It may be considered as quite certain, from reasons based upon their comparative anatomy and their history of development, that Placental animals first developed out of Marsupials, and that this very important development—the first origin of the placenta—probably took place in the beginning of the tertiary epoch, during the eocene period. But one of the most difficult questions in the genealogy of animals is the important consideration whether all Placental animals have arisen out of one or out of several distinct branches of Marsupials; in other words, whether the origin of the placenta occurred but once, or several times.
When, in my General Morphology, I for the first time endeavoured to establish the pedigree of Mammals, I here, as in most cases, preferred the monophyletic, or one-rooted, to the polyphyletic, or many-rooted, hypothesis of descent. I assumed that all Placental animals were derived from a single form of Marsupial animal, which, for the first time, began to form a placenta. In this case the Villiplacentals, Zonoplacentals, and Discoplacentals would perhaps have to be considered as three diverging branches of the common primary form of Placentals, or it might also be conceived that the two latter, the Deciduata, had developed only at a later period out of the Indeciduata, which on their part had arisen directly out of the Marsupials. However, there are also important reasons for the alternative; namely, that several groups of Placentals, differing from the beginning, arose out of several distinct groups of Marsupials, so that the placenta itself was formed several times independently. This opinion is maintained by Huxley, the most eminent English zoologist, and by many others. In this case the Indeciduata and the Deciduata would perhaps have to be considered as two completely distinct groups; then the order of Hoofed animals, as the primary group of the Indeciduata, might be supposed to have originated out of the Marsupial hoofed animals (Barypoda). Among the Deciduata, on the other hand, the order of Semi-apes, as the common primary form of the other orders, might possibly have arisen out of Handed Marsupials (Pedimana). But it is also conceivable that the Deciduata themselves have arisen out of several different orders of Marsupials, Animals of Prey out of Rapacious Marsupials, Gnawing animals out of Gnawing Marsupials, Semi-apes out of Handed Marsupials, etc. As we do not at present possess sufficient empiric material to solve this most difficult question, we must leave it and turn our attention to the history of the different orders of Placental animals, whose pedigree can often be very accurately established in detail.
We must, as already remarked, consider the order of Hoofed animals (Ungulata) as the primary group of the Indeciduata, or Tuft-placentals; the two other orders, Whales and Toothless animals, developed out of them, as two diverging groups, probably only at a later period, by adaptation to very different modes of life. But it is also possible that the animals poor in teeth (Edentata) may be of quite a different origin.
Hoofed animals are in many respects among the most important and the most interesting Mammals. They distinctly show that a true understanding of the natural relationship of animals can never be revealed to us merely by the study of living forms, but in all cases only by an equal consideration of their extinct and fossil blood-relations and ancestors. If, as is usually done, only the living Hoofed animals are taken into consideration, it seems quite natural to divide them into three entirely distinct orders, namely: (1) Horses, or Single-hoofed animals (Solidungula, or Equina); (2) Ruminating animals, or Double-hoofed (Bisulca, or Ruminantia); and (3) Thick-skinned, or Many-hoofed (Multungula, or Pachyderma). But as soon as the extinct Hoofed animals of the tertiary period are taken into consideration—of which animals we possess very numerous and important remains—it is seen that this division, but more especially the limitation of the Thick-skinned animals, is completely artificial, and that these three groups are merely top branches lopped from the pedigree of Hoofed animals, which are most closely connected by extinct intermediate forms. The one half of the Thick-skinned animals—rhinoceroses, tapirs, and palæotheria—manifest the closest relationships to horses, and have like them odd-toed feet; whereas the other half of the Thick-skinned animals—pigs, hippopotami, and anoplotheria—on account of their double-toed feet are much more closely allied to ruminating animals than to the former. Hence we must, in the first place, among Hoofed animals distinguish the two orders of Paired-hoofs and Odd-hoofs, as two natural groups, which developed as diverging branches out of the old tertiary primary group of Primary Hoofed animals, or Prochela.
The order of Odd-hoofed animals (Perissodactyla) comprises those Ungulata in which the middle (or third) toe of the foot is much more strongly developed than the others, so that it forms the actual centre of the hoof. This order includes the very ancient, common, primary group of all Hoofed animals, that is, the Primary-hoofed animals (Prochela), which are found in a fossil state in the oldest Eocene strata (Lophiodon, Coryphodon, Pliolophus). Directly allied to this group is that branch which is the actual primary form of the Odd-hoofed animals, namely, the Palæotheria, fossils of which occur in the upper Eocene and lower Miocene. Out of the Palæotheria, at a later period, the rhinoceroses (Nasicornia) and rhinoceros-horses (Elasmotherida) on the one hand, and the tapirs, lama-tapirs, and primæval horses, on the other, developed as two diverging branches. The long since extinct primæval horses, or Anchitheria, formed the transition from the Palæotheria and tapirs to the Miocene horses, or hipparions, which are closely allied to the genuine living horses.
The second main group of Hoofed animals, the order of Pair-hoofed animals (Artiodactyla), comprises those hoofed animals in which the middle (third) and fourth toe of the foot are almost equally developed, so that the space between the two forms the central line of the entire foot. The order is divided into two sub-orders—the Pig-shaped and the Cud-chewing, or Ruminating. The Pig-shaped (Chœromorpha) comprise in the first place the other branch of Primary-Hoofed-animals, the Anoplotheria, which we consider as the common primary form of all Pair-hoofed animals, or Artiodactyla (Dichobune, etc.). Out of the Anoplotheria arose, as two diverging branches, the primæval swine, or Anthracotheria, on the one hand, forming the transition to swine and river-horses, and the Xiphodonta on the other hand, forming the transition to Ruminating animals. The oldest Ruminating animals (Ruminantia) are the Primæval Stags, or Dremotheria, out of which, possibly, the stag-shaped (Elaphia), the hollow-horned (Cavicornia), and camels (Tylopoda), have developed as three diverging branches. Yet these latter are, in many respects, more allied to the Odd-hoofs than to the genuine Pair-hoofs. The accompanying systematic survey on p. 252, will show how the numerous families of Hoofed animals are grouped, in correspondence with this genealogical hypothesis.
| SYSTEMATIC SURVEY | |||||||||||
| Of the Sections and Families of Hoofed Animals, or Ungulata. | |||||||||||
| (N.B. Those families that are extinct are marked with an asterisk.) | |||||||||||
| Orders of Hoofed animals. |
Sections of Hoofed Animals. |
Families of Hoofed Animals. |
Systematic Name of the Families. |
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| I. Odd-toed Hoofed Animals Ungulata Perissodactyla |
I. Primary Hoofed Animals.* Prochela |
1. | Lophiodonta | 1. | Lophiodontia* | ||||||
| 2. | Pliolophida | 2. | Pliolophida* | ||||||||
| II. Tapir-shaped Tapiromorpha |
3. | Primary Odd-hoofs | 3. | Palæotherida* | |||||||
| 4. | Lama-tapirs | 4. | Macrauchenida* | ||||||||
| 5. | Tapirs | 5. | Tapirida | ||||||||
| 6. | Rhinoceroses | 6. | Nasicornia | ||||||||
| 7. | Rhinoceros-horses | 7. | Elasmotherida* | ||||||||
| III. Single-hoofs Solidungula |
8. | Primæval horses | 8. | Anchitherida* | |||||||
| 9. | Horses | 9. | Equina | ||||||||
| II. Pair-toed Hoofed Animals Ungulata Artiodactyla |
IV. Pig-shaped Chœromorpha |
10. | Primary Pair-hoofs | 10. | Lophiodontia* | ||||||
| 11. | Primæval pigs | 11. | Anthracotherida* | ||||||||
| 12. | Pigs | 12. | Setigera | ||||||||
| 13. | River horses | 13. | Obesa | ||||||||
| 14. | Primæval ruminants | 14. | Xiphodontia* | ||||||||
| V. Ruminating animals Ruminantia |
A. Stag-shaped Elephia |
a. | 15. | Primæval deer | 15. | Dremotherida* | |||||
| 16. | Pseudo musk deer | 16. | Tragulida | ||||||||
| b. | 17. | Musk deer | 17. | Moschida | |||||||
| 18. | Deer | 18. | Cervina | ||||||||
| c. | 19. | Primæval giraffes | 19. | Sivatherida* | |||||||
| 20. | Giraffes | 20. | Devexa | ||||||||
| B. Hollow-horned Cavicornia |
d. | 21. | Primæval gazelles | 21. | Antilocaprina* | ||||||
| 22. | Gazelles | 22. | Antilopina | ||||||||
| e. | 23. | Goats | 23. | Caprina | |||||||
| 24. | Sheep | 24. | Ovina | ||||||||
| 25. | Oxen | 25. | Bovina | ||||||||
| C. Pad-footed Tylopoda |
26. | Lamas | 26. | Auchenida | |||||||
| 27. | Camels | 27. | Camelida | ||||||||
PEDIGREE OF THE UNGULATES.
| Oxen | Giraffes | |||||||
| │ |
Sheep | │ |
Deer | |||||
| │ |
│ |
│ |
│ |
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| Goats | Musk deer | Horses Equi |
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| │ |
│ |
│ |
│ |
│ |
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| Antelopes | Deer-shaped Elaphia |
Camels and Lamas Tylopoda |
Intermediate horses Hippariones |
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| │ │ |
│ │ |
│ │ |
│ │ |
│ │ |
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| Hollow-horned Cavicornia |
│ │ │ |
│ │ │ |
│ │ │ |
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| │ │ │ |
│ │ │ |
│ │ │ |
Primæval horses Anchitherida |
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| │ |
│ |
│ |
│ |
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| │ │ │ │ |
Single Hoofers Solidungula |
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| Ruminating Animals Ruminantia |
│ │ │ |
│ │ │ |
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| Sea-oxen Sirenia |
│ │ |
Tapirs Tapirida |
│ │ |
│ │ |
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| │ │ │ |
River-horses Obesa |
│ │ │ |
│ │ │ |
Lama-tapirs Macrauchenida |
│ │ │ |
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| │ │ |
│ │ |
Pigs Setigera |
│ │ |
│ │ |
│ │ |
│ │ |
||
| │ |
│ |
│ |
│ |
│ |
│ |
│ |
||
| │ |
│ |
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| Primæval pigs Anthracotherida |
│ │ │ |
Rhinoceros-horses Elasmotherida |
│ │ │ |
│ │ │ |
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| │ │ |
│ │ |
Rhinoceruses Nasicornia |
│ │ |
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| │ |
│ |
│ |
│ |
│ |
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| │ │ │ |
Primæval ruminants Xiphodontia |
│ │ |
│ │ │ |
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| │ |
│ |
│ |
│ |
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| Primary Pair-hoofs Anoplotherida |
Primary Odd-hoofs Palæotherida |
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| │ |
│ |
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| Prochela Primary-hoofed-animals (Lophiodontia and Pliophida) |
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| │ |
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| (Hoofed marsupials? Barypoda?) | ||||||||
It is probable that the remarkable legion of Whales (Cetacea) originated out of Hoofed animals, which accustomed themselves exclusively to an aquatic life, and thereby became transformed into the shape of fish. Although these animals seem externally very like many genuine Fish, yet they are, as even Aristotle perceived, genuine Mammals. By their whole internal structure—in so far as it has not become changed by adaptation to an aquatic life—they, of all known Mammals, are most closely allied to Hoofed animals, and more especially agree with them in the absence of the decidua and in the tufted placenta. Even at the present day the river-horse (Hippopotamus) constitutes a kind of transition form to the Sea Cows (Sirenia), and from this it seems most probable that the extinct primary forms of the Cetacea are most closely allied to the Sea Cows of the present day, and that they developed out of Pair-hoofed animals, which were related to the hippopotamus. Out of the order of Herbivorous whales (Phycoceta)—to which the sea cows belong, and which accordingly, very probably, contain the primary forms of the legion—the other order of Carnivorous whales (Sarcoceta) appears to have developed at a later period. But Huxley thinks that these latter were of quite a different origin, and that they arose out of the Carnaria through the Seals. Among the Sarcoceta, the extinct gigantic Zeuglodonta (Zeugloceta)—whose fossil skeletons some time ago excited great interest, it being thought that they were “sea serpents”—are probably only a peculiarly developed lateral branch of genuine whales (Autoceta), which comprise, besides the colossal whalebone whales, the cachalot or spermaceti whales, dolphins, narwhals, porpoises, etc.
The third legion of the Indeciduata, or Sparsi-placentalia, comprises the strange group of the animals poor in teeth (Edentata); it is composed of the two orders of burrowers and sloths. The order of Burrowers (Effodientia) consists of the two sub-orders of ant eaters (Vermilinguia), to which the scaled animals also belong, and the girdle animals (Cingulata), which were formerly represented by the gigantic Glyptodons. The order of Sloths (Tardigrada) consists of the two sub-orders of the small, still living dwarf sloths (Bradypoda), and of the extinct unwieldy giant sloths (Gravigrada). The enormous fossil remains of these colossal herbivora suggest that the whole legion is becoming extinct, and that the Edentata of the present day are but a poor remnant of the mighty order of the diluvial period. The close relations between the still living South American Edentata and the extinct gigantic forms which are found beside the latter on the same part of the globe, made such an impression upon Darwin on his first visit to South America, that they even then suggested to him the fundamental idea of the Theory of Descent. (See above, vol. i. p. 134.) But it is precisely the genealogy of this legion which is most difficult. The Edentata are perhaps nothing but a peculiarly developed lateral branch of the Ungulata; but it may also be that their root lies in quite another direction.
We now leave the first main group of Placental animals, the Indeciduata, and turn to the second main group, namely, the Deciduata, or animals with decidua, which are distinguished from the former by possessing a deciduous membrane, or decidua, during their embryonal life. We here meet with a very remarkable small group of animals, for the most part extinct, and which probably were the old tertiary (or eocene) ancestors of man. These are the Semi-apes, or Lemurs (Prosimiæ); these curious animals are probably the but little changed descendants of the primæval group of Placentalia which we have to consider as the common primary form of all Deciduata. They have hitherto been classed together in the same order with Apes which Blumenbach called Quadrumana (four-handed). However, I regard them as entirely distinct from these, not merely because they differ from all Apes, much more than do the most different Apes from one another, but also because they comprise most interesting transitional forms leading to the other orders of Deciduata. I conclude from this that the few still living Semi-apes, which moreover differ very much among one another, are the last surviving remnants of a primary group now almost extinct, but which was at one time rich in forms, and out of which all the other Deciduata (possibly with the single exception of Beasts of Prey, and Pseudo-hoofed animals) have developed as diverging branches. The old primary group of Semi-apes has probably developed out of Handed or Ape-footed Marsupials (Pedimana), which are surprisingly like them in the transformation of their hinder feet into grasping hands. The primæval primary forms themselves (which probably originated in the eocene period) are of course long since extinct, as are also the greater portion of the transition-forms between them and all the other orders of Deciduata. However, individual remnants of the latter are preserved among the Semi-apes of the present day. Among these, the remarkable Finger-animal of Madagascar (Chiromys madagascariensis) constitutes the remnant of the group of the Leptodactyla and the transition to Rodents. The strange flying lemur in the South Sea and Sunda islands (Galeopithecus), the only remnant of the group of Pteropleura, forms a perfect intermediate stage between Semi-apes and Bats. The long-footed Semi-apes (Tarsius, Otolicnus) constitute the last remnant of that primary branch (Macrotarsi) out of which the Insectivora developed. The short-footed forms (Brachytarsi) are the medium of connection between them and genuine Apes. The Short-footed Semi-apes comprise the long-tailed Lemur, the short-tailed Lichanotus, and the Stenops, the latter of which seems to be very closely allied to the probable ancestors of man among the Semi-apes. The short-footed as well as the long-footed Prosimiæ live widely distributed over the islands of southern Asia and Africa, more especially in Madagascar; some live also on the continent of Africa. No Semi-ape, either living or in a fossil state, has as yet been found in America. They all lead a solitary, nocturnal kind of life, and climb about on trees. (Compare vol. i. p. 361.)
Among the six remaining orders of Deciduata, all of which are probably derived from long since extinct Semi-apes, the order of Gnawing animals (Rodentia), which is rich in forms, has remained at the lowest stage. Among these the squirrel-like animals (Sciuromorpha) stand nearest akin to the Pedimanous Marsupials. Out of this primary group the mouse-like animals (Myomorpha) and the porcupine-like animals (Hystricomorpha) developed probably as two diverging branches, the former of which are directly connected with the squirrel-like animals, by the eocene Myoxida, the latter by the eocene Psammoryctida. The fourth sub-order, the hare-like animals (Lagomorpha), probably developed only at a later period out of one of the other three sub-orders.
Very closely allied to the Rodentia is the remarkable order of Pseudo-hoofed animals (Chelophora). Of these there now live but two genera, indigenous to Asia and Africa, namely, Elephants (Elephas), and Rock Conies (Hyrax). Both have hitherto generally been classed among real Hoofed animals, or Ungulata, with which they agree in the formation of the feet. But an identical transformation of nails or claws into hoofs occurs also in genuine Rodentia and in certain hoofed Rodentia (Subungulata) which live exclusively in South America. Beside smaller forms (for example, guinea pigs and gold hares) the Subungulata also include the largest of all Rodentia, namely, the Capybara Rats, which are about four feet in length. The Rock Conies, which are externally very nearly akin to Rodents, especially to the hoofed Rodents, were formerly classed among Rodentia by some celebrated zoologists, as an especial sub-class (Lamnungia). Elephants, on the other hand, when not classed among Hoofed animals, were generally considered as the representatives of a special order which were called Trunked animals (Proboscidea). But the formation of the placentas of Elephants and of Hyrax agree in a remarkable manner, and are entirely distinct from those of Hoofed animals. These latter never possess a decidua, whereas Elephants and Hyrax are genuine Deciduata. Their placenta is indeed not of the form of a disc, but of a girdle, as in the case of Animals of Prey; it is very possible that the girdle-shaped placenta is but a secondary development of the discoplacenta. Thus, then, it might be thought that the Pseudo-hoofed animals have developed out of a branch of the Rodentia, and in a similar manner perhaps the Carnivora out of a branch of the Insectivora. At all events, Elephants and Hyrax in many respects, especially in the formation of important skeletal parts, of the limbs, etc., are more closely allied to the Rodentia, and more especially to hoofed Rodentia, than to genuine Hoofed animals. Moreover several extinct forms, especially the remarkable South American Arrow-toothed animals (Toxodontia), stand in many respects mid-way between Elephants and Rodentia. That the still living Elephants and Hyrax are but the last survivors of a group of Pseudo-hoofed animals, which was once rich in forms, is proved not only by the very numerous fossil species of Elephants and Mastodon (some of which are even larger, others also much smaller than the Elephants of the present day), but also by the remarkable miocene Dinotheria (Gonyognatha), between which and their next kindred, the Elephants, there must be a long series of unknown connecting intermediate forms. Taking all things into consideration, the most probable hypothesis which can be established at present as to the origin and the relationship of Elephants, Dinotheria, Toxodon, and Hyrax is, that they are the last survivors of a group of Pseudo-hoofed animals rich in forms, which developed out of the Rodentia, and probably out of relatives of the Subungulata.
The order of Insect Eaters (Insectivora) is a very ancient group, and is next akin to the common extinct primary form of the Deciduata, as well as to the Semi-apes of the present day. It has probably developed out of Semi-apes which were closely allied to the Long-footed Lemurs (Macrotarsi) of the present day. It is separated into two orders, Menotyphla and Lipotyphla; the Menotyphla are probably the older of the two, and are distinguished from the Lipotyphla by possessing an intestinal cœcum, or typhlon. The Menotyphla include the climbing Tupajas of the Sunda Isles, and the leaping Macroscelides of Africa. The Lipotyphla are represented in our country by shrew mice, moles, and hedgehogs. The Insectivora, in the formation of their jaws and their mode of life, are nearly akin to Carnivora, but are, on the other hand, by their discoplacentas and by their large seminal vesicles, allied to Rodents.
It is probable that the order of Rapacious animals (Carnaria) developed out of a long since extinct branch of Insectivora, at the beginning of the Eocene period. It is a natural group, very rich in forms, but still of very uniform organization. The Rapacious animals are sometimes also called Girdle-placentals (Zonoplacentals), although the Pseudo-hoofed animals (Chelophora), in the same way, also deserve this designation. But as the latter, in other respects, are more closely allied to the Rodentia than to Carnaria, we have already discussed them in connection with the former. Animals of prey are divided into two, externally very different, but internally very closely related, sub-orders, namely, Land animals of prey and Marine animals of prey. The Land animals of prey (Carnivora) comprise bears, dogs, cats, etc., whose pedigree can be approximately guessed at by means of many extinct intermediate forms. The Marine animals of prey, or Seals (Pinnipedia), comprise sea bears, sea dogs, sea lions, and walruses. Although marine animals of prey appear externally very unlike land animals of prey, yet by their internal structure, their jaw and their peculiar girdle-shaped placenta, they are very nearly akin to them, and have evidently originated out of a branch of them, probably out of a kind of weasel (Mustelina). Even at the present day the fish otters (Lutra), and still more so the sea otters (Enhydris), present a direct form of transition to Seals, and clearly show how the bodies of land Carnivora are transformed into the shape of a Seal, by adaptation to an aquatic life, and how the steering fins of marine rapacious animals have arisen out of the legs of the former. The latter consequently stand in the same relation to the former as do the Whales to Hoofed animals among the Indeciduata. In the same way as the river-horse at present stands midway between the extreme branches of oxen and sea oxen, the sea otter still forms a surviving intermediate stage between the widely separated branches of dogs and sea dogs. In both cases the complete transformation of the external form, consequent upon adaptation to entirely different conditions of life, has not been able to efface the solid foundation of the inherited internal peculiarities.
According to Huxley’s opinion, which has already been quoted, only the Herbivorous Whales (Sirenia) are derived from Hoofed animals; on the other hand, the Carnivorous Cetacea (Sarcoceta) are derived from the marine animals of prey; the Zeuglodonts would form a transition between the two latter. But in this case it would be difficult to understand the close anatomical relations which exist between the Herbivorous and Carnivorous Cetacea. The strange peculiarities in the internal and external structure which so strikingly distinguish the two groups from all other mammals would then have to be regarded only as analogies (caused by the same kinds of adaptation), not as homologies (transmitted from a common primary form). The latter, however, strikes me as being by far the more probable, and hence I have left all the Cetacea among the Indeciduata as one group of kindred origin.
The remarkable order of Flying Mammals, or Bats (Chiroptera), stands near to the Carnaria as well as to the Insectivora. It has become strikingly transformed by adaptation to a flying mode of life, just as marine animals of prey have become modified by adaptation to a swimming mode of life. This order probably also originated out of the Semi-apes, with which it is even at present closely allied, through the flying lemurs (Galeopithecus). Of the two orders of flying animals, the insect-eating forms, or flying mice (Nycterides), probably developed out of those eating fruits, or flying foxes (Pterocynes); for the latter are, in many ways, more closely allied to Semi-apes than are the former.
We have now still to discuss the genuine Apes (Simiæ) as the last order of Mammals; but as, according to the zoological system, the human race belongs to this order, and as it undoubtedly developed historically out of a branch of this order, we shall devote a special chapter to a more careful examination of its pedigree and history.
CHAPTER XXII.
ORIGIN AND PEDIGREE OF MAN.
The Application of the Theory of Descent to Man.—Its Immense Importance and Logical Necessity.—Man’s Position in the Natural System of Animals, among Disco-placental Animals.—Incorrect Separation of the Bimana and Quadrumana.—Correct Separation of Semi-apes from Apes.—Man’s Position in the Order of Apes.—Narrow-nosed Apes (of the Old World) and Flat-nosed Apes (of America).—Difference of the two Groups.—Origin of Man from Narrow-nosed Apes.—Human Apes, or Anthropoides.—African Human Apes (Gorilla and Chimpanzee).—Asiatic Human Apes (Orang and Gibbon).—Comparison between the different Human Apes and the different Races of Men.—Survey of the Series of the Progenitors of Man.—Invertebrate Progenitors (Prochordata) and Vertebrate Progenitors.
Of all the individual questions answered by the Theory of Descent, of all the special inferences drawn from it, there is none of such importance as the application of this doctrine to Man himself. As I remarked at the beginning of this treatise, the inexorable necessity of the strictest logic forces us to draw the special deductive conclusion from the general inductive law of the theory, that Man has developed gradually, and step by step, out of the lower Vertebrata, and more immediately out of Ape-like Mammals. That this doctrine is an inseparable part of the Theory of Descent, and hence also of the universal Theory of Development in general, is recognized by all thoughtful adherents of the theory, as well as by all its opponents who reason logically.
But if the doctrine be true, then the recognition of the animal origin and pedigree of the human race will necessarily affect more deeply than any other progress of the human mind the views we form of all human relations, and the aims of all human science. It must sooner or later produce a complete revolution in the conception entertained by man of the entire universe. I am firmly convinced that in future this immense advance in our knowledge will be regarded as the beginning of a new period of the development of Mankind. It can only be compared to the discovery made by Copernicus, who was the first who ventured distinctly to express the opinion, that it was not the sun which moved round the earth, but the earth round the sun. Just as the geocentric conception of the universe—namely, the false opinion that the earth was the centre of the universe, and that all its other portions revolved round the earth—was overthrown by the system of the universe established by Copernicus and his followers, so the anthropocentric conception of the universe—the vain delusion that Man is the centre of terrestrial nature, and that its whole aim is merely to serve him—is overthrown by the application (attempted long since by Lamarck) of the theory of descent to Man. As Copernicus’ system of the universe was mechanically established by Newton’s theory of gravitation, we see Lamarck’s theory of descent attain its causal establishment by Darwin’s theory of selection. This comparison, which is very interesting in many respects, I have discussed in detail elsewhere.
In order to carry out this extremely important application of the Theory of Descent to man, with the necessary impartiality and objectivity, I must above all beg the reader (at least for a short time) to lay aside all traditional and customary ideas on the “Creation of Man,” and to divest himself of the deep-rooted prejudices concerning it, which are implanted in the mind in earliest youth. If he fail to do this, he cannot objectively estimate the weight of the scientific arguments which I shall bring forward in favour of the animal derivation of Man, that is, of his origin out of Ape-like Mammals. We cannot here do better than imagine ourselves with Huxley to be the inhabitants of another planet, who, taking the opportunity of a scientific journey through the universe, have arrived upon the earth and have there met with a peculiar two-legged mammal called Man, diffused over the whole earth in great numbers. In order to examine him zoologically, we should pack a number of the individuals of different ages and from different lands (as we should do with the other animals collected on the earth) into large vessels filled with spirits of wine, and on our return to our own planet we should commence the comparative anatomy of all these terrestrial animals quite objectively. As we should have no personal interest in Man, in a creature so entirely different from ourselves, we should examine and criticise him as impartially and objectively as we should the other terrestrial animals. In doing this we should, of course, in the first place refrain from all conjectures and speculations on the nature of his soul, or on the spiritual side of his nature, as it is usually called. We should occupy ourselves solely with his bodily structure, and with that natural conception of it which is offered by the history of his individual development.
It is evident that in order correctly to determine Man’s position among the other terrestrial organisms we must, in the first place, follow the guidance of the natural system. We must endeavour to determine the position which belongs to Man in the natural system of animals as accurately and distinctly as possible. We shall then, if in fact the theory of descent be correct, be able from his position in the system to determine the real primary relationship, and the degree of consanguinity connecting Man with the animals most like him. The hypothetical pedigree of the human race will then follow naturally as the final result of this anatomical and systematic inquiry.
Now if, by means of comparative anatomy and ontogeny, we seek for man’s position in that Natural System of animals which formed the subject of the last two chapters, the incontrovertible fact will at once present itself to us, that man belongs to the tribe, or phylum, of the Vertebrata. Every one of the characteristics, which so strikingly distinguish all the Vertebrata from all Invertebrata, is possessed by him. It has also never been doubted that of all the Vertebrata the Mammals are most closely allied to Man, and that he possesses all the characteristic features distinguishing them from all other Vertebrata. If then we further carefully examine the three different main groups or sub-classes of Mammals—the inter-connections of which were discussed in our last chapter—there cannot be the slightest doubt that Man belongs to the Placentals, and shares with all other Placentals, the important characteristics which distinguish them from Marsupials and from Cloacals. Finally, of the two main groups of placental Mammals, the Deciduata and the Indeciduata, the group of Deciduata doubtless includes Man. For the human embryo is developed with a genuine decidua, and is thus absolutely distinguished from all the Indeciduata. Among the Deciduata we distinguish two legions, the Zonoplacentalia, with girdle-shaped placenta (Beasts of Prey and Pseudo-hoofed animals), and the Discoplacentalia, with disc-shaped placenta (all the remaining Deciduata). Man possesses a disc-shaped placenta, like all Discoplacentalia; and thus our next question must be, What is man’s position in this group?
In the last chapter we distinguished the following five orders of Discoplacentalia: (1) Semi-apes; (2) Rodents; (3) Insectivora; (4) Bats; (5) Apes. The last of these five orders, that of Apes, is, as every one knows, in every bodily feature far more closely allied to Man than the four others. Hence the only remaining question now is, whether, in the system of animals, Man is to be directly classed in the order of genuine Apes, or whether he is to be considered as the representative of a special sixth order of Discoplacentalia, allied to, but more advanced than, that of the Apes.
Linnæus in his system classed Man in the same order with genuine Apes, Semi-apes, and Bats, which he called Primates; that is, lords, as it were the highest dignitaries of the animal kingdom. But Blumenbach, of Göttingen, separated Man as a special order, under the name of Bimana, or two-handed, and contrasted him with the Apes and Semi-apes under the name of Quadrumana, or four-handed. This classification was also adopted by Cuvier and, consequently, by most subsequent zoologists. It was not until 1863 that Huxley, in his excellent work, the “Evidence as to Man’s Place in Nature,”(26) showed that this classification was based upon erroneous ideas, and that the so-called “four-handed” Apes and Semi-apes are “two-handed” as much as man is himself. The difference between the foot and hand does not consist in the physiological peculiarity that the first digit or thumb is opposable to the four other digits or fingers in the hand, and is not so in the foot, for there are wild tribes of men who can oppose the first or large toe to the other four, just as if it were a thumb. They can therefore use their “grasping foot” as well as a so-called “hinder hand,” like Apes. The Chinese boatmen row with this hinder hand, the Bengal workmen weave with it. The Negro, in whom the big toe is especially strong and freely moveable, when climbing seizes hold of the branches of the trees with it, just like the “four-handed” Apes. Nay, even the newly born children of the most highly developed races of men, during the first months of their life, grasp as easily with the “hinder hand” as with the “fore hand,” and hold a spoon placed in its clutch as firmly with their big toe as with the thumb! On the other hand, among the higher Apes, especially the gorilla, hand and foot are differentiated as in man. (Compare Plate IV.)
The essential difference between hand and foot is therefore not physiological, but morphological, and is determined by the characteristic structure of the bony skeleton and of the muscles attached to it. The ankle-bones differ from the wrist-bones in arrangement, and the foot possesses three special muscles not existing in the hand (a short flexor muscle, a short extensor muscle, and a long fibular muscle). In all these respects, Apes and Semi-apes entirely agree with man, and hence it was quite erroneous to separate him from them as a special order on account of the stronger differentiation of his hand and foot. It is the same also with all the other structural features by means of which it was attempted to distinguish Man from Apes; for example, the relative length of the limbs, the structure of the skull, of the brain, etc. In all these respects, without exception, the differences between Man and the higher Apes are less than the corresponding differences between the higher and the lower Apes. Hence Huxley, for reasons based on the most careful and most accurate anatomical comparisons, arrives at the extremely important conclusion—“Thus, whatever system of organs be studied, the comparison of their modifications in the Ape series leads to one and the same result, that the structural differences which separate Man from the Gorilla and Chimpanzee are not so great as those which separate the Gorilla from the lower Apes.” In accordance with this, Huxley, strictly following the demands of logic, classes Man, Apes, and Semi-apes in a single order, Primates, and divides it into the following seven families, which are of almost equal systematic value: (1) Anthropini (Man); (2) Catarrhini (genuine Apes of the Old World); (3) Platyrrhini (genuine American Apes); (4) Arctopitheci (American clawed Apes); (5) Lemurini (short-footed and long-footed Semi-apes, p. 255); (6) Chiromyini (p. 256); (7) Galeopithecini (Flying Lemurs, p. 256).
| SYSTEMATIC SURVEY | |||||||
| Of the Families and Genera of Apes. | |||||||
| Sections of Apes. |
Families of Apes. |
Genera of Apes. |
Systematic Name of the Genera. |
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| I. APES OF THE NEW WORLD (Hesperopitheci), OR FLAT-NOSED APES (Platyrrhini). | |||||||
| A. Platyrrhini with claws Arctopitheci |
I. Silky apes Hapalida |
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| 1. | Brush ape | 1. | Midas | ||||
| 2. | Lion ape | 2. | Jacchus | ||||
| B. Platyrrhini with blunt nails Dysmopitheci |
II. Flat-nosed, without prehensile tail Aphyocerca |
3. | Squirrel ape | 3. | Chrysothrix | ||
| 4. | Leaping ape | 4. | Callithrix | ||||
| 5. | Nocturnal ape | 5. | Nyctipithecus | ||||
| 6. | Tail ape | 6. | Pithecia | ||||
| III. Flat-nosed, with prehensile tail Labidocerca |
7. | Rolling ape | 7. | Cebus | |||
| 8. | Climbing ape | 8. | Ateles | ||||
| 9. | Woolly ape | 9. | Lagothrix | ||||
| 10. | Howling ape | 10. | Mycetes | ||||
| II. APES OF THE OLD WORLD (Heopitheci), OR NARROW-NOSED APES (Catarrhini). | |||||||
| C. Tailed Catarrhini Menocerca |
IV. Tailed Catarrhini, with cheek-pouches Ascoparea |
11. | Pavian | 11. | Cynocephalus | ||
| 12. | Macaque | 12. | Innus | ||||
| 13. | Sea cat | 13. | Cercopithecus | ||||
| IV. Tailed Catarrhini, without cheek-pouches Anasca |
14. | Holy ape | 14. | Semnopithecus | |||
| 15. | Short ape | 15. | Colobus | ||||
| 16. | Nose ape | 16. | Nasalis | ||||
| D. Tailless Catarrhini Lipocerca |
VI. Human apes Anthropoides |
17. | Gibbon | 17. | Hylobates | ||
| 18. | Orang-Outan | 18. | Satyrus | ||||
| 19. | Chimpanzee | 19. | Engeco | ||||
| 20. | Gorilla | 20. | Gorilla | ||||
| VII. Men Erecti (Anthropi) |
21. | Ape-like man, or speechless man |
21. | Pithecanthropus=(Alalus) | |||
| 22. | Talking man | 22. | Homo | ||||
PEDIGREE OF MEN AND APES
| Straight-haired men Lissotrichi |
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| Woolly-haired men Ulotrichi |
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| Speechless men (Alali), or Ape-like men (Pithecanthropi) |
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| Gorilla Gorilla |
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| Chimpanzee Engeco |
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Orang Satyrus |
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Gibbon Hylobates |
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| African Man-like Apes |
Asiatic Man-like Apes |
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| Man-like Apes Anthropoides |
Nose apes Nasalis |
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| Silk apes Arctopitheci |
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Tall apes Semnopithecus |
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Clutch-tails Labidocerca |
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| Flap-tails Aphyocerca |
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Sea cat Cercopithecus |
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Pavian Cynocephalus |
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│ |
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Tailed Narrow-nosed apes Catarrhina menocerca |
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Narrow-nosed Catarrhini |
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| Apes Simiæ |
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| Semi-apes Prosimiæ |
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If we wish to arrive at a natural system, and consequently at the pedigree of the Primates, we must go a step further still, and entirely separate the Semi-apes, or Prosimiæ, (Huxley’s last three families), from Genuine Apes, or Simiæ (the first four families). For, as I have already shown in my General Morphology, and explained in the last chapter, the Semi-apes differ in many and important respects from Genuine Apes, and in their individual forms are more closely allied to the various other orders of Discoplacentalia. Hence the Semi-apes must probably be considered as the remnants of the common primary group, out of which the other orders of Discoplacentalia, and, it may be, all Deciduata, have developed as two diverging branches. (Gen. Morph. ii. pp. 148 and 153.) But man cannot be separated from the order of Genuine Apes, or Simiæ, as he is in every respect more closely allied to the higher Genuine Apes than the latter are to the lower Genuine Apes.
Genuine Apes (Simiæ) are universally divided into two perfectly natural groups, namely, the Apes of the New World, or American Apes, and the Apes of the Old World, which are indigenous to Asia and Africa, and which formerly also existed in Europe. These two classes differ principally in the formation of the nose, and they have been named accordingly. American Apes have flat noses, so that the nostrils are in front, not below; hence they are called Flat Noses (Platyrrhini). On the other hand, the Apes of the Old World have a narrow cartilaginous bridge, and the nostrils turned downwards, as in man; they are, therefore, called Narrow Noses (Catarrhini). Further, the jaw, which plays an important part in the classification of Mammals, is essentially distinct in these two groups. All Catarrhinæ, or Apes of the Old World, have exactly the same jaws as Man, namely, in each jaw four incisors above and below, then on each side a canine tooth and five cheek teeth, of which two are pre-molars and three molars, altogether thirty-two teeth. But all Apes of the New World, all Platyrrhini, have four more cheek teeth, namely, three pre-molars and three molars on each side, above and below: they consequently possess thirty-six teeth. Only one small group forms an exception to this rule, namely, the Arctopitheci, or Clawed Apes, in whom the third molar has degenerated, and they accordingly have on each half of their jaw three pre-molars and two molars. They also differ from the other Platyrrhini by having claws on the fingers of their hands and the toes of their feet, not nails like Man and the other Apes. This small group of South American Apes, which includes among others the well-known pretty little Midas-monkey and the Jacchus, must probably be considered only as a peculiarly developed lateral branch of the Platyrrhini.
Now, if we ask what evidence can be drawn, as to the pedigree of Apes, from the above facts, we must conclude that all the Apes of the New World have developed out of one tribe, for they all possess the characteristic jaw and the nasal formation of the Platyrrhini. In like manner it follows that all the Apes of the Old World must be derived from one and the same common primary form, which possessed the same formation of nose and jaw as all the still living Catarrhini. Further, it can scarcely be doubted that the Apes of the New World, taken as an entire tribe, are either derived from those of the Old World, or (to express it more vaguely and cautiously) both are diverging branches of one and the same tribe of Apes. We also arrive at the exceedingly important conclusion—which is of the utmost significance in regard to Man’s distribution on the earth’s surface—that Man has developed out of the Catarrhini. For we cannot discover a zoological character distinguishing him in a higher degree from the allied Apes of the Old World than that in which the most divergent forms of this group are distinguished from one another. This is the important result of Huxley’s careful anatomical examination of the question, and it cannot be too highly estimated. The anatomical differences between Man and the most human-like Catarrhini (Orang, Gorilla, Chimpanzee) are in every respect less than the anatomical differences between the latter and the lowest stages of Catarrhini, more especially the Dog-like Baboon. This exceedingly important conclusion is the result of an impartial anatomical comparison of the different forms of Catarrhini.
If, therefore, we recognise the natural system of animals as the guide to our speculations, and establish upon it our pedigree, we must necessarily come to the conclusion that the human race is a small branch of the group of Catarrhini, and has developed out of long since extinct Apes of this group in the Old World. Some adherents of the Theory of Descent have thought that the American races of Men have developed, independently of those of the Old World, out of American Apes. I consider this hypothesis to be quite erroneous, for the complete agreement of all mankind with the Catarrhini, in regard to the characteristic formation of the nose and jaws, distinctly proves that they are of the same origin, and that they developed out of a common root after the Platyrrhini, or American Apes, had already branched off from them. The primæval inhabitants of America, as is proved by numerous ethnographical facts, immigrated from Asia, and partly perhaps from Polynesia (or even from Europe).
There still exist great difficulties in establishing an accurate pedigree of the Human Race; this only can we further assert, that the nearest progenitors of man were tail-less Catarrhini (Lipocerca), resembling the still living Man-like Apes. These evidently developed at a late period out of tailed Catarrhini (Menocerca), the original form of Ape. Of those tail-less Catarrhini, which are now frequently called Man-like Apes, or Anthropoides, there still exist four different genera containing about a dozen different species.
The largest Man-like Ape is the famous Gorilla (called Gorilla engena, or Pongo gorilla), which is indigenous to the tropics of western Africa, and was first discovered by the missionary, Dr. Savage, in 1847, on the banks of the river Gaboon. Its nearest relative is the Chimpanzee (Engeco troglodytes, or Pongo troglodytes), also indigenous to western Africa, but considerably smaller than the Gorilla, which surpasses man in size and strength. The third of the three large Man-like Apes is the Orang, or Orang Outang, indigenous to Borneo and the other Sunda Islands, of which two kindred species have recently been distinguished, namely, the large Orang (Satyrus orang, or Pithecus satyrus) and the small Orang (Satyrus morio, or Pithecus morio). Lastly, there still exists in southern Asia the genus Gibbon (Hylobates), of which from four to eight different species are distinguished. They are considerably smaller than the three first-named Anthropoides, and in most characteristics differ more from Man.
The tail-less Man-like Apes—especially since we have become more intimately acquainted with the Gorilla, and its connection with Man by the application of the Theory of Descent—have excited such universal interest, and called forth such a flood of writings, that there is no occasion for me here to enter into any detail about them. The reader will find their relations to Man fully discussed in the excellent works of Huxley,(26) Carl Vogt,(27) Büchner,(43) and Rolle.(28) I shall therefore confine myself to stating the most important general conclusion resulting from their thorough comparison with Man, namely, that each one of the four Man-like Apes stands nearer to Man in one or several respects than the rest, but that no one of them can in every respect be called absolutely the most like Man. The Orang stands nearest to Man in regard to the formation of the brain, the Chimpanzee in important characteristics in the formation of the skull, the Gorilla in the development of the feet and hands, and, lastly, the Gibbon in the formation of the thorax.
Thus, from a careful examination of the comparative anatomy of the Anthropoides, we obtain a similar result to that obtained by Weisbach, from a statistical classification and a thoughtful comparison of the very numerous and careful measurements which Scherzer and Schwarz made of the different races of Men during their voyage in the Austrian frigate Novara round the earth. Weisbach comprises the final result of his investigations in the following words: “The ape-like characteristics of Man are by no means concentrated in one or another race, but are distributed in particular parts of the body, among the different races, in such a manner that each is endowed with some heirloom of this relationship—one race more so, another less, and even we Europeans cannot claim to be entirely free from evidences of this relationship.”5
I must here also point out, what in fact is self-evident, that not one of all the still living Apes, and consequently not one of the so-called Man-like Apes, can be the progenitor of the Human Race. This opinion, in fact, has never been maintained by thoughtful adherents of the Theory of Descent, but it has been assigned to them by their thoughtless opponents. The Ape-like progenitors of the Human Race are long since extinct. We may possibly still find their fossil bones in the tertiary rocks of southern Asia or Africa. In any case they will, in the zoological system, have to be classed in the group of tail-less Narrow-nosed Apes (Catarrhini Lipocerci, or Anthropoides).
The genealogical hypotheses, to which we have thus far been led by the application of the Theory of Descent to Man, present themselves to every clearly and logically reasoning person as the direct results from the facts of comparative anatomy, ontogeny, and palæontology. Of course our phylogeny can indicate only in a very general way the outlines of the human pedigree. Phylogeny is the more in danger of becoming erroneous the more rigorously it is applied in detail to special animal forms known to us. However, we can, even now, with approximate certainty distinguish at least the following twenty-two stages of the ancestors of Man. Fourteen of these stages belong to the Vertebrata, and eight to the Invertebrate ancestors of Man (Prochordata.)
THE CHAIN OF THE ANIMAL ANCESTORS, OR THE SERIES OF THE PROGENITORS, OF MAN.
(Comp. Ch. XX., XXI.; Plate XIV. and p. 22.)
FIRST HALF OF THE SERIES OF THE ANCESTORS OF MAN.
INVERTEBRATE ANCESTORS OF MAN (Prochordata).
First Stage: Monera.
The most ancient ancestors of Man, as of all other organisms, were living creatures of the simplest kind imaginable, organisms without organs, like the still living Monera. They consisted of simple, homogeneous, structureless and formless little lumps of mucous or albuminous matter (protoplasm), like the still living Protamœba primitiva. (Compare vol. i. p. 186, Fig. 1.) The form value of these most ancient ancestors of man was not even equal to that of a cell, but merely that of a cytod (compare vol. i. p. 347); for, as in the case of all Monera, the little lump of protoplasm did not as yet possess a cell-kernel. The first of these Monera originated in the beginning of the Laurentian period by spontaneous generation, or archigony, out of so-called “inorganic combinations,” namely, out of simple combinations of carbon, oxygen, hydrogen, and nitrogen. The assumption of this spontaneous generation, that is, of a mechanical origin of the first organisms from inorganic matter, has been proved in our thirteenth chapter to be a necessary hypothesis. (Compare vol. i. p. 338.) A direct proof of the earlier existence of this most ancient ancestral stage, based upon the fundamental law of biogeny, is possibly still furnished by the circumstance that, according to the assertions of many investigators, in the beginning of the development of the egg, the cell-kernel, or nucleus, disappears, and the egg-cell thus relapses to the lower stage of the cytod (Monerula, p. 124; relapse of the nucleated plastid into a non-nucleated condition). The assumption of this first stage is necessary for most important general reasons.