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CHAPTER VII
THE EARTHWORMS OF OCEANIC ISLANDS

Oceanic islands are islands that have always been islands, a definition that seems tautological until we compare it with some other land masses that may be termed 'islands.' Geology teaches us in fact that from the point of view of their origin islands may be divided into two quite sharply contrasted classes. There are those detached land masses usually lying near to or comparatively near to some continent, which have been in the course of time detached by the action of the waves from that continent, such as for instance the British Isles, which undoubtedly represent a portion of the European continent which was once quite continuous with Europe. On the other hand we have the Hawaiian archipelago, St Helena, Fernando Noronha, and other similar islands, which are more remote in their position from continents and concerning which it seems clear that they have originated de novo by the action of submarine volcanos or of the growth of coral, combined with subsidence, following elevation, or from several of the causes combined. In any case the islands which are termed oceanic islands have never formed part of a continent. They are not relics of previously existing continents. It becomes a matter of great interest to compare the earthworms which are to be found upon oceanic islands with those which inhabit continental islands. Fortunately there are a good many facts at our disposal for this purpose; and we shall compare the earthworms of the Hawaiian archipelago with those which are found upon certain small islands lying to the south of New Zealand, viz. Campbell and Auckland islands and the more southern Macquarie islands.

The earthworms of the Hawaiian archipelago have been studied by a good many persons, and altogether a number of species have been described from that group of islands of which the following is a list: Pheretima hawayana, P. heterochaeta, P. peregrina, P. schmardae, P. hesperidum, P. morrisi, P. perkinsi, P. biserialis (= P. elongata), Allolobophora putris (= Kinberg's Hypogaeon havaicum), A. foetida, A. caliginosa, A. nordenskiöldi, A. limicola, A. rosea, and finally the well-known Pontoscolex corethrurus. Of these species there is only one which is even possibly a form limited to the Sandwich Islands, and that is Pheretima perkinsi, a species which I myself at first described as a new form, but which was afterwards regarded as identical with P. heterochaeta by Michaelsen, and later still resuscitated by Ude. All the others are found in many parts of the world and not only in the nearest mainland to the archipelago which we are now considering. I have had already occasion to speak of some of them as peregrine forms, especially of Pontoscolex corethrurus which occurs all over the world.

The conditions which have been recently revealed by an exploration of the antarctic islands mentioned above are totally different. Dr Benham has enumerated the following species from those islands, viz. Notiodrilus haplocystis, N. fallax, N. aucklandicus, N. campbellianus, N. macquariensis, Plagiochaeta plunketi, Rhododrilus cockayni, Leptodrilus leptomerus, L. magneticus, Plutellus aucklandicus, Diporochaeta heterochaeta, D. brachysoma, D. helophila, D. perionychopsis among the Megascolecidae, besides Phreodrilus campbellianus, Pelodrilus tuberculatus, P. aucklandicus and the Lumbricid Helodrilus constrictus. There were also four species of purely aquatic Oligochaeta which we shall leave aside from the present enumeration, as their range in space is a matter requiring a different explanation from that of the terrestrial forms. Here we have a series of worms, all of which, save the widely spread Lumbricid, are apparently absolutely indigenous to the islands mentioned since they are all different as species from those found elsewhere. Indeed there is a whole genus Leptodrilus, consisting, it is true, of but two species, which is a native of the Campbell and Auckland islands and of those only. The other genera are found in the antarctic region, while Pelodrilus is still more widely spread.

These facts as will be observed contrast about as strongly as they can with those supplied by the fauna of Honolulu and its adjacent islands. Not only are the worms of the antarctic islands different species from those found elsewhere, but the majority of them do not consist of widely ranging peregrine forms. It appears therefore most probable that these islands are not oceanic islands but a portion of the former existing northern portion of the antarctic continent. Were the species identical with those of New Zealand this conclusion would have of course to be reconsidered. The barriers to migration (see chap. VIII) explain the contrast recorded in the foregoing pages.

CHAPTER VIII
MOVEMENT AND MIGRATION AMONG EARTHWORMS

That earthworms can move upon the surface of the ground at a rapid pace is probably well enough known to everyone, and that they can also burrow with considerable celerity. Multiplying the inches of progress in minutes of time by centuries with the resulting miles, it is quite clear that there is no reason to suppose that an individual earthworm might not enormously extend its range under favourable circumstances. Their powers of locomotion are such that they could in the course of comparatively few centuries people a continent. As a matter of fact these animals are frequently very widely spread upon a given land surface; but on the other hand they are sometimes equally limited. It behoves us therefore to enquire the reasons for the possibility of extended migration and the causes which have led to its restriction. We are now, it must be borne in mind, considering these animals as purely terrestrial animals moving over the surface of the land by their own unaided efforts. We leave out of consideration any possible assistance in crossing water, whether fresh or salt. We have to consider in fact in the present section the earthworm inhabitants of larger and smaller tracts of continuous land such as the African continent, which will serve as an excellent example wherewith to test the facts and inferences.

And as a 'control' we can compare this continent with the very different continent of Europe.

As an excellent instance, because of the certitude of specific and in most cases of generic distinctions, we may take the Eudrilidae as illustrative of the facts that are to be considered in the present section. That family consists, as will be remembered, of 33 genera at most, which have the following more exact range on the African continent. The genus Eudriloides occurs in British and German East Africa and has been met with as far south as Mosambique and even Durban, in which latter locality it has been thought that it is really an accidentally introduced stranger. Platydrilus is limited to eastern equatorial Africa, thus not having quite the range of Eudriloides.

The small genera (that is small in numbers of species) Reithrodrilus, Bogertia, Megachaetina, Metadrilus, Notykus have the same limitation of range as the last genus. Metschaina has a wider range from tropical North East to lake Tanganyika. Stuhlmannia has a wider range still being found as it is in the Tanganyika district, in tropical North East Africa, and in British and German East Africa near the coast. Pareudrilus reaches still further north while Nemertodrilus is limited to the Mosambique region and to the Orange River district further south. The only remaining genus of this sub-family of the Eudrilidae is Libyodrilus which is purely West African and equatorial.

Of the remaining genera which are usually grouped together into a second sub-family, five, viz. Malodrilus, Kaffania, Gardullaria, Teleudrilus and Teleutoreutus, are confined to tropical North East Africa. Eminoscolex occurs in the same district but also to the south in the great lake region. The most remarkable fact about this genus is that one species E. steindachneri comes from the Cameroons, and another E. congicus from the Congo, and thus the range of the genus is right across the continent. Neumanniella has much the same range. Polytoreutus is a purely equatorial East and Central genus, reaching from the coast to the lakes. Bettonia known by three species is from British East Africa.

The remaining genera, viz. Hyperiodrilus, Heliodrilus, Alvania, Iridodrilus, Rosadrilus, Euscolex, Parascolex, Preussiella, Buttneriodrilus, Beddardiella, Metascolex, are all West African and the vast majority equatorial. We thus see that with one exception the genera of East Africa are totally different from those of West Africa and that the family as a whole is restricted in its range to a comparatively small part of the vast African continent. It also obviously follows, and it is advisable to state this fact however obvious, that no species are common to the two sides of the continent except indeed the ubiquitous Eudrilus, whose range over the world has been more than once referred to in this book.

On the other hand the genus Dichogaster offers quite different facts, which are in contradiction to those just enumerated. This genus as already said is very characteristic of tropical Africa, and a large preponderance of the known species are confined to that continent. Although there is some variation in structural characters among the many species which compose this genus, there is but little doubt that they are all rightly referred to one genus with perhaps some doubtful, though not very striking, exceptions. In any case the utmost divergence of structure between worms usually placed together in this genus is nowhere near to that which separates the genera of Eudrilidae from each other. Of the African members of the genus the species are pretty evenly divided between the eastern and western halves of the continent; they are, like the Eudrilidae, tropical in range, not occurring to the southward, where their place is taken by the Acanthodrilinae and Geoscolecidae. There are it is true a few species, such as D. gracilis and D. bolavi, which are common to the two sides of Africa; but in these cases we clearly have to do with those rather mysterious species which can apparently unduly extend their range and which are known as peregrine forms; for they also occur in other parts of the world besides Africa. We have therefore in Dichogaster the case of a genus which ranges all over the tropical parts of Africa, but whose species are not common to the Atlantic and Indian shores of that continent.

We will now contrast these conditions, which exemplify certain facts shown by the characteristic Oligochaeta of tropical Africa, with those which obtain in Europe. In this region of the world the prevalent and practically the only genera which need be taken into consideration in surveying the Oligochaetous fauna from the present point of view, are Lumbricus and the genus Allolobophora of Eisen which has been variously rearranged into genera and sub-genera known by the names of Helodrilus, Bimastos, Octolasium, etc. The structural differences which divide these genera and sub-genera are not great; in any case they do not exhibit such a wide range of variation from each other as do two such Eudrilid genera as Stuhlmannia and Hyperiodrilus. We find the genera mentioned not only in Europe but extending themselves over more or less of Asia, even occurring in Japan; while the North American continent contains also representatives of the same. Not only do we find this community of genera over vast extents of country greater in diameter than the African continent, but there are also many species which range as widely or nearly as widely as the case may be as the genus to which they belong. Thus the species of Allolobophora (we do not trouble about the newer sub-divisions as they hardly affect the facts to be emphasised), A. caliginosa, A. longa, A. rubida, A. chlorotica, A. octaedra, A. constricta, A. beddardi, Lumbricus terrestris, L. castaneus, have an enormously wide range over what is generally termed the Palaearctic region, extending also in some cases into the Nearctic. It is true no doubt that the majority, indeed perhaps all, of these are, like certain species of Dichogaster mentioned above, among those forms termed peregrine which have the capability of living in every quarter of the globe to which they have apparently been conveyed by man. But there remain many species which have a very extended habitat in the northern hemisphere, and in any case the genera and the species are there truly indigenous and widely spread.

It would thus appear that the capability for independent migration varies greatly among earthworms. Of the types selected for consideration the Eudrilidae are the slowest movers; the genus Dichogaster comes next, while the power of migration possessed by the genera Allolobophora and Lumbricus is very much greater. Assuming for the moment the correctness of this inference it is clear that it will influence many other propositions connected with the relative age of the families of these worms and with many problems of geographical distribution. It appears to us that this simple explanation is the correct one. But to show this it will be necessary to eliminate other possible explanations. It might be urged that the wider range of the genus Dichogaster and the still wider range of the genus Allolobophora (shown by community of species in widely distant localities) was evidence merely of relative age, that the older groups have had more time to travel and that the newer groups have not had so long a time to spread themselves over their habitat. On this hypothesis the genera of Eudrilidae would be geologically much newer than the genus Dichogaster and similar statements might be made for the other forms here under consideration. As already explained we cannot attempt to answer this question in the only way in which it can be really satisfactorily answered, by a reference to fossil forms; for there are no fossils to refer to. So far as comparative anatomy enables us to arrive towards a solution of the question, it would appear that the genus Dichogaster belongs to a more ancient race than either of the other two groups considered, and that of these latter the Lumbricidae are the most modern. Moreover we associate not only a wide, but also a discontinuous, distribution with an archaic race; and for this reason also we should place the genus Dichogaster in the position of being the most ancient of these Oligochaeta. For the genus occurs in Central America and in certain parts of the East as well as in Africa. So that we can fairly dismiss the view that the Lumbricids by virtue of their greater range over a given area are the most ancient type and that their range is associated merely with their antiquity. Nor does it appear that geographical or meteorological consideration can have had effect in the present instances. For conditions favourable to earthworms prevail in tropical Africa, as in Europe and much of North Asia.

Climate as Affecting Migration

That excessively rigorous climatic conditions affect the range of earthworms as well as fresh-water forms is quite clear from the conditions which obtain in the most northern climes. At any rate in those regions where physical conditions render it impossible for these Annelids to have their being. A perpetual mantle of snow and a temperature far below freezing point are absolute barriers to the extension of range. And yet there are some few Oligochaeta which do not in the least suffer from a somewhat milder taste of such conditions. Thus species of Enchytraeidae have been met with on glaciers and even found in frozen water, while a few earthworms have been brought from the island of Kolguev. These however are quite exceptions to the general sterility as regards earthworms of the excessively cold regions. We have already seen that there are no general facts to be deduced as concerning the relative abundance of terrestrial worms in the tropics and in more temperate climes. Tropical Africa is, it is true, rich in genera and species; but on the other hand tropical East Indies have but few genera inhabiting their numerous islands. Temperate England has very few genera and not a large number of species; temperate New Zealand has a considerable number of different indigenous genera. When however we leave this general aspect of the question and consider separate families and genera, there seems to be some little relation between climate and distribution and thus some effect of climate in acting as a barrier to migration. For example, though continuity of land surface permits of the tropical African Eudrilidae ranging southwards as far as the Cape they are not met with so far as we know in the most southern parts of Africa; nor are the South American Geoscolecidae found in Patagonia or northward beyond Central America. These instances do really look like an influence of climate upon range. On the other hand we must be careful to eliminate the possibility of another explanation and that is the impossibility of successful migration owing to the previous occupation of the ground with abundant other forms. The very same countries would appear to show that this explanation is unnecessary. For the prevalent genus of the southern tracts of South America Notiodrilus extends its way northward as does the same genus from temperate to tropical Africa and Madagascar.

It looks very much, therefore, as if certain Oligochaeta are dependent upon climate for their range, and as if others were at least more independent of climatic conditions. And there are other facts which support this view. The same opinion is supported by the phenomena of involuntary migration, a subject which has been considered also separately under the head of 'Peregrine forms.' The great prevalence of Lumbricidae accidentally imported into many parts of the world shows that temperature is no real bar to their voluntary migration. On the other hand the fact that specimens of the East Indian genus Pheretima though commonly imported accidentally into the warmer regions of the world have not been able to make good a footing in Europe, save in greenhouses, shows that this genus is affected in its range by questions of climate. These facts suggest another inference of great interest which can only be mentioned tentatively, and not put forward as a demonstrated conclusion. Seeing that Lumbricus (sensu lato) can comfortably take up its home in warm extra-European countries, but yet that it has evidently not spread to those countries in the course of nature but by man's interference, it seems possible that time alone has prevented this; and that therefore this family Lumbricidae is one of the most recently evolved families of Oligochaeta. Certain structural features support this way of looking at the matter. The same arguments precisely apply to the genus Pheretima, which is also regarded by most systematists as a recently developed race of earthworms. Anyhow the conclusion which the facts seem to warrant is that the effects of climate in influencing distribution are seen to have an unequal effect upon earthworms, some genera being debarred by climatic conditions while others are indifferent to the same.

Mountain Ranges and the Migrationof Earthworms

In many groups of animals the interposition of a lofty chain of mountains presents an insuperable barrier to migration. The barrier is effective for more than one reason. Lack of vegetation and a differing climate are among the more obvious causes which render Alpine chains important as affecting distribution. There is plenty of evidence in the way of positive fact that mountains are not necessarily barriers to the spread of earthworms. The recent explorations of the Ruwenzori chain of mountains in Africa have resulted in the collection of a considerable number of species, some of which come from great altitudes (e. g. 4000 metres and slightly upwards), and one species, viz. Dichogaster duwonica, which Dr Cognetti de Martiis described from the foot of the glacier Elena. I have in my temporary possession a number of examples of the eastern genus Pheretima, some of which are new species from lofty areas in the Philippine Islands. There are plenty of other examples pointing to a like conclusion. It is noteworthy that these forms which have been met with at lofty heights are not essentially different from the plain living forms. One cannot exactly speak, at any rate in the present state of our knowledge, of anything like an Alpine fauna.

It is in fact clear enough that whatever may prove to be the case with regard to particular species, a mountain range is not necessarily a barrier to the dispersal of generic types.