The three other classes of Echinoderma evidently arose at a later period out of the class of Sea-stars which have most faithfully retained the original form of the stellate colony of worms. The Sea-lilies, or Crinoida, differ least from them, but having given up the free, slow motion possessed by other Sea-stars, they have become adherent to rocks, etc., and form for themselves a long stalk. Some Encrinites, however (for example, the Comatulæ, Fig. B, on Plates VIII. and IX.), afterwards detach themselves from their stalk. The original worm individuals in the Crinoida are indeed no longer preserved in the same independent condition as in the case of the common star-fish; but they nevertheless always possess articulated arms extending from a common central disc. Hence we may unite the Sea-lilies and Sea-stars into a main-class, or branch, characterized as possessing articulated arms (Colobrachia).

In the other two classes of Echinoderma, the Sea-urchins and Sea-cucumbers, the articulated arms are no longer present as independent parts, but, by the increased centralization of the stock, have completely fused so as to form a common, inflated, central disc, which now looks like a simple box or capsule without arms. The original stock of five individuals has apparently degenerated to the form-value of a simple individual, a single person. Hence we may represent these two classes as a branch characterized as being without arms (Lipobrachia), equivalent to those which possess articulated arms. The first of these two classes, that of Sea-urchins (Echinida) takes its name from the numerous and frequently very large thorns which cover the hard shell, which is itself artistically built up of calcareous plates. (Fig. C, Plates VIII. and IX.) The fundamental form of the shell itself is a pentagonal pyramid. The Sea-urchins probably developed directly out of the group of Sea-stars. The different classes and orders of marine lilies and stars which are given in the following table, illustrate the laws of progress and differentiation in a striking manner. In each succeeding period of the earth’s history we see the individual classes continually increasing in variety and perfection. (Gen. Morph. ii. Plate IV.)

The history of three of these classes of Star-fish is very minutely recorded by numerous and excellently preserved fossils, but on the other hand, we know almost nothing of the historical development of the fourth class, that of the Sea-cucumbers (Holothuriæ). These curious sausage-shaped Star-fish manifest externally a deceptive similarity to worms. (Fig. D, Plates VIII. and IX.) The skeletal structures in their skin are very imperfect, and hence no distinct remains of their elongated, cylindrical, worm-like body could be preserved in a fossil state. However, from the comparative anatomy of the Holothuriæ, we can infer that they have arisen, by the softening of the cutaneous skeleton, from members of the class of Sea-urchins.

From the Star-fish we turn to the fifth and most highly developed tribe of the invertebrate animals, namely, the phylum of Articulata, or those with jointed feet (Arthropoda). As has already been remarked, this tribe corresponds to Linnæus’ class of Insects. It contains four classes: (1) the genuine six-legged Insects, or Flies; (2) the eight-legged Spiders; (3) the Centipedes, with numerous pairs of legs; and (4) the Crabs, or Crustacea, whose legs vary in number. The last class breathe water through gills, and may therefore be contrasted as the main-class of gill-breathing Arthropoda, or Gilled Insects (Carides), with the three first classes. The latter breathe air by means of peculiar wind-pipes, or tracheæ, and may therefore appropriately be united to form the main-class of the trachea-breathing Arthropoda, or Tracheate Insects (Tracheata).

In all animals with articulated feet, as the name indicates, the legs are distinctly articulated, and by this, as well as by the strong differentiation of the separate parts of the body, or metamera, they are sharply distinguished from Ringed worms, with which Bär and Cuvier classed them. They are, however, in every respect so like the Ringed worms that they can scarcely be considered altogether distinct from them. They, like the Ringed worms, possess a very characteristic form of the central nervous system, the so-called ventral marrow, which commences in a gullet-ring encircling the mouth. From other facts also, it is evident that the Arthropoda developed at a late period out of articulated worms. Probably either the Wheel Animalcules or the Ringed worms are their nearest blood relations in the Worm tribe. (Gen. Morph. ii. Plate V. pp. 85-102.)

Now, although the derivation of the Arthropoda from ringed Worms may be considered as certain, still it cannot with equal assurance be maintained that the whole tribe of the former has arisen out of one branch of the latter. For several reasons seem to support the supposition that the Gilled Arthropods have developed out of a branch of articulated worms, different from that which gave rise to the Tracheate Arthropods. But on the whole it remains more probable that both main-classes have arisen out of one and the same group of Worms. In this case the Tracheate Insects—Spiders, Flies, and Centipedes—must have branched off at a later period from the gill-breathing Insects, or Crustacea.

The pedigree of the Arthropoda can on the whole be clearly made out from the palæontology, comparative anatomy, and ontogeny of its four classes, although here, as everywhere else, many details remain very obscure. Not until the history of the individual development of all the different groups has become more accurately known than it is at present, can this obscurity be removed. The history of the class of Gilled Insects, or Crabs (Carides), is at present that best known to us; they are also called encrusted animals (Crustacea), on account of the hard crust or covering of their body. The ontogeny of these animals is extremely interesting and, like that of Vertebrate animals, distinctly reveals the essential outlines of the history of their tribe, that is, their phylogeny. Fritz Müller, in his work, “Für Darwin,”(16) which has already been referred to, has explained this remarkable series of facts in a very able manner.

The common primary form of all Crabs, which in most cases is even now the first to develop out of the egg, is originally one and the same, the so-called Nauplius. This remarkable primæval crab represents a very simple form of articulated animal, the body of which in general has the form of a roundish, oval, or pear-shaped disc, and has on its ventral side only three pairs of legs. The first of these is uncloven, the two subsequent pairs are forked. In front, above the mouth, lies a simple, single eye. Although the different orders of the Crustacean class differ very widely from one another in the structure of their body and its appendages, yet the early Nauplius form always remains essentially the same. In order to be convinced of this, let the reader look attentively at Plates X. and XI., a more detailed explanation of which is given in the Appendix. On Plate XI. we see the fully developed representatives of six different orders of Crabs, a Leaf-footed Crab (Limnetis, Fig. A c); a Stalked Crab (Lepas, Fig. D c); a Root Crab, (Sacculina, Fig. E c); a Boatman Crab (Cyclops, Fig. B c); a Fish Louse (Lernæocera, Fig. C c); and, lastly, a highly developed Shrimp (Peneus, Fig. F c). These six crabs vary very much, as we see, in the entire form of body, in the number and formation of the legs, etc. When, however, we look at the earliest stages, or “nauplius,” of these six different classes, after they have crept out of the egg—those marked with corresponding letters on Plate X. (Fig. A n-F n)—we shall be surprised to find how much they agree. The different forms of Nauplius of these six orders differ no more from one another than would six different “good species” of one genus. Consequently, we may with assurance infer a common derivation of all those orders from a common Primæval Crab, which was essentially like the Nauplius of the present day.

PEDIGREE OF GILLED INSECTS.

The pedigree on p. 177 will show how we may at present approximately conceive the derivation of the twenty orders of Crustacea enumerated on p. 176, from the common primary form of the Nauplius. Out of the Nauplius form—which originally existed as an independent genus—the five legions of lower Crabs developed as diverging branches in different directions, which in the systematic survey of the class are united as Segmented Crabs (Entomostraca). The higher division of Mailed Crabs (Malacostraca) have likewise originated out of the common Nauplius form. The Nebalia is still a direct form of transition from the Phyllopods to the Schizopods, that is, to the primary form of the stalk-eyed and sessile-eyed Mailed Crabs. The Nauplius at this stage gives rise to another larva form, the so-called Zoëa, which is of great importance. The order of Schizopoda, those with cloven feet (Mysis, etc.), probably originated from this curious Zoëa; they are at present still directly allied, through the Nebalia to the Phyllopoda, those with foliaceous feet. But of all living crabs the Phyllopods are the most closely allied to the original primary form of the Nauplius. Out of the Schizopoda the stalk-eyed and sessile-eyed Mailed Crabs, or Malacostraca, developed as two diverging branches in different directions: the former through shrimps (Peneus, etc.), the latter through the Cumacea (Cuma, etc.), which are still living and closely allied to the Schizopoda. Among those with stalked eyes is the river crab (cray-fish), the lobster, and the others with long tails, or the Macrura, out of which, in the chalk period, the short-tailed crabs, or Brachyura, developed by the degeneration of the tail. Those with sessile eyes divide into the two branches of Flea-crabs (Amphipoda) and Louse-crabs (Isopoda); among the latter are our common Rock-slaters and Wood-lice.

The second main-class of Articulated animals, that of the Tracheata, or air-breathing Tracheate Insects4 (Spiders, Centipedes, and Flies) did not develop until the beginning of the palæolithic era, after the close of the archilithic period, because all these animals (in contrast with the aquatic crabs) are originally inhabitants of land. It is evident that the Tracheata can have developed only after the lapse of the Silurian period when terrestrial life first began. But as fossil remains of spiders and insects have been found, even in the carboniferous beds, we can pretty accurately determine the time of their origin. The development of the first Tracheate Insects out of gill-bearing Zoëa-crabs, must have taken place between the end of the Silurian and the beginning of the coal period, that is, in the Devonian period.

Gegenbaur, in his excellent “Outlines of Comparative Anatomy,”(21) has lately endeavoured to explain the origin of the Tracheata by an ingenious hypothesis. The system of tracheæ, or air pipes, and the modifications of organization dependent upon it, distinguish Flies, Centipedes, and Spiders so much from other animals, that the conception of its first origin presents no inconsiderable difficulties to phylogeny. According to Gegenbaur, of all living Tracheate Insects, the Primæval Flies, or Archiptera, are most closely allied to the common primary form of the Tracheata. These insects—among which we may especially mention the delicate Day flies (Ephemera), and the agile dragon-flies (Libellula)—in their earliest youth, as larvæ, frequently possess external tracheate gills which lie in two rows on the back of the body, and are shaped like a leaf or paint-brush. Similar leaf or paint-brush shaped organs are met with as real water-breathing organs or gills, in many crabs and ringed worms, and, moreover, in the latter as real dorsal appendages or limbs. The “tracheate gills,” found in the larvæ of many primæval winged insects, must in all probability be explained as “dorsal limbs,” and as having developed out of the corresponding appendages of the Annelida, or possibly as having really arisen out of similar parts in Crustacea long since extinct. The present tracheal respiration of the Tracheata developed at a later period out of respiration through the “tracheate gills.” The tracheate gills themselves, however, have in some cases disappeared, and in others become transformed into the wings of the Flies. They have disappeared entirely in the classes of Spiders and Centipedes, and these groups must accordingly be conceived of as degenerated or peculiarly developed lateral branches of the Fly class, which at an early period branched off from the common primary form of Flies; Spiders probably did so at an earlier period than Centipedes. Whether that common primary form of all Tracheata, which in my General Morphology I have named Protracheata, did develop directly out of genuine Ringed worms, or at first out of Crustacea of the Zoëa form (Zoëpoda, p. 212) will probably be settled at some future time by a more accurate knowledge and comparison of the ontogeny of the Tracheata, Crustacea, and Annelida. However, the root of the Tracheata, as well as that of the Crustacea, must in any case be looked for in the group of Ringed worms.

The genuine Spiders (Arachnida) are distinguished from Flies by the absence of wings, and by four pairs of legs; but, as is distinctly seen in the Scorpion-spiders and Tarantulæ, they, like Flies, possess in reality only three pairs of genuine legs. The apparent “fourth pair of legs” in spiders (the foremost) are in reality a pair of feelers. Among the still existing Spiders, there is a small group which is probably very closely allied to the common primary form of the whole class; this is the order of Scorpion-spiders, or Solifugæ, (Solpuga, Galeodes), of which several large species live in Africa and Asia, and are dreaded on account of their poisonous bite. Their body consists—as we suppose to have been the case in the common ancestor of the Tracheata—of a head possessing several pairs of feelers like legs, of a thorax, to the three rings of which are attached three pairs of legs, and of a hinder, body, or abdomen, consisting of many distinct rings. In the articulation of their body, the Solifugæ are therefore in reality more closely related to flies than to other spiders. Out of the Devonian Primæval Spiders, which were nearly related to the Solifugæ of the present day, the Long Spiders, the Tailor Spiders, and the Round Spiders probably developed as three diverging branches.

The Long Spiders (Arthrogastres), in which the earlier articulation of body has been better preserved than in Round Spiders, appear to be the older and more original forms. The most important members of this sub-class are the scorpions, which are connected with the Solifugæ through the Tarantella (or Phrynidæ). The small book scorpions, which inhabit our libraries and herbariums, appear as a degenerate lateral branch from the true scorpions. Mid-way between the Scorpions and Round Spiders are the long-legged Tailor-spiders (Opiliones) which have possibly arisen out of a special branch of the Solifugæ. The Pycnogonida, or No-body Crabs, and the Arctisca, or Bear Worms—still generally included among Long Spiders—must be completely excluded from the class of Spiders; the former belong to the Crustacea, the latter to Ringed worms.

PEDIGREE OF TRACHEATA

Fossil remains of Long Spiders are found in the Coal. The second sub-class of the Arachnida, the Round Spiders (Sphærogastres), first appear in the fossil state in the Jura, that is, at a very much later period. They have developed out of a branch of the Solifuga, by the rings of the body becoming more and more united with one another. In the true Spinning Spiders (Araneæ), which we admire on account of their delicate skill in weaving, the union of the joints of the trunk, or metamera, goes so far, that the trunk now consists of only two pieces, of a head-breast (cephalo-thorax) with jaws, feelers, and four pairs of legs, and of a hinder body without appendages, where the spinning warts are placed. In Mites (Acarida), which have probably arisen by degeneration (especially by parasitism) out of a lateral branch of Spinning Spiders, even these two trunk pieces have become united and now form an unsegmented mass.

The class of Scolopendria, Myriapoda, or Centipedes, the smallest and poorest in forms of the four classes of Arthropoda, is characterized by a very elongated body, like that of a segmented Ringed worm, and often possesses more than a hundred pairs of legs. But these animals also originally developed out of a six-legged form of Tracheata, as is distinctly proved by the individual development of the millipede in the egg. Their embryos have at first only three pairs of legs, like genuine insects, and only at a later period do the posterior pairs of legs bud, one by one, from the growing rings of the hinder body. Of the two orders of Centipedes (which in our country live under barks of trees, in moss, etc.) the round, double-footed ones (Diplopoda) probably did not develop until a later period out of the older flat, single-footed ones (Chilopoda), by successive pairs of rings of the body uniting together. Fossil remains of the Chilopoda are first met with in the Jura period.

The third and last class of the Arthropoda breathing through tracheæ, is that of the Flies, or Insects, in the narrow sense of the word (Insecta, or Hexapoda), the largest of all classes of animals, and next to that of Mammalia, also the most important. Although Flies develop a greater variety of genera and species than all other animals taken together, yet these are all in reality only superficial variations of a single type, which is entirely and constantly preserved in its essential characteristics. In all Flies the three divisions of the trunk—head, breast (thorax), and hinder body—are quite distinct. The hinder body, or abdomen, as in the case of spiders, has no articulated appendages. The central division, the breast or thorax, has on its ventral side three pairs of legs, on its back two pairs of wings. It is true that, in very many Flies, one or both pairs of wings have become reduced in size or have even entirely disappeared; but the comparative anatomy of Flies distinctly shows that this deficiency has arisen only gradually by the degeneration of the wings, and that all the Flies existing at present are derived from a common, primary Fly, which possessed three pairs of legs and two pairs of wings. (Compare p. 256.) These wings, which so strikingly distinguish Flies from all other Arthropoda, probably arose, as has been already shown, out of the tracheate gills which may still be observed in the larvæ of the ephemeral flies (Ephemera) which live in water.

The head of Flies universally possesses, besides the eyes, a pair of articulated feelers, or antennæ, and also three jaws upon each side of the mouth. These three pairs of jaws, although they have arisen in all Flies from the same original basis, by different kinds of adaptation, have become changed to very varied and remarkable forms in the various orders, and are therefore employed for distinguishing and characterizing the main divisions of the class. In the first place, we may distinguish two main divisions, namely, Flies with chewing mandibles (Masticantia) and Flies with sucking mouths (Sugentia). On a closer examination each of these two divisions may again be divided into two sub-groups. Among chewing Flies, or Masticantia, we may distinguish the biting and the licking ones. Biting flies (Mordentia) comprise the most ancient and primæval winged Flies, the gauzy-winged (Neuroptera), straight-winged (Orthoptera), and beetles (Coleoptera). Licking flies (Lambentia) are represented by the one order of skin-winged (Hymenoptera) Flies. We distinguish two groups of Sucking Flies, or Sugentia, namely, those which prick and those which sip. There are two orders of pricking Flies (Pungentia), those with half wings (Hemiptera) and gnats and blow-flies (Diptera); butterflies are the only sipping Flies (Sorbentia), Lepidoptera.

Biting Flies, and indeed the order of Primæval Flies (Archiptera, or Pseudoneuroptera) are nearest akin to the still living Flies, and include the most ancient of all Flies, the primary forms of the whole class (hence also those of all Tracheata). Among them are, first of all, the Ephemeral Flies (Ephemera) whose larvæ which live in water, in all probability still show us in their tracheæ-gills the organs out of which the wings of Flies were originally developed. This order further contains the well known dragon-flies, or Libellula, the wine-glass sugar mites (Lepisma), the hopping Flies with bladder-like feet (Physopoda), and the dreaded Termites, fossil remains of which are found even in coal. The order of Gauze-winged Flies (Neuroptera), probably developed directly out of the primæval Flies, which differ from them only by their perfect series of transformations. Among them are the gauze-flies (Planipennia), caddis-flies (Phryganida), and fan-flies (Strepsiptera). Fossil Flies, which form the transition from the primæval Flies (Libellula) to the gauze-winged (Sialidæ), are found even in coal (Dictyophylebia).

The order of Straight-winged Flies (Orthoptera) developed at an early period out of another branch of the primæval Flies by differentiation of the two pairs of wings. This division is composed of one group with a great variety of forms—cockroaches, grasshoppers, crickets, etc. (Ulonata)—and of a smaller group consisting only of the well-known earwigs (Labidura), which are characterised by nippers at the hinder end of their bodies. Fossil remains of cockroaches, as well as of crickets and grasshoppers, have been found in coal.

Fossil remains of the fourth order of Biting Flies, beetles (Coleoptera) likewise occur in coal. This extremely comprehensive order—the favourite one of amateurs and collectors—shows more clearly than any other what infinite variety of forms can be developed externally by adaptation to different conditions of life, without the internal structure and the original form of the body being in any way essentially changed. Beetles have probably developed out of a branch of the straight-winged Flies, from which they differ only in their transformations (larva, pupa, etc.).

The one order of Licking Flies, namely, the interesting group of the Bees, or Skin-winged Flies (Hymenoptera), is closely allied to the four orders of biting Flies. Among them are those Flies which have risen to such an astonishing degree of mental development, of intellectual perfection, and strength of character, by their extensive division of labour, formation of communities and states, and surpass in this not merely most invertebrate animals, but even most animals in general. This may be said especially of all ants and bees, also of wasps, leaf-wasps, wood-wasps, gall-wasps, etc. They are first met with in a fossil state in the oolites, but they do not appear in greater numbers until the tertiary period. Probably these insects developed either out of a branch of the primæval Flies or the gauze-winged Flies.

Of the two orders of Pricking Flies (Hemiptera and Diptera), that containing the Half-winged Flies (Hemiptera), also called Beaked Flies (Rhynchota), is the older of the two. It includes three sub-orders, viz., the leaf-lice (Homoptera), the bugs (Heteroptera), and lice (Pediculina). Fossil remains of the first two classes are found in the oolites; but an ancient Fly (Eugereon) is found in the Permian system, and seems to indicate the derivation of the Hemiptera from the Neuroptera. Probably the most ancient of the three sub-orders of the Hemiptera are the Homoptera, among which, besides the actual leaf-lice, are the shield-lice, leaf-fleas, and leaf-crickets, or Cicadæ. Lice have probably developed out of two different branches of Homoptera, by continued degeneration (especially by the loss of wings); bugs, on the other hand, by the perfecting and differentiation of the two pairs of wings.

The second order of pricking flies, namely, the Two-winged Flies (Diptera), are also found in a fossil state in the oolites, together with Half-winged Flies; but they probably developed out of the Hemiptera by the degeneration of the hind wings. In Diptera the fore wings alone have remained perfect. The principal portion of this order consists of the elongated gnats (Nemocera) and of the compact blow-flies and house-flies (Brachycera), the former of which are probably the older of the two. However, remains of both are found in the oolitic period. The two small groups of lice-flies (Pupipara) forming chrysales, and the hopping-fleas (Aphaniptera), probably developed out of the Diptera by degeneration resulting from parasitism.

The eighth and last order of Flies, and at the same time the only one with mouth-parts adapted to sipping liquids, consists of moths and butterflies (Lepidoptera). This order appears, in several morphological respects, to be the most perfect class of Flies, and accordingly was the last to develop. For we only know of fossil remains of this order from the tertiary period, whereas the three preceding orders extend back to the oolites, and the four biting orders even to the coal period. The close relationship between some moths (Tineæ) and (Noctuæ), and some caddis-flies (Phryganida) renders it probable that butterflies have developed from this group, that is, out of the order of Gauze-winged Flies, or Neuroptera.

The whole history of Flies, and, moreover, the history of the whole tribe of Arthropoda, essentially confirms the great laws of differentiation and perfecting which, according to Darwin’s theory of selection, must be considered as the necessary results of Natural Selection. The whole tribe, so rich in forms, begins in the Archilithic period with the class of Crabs breathing by gills, and with the lowest Primæval Crabs, or Archicaridæ. The form of these Primæval Crabs, which were developed out of segmented worms, is still approximately preserved by the remarkable Nauplius, in the common larval stage of so many Crabs. Out of the Nauplius, at a later period, the curious Zoëa was developed, which is the common larval form of all the higher or mailed crabs (Malacostraca), and, at the same time, possibly of that Arthopod which at first breathed through tracheæ, and became the common ancestor of all Tracheata. This Devonian ancestor, which must have originated between the end of the Silurian and the beginning of the Coal period, was probably most closely related to the still living Primæval Flies, or Archiptera. Out of these there developed, as the main tribe of the Tracheata, the class of Flies, from the lowest stage of which the spiders and centipedes separated as two diverging branches. Throughout a long period there existed only the four biting orders of Flies—the Primæval flies, Gauze-wings, Straight-wings, and the Beetles, the first of which is probably the common primary form of the three others. It was only at a much later period that the Licking, Pricking, and Sipping flies developed out of the Biting ones, which retained the original form of the three pairs of jaws most distinctly. The following table will show once more how these orders succeeded one another in the history of the earth.