dresden in 1888, belongs to the same group (Figure 2.266). The Jurassic genus Homoeosaurus (Figure 2.265), of which well-preserved skeletons are found in the Solenhofen schists, is perhaps still more closely related to them.

Unfortunately, the numerous fossil remains of Permian and Triassic Tocosauria that we have found in the last two decades are, for the most part, very imperfectly preserved. Very often we can make only precarious inferences from these skeletal fragments as to the anatomic characters of the soft parts that went with the bony skeleton of the extinct Tocosauria. Hence it has not yet been possible to arrange these important fossils with any confidence in the ancestral series that descend from the Protamniotes to the Sauropsids on the one side and the Mammals on the other. Opinions are particularly divided as to the place in classification and the phylogenetic significance of the remarkable Theromorpha. Cope gives this name to a very interesting and extensive group of extinct terrestrial reptiles, of which we have only fossil remains from the Permian and Triassic strata. Forty years ago some of these Therosauria (fresh-water animals) were described by Owen as Anomodontia. But during the last twenty years the distinguished American paleontologists, Cope and Osborn, have greatly increased our knowledge of them, and have claimed that the stem-forms of the Mammals must be sought in this order. As a matter of fact, the Theromorpha are nearer to the Mammals in the chief points of structure than any other reptiles. This is especially true of the Thereodontia, to which the Pureosauria and Pelycosauria belong (Figure 2.267). The whole structure of their pelvis and hind-feet has attained the same form as in the Monotremes, the lowest Mammals. The formation of the scapula and the quadrate bone shows an approach to the Mammals such as we find in no other group of reptiles. The teeth also are already divided into incisors, canines, and molars. Nevertheless, it is very doubtful whether the Theromorpha really are in the ancestral line of the Sauromammals, or lead direct from the Tocosauria to the earliest Mammals. Other experts on this group believe that it is an independent legion of the reptiles, connected, perhaps, at its lowest root, with the Sauromammals, but developed quite independently of the Mammals—though parallel to them in many ways.

One of the most important of the zoological facts that we rely on in our investigation of the genealogy of the human race is the position of man in the Mammal class. However different the views of zoologists may have been as to this position in detail, and as to his relations to the apes, no scientist has ever doubted that man is a true mammal in his whole organisation and development. Linne drew attention to this fact in the first edition of his famous Systema Naturae (1735). As will be seen in any museum of anatomy or any manual of comparative anatomy; the human frame has all the characteristics that are common to the Mammals and distinguish them conspicuously from all other animals.

(FIGURE 2.266. Skull of a Permian lizard (Palaehatteria longicaudata). (From Credner.) n nasal bone, pf frontal bone, l lachrymal bone, po postorbital bone, sq covering bone, i cheek-bone, vo vomer, im inter-maxillary.)

If we examine this undoubted fact from the point of view of phylogeny, in the light of the theory of descent, it follows at once that man is of a common stem with all the other Mammals, and comes from the same root as they. But the various features in which the Mammals agree and by which they are distinguished are of such a character as to make a polyphyletic hypothesis quite inadmissible. It is impossible to entertain the idea that all the living and extinct Mammals come from a number of separate roots. If we accept the general theory of evolution, we are bound to admit the monophyletic hypothesis of the descent of all the Mammals (including man) from a single mammalian stem-form. We may call this long-extinct root-form and its earliest descendants (a few genera of one family) "primitive mammals" or "stem-mammals" (Promammalia). As we have already seen, this root-form developed from the primitive Proreptile stem in a totally different direction from the birds, and soon separated from the main stem of the reptiles. The differences between the Mammals and the reptiles and birds are so important and characteristic that we can assume with complete confidence this division of the vertebrate stem at the commencement of the development of the Amniotes. The reptiles and birds, which we group together as the Sauropsids, generally agree in the characteristic structure of the skull and brain, and this is notably different from that of the Mammals. In most of the reptiles and birds the skull is connected with the first cervical vertebra (the atlas) by a single, and in the Mammals (and Amphibia) by a double, condyle at the back of the head. In the former the lower jaw is composed of several pieces, and connected with the skull so that it can move by a special maxillary bone (the quadratum); in the Mammals the lower jaw consists of one pair of bony pieces, which articulate directly with the temporal bone. Further, in the Sauropsids the skin is clothed with scales or feathers; in the Mammals with hair. The red blood-cells of the former have a nucleus; those of the latter have not. In fine, two quite characteristic features of the Mammals, which distinguish them not only from the birds and reptiles, but from all other animals, are the possession of a complete diaphragm and of mammary glands that produce the milk for the nutrition of the young. It is only in the Mammals that the diaphragm forms a transverse partition of the body-cavity, completely separating the pectoral from the abdominal cavity. It is only in the mammals that the mother suckles its young, and this rightly gives the name to the whole class (mamma = breast).

(FIGURE 2.267. Skull of a Triassic theromorphum (Galesaurus planiceps), from the Karoo formation in South Africa. (From Owen.) a from the right, b from below, c from above, d tricuspid tooth. N nostrils, NA nasal bone, Mx upper jaw, Prf prefrontal, Fr frontal bone, A eye-pits, S temple-pits. Pa Parietal eye, Bo joint at back of head, Pt pterygoid-bone, Md lower jaw.)

From these pregnant facts of comparative anatomy and ontogeny it follows absolutely that the whole of the Mammals belong to a single natural stem, which branched off at an early date from the reptile-root. It follows further with the same absolute certainty that the human race is also a branch of this stem. Man shares all the characteristics I have described with all the Mammals, and differs in them from all other animals. Finally, from these facts we deduce with the same confidence those advances in the vertebrate organisation by which one branch of the Sauromammals was converted into the stem-form of the Mammals. Of these advances the chief were: (1) The characteristic modification of the skull and the brain; (2) the development of a hairy coat; (3) the complete formation of the diaphragm; and (4) the construction of the mammary glands and adaptation to suckling. Other important changes of structure proceeded step by step with these.

The epoch at which these important advances were made, and the foundation of the Mammal class was laid, may be put with great probability in the first section of the Mesozoic or secondary age—the Triassic period. The oldest fossil remains of mammals that we know were found in strata that belong to the earliest Triassic period—the upper Kueper. One of the earliest forms is the genus Dromatherium, from the North American Triassic (Figure 2.268). Their teeth still strikingly recall those of the Pelycosauria. Hence we may assume that this small and probably insectivorous mammal belonged to the stem-group of the Promammals. We do not find any positive trace of the third and most advanced division of the Mammals—the Placentals. These (including man) are much younger, and we do not find indisputable fossil remains of them until the Cenozoic age, or the Tertiary period. This paleontological fact is very important, because it fully harmonises with the evolutionary succession of the Mammal orders that is deduced from their comparative anatomy and ontogeny.

The latter science teaches us that the whole Mammal class divides into three main groups or sub-classes, which correspond to three successive phylogenetic stages. These three stages, which also represent three important stages in our human genealogy, were first distinguished in 1816 by the eminent French zoologist, Blainville, and received the names of Ornithodelphia, Didelphia, and Monodelphia, according to the construction of the female organs (delphys = uterus or womb). Huxley afterwards gave them the names of Prototheria, Metatheria, and Epitheria. But the three sub-classes differ so widely from each other, not only in the construction of the sexual organs, but in many other respects also, that we may confidently draw up the following important phylogenetic thesis: The Monodelphia or Placentals descend from the Didelphia or Marsupials; and the latter, in turn, are descended from the Monotremes or Ornithodelphia.

Thus we must regard as the twenty-first stage in our genealogical tree the earliest and lowest chief group of the Mammals—the sub-class of the Monotremes ("cloaca-animals," Ornithodelphia, or Prototheria, Figures 2.269 and 2.270). They take their name from the cloaca which they share with all the lower Vertebrates. This cloaca is the common outlet for the passage of the excrements, the urine, and the sexual products. The urinary ducts and sexual canals open into the hindmost part of the gut, while in all the other Mammals they are separated from the rectum and anus. The latter have a special uro-genital outlet (porus urogenitalis). The bladder also opens into the cloaca in the Monotremes, and, indeed, apart from the two urinary ducts; in all the other Mammals the latter open directly into the bladder. It was proved by Haacke and Caldwell in 1884 that the Monotremes lay large eggs like the reptiles, while all the other Mammals are viviparous. In 1894 Richard Semon further proved that these large eggs, rich in food-yelk, have a partial segmentation and discoid gastrulation, as I had hypothetically assumed in 1879; here again they resemble their reptilian ancestors. The construction of the mammary gland is also peculiar in the Monotremes. In them the glands have no teats for the young animal to suck, but there is a special part of the breast pierced with holes like a sieve, from which the milk issues, and the young Monotreme must lick it off. Further, the brain of the Monotremes is very little advanced. It is feebler than that of any of the other Mammals. The fore-brain or cerebrum, in particular, is so small that it does not cover the cerebellum. In the skeleton (Figure 2.270) the formation of the scapula among other parts is curious; it is quite different from that of the other Mammals, and rather agrees with that of the reptiles and Amphibia. Like these, the Monotremes have a strongly developed caracoideum. From these and other less prominent characteristics it follows absolutely that the Monotremes occupy the lowest place among the Mammals, and represent a transitional group between the Tocosauria and the rest of the Mammals. All these remarkable reptilian characters must have been possessed by the stem-form of the whole mammal class, the Promammal of the Triassic period, and have been inherited from the Proreptiles.

(FIGURE 2.268. Lower jaw of a Primitive Mammal or Promammal (Dromatherium silvestre) from the North American Triassic. i incisors, c canine, p premolars, m molars. (From Doderlein.))

During the Triassic and Jurassic periods the sub-class of the Monotremes was represented by a number of different stem-mammals. Numerous fossil remains of them have lately been discovered in the Mesozoic strata of Europe, Africa, and America. To-day there are only two surviving specimens of the group, which we place together in the family of the duck-bills, Ornithostoma. They are confined to Australia and the neighbouring island of Van Diemen's Land (or Tasmania); they become scarcer every year, and will soon, like their blood-relatives, be counted among the extinct animals. One form lives in the rivers, and builds subterraneous dwellings on the banks; this is the Ornithorhyncus paradoxus, with webbed feet, a thick soft fur, and broad flat jaws, which look very much like the bill of a duck (Figures 2.269 and 2.270). The other form, the land duck-bill, or spiny ant-eater (Echidna hystrix), is very much like the anteaters in its habits and the peculiar construction of its thin snout and very long tongue; it is covered with needles, and can roll itself up like a hedgehog. A cognate form (Parechidna Bruyni) has lately been found in New Guinea.

These modern Ornithostoma are the scattered survivors of the vast Mesozoic group of Monotremes; hence they have the same interest in connection with the stem history of the Mammals as the living stem-reptiles (Hatteria) for that of the reptiles, and the isolated Acrania (Amphioxus) for the phylogeny of the Vertebrate stem.

The Australian duck-bills are distinguished externally by a toothless bird-like beak or snout. This absence of real bony teeth is a late result of adaptation, as in the toothless Placentals (Edentata, armadillos and ant-eaters). The extinct Monotremes, to which the Promammalia belonged, must have had developed teeth, inherited from the reptiles. Lately small rudiments of real molars have been discovered in the young of the Ornithorhyncus, which has horny plates in the jaws instead of real teeth.

(FIGURE 2.269. The Ornithorhyncus or Duck-mole. (Ornithorhyncus paradoxus).

FIGURE 2.270. Skeleton of the Ornithorhyncus.)

The living Ornithostoma and the stem-forms of the Marsupials (or Didelphia) must be regarded as two widely diverging lines from the Promammals. This second sub-class of the Mammals is very interesting as a perfect intermediate stage between the other two. While the Marsupials retain a great part of the characteristics of the Monotremes, they have also acquired some of the chief features of the Placentals. Some features are also peculiar to the Marsupials, such as the construction of the male and female sexual organs and the form of the lower jaw. The Marsupials are distinguished by a peculiar hook-like bony process that bends from the corner of the lower jaw and points inwards. As most of the Placentals have not this process, we can, with some probability, recognise the Marsupial from this feature alone. Most of the mammal remains that we have from the Jurassic and Cretaceous deposits are merely lower jaws, and most of the jaws found in the Jurassic deposits at Stonesfield and Purbeck have the peculiar hook-like process that characterises the lower jaw of the Marsupial. On the strength of this paleontological fact, we may suppose that they belonged to Marsupials. Placentals do not seem to have existed at the middle of the Mesozoic age—not until towards its close (in the Cretaceous period). At all events, we have no fossil remains of indubitable Placentals from that period.