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THE INTERNATIONAL SCIENTIFIC SERIES EYOLUTIOJ^J^ BY ATROPHY IN BIOLOGY AND SOCIOLOGY BY JEA]Sr pEMOOE AOR^Qfi OF THE FREE UNIVERSITY OF BRUSSELS JEAN MASSART CHARO& DS COCKS OF THE FREE UNIVESSITY OF BRUSSELS EMILE VANDERVELDE PROFESSOR AT THE INSTITUTE OF HAUTES fiTUDES OF BRUSSELS TRANSLATED BY MES. CHALMERS MITCHELL NEW YORK D. applet'on and company 1899 PEEFACE This treatise, compiled in connection with a scheme for research work in general sociology, elaborated in June 1894, was presented to the Institute of Sociology at Brussels. In drawing up the pro- gramme of the Institute, the founder, M. Ernest Solvay, after having mentioned the questions which especially called for the investigation of his col- leagues, added the following statement : — " The Institute of Sociology will take part in the labours of the modern school of Sociology, the object of which is to ascertain the normal conditions under which societies exist, and the laws which govern their evolution. But the advances of Natural Science in this century have not yet been sufficiently assimilated by those Sciences most closely related to it, and it is from such assimilation that the most important additions to knowledge may be expected." In stating that the results of Natural Science have not been sufficiently assimilated by Sociology, M. Solvay is only apparently at variance with those who rightly protest against exaggerated and vi PKEFACE premature comparisons between social organizations and animal or vegetable organisms. The existence of such exaggerations, which have caused a reaction such as recently induced an emi- nent American economist to declare the bankruptcy of biological sociology, is perhaps due to the fact that, with a few distinguished exceptions, bio- sociological investigations have hitherto been con- ducted either by naturalists with a limited know- ledge of social questions, or by sociologists whose training in biology was incomplete and superficial. To prevent this danger, our researches in the same subject have been made separately from the social side, and from the biological side, and have now been co-ordinated and combined. This work was commenced in May 1893, with the collaboration of our friend M. Dollo, the curator of the Natural History Museum at Brussels. In June 1894, however, M. Dollo's many occupations no longer permitted of his collaboration. The zoo- logical part was therefore completed by M. Jean Demoor, to whom most of the facts quoted in the first book were given by M. Dollo, whose assistance we most gratefully acknowledge. CONTENTS Prepacb PAGE V INTRODUCTION Societies and organisms. 2. Individuals, colonies and societies. 3. Communities and societies. 4. Distinc- tive characters of societies of which the members are united by social contract .... BOOK I UNIVERSALITY OF DEGENERATIVE EVOLUTION Part I. Degeneration in the development of institutions and organs ...... 21 Chapter I. In the evolution of organs all modification is necessarily attended by degeneration ... 22 § 1. Preliminary considerations .... 22 (1) Original formation of organs in ancestral forms , 23 (2) Development of organs in the individual . . 23 (3) Philogeuetio evolution of function . . 25 (4) Individual adaptation .... 26 Section I. Transformation of orgaus of animals . 30 § 2. Transformation of homortynamic organs in the indivi- dual (metameric appendages) , , . 30 Theoray-fish ...... 31 § 3. Transformation of homologous organs in individuals of different species (limbs) .... 41 Ijimbs adapted to an aquatic life (Ceratodus, OrtJia- canthuSj Protoptemis amphiHuSj Protojpterus annec- tem, Lepridouren ..... 43 Limbs adapted to a terrestrial life ... 43 viii CONTENTS PAGR 1. Adaptation to walking on two legs (man, birds). 2. Adaptation to leaping (Dipus agyp- tius, kangaroo, Tarsiua epectrum, Rana eaculenta, 3. Adaptation to running (horse, ruminants). 4. Adaptation to flight (birds, Pterosaurians, bats). 6. Adaptation to arboreal life {Arcfocelms cala- laremis, Chamceleo). 6. Adaptation to swimming (Cetaceans, Sirenia). 7. Adaptation to burrowing (Talpa curopcea, Heterocephalw) ... 46 Section II. Modification of the organs of plants . 68 § 4. Modification of hemodynamic organs in the indivi- dual (basilar and apical leaves) ... 68 1. Rosa rugosa. 2. Serrainla centauroides. 3. Sagittaria sagittifolia. 4. Lathyra Aphaca. 5. Nymphaea dentata ..... 70 § 5. Modification of organs which are homologous in individuals of different species (foliage leaves) . 78 1. Adaptation to climbing {Cobaeea scandals, Vicia pyrenaica, Cummis satims). 2. Adaptation to carnivorous nutrition (Utncularia, Nepenthes, Drosera). 3. Adaptation to an aquatic life (Sagil- taria, Nymphaea, Vallisneria, Potamogeton, Ranun- culus, Ouvirandra fenestralis). 4. Adaptation to defence against ants (Acacia sphae/rocephaUi), 5. Adaptation to drought (Sempendman). 6. Adapta- tion to defence against herbivorous animals (Garagana, Ilex, Mamillaria, etc.) . . 78 Chapter II. In the evolution of institutions all modification is necessarily accompanied by degeneration . . 90 § 1. Modifications of similar institutions in the same society ...... 91 (1) The communal budgets of Belgium . , 92 (2) Budget of the States of the German Empire , 95 (3) The budgets of Germany, France, and England . 97 g 2. Modification of similar institutions in different social groups (the development of landed property) , 98 1. Family property (Montenegro). 2. Village pro- perty (EuBsia). 3. Feudal property (England). CONTENTS ix FAGE 4. Public or collective property (Switzerland). 6. Corporative property (Belgium). 6. Private property (Switzerland). 7. Summary . . 100 Part II. Degeneration in the evolution of orgauisms and societies ....... 115 Chapter 1. All organisms exhibit rudimentary organs , 115 Section I. Budimentary organs of animals . . 117 § 1. Budimentary organs in man .... 117 1. Integumentary system. 2. Skeleton. 3. Mus- cular system. 4. Nervous system. 5. Digestive system. 6. Vascular system. 7. Sense organs. 8. Genito-urinary system . . . 117 S 2. Budimentary organs in various groups . . 121 1. Coelenterates. 2. Worms. 3. Eohinoderms. 4, MoUusca. 5. Arthropodes. 6. Vertebrates . 121 Section II. Budimentary organs in plants . . 145 1 3. Budimentary organs in various groups of plants . 145 1. Algse. 2. Mushrooms. 3. Bryophyta. 4. Pteri- dophyta. 5. Phanerogams , , . . 145 § 4. Beduced organs in the vegetative apparatus of the Phanerogams . . . . . . 149 Chapter II. Survivals exist in all kinds of societies . . 151 § 1 . Instances of svirvival in various groups . . 153 (1) Instances of survival in the most modern social groups (the United States) .... 155 (2) Instances of survival in less civilized social groups (Veddahs, Fuegoes, Australian tribes . . 156 § 2. Survivals of ancient forms of marriage, and of the family in Modern Europe . . . . 161 1. Forvis o/ marriage. — (1) Marriage by capture. (2) Marriage by purchase. (3) Marriage by consent of both parties (marriage by simple consent, mar- riage in facie Mcclesite) .... 161 2. The Family System.— {I) Matriarchy. (2) Patri- archy ....... 167 Part III. Summary and conclusions .... 170 175 178 179 179 X CONTENTS BOOK II THE PATH OF DEaENERATIVE EVOLUTION Part I. The supposed law that degeneration retraces the steps of progress ..... diopter I, The path of degeneration in biology Section, I. The path of degeneration in animals 1. Morphology and embryology ; the law of recapitu- lation. 2. Degeneration of the third eye in lizards. 3. Degeneration of the organs of sight in deep-sea Crustacea. 4. Atrophy of the branchial Tessels in man ....... Section II. The path of degeneration in plants. . 192 1. Rarity of cases of recapitulation in the organ- ogeny of leaves (Vicia, Acacia with phyllodes). 2. Organogeny of flowers {Brassica oleracea var. Botrgtis). 3. Progressive degeneration of the pro- thallus in phanerogams .... 192 Chapter II. The path of degeneration in sociology . . 205 § 1. Investigation of facts ..... 205 1. Tithings, hundreds and counties in England. 2. Order of elimination of various racial elements in a country. 3. The degenerative evolution of polir tical organizations. 4. Degeneration in monetary systems. 5. Degenerative adaptation in colonial' legislation. 6. Degenerative evolution of the cor- porations of Western Flanders . . . 207 § 2. A criticism of the supposed inverse path of degenera- tion ....... 217 Pakt II. The irreversibility of degenerative evolution. . 221 Chapter I. Do institutions or organs which have disappeared reappear? . . " . . . . 222 Section I. Disappeared organs .... 222 1. Plants ...... 222 2. Animals. Teratology of the horse, Hypertrichosis, etc. Swimming limbs in Stomatopoda and De- capoda Maoroura ..... 223 Section II. Disappeared institutions . , , 227 CONTENTS XI (1) Apparent revival of bygone institutions . . (2) Apparent disappearance of institutions . (3) Instances of convergence .... Chapter II. Can rudimentary institutions or organs reassume their primitive functions ? .... Seditm 1. Rudimentary organs 1. AnvmaXs. (1) Muscles of the ear in man. (2) The abdomen and appendages in deep-sea hermit crabs 2. Plants. (1) Hermaphrodite flowers in Mdandryiim. (2) Branches of Colletia cruQiata, Crataegus, Vida Fata, Sem/pereiimm, Veronica, etc. . Section II. Eudimentary institutions . (1) Truck system and clearing-house. (2) Corpora- tions and syndicates. (3) Archaic collectivism and modern collectivism. (4) The survival of elective sovereignty in England Chapter III. Can rudimentary organs or institutions re- develop and assume new functions ? Section I. Eudimentary organs 1. Animals. Eespiratory organs in Birgm latro Mesonephric spaces in the higher vertebrates 2. Plants. Stamiaodes of Pentstemon Section II. Eudimentary institutions . Zevirat ...... Part III. Summary and conclusions . . PAOE 227 229 230 2.32 232 232 233 237 239 242 243 243 244 245 245 247 BOOK III CAUSES OF DEGENERATIVE EVOLDTION Part I. Atrophy of organs and institutions . Section I. The factors of atrophy . (1) Biology (accidental, individual, normal and specific atrophy). (2) Sociology (accidental and nuruial atrophy) ..... Section II. Causes producing atrophy Chapter I. Atrophy of organs .... § 1. Atrophy from lack of space , . , 251 251 251 260 260 261 xii CONTENTS FAOI (1) DeTelopment of the teotli. (2) Atrophy of the superior glume (Loliwm). (3) Degeneration of Palese (composite flowers) and of stamens (ScrophulAriacese) .... 261 § 2, Atrophy from lack of use .... 263 1, Functional inutility : (1) Etiolated plants and immobile limbs. (2) Epicotyl and primary leaf of Nymphaea. (3) iSoots of Utrieularia ; coty- ledons of parasitic plants {Cnseata, Ordbanche) ; leaves transformed to spines in Phyllocactus crenMus. (4) Atrophy of the branchial arches in mammals. (5) Atrophy of ventral hns^PediciUatif Protopterus), (6) Atrophy of muscles {Cetacea^ Sirenia). (7) Atrophy of the tail in man. (8) Degeneration of the hyuid apparatus in man and birds. ..... 263 2. Transference of function : (1) Atrophy of the tail in Batraehia anwra, and the larval gills in some insects. (2) Disappearance of limbs (Slow- worms, Ampbisbaena, Snakes, Eels, and Saccu- lina). . (3) Atrophy of the leaf (acada with phyl- \oiea, Xylophylla). (4) Atrophyof theprotonema in mosses, and of the leaves in some xerophilus plants (MnrMetibechia plafyclados, Oenista, Spartivm, Alhagi). (5) Disappearance of the calyx. (6) Atrophy of roots (Pine, Beech, Corallorkyza, Myrmechii, Tillandsia itmeoides), or of leaves and stems {Tcmiophyllum Zollin- geri, Podostemacece) .... 268 § 3. Atrophy from lack of nutrition . . . 274 1. Parasitic castration (Melandryum album). 2. Severe or prolonged compression of a limb. 3. Atrophy of the genitalia in neuter bees. 4. Atrophy of the superior flowers in Carex. 6. Atrophy of pistils and stamens (Ffitillaria peraica, Vibumam tomeniomm, Vibm-nam Opulm) 274 § 4. Atrophy without apparent cause . . . 278 Atrophyof perianth (i4rtemi9!a, Poferium). Atrophy of the eyes (Myriopodes, Cymotlioe). Correlative atrophy 278 CONTENTS PAGE 281 282 Chapter II. Atrophy of institutions .... § 1. Atrophy from lack of use .... 1. Functional intiiility : (1) Offices of the port of Bruges. (2) The forest-courts of England . 282 2. Transference of fuTiction : (1) Republican institu- tions under the Roman Empire. (2) Special jurisdiction in England .... 284 § 2. Atrophy from lack of resource . . . 287 1. Local administration at the close of the Eoman Empire. 2. Degeneration of societies in general 287 Pabt II. Causes of the persistence of organs or institutions without function ...... 292 Chapter I. Survival of organs ..... 292 § 1. Unfunotional organs that are not rudimentary . 292 Absence of variation (flowers of Ficaria ranmieu- loides, LysiTnachia Swrnwidaria, /Uodeacanaderms, straliotes aloides, cleistogamous flowers, eyes in the male Machaeritea .... 292 § 2. Unfunotional organs which persist as rudiments . 295 1. Absence of variation. Insignificance miral seg- COmplctC appendage. ilie ment. Fin8{l,5/1) : ?r, vrotopodite; . . ,, . , «a:,exopodlte; en,endopoditeCUu.\luy).J)9-050pOaite {pr.) IS tulCK and short and has no gill, degeneration being ex- hibited in the loss of the gill, and by the reduced length of the part. The eocopodite (ex.), and the enciopodite {en.) are modified into two large oval plates which serve as propellers. The actual de- velopment of these two parts is accompanied by degeneration. In a typical appendage, the eocopodite and the endopodite terminate in slender parts divided into several rings by false joints ; in the propellers, which should offer the maximum resistance to the TEA^fSFOKMATI0N OF OEGANS OF ANIMALS 33 pressure of the water, the segmentation has dis- appeared, but across the part corresponding to the exopodite there still remains a transverse groove. The second abdominal segment of the female carries appendages similar to those which have just been described. In the male, the organs of this segment, as also those of the segment in front of it, have become organs used in fertilization. We must consider what new structures have appeared here, and to what extent these new modifications have been attended by degeneration. The ap- pendage of the second segment (fig. 5) is longer than that of the other seg- 1^ ■ ments ; pressed against the ventral surface of the back part of the thorax, it stretches out as far as the space be- jimtaUHs. l ft tween the second and third walking SBdabd.minaueK- , . , , ment of the male legs, the part which corresponds to (fiMitTiew)(i,.yi): the oviduct of the female. It serves as «a;', exopodite; w) 1 1 rt ■ T i» endopodite; a, a channel to conduct the liuid from roiied piate of the p T r. 1 endopodite; 6» the male orifice to that or the female, the jointed ex- . . T / \ tfinity of the It consists of a protopodlte (pr.) same (Hnxley). and an exopodite (ex.) similar to the corresponding parts of the appendages we have just described. The endopodite is profoundly modified. The inner border of its proximal region, which is not jointed, is extended into a thin plate rolled into a hollow horn (a) while the outer border is represented by an annulated part (&). The part in process of 34 UNIVERSALITY OF DEGENEEATIVE EVOLUTION degeneration is the exopodite. On considering the development of the various parts of this appendage, it may he concluded that the exopodite has not undergone the same modification as the endopodite. It now remains to conclude this study of the . abdomiiial appendages by an ex- H' amination of the first segment. o-p*- Immense variation occurs in the female. Sometimes there are two Fio. 6. — Astacus Jhivia- Hiis. Lefi appendage appendages, somctimcs one or both of the 4th abdominal ^-^ .. BPRment of the f n ale are missiug : in any case the (3/l):pr, piotopodite; . . °' ■' en,en, epi- podite (Huxley). are transformed into two long thin plates, and the podobranch is replaced by a mem- braneous plate (the epipodite) (ep.). Behind the maxillipedes are the claws and the four walking legs. In these five pairs of appen- dages the exopodite shows most signs of degeneration. We will Fro. n. — Astacvs fliiviatilis. Second left w Iking lei! (1,5/1) : CO, (oxnponite and now examine the com- ha, bhsipodi e, formine pr, protopodite; ir, gill; is, ischlopodite; /ne, meroi-oditej pOUent partS Of thCSC CO, caipopoiiite; i), prupodite; dj, dacty- lopodite (Huxley). appendages (lig. 11). The protopodite (pr.) consists of two parts (ba. and CO.), and carries a gill (Jr.) 'except in the last walking 4 38 UNIVEKSALITY OF DEGENERA.TIVE EVOLUTION leg. The endopodite consists of five primitive joints, which' are well developed and form the ordinary claw ; these five parts are the ischiopodite (is.), the mesoppdite (me.), the carpopodite (ca.), the propodite (pr.), and the dadylopodite (da.). The exopodite is missing. No vestige or rudiment of it is to be found in any phase of the development of the cray- fish. In the lobster, however, which is closely allied to the cray-fish, the exopodite is still to be found during the larval period. The third funda- mental part of the primitive member persists also in prawns throughout the entire period of life, but the organ is very small. At the extremity of the first and second pairs of walking legs there is an apparatus consisting of a fixed part — an elongation of the proiopodiie — and of a moveable part — the daetylopodite. This furnishes the walking leg with a prehensile organ which is well developed in the first pair of walking legs, and which is enormously increased in the true claws. In this evolution degeneration is exhibited by the disappearance of a joint, for in these appendages the basipodite and the ischiopodite are immovably united. This morpho- logical degeneration corresponds to a functional change in the appendage. So long as the claw was used for locomotion a joint at this point was in- dispensable for progression. It is this joint which, in six or eight-footed beasts allows of the horizontal motion of the member necessary for locomotion, which in six-footed beasts results from the general TRANSFOKMATION OF ORGANS OF ANIMALS 39 structure of the parts and their auricular combina- tions. On the claw becoming prehensile, the joint hinge consolidates, the lever thus becoming much stronger and permitting the claw to be used to greater advantage/ It now remains to examine ^^^'^J? J-'SC the appendages situated in front of the maxillipedes, i.e. the appendages of the head. f,o. n.^Astacm jiuvmmu. In the second maxilla (fig. 1 2) L7™.*„p'„1W":„1'6''a/bia(Ji,i a special transformation may ^i'T'eXTuf ? T^S be observed. The coxopodite »»">ide(Hu.ie,). (co.), and the basipodite (la.), are flat plates ; the endopodite (en.), which is small and undivided, ex- ., hibits signs of degeneration in its ^*~t!!\,V*" size, and in the absence of all '?Vo.-^-^X^ articulations. The exopodite, accord- ^fc^ ing to some authorities on the Tia.is.—Axtacusnuvia. subjcct, no longer exists, while tills. First left max. "' ° . j ., ^ ilia (1,5/1): CO, coxo- accordms to others it constitutes podlte, and ba, basl- , . t • / i i podite, forming pr. With tliB cpipodite fthe analogous protopodlte; en. en- i i • i i dopodite (Huxley), part to that which we regard as representing the gill in the maxillipede), a large peculiarly-shaped blade, the scaphognathide (sc). In the first maxilla (fig. 13), a partial degenera- tion of the organ is very marked : the exopodite and ' See J. Demoor, Secherches sur la Marche des Crmtac4s (Arch, de Zool. exp. et gen., 2° serie, t. iv., 1891). T. List, Bewegungsapparat des Arthropoden, 1. Theil, Astacus Fluviatilis. Morphol. Jahrbuoh., xxii. Bd. 3. Heft, 1895. 40 UNIVERSALITY OF DEGENERATIVE EVOLUTION the epipodite are missing ; the endopodite is reduced to a mere unjointed stem, and only the protopodite retains its two normal component parts. The mandibles (fig. 14), the appendages of the fourth segment, are modified entirely for mastica- tion. They consist of a strong transverse piece (jyr) provided at the extremity with an inner surface {st) for grinding and sawing, and of a three-jointed piece {en) with bristles which point outwards. The first piece is the result of the X^ Fifl. 14. — Astacuf fiuvia- Fio. 13. — A&tacus fiimia- Fio. 16. — Astacwi Jfuvia- tilU. Left niKiiHlt'le tUin. Left ant nna tilts. Left antennu'e (1,5/1): pr, protopodite; (1.5/1) : pr, i lotopodite ; (1,S/1) : pr, protopodite; «j, er-dopi'dite; st^ r«sp- ex, exopodite: en^ eudo- ex^ exopodite; c», CLdo. inc purftice of the proto- podiie(MUXley). podite (Huxley). podite (Huxley), modification of the two parts of the protopodite, which have united to form the organ of mastica- tion ; the second, which represents the endopodite, is the feeler, an organ of sensation. All the other parts of the appendage have disappeared. The antenna (fig. 15), which is a tectile ap- paratus, is formed of two parts representing the segments of the protopodite (pr.). The long-ringed process is the endopodite, while the lateral scale of the antenna represents a much reduced exopodite (ex.). TKANSFOEMATION OF ORGANS OF ANIMALS 41 The antennule (fig. 1 6) consists of a protopodite (pr.), furnished with an annulated endopodite (en.) and exopodite (ex.).. The eye-stalk (fig. 17) consists of a two-jointed protopodite (pr.). This is all that remains, the endopodite and the exopodite being absent altogether. ^c'T^P'' This examination of the appen- ^^^ r.'Zsiacus ju^na- dages of the cray-fish clearly shows 'nfiu-^'*^ rotlT'o^to that all fresh adaptation in the (Huxiei). ' appendage entails the modification of some parts and the degeneration of others. In each case evolution is accompanied by degeneration. § 3. Transformation of homologous organs in individuals of different species. The limitations of our present knowledge make it difficult to determine definitely the origin of limbs among vertebrates, but they are universally supposed to have developed from the lateral folds which still persist in AmpMoxus, and which pro- bably existed in the ancestors of vertebrates.^ ' A. Morphological proofs : {a) Lateral folds of amphioxus. (J) Identity of the skeletons of paired and unpaired fins. (c) The number of spinal nerves passing to the fins. {d) The mode of entrance of these nerves into the fins. B. Emhryological proof s : (a) Continuous lateral folds in the embryoes of fish. (i) The formation of metameric poucnes (ooelomie in- vaginations not only at the point of origin of the limbs, but between them). 42 UNIVERSALITY OF DEGENERATIVE EVOLUTION At first the only skeleton of these lateral rods consisted of parallel rods of a tough material. By the transformation of these rods, the skeleton of the limbs of vertebrates was ultimately formed. The skeleton of limbs then consisted originally of a certain number of parallel rods. One of these rods, lying in the long axis of the future limb, became longer than the others, v^hile the neigh- bouring rods began to slant, so that those nearest to the elongated rod spread out like a fan, and gradually moved outwards along the principal rod. This phenomenon was repeated several times, so that eventually those rods nearest to the principal rod passed towards the free end of it, and as the others followed in the same direction, the fin finally acquired a feather-like structure. This transformation of the continuous folds into limbs — a progressive transformation, since new and more perfectly adapted organs were formed — was accompanied by degeneration, for a considerable part of the folds disappeared. In the same way, although the transformation of the parallel rods into bipinnated fins constituted a development, C. Palaxmtologicdl proofs : (a) The series of paired spines in Diplammfhus and in Clvmaims between the pectoral and ventral spines. (S) The skeleton of the fins of Cladoselaclie. D. Philogenetic proof: The necessity of the bifurcation of the unpaired folds at the anal region {Stemarchiis). K Fhysiologieal proof: The lateral folds are the undifferenti- ated condition of organs of equilibrium in modem fish. TEANSrOEMATION OF ORGANS OF ANIMALS 43 the rods nearest to the free extremity of the principal rod were considerably reduced jn size. Two series of transformations should now be followed — that of limbs which have not yet ceased to be aquatic, and that of limbs adapted to terrestrial existence. Among aquatic creatures, the Pleuracanthides and the Dipneusti (Xenacanthus and Ceratodus) have best preserved the bipinnated fin. In Ceratodus especially it is exhibited in almost the primitive condition. Gradually, however (as in Orthacanthvs), the rods situated along one edge of the principal rod disappeared, and the fin, no longer bipinnated, became unilateral. This pro- gressive transformation entailed the disappearance of nearly half of the fin. With creatures which have become adapted to a terrestrial life, the limbs — so far as can be judged from what is known at present — appear to have undergone the following transformations : The bi- pinnated fin (such as that of Ceratodus) (fig. 18) is always the starting-point. Then the lateral rods of one half almost completely disappear (as in (Protopterus Amphibius). Next, the other half follows (as in P. anneeteus) (fig. 19), and finally only the principal rod remains (as in Zepidosiren) (fig. 20). , At this point the limbs are reduced to mere lopped stems ; they have not, however, atrophied ; the degeneration which has accompanied this de- ISf^. iiMJ B94HmmiiHBBHi Fig. Mj—Ceraloim Forsteri, Kretft. Bipinnate fins (after GUnther). (-See Doi Fig. 20. — Lepidosiren parade et Dipneutlea, Bull. Soc, beige de Giol,, de PaUmt., et SHydrva>Lxa Wiirtemberg • 13-2 % Saxony • 9-7% Baden • '^•1% Prussia ■ 8-4% 4. In the budget of the Empire, which is of recent construction, naturally no traces of the feudal system remain. ^ These lands comprise more than 99 German square miles with over 206,000 inhabitants — about 37 per cent, of the population, L, Beaulieu, i., p. 48. EVOLUTION AND DEGENERATION OF INSTITUTIONS 97 III. The Budgets of Gerynany, France, and England. (a) Germany. — We have just seen that in the principal States of the Empire there is still an extensive amount of collective property. In Prussia, for instance, besides forest land, there are nearly 1,500,000 acres of arable land, an area equal in extent to one of the smaller departments of France. (6) France. — Of the old collective property of France only forest land remains, all grazing and arable land having long since been alienated. The budgets of modern times, however, still exhibit traces of the old feudal system. The State, for instance, up to the last few years, continued to receive quit-rents for properties under the old system.^ The budget shows an annual decrease in these receipts, having fallen from 100,000 francs in 1857 to 32 francs in 1869. In 1876, however, there was a rise to 2000 francs. (c) England. — Here the decline of collective property is still further exhibited. In the Statistical Abstract published in 1877, the net revenue from Crown lands for Great Britain and Ireland figures at £40,000 net. Among the " mis- cellaneous receipts" we find only £200,000 which can be regarded as revenue derived from collec- tive property, i.e. 15 millions in a budget * Leroy-Beaulieu, TraiU de la science des finftnces, i., p. 35, 98 UNIVEKSALITY OF DEGENERATIVE EVOLUTION receipt of 1,956,000,000 francs. Later on, in 1884-1885, these revenues became rather larger (principally owing to the purchase of the shares in the Suez Canal which had belonged to the Viceroy of Egypt), but the proportion remains the same — 22 to 23 millions in ^ budget of 2 milliards 300 millions. We see, then, that in all the financial systems of modern Europe the progressive evolution of the modern system is taking the place of the more or less rapidly- degenerating feudal system. § 2. Modification of similar institutions in different social groups. According to the primitive constitution of things, the land occupied by a tribe or clan was regarded as res nullius, and consequently at the free disposal of all the members of the community (the Feld- Walt- und Weidegemeinsehaft of V. Maurer). With the increase of population, the value of land rose, and the state of things became modified, the rights of groups and individuals becoming con- solidated and at the same time limited. Then arose gradually or simultaneously the following various forms of landed property : — (1) Land held by families ; (2) by villages ; (3) feudal property ; (4) communal or public property ; (5) property belonging to corporations ; (6) private property. I'amily, village, and feudal property represent. EVOLUTION AND DEGENERATION OF INSTITUTIONS 99 among certain peoples, three successive stages in the evolution of property. When the old system of land tenure was abolished, private and communal or public property began to develop simultaneously. While certain lands which were free to alL the inhabitants became transformed into collective property, other such lands lost their public char- acter and became private property. In the first case, the communes, on being called upon to fulfil functions of increasing complexity, proceeded to transform all or part of the properties concerned into patrimonial property or property for the use of the people (communaux, allmenden)} In the second case the property of the old community became the joint but undivided property of the members of the corporation ; when, however, for purposes of cultivation it became necessary to divide it, the corporative property became trans- formed into private property. ^ Giron, Le droit administratif de la Belgique, No. 683. " There were three kinds of eommunal property : "(a) Property directly appropriated to the use of the public such as public squares, streets, churches, &c. " (5) Comnwnal property properly speaking — i.e. the real estate and rights belonging to the tribune and to which the people were entitled to a personal share. These consisted of the forest land, rights of appanage, waste land, moorland, and the rights of pasturing. "(c) Patrimonial property, i.e. that held by the commune, the revenue from which went to the commune to defray the expenses of administration. It included timber land, arable land, house property, market places, &o." 100 UNIVERSALITY OF DEGENEEATITE EVOLUTION On reviewing in succession these various phases in the evolution of landed property, it will clearly be seen that modification has everywhere been attended by degeneration. 1. Family 'property (Montenegro). — Of all the Balkan States, Montenegro — owing to the natural barriers which separate it from the rest of Europe — has best preserved its archaic institutions. Here, side by side with modem institutions, may be found the old system of division into forty-two tribes (pleme) which are sub-divided into clans or confrater- nities (brastvo) and into patriarchal families (zad- rugas and inokosnas).^ The development of modern political and judicial institutions has, however, con- siderably lessened the importance of the plemes and the brastvos, so that progression in this direc- tion has not been effected without accompanying degeneration. With regard to property, the two different forms of family tenure have been substituted for what was formerly the tribal or clan system. Of the former collective property of the clan, there only remain the following traces : — 1. Property rights held over certain portions of land — ^generally forest or waste land. 2. The right of pre-emption in favour of mem- bers of the brastvo or of those related to 'For information about the common or differential characters of the zadniga and the inokosna see Ardent, La FamUle zougo- siave au Montinigro, (Mforme sociale, 17th October 1888.) EVOLUTION AKD DEGENERATION OF INSTITUTIONS 101 a member within the first six degrees of lineal descent.^ 3. The right pf allottiijg to relatives their share in the duties of helping widows and paupers in their work. The workers in this case receive no payment, neither have they any right to demand maintenance. Still rarer are vestiges of the collective property of the pleme. A few portions of land, however, still belong to that body, and it is probably a survival of this ancient condition of things that foreigners are ^ Article 48 of the Civil Code of 1888, drawn up by Bogisic in all possible accordance with "the excellent customs" of Monte- negro, begins with the statement that " the right of pre-emption, a privilege which has so long been enjoyed by the members of the brastvo, by persons whose lands adjoin, and by the members of the village and pleme, still flourishes, and will probably continue to do so." Bogisic adds that, in accordance with this right, "any person desiring to sell his land, or any kind of real estate belonging to him, is constrained, according to the established custom in such cases, to first offer it in legal order to those persons who enjoy the right of pre-emption, in order to give them an opportunity of pur- chasing it at the price at which it is to be offered to the public. Article 49, sec. 1, gives a list setting out the order of precedence of those who enjoy the right of pre-emption. 1. Members of the brastvo within the first six lineal degrees of descent, „ T> ■ J- ■ • 1 J "\ Transference, of recent 2. Persons owning adjoining lands. I . . . , , 3. The other members of the village. V f • )\ • ■ ii 4. The other members of the pleme. I „ f , r^ ^ ■' fined to relatives. If none of those entitled to the first offer desire to purchase, the owner may then sell his property to any other Montenegrin. 102 UNIVERSALITY OF DEGENEKATIVE EVOLUTION not permitted to acquire landed property in Montenegro,^ and that the public are unrestricted in the right to hunt over any ground they choose.^ 2. Village property {Russia). — Village communes and the periodical division of land — the mer of Russia or the dessah of Java, for instance — do not represent types of a primitive system, but are the outcome of a whole series of modifica- tions, Kowalevsky traces the evolution of the present system in Eussia through the following principal stages : — 1. The joint use of land by the members of one family group (pechische), corresponding to the zadruga of the Southern Servians, and sometimes comprised of more than forty persons. 2. The division of the mother-family into separate households, thus forming a village community, and the temporary allotment of the land of the community among the separate families. ^ The old Montenegrin law relating to landed property, which prescribes the purchase of land in Montenegro by any but Montenegrins, is still in full force. No transaction in violation of it is legally binding (Dickel, &ud,e sur le nouveau Code civU rtwriUrUgrin). ' Throughout the Southern Slavonic countries hunting is the free right of all. Anyone may hunt where and how they please, not only on public ground, among the mountains and forests, but upon private property, whether cultivated or not (Diekel, p. 86). EVOLUTION AND DEGENERATION OF INSTITUTIONS 103 3. The alienation of all or part of the allot- ments assigned to the families and the constitution of agricultural communes no longer necessarily consisting exclusively of persons akin. 4. The periodical division of land, which, as the population increased, was instituted with a view to re-establishing an equal distribu- tion. This system of division, which was established gradually, only dealt with the more valuable sort of land, such as meadows and arable land. The forest land and pas- turage, that at least which was not already annexed by the Lords of the Manor, was free to all. This transformation of family communities into village communities was not effected without accompanying degeneration. The administrative institutions of the family group disappeared, and the rights of pre-emption in favour of blood rela- tions were gradually replaced by village rights. The importance of the family, regarded as an economic group, decreased in proportion with the increase of the importance of the village. In some places, however, and especially among the Ossetes who inhabit the valleys of the Caucasus, the old system may still be found. There, at any rate up to within the last few years, the aouls (villages) are principally com- prised of families holding land in joint tenure, 104 UNIVERSALITY OF DEGENERATIVE EVOLUTION frequently sharing all things in common. These aouls are very rarely met with nowadays.^ Besides these family communities, there yet remain among the Ossetes, as in Montenegi'o, numerous vestiges of the primitive system of clan property, i.6. the appropriation of certain portions of land by the members of the clan, the common use of pasturage and forest land, the enforced participation in certain public works, and the rights of heritage over unclaimed land, or un- appropriated property which had become so owing to the lapse of some " feu " or by the extinction of a family community.^ 3. Feudal property {England). — The introduction of the feudal system into England resulted in the substitution of a new method of grouping, in place of the old agrarian communities. These new groups, like the townships of earlier times, con- sisted of a complete organization occupying definite boundaries. Instead, however, of being a family group administrated by a democratic organization and government, this new system, which Sumner- Maine calls a " manorial group," consisted of a tenantry autocratically grouped together and governed by a feudal chief — the Lord or Seignior.' ' See KowaJevsky, C) With regard to meadow land, sometimes the Lord of the Manor put up enclosures for his own benefit from Candlemas till Midsummer, the rights of the community being established during the remainder of the year only; sometimes it was the community who put up the enclosures, when the Lord of the Manor was only entitled to the use of the land during the intervals ; sometimes pasturage was held as the joint property of the old community, or rather of their descendants the tenants ; but as a rule it was regarded as more or less common pro- perty. The best meadow land was divided up into what were termed " deals " and apportioned by drawing lots. (c) With regard to arable land, the method of appropriating and cultivating the land occupied by the tenants retained many traces of the village system of collective property. For instance : the enforced rotation of crops; the periodical division of land in certain parts of the country; the division of land into three breaks in other places ; and the destruction after the harvest of the en- closures surrounding the crops, after which the land was used for the herding of cattle. These survivals may yet be found in some districts of modern England, in spite of all the great changes in the English system of property ; changes such as the disappearance of the serf and the appearance of yeomen in the course of the thirteenth to the six- teenth century, and the dispossession of the yeoman EVOLUTION AND DEGENERATION OF INSTITUTIONS 107 in favour of the growth of large properties in the course of the sixteenth to the eighteenth century. 4. Public or collective propei'ty (Switzerland). — It is easy in Switzerland to follow the course of the transformation into political communes of the old system of economic communes, whether village or manorial. In the mountainous parts of Switzerland this transformation is still incomplete, and side by side with the modern commune may be seen the old form of collective property, though in a more or less advanced stage of degeneration. The successive stages of this evolution may be enumerated as follows : — 1. The village communities (the Feld-Walt-und- Weidegemeinschaft of von Maurer). 2. The collective property of the inhabitants, whether feudal, free, or partly both {Gemischte Gemeinde). The Feldgemeinschafi completely disappeared after the Eeformation, the collective land of the community, the joint use of which was the right of all the inhabitants, being restricted to mere waste land, forest land, and pasturage (Allmend). ' The Allmend, in the primitive sense of the word, meant that part of the old collective property held in joint tenancy by a community of inhabitants or any other public body, the use of which was limited to those who had a personal title to it. This primitive meaning has changed in Switzerland — excepting in the 108 UNIVERSALITY OF DEGENERATIVE EVOLUTION 3. Thd institution of JBurgergemeinde — public corporations — the members of which enjoyed the sole use of the Allmend, and upon whom fell the costs of local administration ; as the population increased, the conditions of admission into the Bilrgefr- gemeinde became more and more strict. 4. The creation of Einwohnergemeinde, or political communes, established side by side with the old system, and in many localities eventu- ally taking its place. In these the responsi- bilities of the former system were assumed, and part or the whole of the collective property was appropriated. Where the old communities still survive, they have, as a rule, ceased to exercise the greater part of their original functions. In the Cantons of Berne and Saint-Gall, for instance, the old communities have de- livered up the greater part of their possessions to the political communes to provide for the expenses of general ad- ministration. Their only now remaining function is the administration of the small remainder of their patrimony and the main- tenance of the indigent members of the community. Canton of Sohwyz — owing to the changes in the institution itself (see Miaskowaki, Die Schweiizerische Allmend in ihrer geschicht- lichen Entwickelung, von xiii. Jahrhundert Ms zum Gegenwart). EVOLUTION AND DEGENERATION OF INSTITUTIONS 109 Independently of this decay of the old system of communities, the formation of political com- munes was attended by other phenomena of degeneration : — 1. The suppression of all or part of the " rights of usage" enjoyed by the inhabitants. The EinwohnergemeiTuie, being called upon to dis- charge more and more onerous and complex func- tions, were obliged to either partially or wholly transform the communal possessions, to the per- sonal use of which the people were entitled, into property appropriated to the use of the public, either directly {i.e. into churches, teaching insti- tutions, etc.) or indirectly, as a means of obtaining a revenue (UrwerbsqtieUe). 2. The decrease in collective property. Many of the BUrgergemeinde, although no longer discharging public functions, retained part of their estates, which were held by the members in joint tenancy. On the other hand, as the increased population necessitated the cultivation of the Allmend, the original " right of usage " re- sulted in many instances in a transformation of the land into private property. This transformation, however, was not always complete, and all the intermediate stages may be traced between the old collective tenure and the appropriation by individuals. 5. Corporative property (JBelgiiimy—^Here we will limit the sphere of our observations to Belgium, in 110 UNIVKRSALITY OF DEGENERATIVE EVOLUTION order to avoid repetition, similar examples being almost universally exhibited throughout Europe.^ Our information is obtained from the work of Paul Errera, entitled Les Mamirs, recherches his- toriques et j'uridiques stir quelques vestiges des formes anciennes de la propri4td en Belgique, The masuirs (the amlorgers of Flanders) were the mansuarii or maiisoarii of the Merovingian period, originally serfs, afterwards tenants and copy-holders, and finally freemen. Their history exhibits the following stages : — 1, The feudal epoch in which the masuirs — i.e. all the members of the manorial group — enjoyed "rights of usage" over all Vaste lands, forests and pasturage adjoining their holdings. These rights appear to have been conferred by the Lord of the Manor, but they, really dated from a much earlier period. 2. As the increasing population necessitated the regulation and limitation of these rights, certain conditions of property and residence were stipulated for in those seeking admis- sion to the rights of the masuirs, and these privileged persons organized themselves into corporations which were more or less ex- clusive and separate from the general community. * With regard to corporative property in Switzerland, see von Miaskowski, Die achweiaerisehe Allmend, pp. 87 and rollowing. EVOLUTION AND. DEGENEKATION OF INSTITUTIONS 111 3. By degrees — by means of cantonments, pur- chases, prescriptive claims, &c. — these cor- porations absorbed the best part of the land, and became almost independent of the Lord of the Manor; as a rule, the latter gave up half of the common territory to them, and freed the surplus from all rights of usage. In the corporations of masuirs, however, there were still a few remaining vestiges of some of the institutions of the old manorial group from which they had gradually developed. The Lord of the Manor, for instance, himself being an in- habitant and a masuir, had a right to a share in the property of the conmiunity, and further, in his seigniorial capacity, certain privileges accrued to him such as " la haute ileur des bois," i.e. tithes and pannage (crops of acorns). 4. The Eevolution put an end to all feudal rights, and removed the last remaining traces of the origin of the masuirs. Throughout this long series of transformations, it is evident that degeneration has followed in the track of progress. Besides the disappearance of the manorial group and its attendant institutions, the rights of the masuirs may be said to have become more restricted as they became more defined and secure. In the early days, all the inhabitants enjoyed joint rights over a vast common territory. 112 UNIVERSALITY OF DEGENERATIVE EVOLUTION at the close of the old system this territory had become much reduced in extent, and had become the property of a more or less large group of privileged persons. 6. Private ^property (Switzerland). — After the Eevolution, the communities of masuirs and other similar corporations ceased to have any recognized legal existence. Those which still survived in spite of the irregularity of their legal position, owed their existence to their insignificance. The others dispersed themselves, or were dispersed, and the property which had belonged to them was either incorporated with the communal estate, or divided up among the members of the old com- munity. In each of these cases the transformation was attended by degeneration, for the archaic adminis- trative organization disappeared. We saw in the cdlmend, of Switzerland, this same divergent evolution of public and private pro- perty, part of the common land being transformed into communal property, while the use of the sur- plus ended in some instances in the land becoming ultimately the private property of individuals. This frequently occurred where land was cultivated as orchards. In early times both fruit and fruit- trees belonged, like the land itself, to the community, . and in certain parts of the Cantons of Uri and Schwyz this is still the case. By degrees, however, individual rights over fruit-trees planted on the EVOLUTION AND DEGENERATION OF INSTITUTIONS 113 allmend came to be recognized. These rights, whether temporary or held for a life-time, eventually became perpetual, and finally this right to the private acquisition of trees led to a right to acquire the land itself. This last transformation was not effected without a struggle and occasionally the land was reclaimed by the community, the pro- prietor of the trees receiving compensation. Now- a-days the possession of trees and land usually go together. Duality of this kind, however, is still to be met with in certain localities. In the Sernfthal (in the Canton of Claris) a still stranger custom prevails with regard to the maple forests. There, the soil, the trees, and the fallen leaves (the latter being used as litter for cattle) all belong to different persons.1 With regard to house property there are more intermediary conditions between use and possession. In some villages, the chalets as well as the ground upon which they are built, belong to the whole community ; in other villages, both are part of the collective property. Sometimes private possession is restricted to the house or chalet, the right to the ground upon which it is built lapsing with the existence of the house. In order to limit the number and durability of these buildings, many restrictions are imposed, such as the prohibition to build houses of stones, or chalets of wood cut from trees not belonging to the builder himself or to the corporation to which he belongs, etc. * Miaskowski, Die schweizerische Allmend, pp. 18 and following. 114 UNIVEKSALITY OF DEGENERATIVE EVOLUTION 7. Summary. — This long series of modifications, the result of which was the transformation of primitive communities of goods into the modern forms of public and private property, was ac- companied throughout by degenerative changes. The establishment of family property entailed the curtailment of tribal and clan rights. Family property passed into property held by the village ; next the development of feudal tenure involved the degeneration of the old agrarian communities ; finally, the primitive organization of property with the administrative and political institution depen- dent on it, disintegrated and disappeared as the primitive community of goods lapsed into the personal enjoyment of these by individuals, and as the primitive method of land tenure passed into the rights of private property. We see then that degeneration has always accompanied evolution : the destruction of old institutions is involved in the formation of new institutions. PART II DEGENEEATION IN THE EVOLUTION OF OEGANISMS AND SOCIETIES We have seen that modification of organs and of institutions is always associated with partial de- generation. We have now to show that, similarly, when organisms and societies become modified, de- generation is shown in some of their organs or institutions. This shows again the universality of degenerative evolution. CHAPTEE I ALL OEGANISMS EXHIBIT EUDIMENTAEY OEGANS All existing organisms have lost some organs in the course of their phylogenetic development. This may be proved in two ways : either there are Remaining vestiges of these organs, or else they are to be found in other creatures which may be regarded as ancestors. "5 116 UNIVERSALITY OF DEGENERATIVE EVOLUTION 1. Eudimentary organs, signs of a degenerative transformation in the organism itself, are either organs which have ceased to be functional, or which have so diminished in importance that their total disappearance would be unattended by any appreciable loss to the organism. In the majority of cases this cessation of function is attended by a corresponding structural decay. 2. The system of comparing living organisms with their presumptive ancestors equally demonstrates the retrogression of certain organs. Among the Orobanchacese for instance, parasitic plants derived from normal green plants, no trace of cotyledons is observable from the period of germination.^ Among animals, taking the horse as an example, several organs have wholly disappeared. In the genealogy of the horse, which is well known, the earliest ancestor Uohippus, possessed five functional fingers on the fore-feet, and four toes on the hind- feet. The horse still possesses one functional finger and one functional toe, two rudimentary fingers and two rudimentary toes. Two fingers and two toes have entirely disappeared. It is hardly necessary to point out that this system of comparison does not demonstrate the ' L. Koch, Die EntwicielungsgeachiciUe der Orohanchen. Heidel- berg, 0. Winter, 1887. RUDIMENTARY ORGANS OF ANIMALS 117 degenerative changes attending the phylogenetic development of an organism with such incontestable certainty as does the existence of vestiges of rudi- mentary organs. It is our belief that all organisms contain vestiges of organs, either more or less apparent. In our present condition of knowledge, however, it is quite impossible — particularly as regards plants — to prove this theory universally. It is to be hoped, however, that future researches will ultimately succeed in establishing it. In the meantime, we will point out the most typical among the cases known to us. With animals, as with plants, our investigations have extended not only to every kind of group, but to the most varied systems of organs, thus giving our theory an extremely wide application. Section I. Rudimentary Organs of Animals. § 1. Budimentary Organs in Man. Throughout the whole human organic systems signs of degeneration abound. 1. The Integumentary System. — In the ancestors of man, the entire surface of the skin was covered with hairs. Man's clothing of hair is far from perfect, the hairs of which it is composed being rudimentary. 118 UNIVERSALITY OF DEGENERATIVE EVOLUTION According to Hertwig, the teeth should be regarded as part of the tegumentary system, as they really represent the scales of the skate, situated within the buccal cavity. In man, the last molar, or wisdom tooth, is a rudimentary tooth. The small-sized shallow crown, the diminished number of tubercles, the fusion of the roots, the tardy appearance and occasional absence altogether, are all indications of a rudi- mentary condition. 2, The Skeleton. — With few exceptions, the articular surfaces of the bodies of mammalian vertebrates are covered in youth with bony plates. These sometimes become very thick, and are called terminal epiphyses. In some mammals — the Sirenians, for instance — the terminal epi- physes have disappeared. In man they stiU exist, but in an advanced stage of degeneration. In the lower vertebrates, such as the crocodile, many more ribs are functional than in man. In the crocodile all the ribs connected with the cervical vertebrae are functional; whereas in man they have degenerated. Of one entire section of the human vertebral column — the tail — so fully developed in the majority of other vertebrates, only a vestigd now remains. Other rudimentary skeletal pieces are the lesser horn of the hyoid bone, the stylo-hyoidean liga- ment, the coracoid process, and the Interclavicular ligament. EUDIMENTARY ORGANS OF ANIMALS 119 3. The Muscular System. — The cutaneous muscles, those of the shell of the ear, and those that move the tail, which in most mammals are well developed, are still present in man, but have degenerated. Further, there is to be found in man the intra-aeetabular part (the round ligament) of the deep flexor muscle of the toes which is functional in some animals — in young ostriches, for instance. In the adult ostrich the intra-acetabular part is separated from the rest of the muscle, which is attached to the pelvis. Traces remain in the horse of a connection between the intra- and the extra-acetabular parts ; the muscle itself is divided into two parts, the pectineal muscle in the thigh, and the deep flexor muscle of the toe situated in the leg. In the orang-outang, this degeneration has made further advances than in man, the intra- acetabular part of the muscle having entirely disappeared,^ 4. The Nervous System. — Here we find numerous signs of degeneration, of which the following are a few examples : In the brain the pineal gland, the last remaining vestige of what was formerly a functional eye, is present. In the spinal cord the fllum, terminale still exists. We know that the spinal cord in man does not retain its normal thickness to the extreipity of * See Sutton, Ligaments, their nature and morphology. London, 1887. 120 UNIVEKSALITY OF DEGENERATIVE EVOLUTION the vertebral column but is arrested at the first lumbar vertebra. There a considerable number of special nerves leave it, forming a mass of branches like a horse's tail. Along the centre of these nerves, in the middle line, a slender filament represents the spinal cord to the extremity of the coccyx. This filament is the flam terminale, the spinal cord in a condition of degeneration, 5. The Digestive System. — The caecum and its vermiform appendage, are well known to be organs which have degenerated. 6. The- Vascular System. — In quadrupeds the intercostal veins are vertical, the blood consequently flowing against gravity. These veins contain valves which indirectly facilitate the upward and onward flow of the blood by preventing it from running back. Man, being a biped with a vertical thorax, is provided with intercostal veins that are almost horizontal. The ancestral valves being no longer indispensable are in a condition of degeneration. 7. Sense Organs. — In the olfactory organ there remains a degenerate Jacdbson's organ. In the organ of sight there is a third eyelid in a state of degeneration. In the organ of hearing there remains on the shell of the ear a kind of point (Darwin's point) which is the last remaining vestige of the ancestral elongated and pointed ear. 8. Genito-urinary System. — There is a whole series of rudimentary organs in the genito-urinary system of the higher animals. As is well known, RUDIMENTARY ORGANS OF ANIMALS 121 the Wolffian body plays a considerable part in the t'oripation of the system. This body, the primitive kidney, loses its urinary function at a certain stage of embryonic development, and the permanent kidney which gradually develops alongside, assumes the urinary function. Later on, the Wolffian body assumes new functions, becoming an important part of the genital apparatus. In this transformation partial degeneration occurs, resulting in such reduced structures as the epi- didymis, the organ of Rosenmiiller, the vas aberrans, etc. (see fig. 57). § 2. Budimentary org'ins in various groups. 1. Coelenterates. — The Ooelenterates comprise three great groups.^ The Anthozoa, of which the coral is a type, the Hydrazoa, which include fresh water Hydra and the common jelly-fish of our seas, and the Ctenophora, of which the chief representative in our seas is Cydippe, a globular transparent animal frequently to be found floating in large numbers on the surface of the water. The colonies of Anthozoa are usually composed of individuals all exactly alike. In some species, however, in the Pennatulidse and the Alcyonaria for instance, there is a distinct differentiation amongst the numerous individuals composing the colony. Side by side with sexual individuals 1 See C. Vogt and Emiie Yung, Traits d'anatomie comparie, vol. L 9 122 UNIVERSALITY OF DEGENERATIVE EVOLUTION provided with tentacles and the eight mesenteric folds, are other far simpler individuals : the zooides, the function of which is respiratory and of which the greater part of the organs have degenerated ; the generative organs are lacking, the tentacles are very small, and the mesenteric folds only number two instead of eight. Degeneration, then, is exhibited side by side with specialization. Among the Hydromedusce similar examples abound. It is known that the polyp-like or medusa-like forms of this group which may live independently, frequently associate themselves together to form colonies, sometimes predominantly polypoid, sometimes completely medusoid, and occa- sionally a mixture of the two. In these cases a marked polymorphism is often apparent. The different individuals become adapted to definite functions, and the corresponding organs undergo special development ; the other parts of the body having become either unnecessary or merely accessory, begin to degenerate and finally disappear. Thus we see in Hydroid colonies, not only the hydra-like members, nutritive, fixed and sterile, and the medusa-like members which are reproductive and become free from the colony, but also certain individuals which are termed gono- phores. These gonophores are really medusa-like members which have lost their independent move- ment, and have consequently more or less lost both their tentacles and their umbrella-like discs, RUDIMENTARY ORGANS OF ANIMALS 123 i.e. their organs of locomotion. In some colonies of Hydroids, polymorphism has made such advances that there are tactile individuals of which the digestive tube lacks both mouth and tentacles, and other purely defensive individuals of which the internal organs are almost all in a state of atrophy. Opinions differ regarding the complicated question of the structure of the Siphon- ophora. The organism (fig. 52) may be regarded as a simple medusa of which the different appendages — the pneumatophore (pn.), the swimming bells (d.), the siphons (.s.), the shield (&.), the tentacle (t), the palp (pa.), the gonophores (ffo.), etc. — constitute the organs, or as a colony each part of which is represented by an individual polypoid adapted to fulfil a special function. Whatever theory is accepted, it is clear that a whole series of parts of the creature must be regarded as organs in a condition of degeneration. We accept the second of these theories,^ and 1 Haeokel, System der Medusen : Jena, 1880-1881. A. Lang, Traiti d'anatomie compart. Fig. .is. — Diagram of the structure cf one of the Sipk(mophora. pn. pneumatophore or float ; cl. swimni ng bil ; 6, protective pilyp i t. tentaile; pa, palp; g, gonuDhore; p. peouncle bearing the iriouth; s, i_diTiunals for nu- trition. 124 UNIVERSALITY OF DEGENEEATIVE EVOLUTION regard the whole creature as formed from a Craspedote Medusa which has become mother of a colony and of which the umbrella, developed into the pneumatophore (pw.) has had its radial canals greatly simplified and its tentacles reduced to one during the growth of the colony. The stalk-like stomach (p.) of this medusa has increased in length, but this development is attended by corresponding degeneration, the buccal aperture, which is situated at the free end of the peduncle in normal Craspedote Medusae, having entirely disappeared. The stalk, formed in this way, serves as a support to the number of other individuals of which the colony is constituted, and which are remarkable for the great morphological variation they exhibit. Among these individuals, those at the top, i.e. those nearest to the air-sac, fulfil the function of locomotion. They become transformed into swim- ming bells (cl.) and contain no organs whatever. Below these locomotory organs are the sexual individuals, or gonophores (ff.), and the sterile individuals (s.). The former are of medusoid structure, the umbrella is more or less perfect, and they are sometimes provided with tentacles, and possess a peduncle or manubrium which some- times has a buccal aperture. The sterile individuals provide nutrition for the others. The organs no longer essential to them atrophy in a variable degree. In the case of some the umbrella is present, in others it is absent, and between these EUDIMENTAEY ORGANS OF ANIMALS 125 two extreme types come intermediate types which exhibit every possible stage of degeneration. In the Gtenophore group development and de- generation are exhibited simultaneously in the organs of locomotion. The fundamental and typical shape of the Gtenophore is round or oval, and the eight sides are provided with swimming plates, originally uniform — as in Bero'i. The individuals belonging to this group exhibit important evidences of modification in their external morphology. The body being sometimes com- pressed in various directions, the shape is altered from, the original, and assumes a more or less irregular appearance. mi £ T J. ■ ^iG. 53. — Bolina S'orvegica, ihe organs of locomotion seen from the bioad side. dT . C, Shore rows of swimming ergo a corresponding piatps; a, long rows; l, 1 m 1 1 lobfcs. (After Vogt and change. Take for example Yung, Traiu d'matomie an adult specimen of Bohna norvegiea (fig. 53); the body is lobate, although it was round during the larval period ; the swimming plates are not uniform, four being long and reaching along the whole length of the body, the other four being only developed in the upper half of the body as far as where the lobes are inserted, where they end as degenerate hair-like processes. By referring to the Cestidas, which are ribbon-like in shape, it will be seen that by means of compression the body 126 UNIVERSALITY OF DEGENERATIVE EVOLUTION of the animal is lengthened out into the shape of a narrow ribbon. Of the primitive lateral row of plates, four are represented by mere vestiges, and I the other four, which continue to be functional, are situated, closely coupled together, on the two edges of the animal. 2. Worms. — We will next take in succession the Plathelminthes or flat worms, the Eotifers, the Nemathelminthes, or round worms, the Annelids and the Gephyreans. Among the Plathelminthes, the group of Cesfodes contains the common tape-worm of man {Tcenia solium). In the course of its parasitic existence this worm has undergone considerable morphological changes. The digestive tube is lacking, and the whole nervous system has become greatly simplified. The degeneration of the nervous system is not, however, complete, for important vestiges still per- sist. The degeneration of the digestive tube is much more thorough. In Tcenia solium it is alto- gether absent. In species closely allied to the Tcenia some slight vestiges of the digestive apparatus yet remain. The head or scolex of some species of Tetrarhynchus contain glandular cells which have been homologized with the salivary glands of other flat worms (Trematodes). In other species of Tetra- rhynchus there is a rudimentary organ which repre- sents the oval sucker of the Trematodes, and in Anthroeephalus elongatus the orifices of the salivary glands are in the region of tliis vestige of the digestive tube. EUDIMENTAEY ORGANS OF ANIMALS 127 Many facts in support of our argument may be drawn from a study of the excretory organs of the Cestodes. . The condition of these organs in Caryophylleus 7/iutabilis best represents the primitive condition. Within the body of this worm are a large number of narrow ducts with ciliated funnels communicating with the spaces in the parenchyma (Fraipont). These organs communicate with canals which gradually reunite and anastomose to find a vent in one single aperture, the foramen caudale, which is situated in the posterior part of the body where there is a bladder.. In the Cestodes, however, where the body is very long, the action of the bladder is insufficient to secure a complete evacuation. Secondary apertures are therefore formed at in- tervals along the main ducts. This new structure entails the degeneration of the terminal bladder which has become superfluous. In Botryoce-phalus punctatus, which possesses a great number of excretory apertures, the primitive evacuatory appa- ratus — i.e. the contractile cavity — ^has completely disappeared. Rotifers are minute animals, usually living in fresh water, a few being marine. One of them, an inhabitant of damp eayth or moss, has been supposed to possess the power of revivifying after complete dessication. At the anterior end of the body, a rotifer possesses a complicated ciliary apparatus which fulfils the function of locomotion, and from 128 UNIVEESALITY OF DEGENEKATIVE EVOLUTION the rotatory movements of which the name of the group have been given. This organ is well developed in those types which lead an independent existence, but in those where movement is more restricted or where the character of the organ has changed, it is considerably modified and reduced in size. In PhilodiTWP. (crawling Eotifers) the organ of rota- tion has lost the central part, and in its place is substituted a very complicated organ of prehension. In sessile forms such as Floscularia, the organ is modified and only the primitive character of the inner ring is retained, while the outer ring is segmented and becomes a series of arms or lobes, furnished with stiff bristles. In Apdlus, another sessile form, the organ of locomotion has disap- peared ; this is obviously an instance of true retro- gression, for in young specimens a vibi'atory crown still persists. The Nemathelminthes contain such round worms as these common intestinal parasites : Ascaris, Oxyuris, Strongylus, etc. These all belong to the Nematode group, and possess a complete digestive tube. Gordius, however, exhibits organs which are reduced in a marked degree ; in the adult animal the buccal orifice of the digestive tube is closed, and the posterior part of the intestine has disappeared, although in the young worm the alimentary canal is complete. This modification, though incomplete and appearing only in the adult life of Gordius, is complete EUDIMENTAEY ORGANS OF AKIMALS 129 and permanent in some other Nematodes. In UchinorJbynchiis, for instance, the digestive tube is absent, and nourishment is obtained by means of osmotic soaking through the body walls. The Annelids comprise the annulated sea-worms and forms like the common earth-worm {Lumbricus terricola). In these creatures we will take, first, the development of the eyes. In Oligochcetes, which for the most pari live in soil or mud, the organs of sight are greatly reduced. The Maidomorphce alone have eyes. The Archiannelida — Histrior (a para- site), for instance — possess eyes when young, but in the adult state the eyes have greatly degener- ated. As a rule, the eyes of the Polychcetes are well developed, and in some of them quite re- markably so. In species, however, which do not move about much, the eyes are merely represented by small pigmented spots. We may mention, too, the Qephyreans, without pledging ourselves as to their exact relationship. Bonellia viridis, the history of which is well known and of great interest, belongs to this group. The male Bonellia lives as a parasite on the proboscis, or in the gullet or the nephridium of the female. It is flat and small, and has neither mouth, arms, nor circulatory system. All the organs remain as in the larval condition, with the exception of the genital organs, which are fully developed. Bonellia and Binophilus, a rotifer, of which the male is degenerate, exhibit a progressive degenera- 130 UNIVERSALITY OF DEGENEKATIVE EVOLUTION tion of all the organs not connected with reproduc- tion. Degeneration has made furthest advances in JBonelUa, which affords a striking example of retro- gression side by side with development. Investigations of animal series such as these might well be continued throughout the various classes and groups, showing the existence of rudi- mentary organs in all. We will restrict ourselves here, however, to mentioning the larger subdivisions only, taking one example from each group. 3. The Hchinoderms. — This order comprises star-fish or asterids, sea-urchins, Crinoids and Holothuria or trepangs. Of these we will take the star-fish, and proceed to examine its digestive tube. Under normal circumstances, the intestine terminates in a dorsal anus, centrally, or slightly excentrically placed, pre- ceded by a very short but well-developed rectum. Asteracanthion and Solaster furnish good examples of this. In some kinds of Asterids — in Astropecten aurantiaeus, for instance — the anus no longer exists, and the rectum, having become useless, is greatly reduced, though still exhibiting signs of its original condition. 4. Mollusca. — Of the group of Mollusca we will take the Gastropods and the Cephalopods as examples. There are two kinds of Gastropods, straight and twisted, the former representing the primitive type. The straight types — such as Chiton, Patella, EUDIMENTAEY OEGANS OF ANIMALS 131 Saliotes, and Fissurella — are bilaterally symmetri- cal, while in other Gastropods the spiral twisting of the body causes a progressive diminution of the organs situated on the side towards which the twisting occurs; the organs of the left side may therefore become smaller, and finally atrophy almost completely. The internal shell of the Cephalopods furnishes a striking example of a rudimentary organ.^ The Nautilus (fig. 54) has a shell, the spiral coils of which are pressed tightly against one another. The spiral is divided into a series of chambers by means of partitions, each par- tition being provided with an aperture for tVio nf)Tni«. Very degenerate pineal eye as In C^c?oci«s and like the earliest stace in the slow-worm ; there is no differentiation of the divertialum from the thalamencephalon. E, F, G. Other modes of degeneration of the pineal eye. The eye lies within the skull and there is no parir'til foramen ; cr, cranial membranes; £. Ceratophora. F. Birds; ^, mammals. (After Baldwin Spenct.r.) and becomes divided into a proximal and distal portion. The cells lining the distal 184 THE PATH OF DEGENEKATIVE EVOLUTION part, that farthest from the brain, become differentiated into the cells which will form the lens, and the cells which will form the retina. (3) The distal part becomes specialized, the lens, the retina, and the stalk of the optic nerve are mapped out. (4) The lens, the retina, and the optic nerve become fully formed (fig. 62, a). At this stage the third eye has reached its limit of development. There is a. well-formed retina connected with the brain by a special optic nerve. The organ projects strongly from the surface of the head, but from this point, owing to the development of the cerebral hemispheres, degeneration begins. The nerve (fig. 52, c), becomes broken and fatty, and pigmentary degeneration occurs in it. At the same time, the pineal eye having become useless or even harmful to the animal possessed of it, before the power of receiving perceptions of light has been lost, and before the organ has been far reduced by phylogenetic destruction, a veil of black pigment is formed over it, completely shut- ting it off from the outer light. The nerve disappears completely before birth, its degenerate cells becoming lost in the mesoblastic skeletal tissue of that region. At the time of birth the whole eye is enclosed in a thick membrane which isolates it. The deposition of pigment has THE PATH OF DEGENEKATION IN ANIMALS 185 destroyed any functional activity in the lens and the retina, but these parts none the less retain traces of a complicated structure recalling their condition when functional. In the Ehynchocephala and Lizards examined by W. B. Spencer,^ there is to be found a series of types representing the various stages of the degeneration of the eye in the slow-worm. In the type Sphenodon {Hatteria, fig. 62, a), the organ in the adult is in the complete form. The eye possesses a lens (c), a retina, (r), with com- plicated histological structure. A nerve {n) places the retina in communication with the brain. In Chamceleo (fig. 62, b), the degradation of the organ has reached the following stage : the epi- physis has a distal portion corresponding to the eye, but the histological differentiation of this is incomplete, neither the retina nor the lens being distinct. Netvous fibres connect this with the proximal portion which is hollow and in com- munication with the brain. It thus represents the second stage in the formation of the eye in the slow-worm. In the types Leiodera and Calotes (fig. 62, c), the chief degeneration is in the optic nerve, which has partially disappeared, and no longer connects the eye with the brain. The eye itself is not quite ' W. B. Spenoer, On the presence and structure of the pineal eye in Lacertilia {Quarterly Journal of Micrasoopicat Hoienee, 1886). 13 186 THE PATH OF DEGENEEATIVE EVOLUTiOK SO degenerate : the retina has not a complex structure, but both retina and lens are present. In the type Cyclodus (fig. 62, d), the degeneration is still greater : the epiphysis is a vesicle attached to the thalamencephalon. The walls of this vesicle show only the smallest symptom of primi- tive differentiation into lens and retina. The proximal part remains hollow, and shows no trace of differentiation at all. This, however, is not the only fashion in which degeneration of the pineal eye proceeds. In another series of creatures it retains its connec- tion with the thalamencephalon, but remains inside the skull. The parietal foramen closes, thus completely shutting off the eye from the light; the eye becomes useless, degenerates, and the optic nerve loses its function as a conducting channel, Ceratophova (fig. 62, e). The pineal organ then becomes a degenerate structure in which it is exceedingly difficult to see traces of its original condition, and which is usually marked by an abundance of blood-vessels : Birds (fig. 62, f), Mammals (fig. 62, g). Thus, the degeneration of the pineal eye shows that the optic nerve, the last organ to be completed, is the first to disappear.^ In studying the d^eneration of the pineal eye, ' The same happens in the case of the ordinary paired eyes. In the degenerate eyes of the mola, tl\e optic nerve is more reduced than arc the other structures. THE PATH OF DEGENERATION IN ANIMALS 187 we have seen the path of progressive evolution being retraced, at least in the case of these reptiles. The present state of knowledge does not permit an extension of the enquiry to the conditions found in higher animals. HoWever, according to Flesch, traces of a sensory epithelium have been found in the pineal^irody ^i -man, the horse, the sheep, and the bat : that is to say, that in these creatures too the oldest parts of the structure have resisted degeneration longest. We cannot refrain from the conclusion that in this series degeneration retraces to a large extent the steps of original advance. 2. Degeneration of the organs of sight in deep- sea Crustacea. — We cannot however establish the conclusion of the last paragraph as a general principle. The fauna of the deep sea includes a large number of Crustacea, and in these the eyes, which are relatively useless, are often degenerate. The course of the degeneration is generally definite, and of all the structural parts the most long-lived are the eye-stalks, although we know that these are a recent formation. A number of examples chosen from Decapod Crustaceans, which are specially abundant, will illustrate this point.i Nephropsis, which lives in the Atlantic and Indian Oceans at moderate depths, is a relative of the Lobster. The optic stalk is short and carries ' See Pelseneer, V Exploration mologique des mers profamdei (Omifireruies Universitaire de Bruxelles). 2 Annee, 1890. 188 THE PATH OF DEGENEKATITE EVOLUTION a rudimentary eye which has neither pigment nor cornea and is coloured like the general surface of the body. Eryonicus (fig. 63) belongs to the same group, and comes from the region of Saint-Thomas in the Antilles, where it lives at a depth of about 825 yai'ds. This animal has a reduced optic stalk, but at the extremity of this, where in littoral forms the eye is borne, there is only a de- pression as if the eye had been carefully scooped out. Willemcesia '(fig. 64), a re- lative of the marine cray- fish and an in- habitant of the Atlantic at a depth of about 3500 yards, is com- pletely devoid of eyes in the adult condition, although it possesses them in the larval stage. ' Scolophthalmus (fig. 65), which lives' down to 4000 yards, is quite devoid of eyes, but possesses eye-stalks which terminate in spines. It seems, then, that different species' of deep-sea Crustacea may present different degrees of degenera- tion of the eye. One species in itself exhibits all Fio. 63. Eryonicus caecus. Bate? (After W. Faxon, The Stalk-eyed Crustacea, Mem. of Mus. of- Comp.. Zuol. Hurvui'd College, vol. xviii., 1895.) mm- i-m; 190 THE PATH OF DEGENEEATITE EVOLUTION grades of degradation according to the depth at wliich it lives. This creature — Gymonomus — which, when near the surface, has fully formed eyes upon movable stalks, at a depth of a few hundred yards exhibits movable stalks without eyes; and at 1500 yards the stalks are fixed and end in spines. Isopod Crustacea, which live in the deep sea, present similarly degenerate eyes. Many are blind Fig. 65 Scolophthalmus htcffugus. Fax. a, optic peduncle transfoi-med to a i-pine. (Afttsr W. Faxon, The Stalk-eyed Crustacea, Mem. of Mus. ofComp. Zool. Harvard College, vol. xviii., 1S95.) and display all kinds of optic degeneration. Noesa, for instance, simply has eyes devoid of pigment. Thus, in abysmal Crustacea, the degeneration of the eyes is in no sense a retracing of developmental stages. Another instance chosen from examples of the atrophy of organs in individuals, shows that the supposed law of retracing cannot be made uni- versal. 4. Atrophy of the branchial vessels in man. — Examination of a human embryo of about three THE PATH OF DEGENERATION IN ANIMALS 191 weeks old shows the presence of a series of slits on the sides of the neck, the slits not being parallel, but converging towards the ventral surface. Be- tween these slits are swellings, or pads, which pass up towards the dorsal surface and appear like the beginnings of hoops or ribs enclosing the visceral cavity ; the elevations are the branchial arches, the slits are the gill-slits. In the human embryo (fig. 66, A), as in fish, these slits appear from above downwards, and as they are formed, the cor- responding blood- vessels arise. These vessels, or aortic arches, arise from a ven- tral aorta (a.) which gives off six lateral branches (c.) at each side. These lateral branches run up between the gill-slits and form two main trunks on the dorsal side which converge to form the descending aorta (ad.). In man the branchial arches are transformed, parts of them entering into the structure of the face, and during the transformation parts of the aortic arclies atrophy (fig. 66, b). But the order Fig. GG Diagram of brandiial arches in roammalS'. A. Embryonic sta^e. a, anrta; c. aortic arcties; AD. uorsal anrta. B. Adult Btage. Itie parts represented- bv dntted lines have degenerated. At aorta; v, carotid; ap^ pulmonaiy arteiy. 192 THE PATH OF DEGENERATIVE EVOLUTION of this atrophy does not correspond in any way to the order of the formation of the vessels. The median parts of the anterior two lateral branches {m} and m^) disappear, and the vertical parts remain as the internal and external carotid vessels. The vertical piece which joined the posterior parts of the third and fourth arches dis- appears : the internal and external carotids thus acquire a stem of their own. The parts of the fourth arch remain ; the fifth arch disappears at each side, and the sixth arch forms the pulmonary artery (a.p.). Thus the degeneration of these vessels represents in no way whatever a retracing of their develop- mental history. All that occurs is that the useless parts disappear and the useful parts persist. A comparative study of this example would only enforce our conclusion. In ontogeny the neurapophyses are more ancient than the vertebral centres. None the less, as we have already seen, the examination of any vertebral column from head toward tail shows a gradual disappearance of all parts except the centra, although the centra are the last to be formed. Section II. The path of degeneration in plants. 1. Rarity of cases of recapitulation in the organogeny of leaves. — We have already said that THE PATH OF DEGENEKATION IN PLANTS 193 recapitulation seldom occurs in plants, the develop- ment of the whole and of its organs being usually direct. When it does occur, it is generally limited to characters coming from comparatively recent ancestors and not even in the most transitory form entering into the formation of the fundamental parts of the plaut.^ ' ■■ The rarity of recapitulation among vegetables is the result partly of their fixed condition in the soil, and partly of the more rigid nature of their cells. The immobility of a plant forces the adnlt to live in the same place as the embryo. Among animals, on the other hand, it frequently happens that the young pursue a manner of life different from that of the adult and resembling that of the ancestor. Young Cirrepedes are vagrant and have the same needs and use the same organs as other vagrant Crnstaoea ; larval frogs inhabit the water like their fish-like ancestors. In plants there is nothing similar; all the aqnatic flowering plants are derived from terrestrial ancestors, bnt if at the beginning of their existence these aqnatic plants were to bear leaves adapted to aerial life they would ensure their own destruction. The ex- ceedingly rare ancestral traits to be found in a few species are naturally of a kind not to incommode their possessors. It is improbable that these are a legacy from distant ancestors ; they would not have been spared by natural selection had they not come from-ancestors of very much the same habit. The absence of locomotion in plants has also produced a greater adaptability than among animals. Animals, when conditions are unfavourable can remove in search of more suitable localities, plants being fixed in the soil must become modified or perish. Plants, therefore, offer numerous cases of individual adaptation. We do not know if these adaptations are transmitted by heredity, but natural selection has at least secured the widest range of plasticity. Thus plants rapidly rid themselves of ancestral legacies which have become useless. The transitory organs of animals arc employed for the service 194 THE PATH OF DEGENEKATIVE EVOLUTION In consequence, vegetable embryology is of little use for investigation of the supposed backward path of degeneration, for the rudimentary or reduced organs of plants do not generally represent ancestral stages. The seedling of LatJiyrus tenuifolius (fig. 67), a vetch, possesses rudimentary organs which . cannot be ancestral stages as their development is direct. In this plant a whole series of leaves are formed between those arising at germination and the adult leaves. This intermediate series displays many arrests of development. The adult leaf, has a pair of stipules, foliage leaflets, and tendrils (fig. 67, J). The leaves just before these, have a pair of stipules (fig. 67, l), which are absent in the leaves next before (fig. 67, h). Still earlier leaves are produced with fewer leaflets and tendrils (fig. 67, d-g), leaves" without leaflets and with a single tendril (fig. 67, c), and leaves entirely without tendrils (fig. 67, b). Lastly, at germination very rudimentary leaves are pro- of the whole hody, the branchial arches of mammals are employed in the formation of important parts of the head and neck. The tail of the tadpole is reabsorbed by phagocytes and its substance used for the nutrition of the body. In the case of plants, such occurrences are rare and limited ; the cells are enclosed in a rigid wall which resists displacement or alteration ; the protoplasmic contents may be absorbed and used as nutritive material by another part, hut the cellulose cell-wall remains. A useless organ can be eliminated only at the expense of loss of material. J. Massart, La JUcapitulation et I'lnrwvation en embryologie vigitale {Bull. Soc. Soy. Bot. Belg., t. xxxiii., p. 150, 1894). Fio. St .—Lathynia tenuifolius. A. B, seedlings in two stages of giowili. C to J, different forms pt successlye leaves. 196 THE PATH OF. DEGENERATIVE EVOLUTION duced, the sole function of which is to protect the axillary bud (fig. 67, B, 1, 2). We have now to show that this series is by no means a retracing of ancestral stages. All, or nearly all, the Papilionace- ous plants have stipulate leaves ; this shows that the exstipulate leaves of L. tenui- folius do not re- present an ances- tral stage. More- over, before the acquisition of tendrils, vetches had a terminal leaflet (see Vicia Pyrenaica, fig. 44) ; none of the reduced leaves in L. tenuifolius re- produce this Fig. C8. — Seedling Vicia of monantlios, stao'e ' moreOVCr the simple leaves without leaflets do not represent an ancestral condition ; the winged petiole is not an ancestral character. The primary leaves of another vetch Vicia monanthos confirm our conclusion. In this case THE PATH OF DEGENERATION IN PLANTS 197 the first few leaves formed in the seedling remain rudimentary, aud serve only to protect the axillary buds (tig. 68, leaves 1 and 2). Contrasted with the condition in L. tenuifoUus, although all the vetches had probably a common ancestor, these primary leaves have three little projections, the two lateral of which are reduced stipules. We do not know of any vegetable example of recapitulation in the case of an organ reduced by arrest of development. Plants exist, however, which after having produced leaves of typical structure begin to produce leaves the development of which remains arrested. Such a plant is the Acacia which bears phyllodes. At first the leaves are like those of other Acacias ; next it bears leaves, the blades of which are rudimentary ; and finally leaves with normal stipules, but with no trace of lateral leaflets on the petiole. In this case the ancestral conditions are known and are quite different.^ 2. Organogeny of flowers. — What we have said about leaves applies to the organogeny of flowers. Here also in the cases of atrophy produced by arrest of development there is no indication of ^ Many other reduced plant organs might be instanced, such as the stipules of SamhvAsus or the teeth of the calyx, in many Com- positse and Umhelliferse. After then' formation such organs grow very slowly and exhibit no trace of recapitulation. This happens with the leaves of Sempervwum (fig. 49). Without doubt these leaves are derived from leaves normally divided into hypopodium and epipddium, but they show no trace of this division. •198 THE PATH OF DEGENERATIVE EVOLUTION ancestral stages, and thus there is no evidence that degeneration retraces the path of progressive evolu- tion. In the cauliflower {Brassica oleracea, var. Botry- tis), a cultivated variety, the inflorescence branches ex- uberantly; most of the flowers produced on these branches are arrested in their develop- ment. Of the immense num- ber of flowers produced on each plant only a few attain sexual maturity and produce seeds; the others abort at differ- Most of these remain in a very primitive stage and do not develop sepals ; but, scattered among them, may be found more fully developed flowers, so that the same plant presents almost every possible stage of flower development. The organogeny of these flowers is quite like FiQ. 69. — Development of flowera of ihn Cauliflower. sa, anterior sepal; sp^ postjiiui* sepal; si, lateral sepal; p, petal ;,.'«, lu^igiitianien ; ee, short stamen ; 0, ovaiy. A, very yQung flower with only rudi- ments of two sepals; 13, flower with rudimentu of four sepals ; C, older flower with rudiments of petals; D, still older flower with rudiments of stMmens and ovary. (The flowers A — D are seen ft nm the t-ide. In the flgures C and D the lateral sepal has been removed.) E, flower seen from behind; the posterior sepal has been removed. ent stages. THE PATH OF DEGENEEATION IN PLAJS'TS 199 that of other Cruciferse. If in the cauliflower the arrested flowers corresponded to different ancestral stages the case would be striking; but this does not occur. Primitively the flower contained only the stamens and pistils, the essential organs of reproduction. But we see that the cauliflower produces first the anterior and posterior sepals (fig. 69, a), then the lateral sepals (fig. 69, b), then successively the petals (fig. 69, c), the four larger stamens (fig. 69, d), and the two shorter stamens (fig. 69, e). Moreover, the flowers display a special readiness to the suppression of certain parts such as the petals, as in the flower D (fig. 69). 3. Progressive degeneration of the prothallvs in phanerogams. — Although embryology gives us few examples, morphology proves clearly enough that in plant degeneration there is no return to ancestral types. This appears clearly from a comparison of the progressive evolution of the prothallus in crypto- gams with its degeneration in phanerogams. Terrestrial vegetation has been derived entirely from aquatic life. The Bryophyta (mosses and liver-worts), the Vascular Cryptogams (ferns), and the flowering plants all have sprung from aquatic algse probably not very different from Goleochcete. Such aquatic forms are reproduced by means of true ova and spermatozoa. The terrestrial plants which were derived from them had also spermatozoa with vibratile locomotor hairs and impregnation 200 THE PATH OF DEGENERATIVE EVOLUTION took place in a fluid. A special organ, the arche- gonium, was developed ; this contained the ovum and made the approach of the spermatozoa more easy. For the present purpose we may omit considera- tion of the mosses and liver-worts as it is improb- ahle that they are in the line of ancestry of the flowering plants. It is necessary only to say that in them, while a single egg is produced in the archegonium a large number of spermatozoa are produced in the antheridium. The same condition is found among the fern-like plants, but in their case, owing to the development of special channels for the passage of nutritive materials, it is possible for a much greater size to be reached. In these circumstances it would be unlikely that the spermatozoa should find at the summit of a comparatively lofty plant the drop of water necessary for the task of fertilization.^ Accordingly the sexual cells are produced on prothalli, which hardly reach above the surface of the ground, and are in a favourable position for the necessary moisture. In the less specialized ferns (fig. 70) a single prothallus bears both male and female organs (antheridia and archegonia). In the Equisetums ' The smaller forms like the Selaginellidse and the cUib-mosses were represented in the past by plants of much greater size, as is seen from fossil remains, and it is probable that the modem forms have descended from these giants. THE PATH OF DEGENERATION IN PLANTS 201 the prothalli are unisexual, but the spores from which they are produced are alike. Finally, in the allies of Selaginella which are the Cryptogams most nearly approaching flowering plants, the prothalli which bear male organs are quite distinct from the prothalli which bear female organs ; more- over, the spores which give rise to the two kinds of prothalli are quite different. A small number of large spores arise in special sporangia, termed macrosporangia, and it is from these that the female prothalli arise. A larger number of smaller spores are pro- yw. to.— seeaiinK of rem (Aspimmm duced . m miCrOSpor- ^1,6 prothallus (J) beai-s anlheridia at its nnmn nnrl fVif«5P minrn edees and towards the centre an aiche- angia anu tnese micro- goniam -which has teen fetUized. The prfv theleaves are small and applied to the stem; Ajtiiiucu jjiujjciu^y, in the youncer parts (A), the leaves are larger formerly collective ondprotmde from the siem. and now individual, seems to be again tending towards collectivism. The same phenomenon occurs in the evolution of matters relating to maritime rights. The sea, according to Eoman law, was equally open to all maritime nations. Later on it has been from time to time practically in the hands of a few nations, and we have now returned to a condition iji which it is equally open to all.^ ^ Tarde, TrcMs/ormation du droit, pp. 161.-162. Paris, F. Alcan, 1890. 238 THE PATH OF DEGENERATIVE EVOLUTION These,' however, are not cases of true revival. The resemblance goes little farther than the name. A comparison of modern institutions with such survivals of primitive institutions as continue to exist will demonstrate this point. The difference between them is so wide that it would be hardly possible to utilize the old as a basis upon which to form the iiew.^ ' Cf. Diirokheim, Les Rigles de la mifhode aociologique. Paris, F. Alcan, 1895. In sociology, dealing as it does with things familiar to us all, such as the family, property, crime, etc., it is useless to attempt to adhere to exact definitions. The exact meaning of some words in common use in conversation cannot he defined with any pre- cision ; the common acceptation of these w^ords cannot be avoided. Now this common acceptation is frequently very ambiguous, so that two totally different things are often referred to under the same name, causing hopeless confusion. There are, for instance, two different kinds of monogamous unions— those so only in point of fact, and those which are legally so. In the first case, a man has only one wife, though legally entitled to several ; in the second he is only legally entitled to one. These two kinds of conjugal conditions are quite different, and yet the same word serves to express both ; it is commonly said of some animals that they are monogamous, although there can be nothing approaching to a legal contract between them. Spencer, when dealing with the subject of marriage, makes use of the term monogamy without defining it in its common and equivocal sense. The result of this is that the evolution of marriage seems to him to represent an incomprehensible anomaly. It seems, according to him, that the superior or monogamous form of union was prevalent during the primitive phases of historic development ; that it then dis- appeared during an intermediate period, to subsequently reappear. From this he concludes that there is no regular connection between CAN ORGANS RESUME THEIR PRIMITIVE FUNCTIONS? 239 It may be definitely asserted then that a re- duced, but still persistent, institution never again becomes actively functional. The following are a few examples which will serve to illustrate this point: — 1. The truck si/stem and dearing-house. — Some forms of the primitive system of exchange sur- vive, not only in countries where money is unknown, but in certain industries where the workers continue to be paid in kind (the truck system). On the other hand, there seems to be a modern tendency towards the elimination of money as an instrument of exchange. The clearing-house system is singularly analogous to the old exchange system. "The truck system," says Stanley Jevons, "represents the first and the last stage ; but it appears for the second time in a very different form. Gold and silver money continue theoretically to be the instrument for buying and selling, but practically metal no longer constitutes the real medium of exchange, and has ceased to pass from the hands of the purchaser into those of the vendor." In this transformation there is obviously no return to primitive systems, the last vestiges of which, far from being revived, are rapidly dis- appearing. social development in general and a progressive advance towards an improved system of family life. A more exact definition would have prevented this erroneous conclusion. 240 THE PATH OF DEGENEKATIVE EVOLUTION The suppression of the truck system coincides with the development of the clearing-house system. 2. Corporations and syndicates. — The radical differences existing between the corporations of former days and the greater part of modern pro- fessional associations has already been pointed out. The ecclesiastical associations, however, of the present day are modelled as closely as possible upon mediaeval institutions. It does not follow that the last remaining vestiges of the latter have been revived. There seems to be evidence that quite the contrary has taken place. At Bruges no attempt was made by the founders of the guild of ambachten to resuscitate such mediaeval corporations as continue to exist in a state of decline. At Iseghem, a small town in the west of Flanders, we have already seen that the corpora- tion of shoemakers was divided up into six or seven guilds at the time of the Eevolution. An attempt was made a few years ago to reconstruct and modernize these guilds, but the scheme fell through. A new corporation — wholly disconnected with the guilds of Saint Crispin, and with no structural resemblance to them — was established instead.^ 3. Archaic collectivism and modern collectivism. — Societies of the present day exhibit numerous ' Emile Vandervelde, EnquUe sw lea Associations professionelles d'ouvriers el d'artisans en Belgique, i., p. 17, Bnixelles, 1891. CAN ORGANS RESUME THEIK PRIMITIVE FUNCTIONS? 241 vestiges of archaic collectivism. The question arises as to whether there is a tendency in the modern school of collectivism to resuscitate such vestiges as remain of the old archaic form of col- lectivism. Far from this being the case, collective property, as conceived by the modern socialist, implies the suppression of the few existing remnants of archaic collectivism. Inheritance, ah intedat, for instance, is a survival from the days of the family community, which itself arose, as we have already seen, from the primitive community. If the modern coUectivist school had any desire for a return to the old primitive community, it would make for the re- construction of the family community by re-estab- lishing the law of collateral succession. Now it is just the opposite with the coUectivists. In order to establish a universal system of collective property they demand among other things, the suppression of inherited succession, ah intestat, at least as regards the collateral line of descent. 4. The survival of elective sovereignty in Eng- land. — The above examples apply to institutions which have degenerated without having completely ceased to be functional. It very rarely happens, however, that having arrived at that condition, they renew their vitality and all their former functions, and this still more rarely occurs in cases of genuine survival. In the English coronation ceremony vestiges 242 THE PATH OF DEGENERATIVE EVOLUTION remain of the old democratic system in which the king was elected by the people.^ The English sovereignty of the present day is merely a decorative institution, the real head of the Government being the Prime Minister, who is nominated in fact if not in theory by the public. This system may almost be regarded as a return to bygone democracy. Nobody would wish, how- ever, to revive the old system of elective sovereignty, and to retrace in an inverse direction the various stages of its degeneration, CHAPTER III CAN RUDIMENTARY ORGANS OR INSTITUTIONS RE- DEVELOP AND ASSUME NEW FUNCTIONS ? The few facts which we are able to cite on this subject must be received with considerable caution. ^ The formality of an election disappeared during the Tudor period. The coronation of Henry VIII. was the last occasion on which the formula was read which set forth the national agree- ment with and recognition of, the succession. The king was, in fact, declared chosen and elected. This formula of election, which disappeared after the coronation of Henry VIII., is recalled to mind by the conclusion of the coronation ceremony of the present day. The archbishop, walking in succession to all four corners of the platform upon which the throne is placed, addresses the people in the following terms : "Gentlemen, I herewith present to you the undisputed sovereign of the realm. Come all who are present and offer homage to him. Are you prepared to offer it?" and the people signifying their assent by acclamation, cry, ' ' God save the Queen " or "God save the King." (De Kranqueville, Le gouverM- memt et lepwrlement JBrUmmiques, i., p. 291.] CAN OKGANS EEDEVELOP NEW FUNCTIONS ? 243 Section I. Bicdimentary organs. 1. Animals. — In Birgvs latro (a land-crab of the Philippines), the gills are atrophied and the bronchial chamber is very richly supplied with blood vessels, while a kind of incipient lung is formed from the lining membrane of the reduced bronchial chamber.-^ However, it is by no means certain that the atrophy of the bronchial apparatus has preceded this development of a pulmonary apparatus. In the following case it rather seems to be one in which a rudimentary structure has redeveloped in order to assume a new function. In the develop- ment of the urinary organs, it appears that the ducts of the mesonephros are quite independent of those of the pronephros, although these mesonephric ducts become functional later in the embryonic life than the pronephric ducts. They are, neverthe- less, formed at an earlier stage, and their rudiments have appeared before there is any trace of the others. From this fact it would appear that in some ancestors of existing vertebrates there existed simul- taneously mesonephric canaliculi and canals homo- logous with them, but exercising a different function. Such a condition actually exists in Amphioxiis : in the branchial region of that animal there are pro- nephric urinary canals and genital chambers which 1 Semper, "The Natural Conditions of Existence as they affect Animal Life.'' (International Scientific Series.) 244 THE PATH OF DEGENERATIVE EVOLUTION are homologous with mesonephric spaces ; but these latter do not exist as genilal chambers in higher vertebrates.^ It must be noted that the homology between mesonephric spaces and the genital spaces of AmphioaMS, as made by these writers, is not uni- versally accepted. 2. Plants. — The Scophulariacese, which have usually four stamens, are derived from ancestors which possessed five. Usually the fifth stamen is only represented by a tiny process which rapidly atrophies. However, in Pentstemon the fifth or posterior stamen is developed, not as a functional stamen, but as a staminode, the function of which is to stretch open the flower to make it accessible to hymenopterous insects with short probosces. Can it be said that in such cases a rudimentary organ has really become redeveloped to assume a new . function ? To establish this it would be necessary to show in the case we have just mentioned, that the stamen did not become trans- formed directly into a staminode, but that it first became rudimentary and then developed afresh into a staminode. An interesting fact is, that in some hybrid varieties of Pentstemon the staminode becomes fertile ' Boveri, Die Nierencanalchen der Amphiooeua. {Zool. Jdhrhuch. AUh. Anat. und Ontogenie der Thiere, vol. v., 1892.) Wiedersheim, Orundziige der Vergleidienden Anatomie der Wir' belthiere. Jena, 1893. CAN ORGANS EEDEVELOP NEW FUNCTIONS ? 245 again. In some flowers sent to us by Mr Cannell of Swanley, the number of petals was increased to six, seven, eight, or nine. In some of these the posterior stamen was sterile and like a staminode ; in others there were five fertile stamens. It is obvious that in this case the staminode had resumed its original function after having lost it. Section II. Mvdimentary institutioiis. The Zevirat. — In his work entitled Tableau des origines et de revolution de la famille et de la propridU, Kovalewsky mentions an instance of a reduced institution which, without having first ceased to be functional, became transformed into another institution. " The custom mentioned in the Bible of alloting a woman to the brother of her deceased husband, is explained by the primitive condition of things with regard to the relations between the sexes ; all the women were the common property of the men belonging to one group of relations. Under the name of levirat, this custom survived for several centuries, owing to the idea which arose later on that a wife was property. Consequently, on the death of the husband, the widow, along with his other belongings, was treated as the inheritance of the person whom the death promoted to the rank of chief or head of the family community." 246 THE PATH OF DEGENERATIVE EVOLUTION The levirat, a family institution, thus derived from the old system of marriage by groups, was transformed by degrees into an economic institution. It is important to notice, however, that this trans- formation was effected without the institution having even been reduced to the condition of a mere survival. PABT III SUMMARY AND CONCLUSIONS From all the facts that we have brought together, the general conclusion becomes plain that retrogres- sion, notwithstanding the etymology of the word, does not imply a return to the ancestral condition. Eudimentary organs and institutions resemble the primitive states of these, in so far as they no longer possess certain parts which the primitive stages did not yet possess. None the less, profound differences exist between the primitive and the reduced forms. In the primitive condition the institution or organ is capable of varying in the direction of new uses ; in the reduced form, after a certain ■ degree of atrophy, there is no longer the possibility of redevelopment to resume old or to acquire new functions. These observations apply equally to biology and to sociology. Magnan and Legrain, in their work on de- generate persons, came to similar conclusions. They came to regard degenerate persons as abnormal, chiefly because they were devoid of the power to reacquire the normal condition and quite unlike their primitive ancestors, who, although "47 248 THE PATH OF DEGENERATIVE EVOLUTION possibly brutal and unintelligent, were normal beings with the activity and stamina necessary for future progress. The following two diagrams, borrowed from these authors, represent clearly the differences between the initial and reduced condition of an organ or institu- tion : — aO z z In the diagrams the ascending lines represent the progressive evolution of an organ or institu- tion ; the descending lines represent the degenera- tive evolution. From the point a, representing the primitive condition progressive evolution passes towards o, an imaginary perfect condition of the organ. Along the upward line, however, the points a, 6, c, d, etc., represent obstacles to further pro- gress — that is to say, factors tending towards degeneration. From these points lines of de- generation pass towards z, and the condition at z, although representing that at a, is not identical SUMMARY AND CONCLUSIONS 249 with a, and is not reached by a sliding backwards down the line o, a. Thus, although the most recently acquired features may disappear first, degeneration is not an actual retracing of steps until the point of departure is reached. The degenerate condition is a new point, and really the term retrogressive evolution is misleading. It BOOK III CAUSES OF DEGENERATIVE EVOLUTION PART I ATROPHY OF OEGANS AND INSTITUTIONS The factors of atrophy The causes which are active in producing degenera- tion are various, but they may all be referred to the limited nature of the means of subsistence, that is to say, of nourishment in the case of organisms, and of capital and labour in the case of institu- tions. This limitation produces a struggle between the individuals (societies or organisms) and between their component parts. In the course of the perpetual struggle for existence among the different parts of an indivi- dual, the institutions or organs which have ceased to be functional tend to disappear, their nourish- ment being absorbed by the active parts, 1. Biology. — In biology the struggle for existence among component parts appears clearly as a factor of degeneration in the case of accidental atrophy. This is to be seen, for instance, in the atrophy of 252 CAUSES OF DEGENERATIVE EVOLUTION the leaves of an etiolated plant ^ or of the muscles of a limb which has been immobile for long, or in the case of muscles which have become inactive from disease of the central nervous system. The results are similar in cases of normal atrophy. In frogs, toads, and other Batrachia Anura,^ the disappearance of the tail before the adult state is reached is the result of a struggle amongst the cells. The active protoplasm of the muscular fibres develops specially, and gives rise to many cells, which enter the contractile material and separate its elements. Gradually all the contrac- tile material is absorbed by these isolated cells. Many plants, especially Sempervivum (see fig. 73, A, p. 236) possess a reduced stem with the leaves closely massed upon it. This reduction of the stem, which is nearly constant in the ^ When a cutting from a potato or a seed (fig. 75) is allowed to sprout in the dark, the young stems assume characters different from those of plants grown in light. The absence of chlorophyll produces important modifications of growth. In light the stem is short, and the leaves are large and expanded ; iu darkness the stem is very long, and the leaves are much reduced. This atrophy of the leaves is the result Of the struggle for existence amongst the organs of the plant. Light increases the rate of transpiration, which is chiefly due to the presence of chlorophyll. As chloro- phyll is most abundant in the leaves, the transpiratory current sets strongly towards them, carrying in it the nutritive materials for the formation of new cells. On the other hand, in etiolated plants, transpiration is slower, and the nutritive materials delayed in the stem give that the opportunity for specially active growth, which takes place at the expense of the leaves. ' Metchnikoff, Amnalea Inst. Pasteur, January 1892. THE FACTORS OF ATKOPHY 253 species, is the result of a struggle amongst the organs produced by a scanty water supply. The leaves attract to themselves the greater part of the water absorbed by the roots, and thus retard the growth of the stem. But if the plant be cultivated in an atmosphere satur- ated with water, the struggle be- tween the leaves and stem is stopped, and the stem grows to a much greater extent (fig. 73, B, p. 236). Vicia Faha, like most vetches, produces at germination rudi- mentary leaves, and similar leaves are borne at the base of each branch. Goebel has shown that these rudimentary leaves may be made to grow by cutting away the terminal buds at a young stage. In this way the struggle between the first formed and later leaves is suppressed. In the case of the individual, '''i;rfanre"?ij;'r''l! atrophy results from the struggle for existence amongst the organs. In atrophy throughout a species it is the struggle for existence amongst individuals that plays the chief part. Clearly in this struggle, useless organs become impediments and burdens. If any organs Fio. 75. —Two 8<>edllngs of Cicer arietinum. seedliap grown in ^lf;ht. B, seedling grown in darknesit. 254 CAUSES OF DEGENEEATIVE EVOLUTION are useless they are harmful as they use nutrition without conferring any advantage upon the whole organism. Darwin pointed out that when through changed environment a structure became useless, its degeneration became certain, as it was a dis- advantage to the individual to squander nourish- ment upon a useless part. Weismann has shown that such a reduction or disappearance of a useless organ is the result of variation and natural selec- tion.^ Variation results in the appearance of individuals with the useless organ in various stages of imperfect development ; natural selection perpetuates these advantageous stages by giving advantage to indi- viduals which tend to produce the organ in the most degenerate condition.^ ^ See, however, Herbert Spencer's Social and Moral Problems, as he differs from Weismann on this point. ' The following are good examples of the operation of variation and selection in producing atrophy in a species : — (1) Loss of constaTU colour among domestieaied animals. — Wild animals, especially birds and mammals, have a colour which is constant for a whole species. Frequently the colour is protective in rendering the animal little distinguishable from the environ- ment in which it lives. As soon as such a species has been domesticated man becomes its protector, and protective colouration is no longer necessary, and soon disappears. In the wild state, the colour is quite as variable as in the domesticated state, but the abnormal individuals become the prey of enemies and are removed from the species. This applies not only to animals which are preyed on by others, but to predatory animals them- selves. The wild-cat, for instance, will have less diflBoulty in stalking its prey if its colour makes concealment easy. In the THB FACTORS OF ATKOPHY 255 2. Sociology. — With societies this elimination of useless structures is effected much more easily than with organisms for several reasons : — In biology a special factor, heredity, gives to specific characters a force which does not exist in the same degree with social institutions. Now functional organs common to a whole line of descent are not easily effected by the influence, of individual surroundings. Further, the trans- domesticated condition man provides food and the colour being unimportant all variations may survive. (2) Loss of spines inplamts on oceanic islands. — It is well known that the presence of spines protects plants from the ravages of herbivorous animals, particularly mammals. But in oceanic islands bats are generally the only mammalian inhabitants, and so, accord- ing to Wallace (Darwinism), there are no spiny plants in the indigenous flora of St Helena. The much richer flora of the Hawaian islands includes only a very few prickly plants. All the endemic genera are unarmed, as also are most of the endemic species of other genera ; even genera like Xwn.thoxyVwm, Acacia, Xylosoma, Lycium, and Solanum, which are so frequently armed in other countries, are there represented by unarmed species. The two species of JRubus bear prickles reduced to the merest points and . the two palms are devoid of spines. How is the absence of spines to be explained in these plants? The plants have been derived from the mainland, the seeds being brought by the wind, by currents, or by birds, and having found soil have germinated. In their new country they are not attacked by herbivorous animals, and it is immaterial to them whether or no they bear spines. The individuals badly armed are at no dis- advantage compared with those possessing the normall armature ; on the other hand, they have the advantage of being without useless organs to support. Spines, ip consequence, gradually disappear. This struggle for existence may cause the disappearance of some organisms themselves, and not only the atrophy of parts of them. 256 CAUSES OF DEGENERATIVE EVOLUTION mission of acquired characters is, to say the least, doubtful. There is no proof of individual atrophy- being hereditary, while with societies modification may be transmitted by imitation. Institutions which have fallen into disuse rarely recur in freshly-formed societies. Natural selection plays an all-important part in biology, but it is artificial selection which almost exclusively governs the social domain. Many vegetables, as for instance the carrot {Daiicus Carota) are natives of France. The seeds of the cultivated carrot must fre- quently be carried to waste lands or uncultivated soiL The domesticated variety, however, is never found wild although the wild variety is abundant. This vegetable has lost the power of struggling against weeds ; it flourishes only when it is protected by man and when by repeated weedings its wild competitors are removed. When it is returned to its original wild haunts the plant dies out at once. Most cereals, although we may not know their wild ancestors, are in a similar condition. For instance, if man were to cease cultivating Wheat {^Triticv/m, saMvum), or Eye (SecaZe cereaie), there is no doubt but that these would completely disappear. Their fate would be shared in Belgium at least by many species which are reaped with them at harvest, such as Centaurea cyamts the Corn-flower, Agrostenima Githago the Rose-Campion, Specu- alaria speculum, and others. If a corn-field were left to the free operation of nature, weeds would soon intrude and cause the dis- appearance of the plants usually present in it. What would happen in Belgium would happen with other plants in other countries. Thus, near Bergen in Norway, some plants, such as Melandryum album, Sihne inflsta, Vida cracca, etc., occur only in cultivated fields. In Java, many aquatic plants such as Limno- chwHs Plumieri, I/udwigia perennis, Jussiaea suffruticosa, etc., live only in the rice-fields which are artificially watered and manured. The cessation of tillage would cause the disappearance of all these plants from those localities. THE FACTOES OF ATROPHY 257 An institution which has become useless and burdensome is generally suppressed before its complete degenerative evolution is accomplished. This suppression may be either voluntary — as in the liquidation of a commercial company, for instance — or it may be enforced. By the terms of Article 73 of the Belgian Company's Act, " the dissolution must be declared upon the demand of all those interested at the termina- tion of six months from the time when the number of shareholders has been reduced to less than seven." The downfall is generally effected in this sudden way, either voluntarily by the interested parties, or by the intervention of legislative means.-^ Sometimes, however, artificial selection does occur, ' The occurrence of autotomy or self-mutilation in animals, as in crabs, has analogies with what we have been discussing. Similarly some plants brought into a new locality suddenly shed their leaves, lianuncuhcs aquaiilis, cultivated in water, produces long divided leaves without stomata and with chloro- phyll in the epidermic cells. If, from some chance, the water falls below the level of the plants, the adult leaves become dry and perish. The very young leaves growing unsubmerged are still divided, but to a lesser extent ; they have stomata, and the epidermis is devoid of chlorophyll. If the plants be again sub- merged, this form of leaf dies, and there is a new development from the youngest leaves of the normal aquatic type. Other plants show similar occurrences. Thus, when a Fuchsia that has been cultivated in a conservatory is exposed to the air, all its leaves are shed and are replaced by new leaves. These new leaves again fall at once if the plant be brought back into the conservatory. This is a real case of autotomy in plants. 258 CAUSES OF DEGENEBATIVE EVOLUTION and then the degenerative evolution of useless insti- tutions is brought about in a similar way as that of non-functional organs. Atrophy of this kind may be, as in biology, either accidental or normal.^ In a besieged town cut off from all outside com- munication, all train service is necessarily stopped and the railway staff rendered useless. On the other hand, the defence of the city requires both men and money. Under these circumstances the resources of the railway naturally come to be absorbed in the service of the defence. As a good example of normal atrophy may be mentioned the disappearance, at a certain point ^ See Durekheim on the difference between normal and acci- dental sociological phenomena in Xes Regies de la Tn&hode sociologique. "All sociological phenomena, like biological phenomena, are liable, while remaining the same individuals, to revert to different forms. Now, of these forms, there are two kinds of reversion : — ■ "The one is common to the whole species, and is to be found, if not in each individual, at least in the greater part of them. The cases are not always identical, varying slightly with the individual, but individual variation is restricted to very narrow limits. " The other kind of reversion is exceptional, being of a nature rarely met with, and, when occurring, is seldom permanent throughout the life of the individual. Cases of this kind are excep- tional in point of duration as in other respects. "Here then are two distinct varieties of phenomena which should be distinguished from one another by different terms. Individuals exhibiting only common characters are called 'nor- mal,' while those exhibiting exceptional characters are designated as 'morbid' or 'pathological.'" THE FACTOES OF ATKOPHY 259 of social development, of the popular assemblies in which lay the origin of future societies, i.e. the comitia, assemblies at the market-place, the witenagemot, May Day games, rustic assemblies, the Landsgemeinde of the Swiss cantons, the parochial assemblies of the Andora Valley, and the town meetings of New England, etc. Some of these were suppressed, but some of them merely fell into disuse as other institutions arose which were better adapted to more modern conditions of society. This happened with regard to the comitia curiaia of the Eomans during the period of the Empire, which were gradually supplanted by the comitia centuriata and tribunal comitia. At the time when they are first mentioned in history, they fulfilled only one function: that of ridding the laws of all traces of extrinsic customs.^ To sum up then, it is plain that although social degeneration is brought about by the same general causes as organic degeneration, the comparative importance of the factors in degenerative evolution is far from being identical in the two cases. The autotomy of organs, a protective self-mutilation, exhibits only a far-fetched analogy with the conscious and voluntary suppression of social structures which have become either useless or prejudicial. Direct individual adaptation, which plays a part 1 Mommsen, Droit public romain. Lepeupte et h Sdnat, vol. i., p. 364. 260 CAUSES OF DEGENERATIVE EVOLUTIOK of only secondary importance in the development of animals and plants, is a dominating element in sociology. Institutions are able to modify their structure by assimilating new inventions and im- provements, and by getting rid of the useless parts. On the other hand, indirect adaptation spread over a species plays no part whatever in sociology, for societies seldom reproduce the structures of the societies from which they sprang, when the latter have ceased to exist ; whereas, in animals, when a useless organ is reproduced by hereditary repetition, variability and natural selection become agents in its suppression. CHAPTER I ATROPHY OF ORGANS i Part II Causes producing atrophy The ultimate cause of the atrophy of organs is the limitation in the quantity of nourishment. We have shown that if there were an indefinite supply of food there would be no struggle, and, as a result, no degeneration. We have now to examine more minutely the course of atrophy and the nature of its immediate causes. The atrophy of an organ is a reduction in ATROPHY OF ORGANS FKOM LACK OF SPACE 261 structure, in nutrition, and in functional activity, but the succession of these three events varies with the nature of the exciting cause. Eeduction begins with structure when the exciting cause is lack of space, due, for instance, to increase in another organ {atrophy from lack of space). Atrophy begins with function when an organ has become useless {atrophy from lack of utility). This uselessness may arise from two causes ; the function may be no longer useful to the individual or to the species, or it may be assumed by another organ. Lastly, atrophy may begin with a diminution in the supply of nutritive materials (atrophy from lack of nutrition). This defective nutrition may be the result of a general cause such as feebleness of the whole organism, or it may be due to the hypertrophy of another organ. § 1. Atrophy from lack of space. Cases of this kind are rare. 1. Development of the teeth. — Among animals, the development of the teeth furnishes an excellent example. The number of the teeth in human beings is reduced compared with the number present in Lemurs and in Platyrhine Monkeys. These have six grinding teeth while in man five is the maximum number. Our posterior molar, however, appears late in life ; it is smaller than the others, so that it may be useless for chewing ; 262 CAUSES OF DEGENEKATIVE EVOLUTION frequently it is absent, in the lower races in nineteen per cent, of cases examined ; in the higher races in forty-two per .cent. The reduction in num- ber and size of these teeth is due to a re- duction in size of the jaw-bones, a cause which also frequently produces a distortion in the arrangement of the other teeth. The rudiments of the wisdom teeth appear on the maxillary tuberosity and on the coronoid process ; it is only after eruption that they come into normal connection with the jaw- bones. 2. Atrophy of the ■Among Fig. 76, — ^Tip of an ear otLolium perenne^ with two earlets, the superior bearing two glumes, the lateral earlet with only the superior Rlume. In the lateral earlet the lower flower is open . , and has two glumules ; all che other SVUp&rwr glume. flowers are closed and exhibit only the i , j.i t; inferior giumuie. plants tnere are tew cases GI, inferior glume ; GS, superior glume; o . i_ ii. c gi, inferior glumulei gi, superior 01 atrOphy aS a reSUlt 01 ^ """ "' lack of space. In grasses of the genus Loliwm, the earlets are arranged in a spike, but in such a fashion that only the terminal earlet has space for both glumes (fig. 76). The ATROPHY OF ORGANS FROM LACK OF USE 263 superior glumes remain ; the inferior, pressed against the axis, disappear after the embryonic develop- ment of the flower. 3. Degeneration of palece and of stamens. — Lack of space is probably the immediate cause of the disappearance of palese in the receptacles of some composite flowers and of the posterior stamen in the flowers of some Scrophulariaceae and Labiates. In normal racemose inflorescences each floret grows in the axil of a reduced leaf called a bract. When the axis of the inflorescence is shortened and the florets crowded, as in the capitula of composite flowers, it frequently hap- pens that the bracts of the florets (termed palese) disappear. This absence is most usual when the capitulum is small and the florets are large. In Labiates and most Scrophulariaceae, although the ancestral number of stamens was five, there are not more than four present ; when only one is absent, it is the original posterior stamen which was pressed against the axis of the inflorescence. § 2. Atrophy from lack of use. 1. Functional Inutility. (1) Etiolated plants and immohile limbs. — We have already quoted as instances of accidental atrophy, cases of degeneration of leaves in etio- 264 CAUSES OF DEGENERATIVE EVOLUTION lated plants, and of muscles in unused limbs (see fig. 75, p. 253). Fia .77.— AympAoeo alba sown on tlie mud and at different depthg In It 1, 2, S, successive stages of the same seedling. £au, water ; llmon, mud. ATROPHY OF ORGANS TKOM LACK OF USE 265 (2) Upicotyl and primary leaf of Nymphaea. — These structures in the water-lily are good examples of normal atrophy in individuals. Dur- ing germination ^g. 77) the cotyledons of the water-lily remain inside the seed, and a new organ (at right angles to them) grows vertically upwards. The lower part of this is the first internode of the stem (epicotyl), and the upper part is a primary acicular leaf. It grows up- wards through the mud until the summit of the leaf reaches light. The growth of the epicotyl is then much slower, and its terminal bud begins to shoot out horizontally. The use of this growth of the epicotyl and primary leaf is to carry the bud to the light. When that purpose is achieved, these structures atrophy. A similar occurrence may be found in SagUtaria sagittifolia (fig. 40, H, I, p. 72). In that case, however, it is the hypocotyl which elon- gates, until light is reached, and then degenerates. (3) Moots of Uiricularia, cotyledons of parasitic plants, leaves transformed to ^ines in Phylloeactus crenatios. — As instances of atrophy throughout a species produced by inutility of the parts con- cerned, we have already mentioned the roots of Utricfidaria and the cotyledons of the parasitic plants Guscuta, Orohanche, etc. The spines of Fhyllocadus crenatus produced from modified leaves are another example (fig. 78). Above the rounded base by which a branch of Phylloeactus is attached to older branches, there 18 266 CAUSES OF DEGENEKATIVE EVOLUTION are \, /' N' is an angular region, the sides of which prominent, and bear leaves modified into spines to serve as protecting organs, as in the similar case of Cereus. Higher up the branch the pro- minent sides become flat, and the spines are replaced by minute scales. This degeneration is the result of loss of utility. The Phyllocacti are epiphytes, and their situation consequently places them out of the reach of crop- ping animals. The spines near the basis of the branches are a survival from the terrestrial an- cestors of Phyllocactus. The spines higher up have degenerated. Animals offer many instances of atrophy as a result of inutility, both in individuals and in species. (4) Atrophy of the branchial arches in mammals. — As they are no longer functional, most of the mammalian branchial arches atrophy. Three pairs alone persist, and of these it is only those parts which are useful. (5) Atrophy of ventral fins. — Instances of atrophy through uselessness in species are to be found in the ventral fins of fishes like the Pediculati, which live in the mud, or in Frotopterus, which for a part of the year is completely buried in mud (fig. 19, p. 44). Fro. 78.— Branch of Phyllocactus crenatus. ATKOPHY OF ORGANS FROM LACK OF USE 267 (6) Atrophy of muscles. — Cessation of use is also the cause of the degeneration of the flexor and extensor muscles of the fore-limh in Cetacea, and of the imperfection of the finger joint articulations in Cetacea and Sirenia (fig. 79). In the latter cases the surfaces of the articu- lation which make flexor and extensor movements easy dis- appear. When a limb becomes a paddle, it is necessary that it should be flexible, but that the articulations should be immobile. (7) Atrophy of the tail in man. — The caudal region of the human vertical column is com- posed of four or five very de- generate vertebrae. The whole of this organ is degenerate. When the tail is formed at an early stage of embryonic life, no. 79.— a!otiocephaiu$ ,1 i. T_ 1 1 • J. p melas. Kigiit anterior the vertebral column consists of nn suowirg absence of , 1 - . • 1 . i. 1 j_i 1 artifular fa'-ets f r the thirty-eight vertebrae ; the lesser joints of the angers. number of vertebrae in the adult ' is due to reduction of the tail, which in man is quite useless. Later on in life a further instance of atrophy may occur in individuals. In old men the caudal vertebrae are frequently fused, and the whole region is smaller. (8) Degeneration of the hyoid apparatus in man and birds' — This case shows a close correspon- 268 CAUSES OF DEGENEKATIVE EVOLUTION dence between atrophy and loss of function. The second arch becomes connected with the third, the parts of which are, in the adult, the styloid pro- cess, the stylo-hyoidean ligament, and the lesser horn of the hyoid bone. According to the weight of the tongue, the parts of the second arch become more or less developed. In man the suspensory apparatus of the hyoid bone is extremely simple, and it is still more reduced in birds. " The tongue, reduced to a minute cartilage, no longer requires the support of a bony base so that the hyoidean apparatus might almost be removed from the anatomy of a bird. It is present, but in a rudimentary condition " (Geoffroy-Saint-Hilaire). In the horse, which has a heavy tongue, the second arch is strong and completely bony. In fish the hyoidean system is still more im- portant, although in them this is associated not with any importance of the tongue, but with the branchial apparatus. The parts of the second arch are very strong, as they form a fulcrum against which the branchial system works, but its main parts are recognizable. " The hyoidean apparatus is the same in all vertebrates ; its functions are at a maximum in fish, and at a minimum in birds, while in mammals the condition is intermediate" (Geoffroy-Saint-Hilaire). II. Transference of function. — Among plants and animals there are many instances of organs well developed in younger stages of life, hut which ATROPHY OF ORGANS FKOM TKANSFEEENCE 269 become rudimentary in later stages on account of their functions being assumed by other organs. (1) Atrophy of the tail in Batrachia, Anura, and of the larval gills in some insects. — The tadpole of the frog has a well-developed tail which acts as the organ of locomotion ; the adult animal moves by its limbs, and the tail, useless in the adult, has been removed by phagocytosis. The aquatic larvse of many terrestrial insects possess tracheal gills, that is to say, membranous expansions of the skin, within which tracheae ramify. In the adult insect respiration is con- ducted by normal tracheae communicating with the air, and the larval organs of respiration atrophy. In addition to such cases of atrophy occurring normally in the life-history of individuals, there are known many cases where the organs of a species have disappeared on account of the trans- ference of their functions. (2) DisappearaTice of limbs. — ^Vertebrates which move by general undulations of the body have lost their limbs for this reason. Such cases are Slow-worms, Amphisbsena, Snak'es, Eels and Cse- cilians. Many parasitic creatures have similarly lost their organs of progression, as they depend upon their host for movement from place to place. Sacculina, a parasite on the carapace of crabs, has completely lost its organs of locomotion. More- over, as it takes its food by processes passing into 270 CAUSES OF DEGENEEATIYE EVOLUTION the tissues of the crab, its digestive canal has been lost. (3) Atrophy of the leaf. — In many plants the leaves have disappeared, their function having been assumed by some other part of the plant, as, for instance, by the phyllodes in Acacia and in PhyllaTdhus (fig. 84). (4) Atrophy of the protonema in mosses, and of the leaves in some xerophUoiis plants. — At germina- tion, mosses produce a much branched filamentous structure which serves as the organ of nutrition, and is termed the protovsma. Later on this gives rise to buds which develop into the normal leafy shoots of the plants. As soon as the leaves are large enough to manufacture food for the plant, the protonema begins to degenerate, and disappears completely, except, in a few rare forms (Ephe- meracese) where the leafy shoots remain very small. In MuehlenbecMa platyclados (fig. 80), which has become adapted to arid regions, and in consequence has the surface from which evaporation may take place much reduced, the branches which do not bear assimilating leaves are rounded at their bases, but higher up flatten into broad blades. These blades contain chlorophyll and fulfil the functions of leaves, these latter being present only as minute scales. Similar phenomena occur in many Papilion- aceous plants belonging to the genera Genista ATKOPHT OF ORGANS FKOM TKANSFEKENCE 271 (broom), Spartium, Alhagi, etc. The leaves dis- appear, and the stems assume their functions. (5) The reduction or disappearance of the calyx. — The disappearance of the calyx in many com- posite flowers and of roots in epiphytes are instances of species losing organs because of the transference of function to other organs. In the simplest Compositae, the fruits (achenes) are dissemi- nated by the wind, the calyx usually being modified into a feathery tuft. In other Com- positae, although wind dispersal still occurs, the feathery calyx is lost, its purpose being fulfilled by wings on the sides of the fruit {Florestinia pedaia), or it is united to a large membranous scale developed from the receptacle (Dahlia), or the sterile florets may be turned into wings {Zind- heimeria teocana). In other cases the fruits are dispersed, not by the wind, but by animals which eat them (Clibadium asperuTn), or they adhere to the fur of animals by hooks formed on the achenes {Calendula), or by hooks formed from the involucre of bracts {La.ppa). In all these cases, and the list might have been made longer, Fia. 80. lenbeckia plaiyclados. 272 CAUSES OF DEGENERATIVE KVOLUTION the function of the calyx has been reduced, or has disappeared on account of the transference of its function. (6) Atrophy of roots. — Eoots in most plants per- form two functions : they fix the plant in the soil, and, chiefly by means of the delicate hairs on their youngest parts, they absorb water and dissolved mineral substances. Sometimes, however, instead of having root-hairs, the rootlets enter into a kind of partnership with a fungus, which lives in their tissues, and absorbs by its processes the necessary food materials from the soil. The pine and beech are examples of this. In some orchids {Gorallorhyza, Myrmechis) the fungi lodge in the subterranean part of the plant, and the branches, having no function, disappear. In most of the epiphytic Bromeliacese the roots are useful only to anchor the plants ; the absorption of water takes place entirely through new organs developed from the leaves. The roots are few in number and small, and after the plant has obtained a firm position their growth almost ceases. In another species {Tillandsia usneoides) of the same family the roots have completely disappeared.^ This plant, called by the Brazilians', " the plant of the air," fixes itself to branches of trees by its leaves, and is easily transported by the wind. Its * For further details oonceming the Bromeliacese see Sohimper, Die epiphytische Vegetation Amerikas, in Schimper's Botanisehe Mittheilungen aus den Tropen, Jena, 1891. ATROPHY OF ORGANS FROM TRANSFERENCE 273 absorption takes place entirely through hairs de- veloped upon the leaves. In addition to such examples of plants which have lost their roots on account of the functions of the roots being assumed by other organs, there are also plants in which the roots represent the sole Fia. 81. — TtBniqphyUum Zollingeri with branches adhering to bark. The plant bears a flower and a bud. vegetative organs. In Tceniophyllium Zollingeri (fig. 81), an epiphytic orchid, the leaves are reduced to minute scales and are of no importance in the nutrition of the plant, that function being trans- ferred to the roots which in the form of flat green ribbons, apply themselves closely to the bark of trees. In this case the roots have assumed the 274 CAUSES OF DEGENERATIVE EVOLUTION functions of leaves, while in Tillandsia usneoHdes the leaves have taken on the functions of roots. The Podostemacese are aquatic plants which live in warm regions attached to rocks in cascades. The stems and leaves are completely absent, the flowers even being produced on the roots. Some of the roots become closely attached to the stones. Others which are green and ribbon-like, float in the stream, and serve for assimilation. § 3. Atrophy from Iodic of nutrition. In some cases of degeneration, the organs do not lose their functions, but become reduced, merely because adjacent parts rob them of their nourish- ment. This kind of lack of nutrition which results in local atrophy, must be distinguished from the general limitation of food-supply which is the ultimate cause of all degeneration. The possible amount of food within the reach of any organism is limited, but besides this, the share of the absorbed food obtained by any particular organ or part of an organ may be limited with a resulting atrophy of that organ or part. Atrophy of this kind may be accidental or normal. 1. Parasitic castration. — A good instance of acci- dental atrophy of this kind is found in Melandryum album where the ravages of a fungus UstUago antheridarum may produce parasitic castration.^ ' See Giard, La Castration paraHtaire in the Bull. Sdent, de la France et de la Belgiqm. 1887, 1888, 1889. ATROPHY OF ORGANS TEOM LACK OF NUTRITION 275 Under the influence of this parasitic fungus which makes its way to the anthers, the stamens of the female flowers assume the form of those in male flowers and in consequence the pistils of these flowers abort from defect in nutrition. 2. Severe or prolonged compression of a limb, — A long continued or forcible compression of a limb results in atrophy of its extremity on account of lack of nutrition from the compression of the blood- vessels. 3. Atrophy of the genitalia in neuter tees. — ;Lack of nutrition is also the cause of the arrested de- velopment of the genitalia normal in neuter bees. The neuters of bees and of some of their allies are females in a state of arrested development. In wasps, humble-bees, and hive-bees, it sometimes occurs that the genitalia of these forms develop sufficiently to be functional, thus resulting in the appearance of small females. In most honeycombs two kinds of cells are formed : in the smaller and more numerous cells are placed the la,rvse destined to become neuters, in the larger and less numerous those destined to become queens or perfect females. The food of the two sets of larvae is different; those in the larger cells are given " royal food " a more nutritious substance. When some of the royal food by an accident gets into a worker cell the sexual organs of that larva are developed so that a small female is formed. In this way as many females as may be desired can be produced, 276 CAUSES OF DEGENERATIVE EVOLUTION and when a hive has lost its queen, the worker bees produce another^ (Lacordaire). Plants afford many instances of degeneration due to defect of nutrition. 4. Atrffphy of the superior flowers in Carex. — In the tall spikes of Carex, it frequently happens that the flowers towards the summit are rudimentary, and authorities are agreed in regarding this condi- tion as the result of defect in nutrition. 5. Atrophy of pistils wnd stamens. — Cases of atrophy of the stamens or pistils normal in species may be given. In Fritillaria persica the flowers are disposed in bunches. The lower flowers possess six perianth members, two cycles of three stamens each, and a pistil. In the median flowers the pistil is smaller, and rarely capable of being fertilized. In the superior flowers the degeneration is complete, the pistil hardly being formed. It might be shown ' The transference of a function is not invariably accompanied by degeneration. Thus, in the functional development of an individual's nervous system voluntary acts which have been re- peated frequently become reflex actions, and have their seat in a diflerent region of the nervous system — as, for instance, walking and acquired professional dexterities. It has been sought to ex- plain the development of the instincts of species in this way by supposing that frequently repeated voluntary acts have become inherited reflexes. As pathological degeneration in man frequently aS'ects the higher regions of the brain, reflexes and instincts may persist after loss of voluntary action. In the cases of transference of nervous functions to lower centres the higher centres do not degenerate but remain able to acquire new voluntary functions. ATEOPHY OF OEGANS FKOM LACK OP NUTRITION 277 that this degeneration is due to lack of nutrition, by removing the inferior flowers from a young bunch, but we are unaware that the experiment has been made. In Viburnum tomeniosum (fig. 82) the flowers are arranged in a compound corymb and occur in two forms. The central flowers have a small corolla, five stamens, and a well-developed pistil Fio. 83 — Flowers from the same inflorescence of Tibumum tomentosum. A, stenie flower with rudimentary piHtii and no s:amen«. — B, sterile flawer witll very rudimentary pisJil and stamens. — C, flower with two ferti.e stamens; the other three stamens and the pistil are rudimentary. — D, fertile hermaphrodite flower. (fig. 82, d). The eight or nine peripheral flowers of each inflorescence have the corolla much larger, but the stamens are absent and the pistil is rudi- mentary (fig. 82, a). The petals turned towards the outer side are much the largest, and it is not rare for the petals turned towards the inner side to be hardly larger than those of the central flowers. What is important to notice, as it bears out the relation between degeneration and lack of nourish- 278 CAUSES OF DEGENERATIVE EVOLUTION ment, is that a smaller petal is always associated with a larger stamen (fig. 82, B, c). In Viburnum opulus, the wild guelder-rose, there are also two kinds of flowers, the central flowers which are sexual and hermaphrodite, and the peripheral flowers which are sterile and possess a very large corolla. The five petals are, however, equal in size, and there are no forms transitional between the sterile and sexual flowers. The cultivated guelderTrose is a variety of the wild form in which all the flowers are sterile and possess a large corolla : the plant has completely lost the power of sexual reproduction. § 4. Atrophy without apparent cause. In many cases it is impossible to determine the cause of atrophy. Why, for instance, has half the inflorescence disappeared from a unilateral cyme ? Why have some composites like Artemisia or Eosaceae like Poterium lost their habit of being fertilized by the agency of insects, and become anemophilous without conspicuous perianth ? Nor can we explain why many Myriapods are blind, al- though they live in association with species possess- ing eyes and in conditions where vision apparently would be useful. Semper discovered, in brackish water in the Philippines, Crustacea (Cymothoe) which were completely blind, although they lived in light. The cause of such atrophies is unknown. ATROPHY OF ORGANS WITHOUT APPARENT CAUSE 279 Moreover, we know that organs may atrophy through correlation with other degenerating organs, or because the whole organism is degenerating although there are no obvious anatomical bonds present between the related parts. As instances of such atrophies we may mention degeneration of the brain resulting on degeneration of the supra-renal capsules ; or of the condition of myxodoema resulting from mechanical or functional disturbance of the thyroid gland. When the essential sexual organs of the male or female are congenitally or accidentally absent, the secondary sexual characters — the beard, voice, hair, and whole male or female aspect — of the body degenerate. When one eye becomes injured or diseased, a frequent consequence is sympathetic degeneration of the undamaged eye. To these cases of correlative atrophy we may add the cases of leaves on many plants with drooping or horizontal branches. On these, the leaves on the upper aspect are frequently rudi- mentary. The appearance is most striking where the leaves are opposite the pairs being at right angles to one another. In a branch of Acer campestre which is upright, all the leaves are of the same size. In a drooping branch the leaves turned towards the sky are small. In plants belonging to very different families (Acanthacese, Melastomacese Urticacese, etc.) oblique branches exist, and the failure in development of the dorsal upper leaves is invariable. In Procris laevigata (fig. 83) the reduc- 280 CAUSES OF DEGENERATIVE EVOLUTION tion of these leaves has gone very far, but in Klugia notoniana the leaves in the corresponding Fig. 83. — Branch of Procris laevipala. F'f, F"/") V"f". The successive pairs of leaves in the axil of a lar^e leaf F' is a female inHoresct-nce ; in the axil of a Bm»ll leaf /" is a male inflnrescence. (The original drawirg was maue at Buitenzorg in Java by Mas Kromobardjo, a Malay draughtsman.) position are absent. In another representative of the same family (Streptocarpus monophylleia), the degeneration has gone still further ; all the leaves ATROPHY OF INSTITUTIONS 281 are absent and a single greatly enlarged cotyledon is the sole assimilating organ.^ It is evident that in these oblique and horizontal branches, the leaves directed vertically towards the sky are in the most unfavourable position for assi- milation, and that, in addition, their presence would shade the leaves lying under them. In these species the more or less complete reduction is an inherited fixation of a sacrifice of these particular leaves for the benefit of the whole organism. CHAPTER II ATROPHY OF INSTITUTIONS The causes of atrophy in institutions are more or less analogous to those which bring about atrophy of organs. First, there is atrophy from want of use, when function either becomes useless or is transferred to another institution. Atrophy from lack of resources corresponds precisely with atrophy from lack of nutrition. There is no- thing, however, in the atrophy of institutions quite analogous to that which, in organisms, results from lack of space. But if the development of an institution cannot be actually impeded by the co- ^ See the figures of plants of this family in Fritsch, Gesneriaceee. {Engler mid PrardVsTwMrlichen PflomsmfoimiUm. Leipzig, 1891.) 19 ' 282 CAUSES OF DEGENERATIVE EVOLUTION existence of another institution, certain instances of atrophy do occur which cannot be said to arise from either want of use or lack of resources. The institution degenerates merely from hindrance offered to the exercise of its functions. Thus, for instance, the laws of exclusion voted in Germany against socialists brought about the decadence, and, ultimately, the downfall of a number of professional and trade associations which cannot be said to have been actually suppressed by law. It is the same with the corporations of Western Flanders ; these have survived the revolutionary laws, but are no longer legally recognized, a state of things leading to many difficulties ; the properties of several of them have been confiscated, as having no owners, and allotted to benevolent institutions. This want of legal recognition must inevitably lead in the near future to the entire disappearance of these vestiges of the corporative system. Cases such as these, however, must be regarded as exceptional, and it may be said that, on the broad average, institutions atrophy either from want of use, or from lack of resources. § 1. JLtrophy from want of use. I. Functional inutility. (1) Offices in connection vnth the Port of Bruges. — The disorganization of the train service in a besieged city has already been alluded to as ATEOPHY OF INSTITUTIONS FROM LACK OF USE 283 furnishing an example of degeneration by susr pension of function. Another incidence of accidental degeneration is furnished by the oBBce of wharf-porter, which was formerly exercised in the port of Bruges. When Bruges, consequent upon the blocking up of the Zwyn, ceased to be a sea-port town, the wharf- porters who were formerly employed to carry grain, lime, and coal, etc., were no longer required and so abandoned their calling. (2) The forest-courts of E'Mjlwnd. — As an instance of normal degeneration due to the transformation of an agricultural country into a commercial country, take the old English forest-courts. In the middle ages there still existed in England great tracts of forest land which were Crown property, and sub- jected to special legislation conducted by three separate courts of justice : (a) The Court of Attach- ment ^ which instituted proceedings ; (&) the Court of Swainmote ^ before which the culprits were tried, and the Judge's Court presided over by the Lord Chief Justice, who pronounced sentence, and from whose decision there was no appeal. These courts have lost all importance since the seventeenth century, and the forest laws are now only functional ^ When a forest law was infringed, it was the duty of the forester to "attach" the culprit — i.e. constrain him to appear either by seizing his person or his goods. These attachments were then submitted to the Court of Attachment. '^ The judges were called verderers; the jury was composed of foresters of the reeve, and of four men out of each forest hundred. 284 CAUSES OF DEGENERATIVE EVOLUTION with regard to the Forest of Dean and the New Forest. The office of Forest Judge has disappeared, and but few vestiges remain of the courts of attach- ment and Swainmote. Those persons who are con- nected with either the Forest of Dean or the New Forest, meet together still to transact business in connection with them, but these meetings are of no great importance. II. Teansfeeence of function. Instances abound of the transference of a function from one institution to another ; but as a rule, when this change is effected, it is attended by the sup- pression of the old institution. Sometimes, how- ever, this is not the case, and the institution merely atrophies without having been suppressed. (1) Republican institutions under the Roman Empire. — As instances of this form of degenera- tion take the institutions of the Eoman Eepublic, after the establishment of the Empire — or the decadence of the functions formerly discharged by the Privy Council in England, the political functions of which have been transferred to Parlia- ment, and the judicial functions to the Court of Common Pleas and the Court of Exchequer.^ (2) Special jurisdiction in England. — It is the 1 De FranqueviUe, Le Oouivemement et le ParlemerU brUlanique, i., p. 431. ATKOPHY OF INSTITUTIONS FROM LACK OF USE 285 same with special jurisdiction, such as the Eccle- siastical Court, and the University Courts ; they have completely degenerated, their functions having Jbeen transferred to the jurisdiction of the Common Law Courts. In England, however, special juris- diction has not wholly ceased to be functional, for there still exist, side by side with the modern Courts of Justice, a few local Courts and other exceptional forms of jurisdiction which are still maintained in support of certain ancient acquired rights and traditions.^ By far the greater part of these special jurisdictions have, however, fallen into disuse, owing to the creation of the modern Courts of Justice. This happened, for instance, with the following institutions : — (a) The Local Courts of Fevdal origin. — These Courts have decreased both in number and impor- tance since the close of the thirteenth century ; the only vestiges now remaining of them are the Court- leets of certain manors. Sir James Stephen makes special mention of the Court-] eet of the Manor of Savoy, which extended from near the old city gate of London (Temple Bar) up to Cecil Street. Some of the old functions of this Court are maintained in the present day. (6) Country Town Courts. — Twenty-seven local courts are mentioned in modern judicial statistics before six of which no case has been heard for twelve years. On the application of a litigant, ' Idem, Le si'stime Judiciaire de la Grande- Bretagne, i., p. 216. 286 CAUSES OF DEGENEEATIVB EVOLUTION however, any one of these would resume its functions.^ (c) The Gov/rt of Stannaries, the vice- warden of which is appointed by the Prince of Wales. This Court has greatly degenerated in importance and only a few minor cases are now heard before it. (d) The Court of Piepowder . — This was a Court of summary jurisdiction, dealing promptly with disputes arising during fairs and markets. It has almost disappeared from having fallen into disuse. Practically only one example remains ; that is at Bristol, and is becoming merged in another local court called the Tolzey Court. (e) The Husting's Court (or folkmote, scirmote, a kind of County Court of the city of London). — The old Husting's Court has completely lost all its former attributes, in favour of the Court of the Lord Mayor and Sheriffs. Up to 1860, however, it reserved to itself the right- of jurisdiction with regard to matters relating to landed property in the city; the only cases which come before this Court nowadays are those of replevy.^ Finally, there is the House of Lords which ' According tn jurispradence, the fact that a Court still existing by virtue of Eoyal Charter, has ceased to be functional for two hundred years, is no reason against a citizen having his case tried before it, if he so chooses, even though the town authorities declare the funds to be insufficient for payment of the judge. (Case of Rex v. Mayor of Wells, DowHng PrcuMse Oases, p. 562), mentioned by Franqueville. " Franqueville, SyaUmejvd., i., pp. 235, and following. ATROPHY OF INSTITUTIONS FROM LACK OF USE 287 formerly held full powers of jurisdiction with regard to matters relating to members of the peerage ; this jurisdiction is now limited to cases flf high treason or crimes committed by peers to the exclusion of mere acts of misdemeanour which are tried before the ordinary Courts. § 2. Atrophy from lack of resources. The instances we have mentioned are those of institutions decaying because they had become useless, and their resources were transferred to other objects. The lack of resources was the result of the lack of function. Sometimes, however, it is the cause and not the result, in which case atrophy may be due either to an abnormal development, such as the hypertrophy of another institution, or to poverty ensuing on the general decline of society in general. Of course both factors may act simultaneously. 1. Local administration ai the close of the Roman Empire. — The decline of local government at the close of the Eoman Empire is an instance of atrophy ensuing from the ultra-development of another institution. As the demands of the central powers grew more and more excessive, the fiscal rates had to be augmented in order to meet them, and the curiales, which consisted of members of the City Council who were made responsible for the payment of 288 CAUSES OF DEGENERATIVE EVOLUTION taxes, finally found it was quite impossible to meet their engagements, and made every effort to leave the curia.^ 2. The degeneration of Societies in all their parts. — A number of instances might be mentioned of general social degeneration bringing about the atrophy of some one or other institution in particular. Besides giving classical examples, such as the Eomans, Peruvians and Astecs, V. Lilienfeld men- tions the decline of the Negro kingdoms which existed during the sixteenth and seventeenth centuries in Southern and Western Africa, and which are merely represented nowadays by wretched little tribes.* There are, according to Waitz, at some distance from Carimango (the equatorial Eepublic) some people of pure Spanish blood who have fallen back into absolute barbarism. Their language is de- formed past recognition, and their manners and customs exhibit no traces of their former condi- tion.* Space precludes us from dwelling further upon the various causes — often complex and obscure — which bring about the downfall of societies, suffice it to say that they are connected with 1 Laviase and Rambeaud, Sistoire gindrdle, I., ch. i., pp. 14aDd following. ^ Von Lilienfeld, Gedanke iiber de Sozialimssensch({ft der Zuhmft, ii., p. 241. ' Waitz, Anthropologic der Naiurvolker, 1. B., p. 86D, ATKOPHY OF INSTITUTIONS lEOM LACK OF USE 289 territory br with population, the two factors of social evolution. Either the physical surroundings of institutions undergo unfavourable transformations, or else the population itself degenerates. (1) The almost complete disappearance of the great family communities {zadrugai) of Montenegro is a characteristic instance of atrophy from lack of resources caused by the impoverishment of physical surroundings. The persistent cutting down of trees in the Black Mountain has had a disastrous effect on the water supply, and consequently upon the fertility of the ground. Most of the zadrugas, having found it impossible to continue their existence in common upon the same territory, have split up into small families (inokosnas). These latter represent, in a reduced state, the old family system from which they have sprung. Bogisic has shown that these in no way resemble, our modern families, but are to be regarded, from the judicial point of view, as reduced family com- munities, each compriatfd of only a few persons. (2) Other cases occur where the atrophy of an institution — of an artistic or scientific society, for instance — is brought about by the degeneration of a population which ceases to be interested in the society and no longer contributes to its support. A large number of cases of this kind might easily be mentioned, especially as occurring during the period of the Byzantine Empire, but it is difficult 290 CAUSES OF DEGENERATIVE EVOLUTION to account for the sudden degeneration of a people where the physical surroundings had re- cently undergone no special modifications, and when there had been no sudden and violent check upon social development. According to Lapouge and other sociologists of the Darwinian school, this social degeneration was merely the outcome of hereditary influences. The destiny of a nation is dependent upon the quality of the elements of which it is composed and by which it is directed. If a nation is rich in energetic and intelligent qualities, the greatest of disasters can only have a transitory and limited influence. When the contrary is the case, the same circum- stances may produce an arrest in development or a complete decline and fall. Up to the present time, and especially in antiquity and the middle ages, these favourable qualities were gene- rally supplied by a dominating minority establish- ing itself in a conquered country. In the common course of evolution, these superior elements, which are indispensable to social progress, are eventually eliminated. The inferior elements regain greater power, and each step of their progress is attended by a backward step towards barbarism. Although, at first sight, this seems contrary to the Darwinian theory, it is strictly in accordance with it. The superior individuals are relatively inferior when their chances of success or of posterity diminish. The superior individuals may not only be swamped ATKOPHY OF INSTITUTIONS FKOM LACK OF USE 291 by a diminution in their birth-rate, but in some cases there may be a direct elimination of them.^ The tendency of decadence is always towards the degenerative and eliminatory selection of superior elements. It may be said in conclusion that there are constant calls upon the capital and labour of a society from its various institutions, and the conse- quence is that, the resources not being unlimited, a regular struggle for existence goes on amongst the institutions. In the course of this struggle, the decline of an institution may be brought about in two different ways. It either begins to de- generate from lack of sufficient means of support, or degeneration sets in consequent upon the insti- tution having ceased to be functional by inutility, by transference of function to another institution, or by obstacles placed in the way of exercising that function. In either case the institution dis- appears. It is only in exceptional cases, which will be alluded to further on, that existence is still maintained. ^ 6. de Lapouge, La Vie et la Mart des nations (S^mie ini. de Sociologie, 1894, pp. 421 and following). Several terms used in this treatise were borrowed from tlie above article. See also Hovelacque and Herv^, Precis d' Anthropologie, p. 189 : ' ' War, in its double consequence of the elimination of the strong and the survival of the weak, is for the more civilized races a powerful factor in the cause of degeneration and downfall." PART II THE CAUSES OF THE PEESISTENCE OP ORGANS OR INSTITUTIONS WITHOUT FUNCTION CHAPTEE I SURVIVAL OF ORGANS We have shown how and why oi-gans may become rudimentary and tend to disappear. In many cases the disappearance is complete ; and the organ may not even reappear temporarily in the course of the individual development. This disappearance is, however, by no means universal. Even apart from the phenomena of recapitulation, rudimentary organs may persist in the adult, and sometimes, even although organs have ceased to be functional, they persist without degenerating. We have now to con- sider why in such cases degenerative evolution does not result in complete obliteration of such organs. § 1. JJnfunctional orgains that are not rudimentary. Absence of Variations. — There are some plants such as Ficaria^ ranunculo'ides and Zysimachia 1 Lysimachia Nwnmularia occasionally produce seeds in some valleys of the Pyrenees, and Errera has shown us specimens grown from seeds coming from the shores of the lake of Quatro-Cantons. SURVIVAL OF ORGANS 293 Nummularia, the flowers of which hardly ever produce seeds. How is it that in such species flowefs are still produced ? The probable ex- planation of this anomaly is, that for the disappear- ance of flowers there would have to be produced individuals with this advantageous variation. It is the case, however, that the Ficaria and the Lysimachia reproduce most actively by asexual methods, and variations are extremely rare in cases of these modes of reproduction. The result is that these species having begun to form sterile flowers continue to produce them through simple lack of variation. An analogous case is presented by Modea Canadensis. This unisexual plant is represented in Europe by only female plants. These plants have multiplied asexually so luxuriantly that in Holland they began to choke up the canals, and it became necessary to make provision in the budget of that country for the extermination of the pefet. The plants are, of course, able to multiply only asexually, as the female flowers cannot be fertilized, and these useless flowers have been maintained simply from the absence of variations. Stratiotes alo'ides, a plant belonging to the same family as Modea, is practically only repre- sented by male individuals. Females are extremely rare, and none the less the male flowers are pro- duced, although in the vast majority of cases they must be useless. 294 CAUSES OF DEGENEKATIVE EVOLUTION W. Burck has called attention to, without endeavouring to explain, other instances of the persistence of functionless organs. A ■ large number of Anonaceae bear flowers which do not open, and which are self-fertilizing (cleistogamous flowers). None the less, they have retained the corolla, the original purpose of which was to attract insects.^ Burck has called our attention to the circum- stance that several species belonging to the same genus produce cleistogamous flowers, and that it is improbable that this condition has been acquired independently by these species. One would thus have to admit that the original type must be very remote, as it has given rise to descendants of specific distinction, and yet the useless corolla has persisted through the long series. Parallel examples may be found among animals. Maehaerites is an insect which inhabits caves in North America. The females are quite blind ; the males, on the other hand, have preserved, or seem to have preserved, well-developed eyes ; but are these eyes real? An abyssal fish, Ipnops,^ seems as if it had enormous eyes extending from the corner of the snout some distance along the neck, but these organs are not really eyes; they ' W. Burck, Ueber Kleistogamie im weiteren Sinne und das Knight- Darwinsehe Gesetz {Ann. Jard. Bot. Buitemorg, viii., p. 122, 1889). ^ Dollo, La Vie au sein des men, p. 242, Paris, Bailliire, 1891. SURVIVAL OF OEGANS 295 are light-producing organs, and the fish are in reality blind. This may he the case also in the males of Machaerites. It may also be the case that the male has an opportunity for using eyes absent in the case of the female, the males some- times leaving the caves, the females remaining within theni. Something analogous to this occurs in the case of eels : the males remain always in the sea while the females rejoin them only for purposes of reproduction. Moreover, there is still a third hypothesis, that the male of MachaeHtes be- came an inhabitant of caves later than the female, and has not yet had time for the loss of its eyes. § 2. Uhfimctional organs which persist as rudiments. It is outside our purpose to discuss here the numerous cases of organs reduced through adapta- tion, such as the leaves reduced to serve as protectors of young buds {Phyllocactus, fig. 78), or the wings of the ostrich which, although much reduced, are supposed to assist the bird in running. The utility of such organs explains their persist- ence; we are concerned here with organs which, although useless, persist in a reduced form. 1. Absence of Variations. — In discussing organs which, although without function, have persisted in a complete state, we attributed the persistence to absence of variations. It is probable that the same cause operates in maintaining useless vestiges. 296 CAUSES OF DEGENEEATIVE EVOLUTION It is to be noticed, however, that the variability of vestiges is frequently considerable. The flowers of Asparagus^ officinalis are sometimes, although rarely, hermaphrodite. Usually they are uni- sexual and exhibit the organs of the other sex in every conceivable stage of degeneration. It is probable that the unisexual condition has been acquired recently, and that there has not yet been time for the operation of natural selection to cause the disappearance of the useless organs.^ ' We have actual knowledge of the mode of disappearance of an organ in one case, and can see the part played by variability. In Phyllanthus speciosus {Xylophylla arbuseida) the adult plant has three kinds of branches : vertical branches, with rudimentary leaves ; oblique branches which spring from the axils of these leaves and themselves bear in two rows very rudimentary leaves ; and flat branches which are the chief organs of assimilation of the plant and which also bear rudimentary leaves in two rows. In the seedling, on the other hand, there are formed after the two coty- ledons, one or two completely developed assimilating leaves upon an upright stem (iig. 84, a). The flat branches grow from the axils of these, and bear, unlike the fiat branches of the adult, assimilating leaves ; higher up the vertical stem Dears only rudimentary leaves with flat branches in their axils. It may happen, however, that the seedlings bear, directly after their cotyledons, rudimentary leaves (fig. 84, b) and in this case the reduction of the leaves is present not only in the leaves borne upon the vertical stem but in those borne on the flat branches. These latter bear only a small number of assimilating leaves. Thus, we have in this plant a remarkable example of incomplete recapitulation : the seedling preserves in a functional condition organs that are rudimentary in the adult, but the species furnishes instances where these leaves cease to be functional even in the seedling. SUETIVAL OF OKGANS 297 2. Insignificance of the Rudimentaey Oegan. — It frequently happens that rudimentary organs are preserved simply on account of their insignificance : the absence of organs so small would not be an advantage to the plant sufficiently great to be laid hold of by natural selection. Many species of Tropaeolum bear leaves without Fig. 84. — Two seedllrgs of Phyllanthus speciosu^. 1, 2, 3, 4, successive assimilatmg brauchos ; c, cotyledons. stipules. In Tropaeolum inajus there are stipules only in the case of the first two leaves of the seedling, and the position of these stipules is very variable. Sometimes both stipules are at the base of the petiole ; sometimes only one is present ; sometimes both are several millimetres from the base — a position in which functional stipules never occur. 20 298 CAUSES OF DEGENERATIVE EVOLUTION In the same way may be explained the persist- ence of accessory rudiments of enamel organs in the development of teeth. Besides the rudiments of the enamel organs for the milk teeth and permanent teeth, there are additional organs present in a very variable condition and number, nearer the external surface. They are, however, very generally present, and are exceedingly similar to the youngest stages of the normal enamel organs. Kollmann and Gegenbaur believe that they are abortive rudiments surviving from an ancestral condition in which teeth were more numerous. CHAPTEE II THE SURVIVAL OF INSTITUTIONS We have seen that an institution which ceases to be functional without dissolution — either voluntary or enforced — ensuing, tends to atrophy and dis- appear, while its resources are appropriated by other institutions. It now remains to account for the fact that this atrophy does not generally end in total disappearance. The two following hypotheses may be made : — 1. The structure of a non-functional institution may remain intact. 2. The institution may survive, but in a rudi- mentary condition. SURVIVAL OF INSTITUTIONS 299 § 1. The ijdegral persistence of an institution. A non-functional institution may survive and retain its structure and resources in the following cases : — 1. By the intervention of some superior authority to prevent its suppression. 2. When, while ceasing to he functional, it con- tinues to be useful, though indirectly so. 3. When its existence is maintained out of respect for old traditions. We will take these three hypotheses in succession, but it must be borne in mind that when a non- functional institution is maintained out of respect to tradition, or by virtue of an indirect usefulness, it is always by the intervention of legal authority. This legal authority, however, amounts to an ex- pression of the public will, whereas an institution may also be maintained by the exercise of personal influence on the part of some one person. It sometimes happens, too, that a non- functional institution continues to survive because its suppres- sion would entail important changes in other parts of the social organization. 1. Maintenance by compulsion. — A useless in- stitution is frequently maintained by compulsion, when its conservation is advantageous to those con- nected with it, or even to other persons.^ The following are a few examples of this : — ' Spencer, Principles of Sociology, vol. iii. 300 CAUSES OF DEGENEKATIVE EVOLUTION (a) A long list might be made of all the sinecures, now quite useless, that some governments insist upon maintaining for the advantage of those occu- pying the posts. Such were certain offices in con- nection with the Court in former days or the avoueries of the end of the middle ages. " Like the Fief system," says Errera, when writing about the Massuirs, " The avouerie afforded an effec- tive protection — military as well as judicial — against the various dangers arising in a still barbaric age. But, in the course of the last centuries of the middle ages, the obligations of feudal chiefs and the condition of avouerie disappeared ; the reasons were that relative security was attained; militia was established, and the army, under the command of the sovereign himself, became better disciplined ; and that there arose the organization of the justices scdbinales, of bailiwicks, and of superior courts of justice. However, although the ancient offices dis- appeared, the emoluments attached to them con- tinued to be drawn.^ (6) It often happens that institutions which have ceased to be functional are yet maintained as being a source of profit not only to those in direct con- nection with them, but to a considerable number of other persons. Before the Eeform Bill of 1832, when large towns like Leeds, Birmingham, and Manchester were unrepresented in Parliament, the House of ' Errera, Les Mansuira, p. 75. Brussels, Weiasenbruch, 1892. SURVIVAL OF INSTITUTIONS 301 Commons contained seventy members, nominated by thirty-five rotten boroughs in which there were no electors, and ninety members nominated by forty-six boroughs, containing less than fifty electors. The borough of Old Sarum was a mere hillock belonging to Lord Canalford ; Gatton and S. Michael had only seven electors ; the borough of Dumwich had been long since submerged by the encroach- ment of the sea; Beeralston, belonging to Lord Beverley, consisted of one house, and Castlerising of two. In the county of Bute there were twenty- one electors, only one of whom was a resident and who nominated himself. The preservation of this system of representation which had long ceased to be adequate, was eminently favourable to the few persons who benefited by it, and they vigorously resisted the passing of the Eeform Bill. (c) After the provincial states of Normandy and the Dauphiny had been suppressed, the state func- tionaries retained their titles and emoluments. In the Dauphiny the representative institutions ceased to be functional in 1627, but at the close of the eighteenth century the Bishop of Grenoble con- tinued to receive 6000 livres per annum as primate of the dominion. Two barons, delegates of the nobility, shared a similar salary, and the Syndic of the province and two secretaries received lesser emoluments from the province which continued 302 CAUSES OF DEGENEKATrSTE EVOLUTION to make these payments for services no longer rendered.^ In each of the above cases of survival it is plain that compulsion intervened — more or less directly — to secure the maintenance of non-functional institutions. In. the first cases referred to, the privileged persons concerned, took advantage of their influ- ential positions to enforce the maintenance of their sinecures. In the second case — that of the rotten boroughs — the institution was not only advantageous to the member himself but to the whole of his party, so that naturally its maintenance met with the cordial support of the latter. In the third case, those in authority maintained part of the institution — that of the mere titles and emoluments — in order to suppress the remainder more easily. In this case it was not an institution which atrophied, but an institution which was caused to atrophy by compulsion. 2. Indirect usefulness. — It sometimes happens that an institution, although ceasing to be functional, yet retains a certain usefulness. This is so in England, with the office of the Privy Seal. All the functions formerly discharged by the holder of this office have long since disappeared, but the post is reserved as a sinecure for persons who have ^ Babeau, Les assemblies des pays d'Etat sous I'ancien rigime ; B^foi-me sociale, 1893, p. 701. SURVIVAL OF INSTITUTIONS 303 distinguished themselves in politics, but who from advanced age are unable to take an active part in public affairs. A great many sinecures are main- tained for a similar purpose — that of furnishing practical though not nominal pensions to distin- guished persons in art or science. " It may well be," says VioUet, " that an institution, which at first sight seems to be a mere useless wreck, is really of immense service to society. The deep roots of a dead tree may continue to furnish support to a new structure." It is by reason of this negative usefulness that — according to Bagehot — the English monarchy has been preserved, forming as it does a symbol of unity and coherence amidst the electoral changes of power. The English monarchy offers a char- acteristic instance of an institution which has lost nearly all its former functional importance, while nominally retaining its power. According to Bagehot, the Queen would now have to sign her own death warrant if condemned by both Houses of Parliament. The outer form, however, remains almost the same as in the days of absolute monarchy, when the sovereign took an active part in public affairs. At within the last few years, the Queen's signature was required to all official documents. It was only in 1862 that a law was passed deciding that for the future, promotions in the Army or Navy should be signed either by the Commander-in-Chief, or by the Secretary of 304 CAUSES OF DEGENERATIVE EVOLUTION State. At that time, by dint of hard work, the Queen had signed all commissions up to 1858, and there remained 15,931 documents of this kind still unsigned.^ 3. Respect for Tradition. — The persistence of some institutions can only be accounted for by a lack of invention. Mommsen calls attention to a remarkable instance of this kind in the history of early Rome : When a government by praetors — as consuls were first called — was substituted for a govern- ment by kings, the new system remained the same in idea though nominally difl'erent. The old idea of royal authority survived for a long period, and the praetors enjoyed all the old kingly powers, even those in plain contradiction to the temporary character of their office : the king could not be deposed, but neither could the praetor be con- strained to depose himself ; the king, when dying, nominated his successor himself, and this power remained to the praetor — although the system of election by the comitia had been introduced — for the praetor had the right of excluding whoever he chose from the list of candidates, and also of annulling the votes given to those who displeased him. It was only at a later period that a logical 1 Bagehot, The, English Constitution, pp. 57 and following; London, 1891. De Franqueville, Oou/oemement ei Parlement Britanniques, i., p. 251. SURVIVAL OF INSTITUTIONS 305 and consistent conception of the consular authority came to be formed.^ In the same way the maintenance of the insti- tution of sheriffs in modern England can only be attributed to an exaggerated respect for tradition. In the commission which sat in 1888 to inquire into judicial organization, one of the commissioners expressed himself as follows : " I cannot see that sheriffs are of any use whatever, unless it be for show ; there is not one single function attached to the post which a sheriff can fulfil himself. I am a sheriff, but I do not know what a sheriff's duties are." The institution, however, remains intact. The sheriff is credited with the discharge of several functions, which are executed in his name and on his responsibility. All he does is to receive the judges, accompany them on circuit, and preside over parliamentary elections.^ The mode of nominating the sheriffs has remained unchanged since 1340. The Queen, by means of a traditional gold pin, is supposed to prick by chance in the list of candidates which is presented to her, the name of him upon whom she confers " the ' Ferrero, Simbolii, p. 58, Torino, 1893. 2 "Tho two sheriffs of London, who are elected from among the liverymen of the varions city companies, have not to go on circuit, but are supposed to attend at the Central Criminal Court. Their functions chiefly consist in escorting the Lord Mayor to all city ceremonies, and in attending numerous banquets, some of which are given at their expense. " De Franqueville, Sydeim judiaiaire de la Grand- Bretagne, i., p. 611. 306 CAUSES OF DEGENERATIVE EVOLUTION charge and keeping of the county." This ceremony is known as the " Pricking of Sheriffs." § 2. The Survival of Institutions in a keduoed STATE. It has been shown that a number of institutions in a state of decline continue to be maintained, because they are still useful in spite of their reduced condition. This is the case with the symbolic ceremonies which in former days accompanied the drawing up of solemn contracts.^ "We now come to institutions which persist in a reduced condition, but which are not directly use- ful to any one. In this case their persistence may be attributed to one of two causes: either to respect for tradition or to the insignificance of the vestiges which remain. ^ According to VioUet in Histoire du droit dml fr. , p. 607, the primitive assemlily of the people still survives, though in a reduced condition, in the Roman mawApatM, and in Scandinavia in a solemn form of sale called the scotatio. " I helieve," he says,' " that it was the primitive sale of German law a sale concluded and ratified in a popular asseml>ly, that gave rise, in the middle ages, to the Scandinavian scotatio. So also in the case of the Roman mancipalio there has long been believed to exist the remnants of a popular assembly. The dumb witnesses in the scotatio appear to me the petrified representatives of the German tribe or village ; and, in the opinion of good judges, the witnesses in marwipatio are no other than symbolic statues of the five classes of the Roman people. However, this is mere li^'pothesia. SUEVIVAL OF INSTITUTIONS 307 It is difficult, however, to distinguish between these two causes, for insignificant vestiges of institutions are especially numerous in very con- servative environments, and, on the other hand, mere respect for tradition very rarely ensures the maintenance of either a harmful or an expensive institution. 1. Insignificance of the Institution. — Some of these insignificant institutions are reduced to mere vestiges, no longer functional, or of use to any one, others — representing traces of a former system — retain some local vitality, although not in keeping with the new conditions ; these are of such small importance that their very insignificance ensures their survival. (a) The two French laws passed in 1835 and 1849 relating .to entail have never become func- tional at La Martinique. The old law of entail is still active there, and this state of things is allowed to continue without authoritative interference.^ (&) In England, the sovereign, up to the close of the sixteenth century, reserved to himself the right of presiding over the Courts of Justice, and of pronouncing sentence. Since the Revolution, no sovereign has essayed to render justice personally, and any attempt of the kind would be regarded as unconstitutional nowadays. Vestiges of the old system remain, however, in the formula of certain legal proceedings, such as the serving of warrants, > Viollet, Hist, du dr. civ./r., p. 883, 308 CAUSES OF DEGENERATIVE EVOLUTION in which an order is given to appear before the Queen herself, and it has never been suggested that the old formula should be altered.^ The following are a few examples of local sur- vivals which owe their existence to their insignifi- cance : — (a) Although Cambray has belonged to France since the close of the seventeenth century, its diocese includes a small part of Belgium ; the ancient religious organization has in this instance survived political changes.^ (b) It is not generally known that with certain properties situated in Artois — consisting chiefly of marsh land — the law of primo- geniture still holds good.* (c) Public attention has recently been directed to a very curious survival of the political connections which existed during the middle ages between B^arn and Spain. Every year, on the 13 th of July, the inhabitants of the French valley of Baretous in the Pyrenees, solemnly pay a tribute — for the ' De Franqueville, Systime judiciaire de la GrandeSretagne, p. 23. 'VioUet, ffist. du. dr. civ.fr., p. 882. ' Errera, Les Masuirs, p. 290. Decision of the Council of 25th B'ebruary 1779, as regards the eastle-wards of Lille, Douai, and Orohies. " Those portions of land which fall or have fallen to each Inhabitant as the result of division, shall be inalienable; no person shall possess two portions. The eldest male of each family, or in default of the eldest male, the eldest female, shall alone inherit the said portions of land." SURVIVAL OF INSTITUTIONS 309 maintenance of peace — to the inhabitants of the Spanish valley of Eoncal.-' It is plain that this custom, which has gone on so long as to pass unnoticed, is too inconsistent with the present relations between France and ^ The ceremony commences at nine o'clock in accordance with instructions laid down in a document which, according to the mention made of it in the Procte-verbal, dates from 1375. First, the French mayors don their scarves of office ; next the Spanish mayors, advancing from a group of compatriots, proceed towards the frontier boundary line, accompanied by a peasant carrying a lance with a red flag tied to it— a symbol of justice — and draw up within six yards of the boundary. The French mayors follow suit, but the flag hoisted on the lance which precedes them, carried by a peasant, is white, as signifying their pacific intentions. The Mayor of Isaba then says to them, "Is it peace?" The French mayors reply in the affirma- tive, and, as a proof of their sincerity, couch their lance upon the mile-stone marking the boundary. The Spaniards then first plant their lance in French soil leaning against the stone, and after- wards place it so as to form a cross with the French lance. Next the Mayor of Arette places one hand upon the crossed lances, and the Mayor of Isaba does the same, and together they utter the formal declaration of peace, which all those present swear to observe. After the vow, the Mayor of Isaba cries three times, " Paz davans ! " which means, "May peace con- tinue." Peace being thus declared, the Roncalais, in order to ratify their abandonment of hostilities, order the guards to remove their arms from the French side. The ceremony being over, it only remains to pay the blood-tax. This formerly consisted of three white mares, all exactly alike ; but, owing to the great difficulty of matching them, three unblemished heifers were sub- stituted, all of same colour and with the same markings. These three heifers cost about 600 francs, which is a large sum for that part of the world. 310 CAUSES OF DEGENERATIVE EVOLUTION Spain to be continued much longer. " It is to be expected," says the paper from which this information was obtained, " that the French Foreign Minister will in the near future come to an understanding with the Spanish authorities to put an end to this iniquitous custom, and it is to be hoped that this year is the last occasion upon which a blood-tax will be paid by the valley of Baretous to the valley of Roncal." 2. Respect for tradition. — In his " Essays on Pro- gress, Manners and Customs " ( Westminster Review, 1854), Spencer points out the connection between respect for tradition and custom, and the con- servatism of those in authority. He says that certain customs, which have elsewhere died out, survive in some departments of the government. The Secretary of State in ratifying acts passed in Parliament uses old Norman French,^ and certain legal terms in old Norman French are still used. The wigs now worn by judges and barristers are identical with those seen in old portraits, while the " Beef-eaters " of the Tower of London wear the ^ " For financial Acts the formula is : La Reyne remercie ses bons sujets, acoepte leur b&^volence et ainsi le veult ; for general Acts : La Reyne le veult ; for private bills : Soit fait comme il est d^sir^ ; for petitions : Soit droit fait oomme il est d^sir^. The veto is announced : La Beyne s'avisera. Cromwell had changed these old forms ; he gave his consent to Bills in English ; the old custom was resumed at the Restoration, and the House of Commons in 1706 rejected a Bill passed by the Lords to abolish the French phraseology r" De Franqueville, Oouvernement et Parlemeid Sritan- niques, vol. i., p. 279. SUEVIVAL OF INSTITUTIONS 311 same costume as that once worn by the body-guard of Henry VII.i Two similar examples may be added : — (a) At the coronation of English sovereigns two gentlemen of the Privy Council, chosen " on account of their appearance," and created knights for the occasion, are appointed by the Lord Chamberlain to represent the Dukes of Aquitaine and Normandy. (b) The First Officer of the Crown was formerly the Lord High Steward, which title having, in the course of time, become purely honorary, was hereditary in the family of the Earls of Leicester. The post is now in abeyance, but, on the coronation of a sovereign, or, on the occasion of a peer being placed upon his trial, this dignity is conferred upon some important person nominated solely for the occasion. It is not only in ceremonial — which, according to VioUet, is the museum of history — that reduced institutions, which are completely useless, are tenaciously maintained ; it is the same with iudicial and religious institutions. A few examples will suffice to show that this is so : — (a) In its primitive form, the Assembly of the People included the whole army, and was necessarily held in some large open space. The custom survived the necessity of the choice of some such spot, and up to the sixteenth century, whenever a new Emperor was proclaimed in Germany, it was the ^ H. Spencer, Morals, Science and Art. 312 CAUSES OF DEGENERATIVE EVOLUTION custom for the electors to proceed to some mountain for the purpose, probably because it was the custom in former days to hold a general meeting there, before the election. In Iceland, the althing — consisting of two chambers now — was formerly one large assembly held in the open air upon the Logberg (the Mountain of the Law) near to the Lake of Thingvellir. In the Eepublic of Andora, criminal sentences are still pronounced with great solemnity from the Market-Place.^ (6) Among the Ossetes, where the family com- munity still flourishes, the only inalienable and unsaleable property is not the real property, but certain personal possessions, such as the great cooking-pot with the chain by which it is hung over the fire. " At first," says Kowalewsky, " this may appear a strange custom, but it must be remembered that possessions of that kind were of equal importance to the Ossetic 'family' as were their tombs to the ancient Greeks and Eomans, which ex- plains how any infringement of the * In Montenegro, it was the custom, till recently, for the Prince himself to render justice, sitting under a tree in front of his palace at Cettigne. This mode of jurisdiction, which was probably a survival of the old system of which we have heen speaking, has not completely disappeared. SUKVIVAL OF INSTITUTIONS 313 custom is regarded as an infamy by the Ossetes." ^ We now come to cases of purely religious survival, which offer the strongest resistance to the inroads of change. Spencer instances the custom of circumcising with a knife made of flint, and the vestiges remaining in Catholic worship of former primitive religions. The Eucharist, as we have already pointed out, is reminiscent of real sacrifices, and the symbolic representation by a dove of the Holy Ghost is only a rudimentary form of zoolatry.^ In Belgium there are still traces of the old custom of sacrificing an animal upon the com- pletion of a new building, with the idea that the animal's spirit will protect the edifice from harm,* and if the observer of the following facts is correct in his interpretation of them, there also remain in Belgium traces of the ancient sacrifices to the genius of the earth : — " It is the custom, round about riorenville (in the Belgian Ardennes), to offer a sacrifice to the presiding genius of the road upon the con- struction of a new road or railway. It is usually a fowl, or a rabbit, or even a calf which is sacri- ficed. ... In some parts of Luxembourg animals ' Kowalevsky, Droit coutv/mier Ossetien, p. 105. ^ Spencer, Principles of Sociology. ' Folklore, Wallon, No. 1526, p. 115 (Bulletin de Folklore, ii., 177). 21 314 CAUSES OF DEGENEEATIVE EVOLUTION are also offered in sacrifice to the genius supposed to preside over a newly-purchased field, with a view to ensuring abundant crops.^ Eemains, either undoubted^ or only probable,' of phallic worship, are scattered throughout Europe. In Brittany* and in Belgium,* for instance, strange old customs still exist showing that here, ^ Rivue des traditions populaires, 1893, p. 394. ^ Th. Volkov, Bites et usages nupticmx en Vlcraine {I'Anlhro- pologie, 1891, p. 167). Only a short time ago it was the custom in Tver, on the day dedicated to Yarilo (the phallic God of Spring), for the parents of young daughters to send them to join in games similar to those of the ancient Slavs, with a, view to their getting married. ' Note sur wn, vestige du culte de la terre mire (phallism) en Frovence, hy B6renger-F&aud {Biime d'Anthrcfpologie, 1888, p. 563). " At Luc, in Provence, upon the Ist of May, which is a country holiday, the young girls proceeded to a place where two roads met. Here they assembled around an olive tree, and after each dance they slnick the olive tree three times with their backs. "This fete, a survival of the floral fStes of the month Of May which are still celebrated in Provence and Italy, continued to be held until quite recently, and appears to have been a lingering vestige of the ancient worship of creative Nature, Mother-earth — in short, of phallic worship. " The three knocks given by the young girls to the tree trunk is a survival of the ancient virginal sacrifice to the phallic emblem. The original meaning was not quite lost, for the Pro- ven9als still realized, though vaguely, that the three knocks were \ somehow connected with the idea of marriage." * "Zes MigalUhes de Locmariaqiier et de Camac, et Us amours, by Bonnemfere (Bivue des traditions populaires, 1894, p. 123). "In former days it was the custom for all the young women who wished to get married to climb (on the night of May 1), to the top of the great menhir where they lifted up their clothing that their bodies SURVIVAL OF INSTITUTIONS 315 as elsewhere, a belief in the influence of fetiches once prevailed, and particularly in the form of megalithic monuments relating to the fecundity of women. The custom still prevalent in the African Congo, of driving a nail into a fetich, with the view of reminding it of a request, has not disappeared from Europe. The young men of Couvin (Namur) still stick pins into the wooden saints of the little chapels round about in order to draw a lucky number in the military lottery, and young girls in Brittany do the same with a view to getting inigtt come in direct contact with the stone, and then slid from the top to the bottom." At Carnao, young girls wishing to marry undressed completely upon the same night, and proceeded to rub their abdomen against a' special menhir. In that part of Brittany, where only French is spoken, similar customs have equally prevailed. " File de Notre-Dame de Eide-cul (Jules Lemoine, in the journal Le petit bleu, of October 18, 1896). Similar to the above Breton custom is that of sliding down the Rocher Eide-cul, which is situated near to Landelies, in the valley of the Sambre. Here, as in many other places, old customs have been Christianized, and a Christian chapel now stands close to the ancient shrine. Young people of both sexes used to seat them- selves upon the top of the stone U])on little fagots of boxwood gathered in the neighbourhood, and then slide down to the bottom. According to the old saying an upset meant waiting ; an em- brace signified mutual affection ; a collision, indifference ; and an embrace followed by rolling over indicated matrimonial suitability. Similar customs prevailed at Trou-deux-Trous, situated near to the Rocher Eide-cul. These two megalithic temples disappeared about forty years ago, their materials have been used as fluxes in smelting works. 316 CAUSES OF DEGENERATIVE EVOLUTION married/ and in Belgium,^ as in Brittany,^ if the image prove too hard, the reminding pin is stuck instead into a fissure, or into the door of the niche containing the image. Besides these mere vestiges of pre-historic customs and belief, by referring to certain illustrated documents bearing upon the subject, it would be easy to work out the connec- tion between the worship of saints which is pre- valent in Belgium in the present day, and the pagan worship of the ancient Celts and Germans.* '^Bulletin de Folklore, i., 250-251. * Ibidem. ' The menhir of the Pierre-Frite in the valley of Lunain. In nearly every hole or fissure of this monument, a nail or pin has been stuck by the young people of that part in the belief that it will ensure them a speedy marriage. {Bivue des traditions popu- laires, 1893, p. 448.) ^See for Saint Eloi, Mdlusine, viii., 122-132 ; for Saint Martin, Bulletin de Folklore, i., 309-315; for Saint Hubert, Gaidez, La Ba eet Saint Hubert. PABT III EESUMi AND CONCLUSIONS When an institution or an organ ceases to be functional or in any way useful, it very soon disappears altogether. If, as happens in some exceptional cases, it persists, it is because neither of the chief factors in causing atrophy, variability or selection, have intervened. Sometimes the vestiges are of too insignificant a nature to call for their removal by either artificial or natural selection, and sometimes their existence is ensured by the lack of variability, as in the case of the persistence of flowers in plants which multiply asexually. This absence of varia- tion occurs equally in the social domain, especially in matters connected with religion, wherein ancient customs are credited with a divine origin, Eeligions may pass away, philosophies may be transformed, and old beliefs cease to prevail, but the remnants of old creeds, conveyed by popular tradition through the centuries, defy destruction by modern innovations. The ancient winter festival, on which day the 318 CAUSES OF DEGENERATIVE EVOLUTION dead were supposed to leave their graves aud join the living in a feast around the family hearth, is still celebrated in the keeping of Christmas and in the various customary practices on the first two days of November. The May-Day festivals — pagan festivals held in honour of vegetable and human fecundity — are still held in their early form round about Loemariaquer and in the village of Campine. Traces also remain in the picking and wearing of flowers on the 1st of May, and the same day is selected by the socialists for the celebration of their near approach to a life under freer and happier conditions. This survival of festivals, customs and traditions, while the religions and civilizations which produced them have passed away, is the principal link which connects us with bygone generations. " Their value lies," says Houzeau in his Etvde de la Nature, " in the establishment of a chain between successive generations. The memory of au indivi- dual may be regarded as constituting his personality. Take from him the memory of his past, and he is left at a point in time wherein there is no stability and complete isolation. To be himself, a man requires not only his recollections, but a knowledge of his past habits and traditions. When a savage is removed from his fellows and transported to new surround- ings in a distant country, he loses all knowledge of his former condition. Society itself, made up as it SUMMAKY AND CONCLUSIONS 319 is of customs and prejudices, constitutes history. The mirror of the past is exhibited in the conscious- ness of the collective individual which is called a nation. What link shall we have with former generations if not a heritage of their ideas — i.e. of their discoveries and their mistakes ? Nations, like individuals, are continually modifying, this inherited legacy, but, like the individual, they cannot get away from it without breaking the thread which has made them themselves." GENERAL CONCLUSIONS All evolution is at once progressive and retro- gressive. All modifications of organs and institutions are attended by retrogression. This occurs equally in the modifications of organisms and of societies. All existing forms, whether organic or social, have undergone certain modifications, and, as a result, have lost some parts of their structure. This universality of degenerative evolution may be proved either by the comparative method, or by showing that all organisms contain rudimentary organs, and that all societies contain survivals. II Degenerative evolution follows no definite path, and can in no way be regarded as constituting a return to the primitive condition. In some cases — when one cause of dissolution equally and simultaneously affects all the parts of an institution or an organism — the most complicated and delicate structures are the first to disappear; but it must not be taken as a general principle that the most complicated structures are necessarily 320 GENERAL CONCLUSIONS 321 the most recent, and that consequently degeneration always retraces the path of progress. Evolution is irreversible, and accordingly, with a few more or less obvious exceptions, we draw the following conclusions : — 1. That an institution or an organ which has once disappeared never reappears. 2. That an institution or organ once reduced to the condition of a vestige cannot be re-established and resume its former func- tions. 3. Neither can they assume fresh functions. Ill Degenerative evolution is brought about by a limitation in means of subsistence — either in nutriment, capital or labour. In biology the prin- cipal if not the sole agents in its accomplishment are the struggle for existence between the various organs, and the struggle for existence between the various organisms. In sociology it is artificial selection which is the dominating agent, and natural selection plays only a secondary part. The occasional causes of degenerative evolution are inutility of function, insufficiency of nutri- ment or resource, and (in biology only) lack of space. 322 GENERAL CONCLUSIONS An institution or an organ which has ceased to be fuactionalj and has also ceased to be useful either directly or indirectly, continues to exist if neither variability or selection intervene. THE END D. 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