Part 18

With respect to range, the results arrived at have no particular interest, for the species are not sufficiently numerous; and, what is still more adverse, the genera, with unimportant exceptions, range over the world; so that there is no scale of differences, and it cannot be said that these two regions differ in their genera, and these two only in their species. In all the following remarks, I have trusted exclusively to my own specific identification; and have rejected all assigned localities which appeared from any cause to be doubtful. Sessile cirripedes are found in every sea, from lat. 74° 18′ north to Cape Horn. The area included between the north point of the Philippine Archipelago and the south point of Australia, extending on the right hand to New Zealand, and on the left to Sumatra,--an area, which, though including two distinct Cirripedial regions, is small compared with the surface of the globe, yet includes a greater number of species, especially of peculiar species, than the whole rest of the known world. This is, probably, in chief part due to the broken nature of the land, affording diversified habitats, and to much of the coast being rocky. Cirripedes, from requiring to be attached, cannot live where the shores and adjoining bottom are sandy or muddy or formed of moving shingle; hence, no doubt, it arises, that there is such a remarkable contrast in the great number of the species inhabiting the bold rocky western coast of South America, and the few species living on the sloping, and often sandy or muddy or shingly, eastern shores of this continent. Hence, also, I believe, it is that not many species have been brought from India. Coral-reefs are not favorable to Cirripedes, consequently but few are known to inhabit the islands of the Pacific Ocean. Where Cirripedes can live, though the species in no district are numerous, the individuals abound in infinite numbers: I have walked over the coast-rocks of the Falkland and Chonos Islands, of Chile and Van Diemen's Land, fairly encrusted over wide spaces with a continuous layer of Cirripedes, consisting of only two or three species; in the same manner as may be observed on many parts of the shores of Great Britain, and, I believe, of North America.

With respect to the effects of temperature on the range of Cirripedes, no genus (having more than one species) is confined to the torrid zones. Pyrgoma, from being always attached to corals, is, of course, ordinarily found in the hotter seas; but one species ranges from the Cape de Verde Islands in the torrid zone to the southern shores of England and Ireland. Tetraclita is not found in the colder seas, but is numerous in species and in individuals, on the southern shores of Australia and at the Cape of Good Hope. I may here add, that the two genera with the most confined ranges, are Chamæsipho and Elminius; the former has only two species, one inhabiting Australia, and the other the East Indian Archipelago; Elminius has four species, confined to the southern hemisphere, and inhabiting Australia, New Zealand, and South America. To return to the effects of temperature; in Mr. Dana's great work on Crustacea, an excellent chart is given, in which the _isocrymal_ lines, or those exhibiting the mean temperature of the waters along their course, for the coldest thirty consecutive days in any season of the year, are given; and which lines Mr. Dana has shown are the most influential in governing the distribution of marine animals. At the isocryme of 68°, Mr. Dana divides the torrid and sub-torrid zones from the several temperate zones; and at 44°, the temperate from the sub-frigid and frigid zones; but as no Cirripedes are exclusively confined to these frigid zones, we may here disregard them. From Mr. Dana's[76] table of the areas of the torrid and temperate ocean-zones, on both sides of the equator, it seems that they are nearly as 337 to 278, in relative area; and consequently, he remarks, that the marine species in any class, if distributed equally over the two, would be one fifth more numerous in the torrid than in the temperate zones. Now of Cirripedes, taking all the orders, there are at present known 147 species; of these, 7 have doubtful habitats, leaving 140 for comparison. Of these 140, nearly one quarter, or 37, inhabit both the torrid and temperate zones, as above defined; 46 are found exclusively in the torrid, and 57 exclusively in the temperate zones; so that the temperate zones, though less in area, and having, proportionally, even a considerably lesser length of coastline, nevertheless have a preponderance in the number of species. But it should be borne in mind, that there are _two_ great temperate districts, separated from each other by _one_ great torrid district; and, inasmuch as the number of species in any region seems to depend in some degree on the isolation of the sub-regions, we might have expected (the other conditions now being, and the _past_ conditions having been alike), that the two great temperate areas would have contained more species, perhaps doubly more, than the single great torrid area.

[76] 'Crustacea: United States Exploring Expedition,' p. 1476 (corrected).

The proportional numbers, above given, are not very widely different, whether we take separately the Balanidæ, the Lepadidæ, or all together. Mr. Dana has shown[77] at length, that generally amongst the Crustacea, the species which he considers of highest rank, belong to the extra-torrid zones: there seems to me in all such cases to be some degree of vagueness in the attempt to determine which are highest or lowest, but I have already elsewhere stated that Balanus is, probably, the most eminently Cirripedial form, and exhibits in the strongest manner all the characters by which Cirripedes differ from other Crustacea; as this genus is the largest, containing 36 species, of which the habitats are known, I may state that of these, exactly one third, or 12, inhabit both zones; 9 exclusively inhabiting the torrid, and 15 exclusively the temperate zones. According to the proportions of the whole class, the numbers should have been 9 torrid, to only 11.11 temperate; so that evidently the genus Balanus (in one sense typical) inclines towards the temperate regions more strongly than does either the family or the sub-class to which it belongs.

[77] Ibid., p. 1528.

With respect to the relation between the size acquired by the different species of sessile cirripedes, and the temperature of the localities inhabited by them, the genera Chthamalus, Tetraclita, and Balanus, alone can serve for comparison: in Chthamalus much the largest species is found in the temperate zone: on the other hand, the two largest species of Tetraclita are from the torrid zone, though one of them also sometimes ranges into the temperate seas: in Balanus, the largest species, and six other species having a basal diameter sometimes over two inches, inhabit the temperate regions; and two out of these seven species, flourish even in the Arctic seas; whereas, within the torrid zone, there are only three species with a diameter sometimes exceeding two inches, but two of these frequently become very large and massive; so that Balanus, judging from the size of the species, as well as from their range, does not require for its highest development the temperature of the torrid zones.

The greater number of the species of the Balanidæ have wide ranges, as might be inferred from the fact of between one third and one fourth of the total number inhabiting both the torrid and temperate zones; but it should not be overlooked, that those species, as _Balanus tintinnabulum_, _amphitrite_, _improvisus_, and, in a lesser degree, _B. trigonus_ and _Tetraclita radiata_, which seem to range over nearly the whole world (excepting the colder seas), are species which are habitually attached to ships, and which could hardly fail to be widely transported. Indeed, it appears to me surprising, that such species as _Balanus psittacus_ and _eburneus_, which often become attached to vessels, should still be confined, the one to Southern, and the other to Northern America. But some other Cirripedes, which I have never seen attached to vessels, have likewise immense ranges: thus _Tetraclita porosa_ is found in every tropical and warmer sea, and _Chthamalus stellatus_ ranges round the world in the northern hemisphere, and, along the eastern side of America, far south of the equator: _Balanus spongicola_, and _Acasta spongites_, extend from the shores of Britain to the Cape of Good Hope: _Balanus lævis_ ranges from Tierra del Fuego to California. I may further remark, that the only two other species of Balanus, and the one Chthamalus, inhabiting Tierra del Fuego, are, also, found on the shores of Peru. But to show the powers of endurance in some species, I may specify the case of _Balanus improvisus_, which flourishes on the coast of Nova Scotia, amongst the West Indian Islands, in Southern Patagonia, and near Guayaquil. Even more striking is the case of _B. crenatus_, of which I have seen specimens from latitude 74° 48′ north, from the West Indies, and the Cape of Good Hope! In these two latter localities, however, it seems to be rare, and may have been first transported to them from the shores of Europe, on the bottoms of vessels, to which it sometimes adheres.

The several species of Balanidæ live attached either to coast-rocks, or to objects at various depths, down to, as in the case of _Balanus crenatus_, 50 fathoms. _Balanus balanoides_ sometimes adheres to rocks or wood so high up, that it is not covered by water during the neap tides. Mr. Thompson has informed me, that he once accidentally kept some specimens of this species out of water for seven days in a warm room, and that they were then quite lively. This species, is very easily killed by brackish water, as are some other species, whilst _B. improvisus_ and _eburneus_ can flourish in it; and at the Falkland Islands, I saw _Elminius Kingii_ attached to rocks at the mouth of a fresh-water brook, so as to be covered by pure water during the ebb of each tide. Sessile cirripedes adhere to all sorts of objects, floating and fixed, animal and vegetable, living and dead, organic and inorganic. Chthamalus is, perhaps, more commonly attached to rocks than are the other genera. Living Mollusca are, I think, the most frequent objects of attachment: Mr. Cuming has remarked to me, that shells covered by an epidermis, as Patella, Haliotis, and Mytilus, are the greatest favorites. Acasta is always imbedded in sponges, or in the sponge-like bark of Isis; Pyrgoma and Creusia in corals; Chelonobia is attached to turtles, and one species to crabs or very smooth shells; Coronula, Tubicinella, and Xenobalanus, are imbedded in the skin of Cetaceans; and Platylepas in that of manatee, turtles, or sea-snakes.

If we attempt, with our present not very imperfect materials, to divide the globe into provinces, according to the amount of difference in their Cirripedial inhabitants, including all orders and families, and disregarding entirely, as I think we ought, all probabilities or conclusions deduced from the distribution of other tribes of animals, we find that the globe may be divided into the four following great provinces and one sub-province. I should premise, that in the following remarks and tables,[78] the species of Lepas, Conchoderma, Chelonobia, Coronula, Platylepas, and Tubicinella, are excluded, owing to their being attached to floating or swimming objects, and being consequently widely and irregularly distributed.

[78] As the number of Cirripedes in the whole class is not very great, I have given lists of the species in the four main provinces and in the one sub-province.

The first, or North Atlantic province, is that of Europe and the eastern shores of North America, from the arctic regions to lat. 30°: the island of Madeira, part of the north-west coast of Africa, and the whole Mediterranean being included. In this province (the above-named genera being excluded) we have 31 species, of which 22 are not found in any other distant quarter of the globe. As some few of these species range into the West Indies, I have not, on this account, excluded them from the 22 peculiar forms. Had I included the West Indies[79] in my first province, the total number of species would have been 42, of which 28 would have been peculiar. The coast of Brazil, even as far south as the Rio Plata, might, also, have been included, for I have not seen from it a single species not included in the above 42 West Indian species. So also, by adding a single species, might the west coast of equatorial Africa. The two coasts of South America and Africa, which face each other within the torrid zone, seem to be remarkably barren in Cirripedes. Europe has several more species than the United States, which is inhabited by only ten species, including even the probably imported _Balanus tintinnabulum_ and _amphitrite_. Of these ten United States species only two are not found in Europe; and both these two range into the West Indies, and as far as the northern shores of South America, and therefore cannot be considered as peculiar to the United States.

[79] The total number of species which I have seen from the West Indies, is 19 or 20; of these, only 6 are peculiar to it, or 8, if the United States be likewise included, the other 12 or 14 species being found in other quarters of the world. Six peculiar species out of 19 or 20, has not appeared to me sufficient to institute even a sub-province.

I have formed my single sub-province for the southern extremity of Africa; for although I know of only 11 species from this comparatively short and uniform line of coast, yet I was not able to group these eleven in any of the main provinces: 5 of the species are peculiar, 1 Australian, 3 European and West Indian, and 2 almost universally distributed.

The second province includes the west coast of North and South America, from Tierra del Fuego to Behring's Straits: on this enormously long line of coast, only 22 species are known to exist, but of these no less than 15 are peculiar. Of these 15, four are not found south of the torrid equatorial region, and eight are not known to occur north of this same region; so that this long line of coast might have been divided into two sub-provinces, of which the southern would have been the most peculiar; but as eight species are found both north and south of the equatorial region, I have not made this sub-division. Two of the species occurring on the western coast of North America, seem to represent species found on its eastern coast, and in Europe; thus, _Balanus glandula_ takes the places of _B. crenatus_, and _B. cariosus_ that of _B. balanoides_. Not a single species, excepting a few which are also widely distributed over other parts of the world, is known to be common to the east and west coasts of the two Americas.

The third province is that of the East Indian Archipelago, and includes the Philippines, Borneo, New Guinea, Sumatra, Java, Malacca, and the eastern coast of India. Here we find 37 species, of which 24 are peculiar. I may remark, that I have received no species from Madagascar or the eastern coast of Africa; few from India, or from the coast of China; and I suspect, that on most of these coasts, only few exist. Probably our third province will hereafter be found to include the whole Indian Ocean.

The fourth province is that of Australia, including New Zealand: it has 30 species, of which 21 are peculiar. Had the temperate Australian coasts (_i. e._, those south of the isocryme of 68°) been alone considered, the number of the species would have been probably 25, of which 20 would have been peculiar,--that is, if we admit within the 20, several species which range from the temperate into the torrid zone, but do not extend beyond the Australian shores. Owing to the widely-extended ranges of most Cirripedes, no Arctic or Antarctic provinces can be said to exist.

To recapitulate the above results, bearing in mind that, although the total number of known existing Cirripedes is 147, yet that the habitats of seven are unknown, and that eighteen are excluded owing to their being attached to floating or swimming objects, so that there are only 122 species referred to in the following table:

+--------------------------------------------------+--------+---------+ | | Total | Species | | | number |confined | | | of | to the | | |species.|province.| +--------------------------------------------------+--------+---------+ |(1.) First, or North Atlantic Province, to lat. }| | | | 30° N. (If the West Indies had been included}| 31 | 22 | | the numbers would have been 42 and 28) }| | | |(2.) Sub-province of South Africa | 11 | 5 | |(3.) Second province, or West Coast of North and }| | | | South America }| 22 | 15 | |(4.) Third province, or East Indian Archipelago | 37 | 24 | |(5.) Fourth, or Australian province | 30 | 21 | +--------------------------------------------------+--------+---------+

The least prolific of these provinces contains 22 species, or between 1/5th and 1/6th of the total number of species, and the most prolific between 1/3rd and 1/4th of this same number. In each of these provinces, it is remarkable that the peculiar species are very nearly two thirds of the whole of its inhabitants. These facts, I think, show that the above provinces are natural divisions of the world, as far as their Cirripedial inhabitants are concerned.

As Cirripedes belong to the great class of Crustacea, and as the distribution of the latter has lately been fully discussed by Mr. Dana, it may be worth while briefly to compare my results with his; more especially as they are so very different. I should premise, as perhaps accounting to a certain extent for this difference, that, owing to the wide range of many species, and the almost universal extension of the same genera, my provinces are founded merely on a certain proportion of the species, namely, two thirds, being peculiar or confined to a region of considerable dimensions: whereas, in the case of ordinary Crustaceans, the greater number of the species are distinct even in the sub-provinces, and the provinces are founded mainly on generic differences. Mr. Dana divides the surface of the globe into three great sections, or provinces, the _Africo-Europæan_, the extent of which is shown by its title; the _Occidental_, which includes both the east and west coast of both Americas; and the _Oriental_, including the Indian and Pacific Oceans, with the East Indian Archipelago, and Australia. Thus Mr. Dana entirely separates the Eastern shores of North America from Europe; whereas, as far as their Cirripedial inhabitants are concerned, they are most intimately allied, and form my first or North Atlantic province; and to this, as I have shown, even the West Indies, the coast of Brazil, and equatorial West Africa might have been added. It follows, from this similarity in the Cirripedes on the two sides of the Atlantic, and from their dissimilarity with those on the shores of the Pacific, that the east and west coasts of the two Americas form two quite distinct Cirripedial provinces; though, in the northern half, some connection is shown by a few representative species: on the other hand, Mr. Dana unites both sides of the whole American continent, into his single Occidental province. The South-African province is not brought out by Mr. Dana so prominently, as I have found necessary. Mr. Dana joins the East Indian Archipelago and Australia into his single Oriental province, and makes New Zealand, as a sub-province, apparently as distinct from Australia, as Australia is from the East Indian Archipelago: whereas I find that the Cirripedes of New Zealand clearly belong to Australia; and that the Australian Cirripedes, especially if the temperate shores be alone considered, are as distinct from those of the East Indian Archipelago, as from those of any other quarter of the whole world. I believe that the provinces deduced from the distribution of Cirripedes, accord better with the Molluscan provinces, than with those given by Mr. Dana for the rest of the great class of Crustaceans.

In the following tables, an asterisk means that the species is not found in any other distinct region of the globe. When found in one of the five provinces, a corresponding number, within brackets, is appended, to show in which province or sub-province it has been found.

(1.) FIRST OR NORTH ATLANTIC PROVINCE: _Europe and the Eastern United States, from the Arctic Regions to 30° north latitude._

Balanus tintinnabulum (1 to 5). tulipiformis* confined to Europe. calceolus Europe and India. { confined to North America and West galeatus* { Indies. spongicola (2) and West Indies (?). { confined to Europe, but possibly in the perforatus* { West Indies. amphitrite (1 to 5). eburneus* { confined to North America and West { Indies. improvisus Europe and North America and (3). porcatus* Europe and North America. crenatus " " W. Indies and (2). balanoides* " " Hameri* " " Acasta spongites Europe and (2). cyathus* Madeira and West Indies. { confined to Europe, but ranges as far at Pyrgoma anglicum* { least as the Cape de Verde Islands. Xenobalanus globicipitis* confined to Europe. Chthamalus stellatus Europe and North America and (3 and 4). Pachylasma giganteum* confined to Europe. Verruca Strömia Europe and Red Sea. Spengleri* Madeira. Pœcilasma aurantia* " crassa* " Dichelaspis Lowei* " Oxynaspis celata* " Alepas minuta* Europe. parasita* Europe and Atlantic Ocean. Anelasma squalicola* Europe. Alcippe lampas* " Scalpellum vulgare* " Pollicipes cornucopia* "

Here we have 31 species, of which 22 are not found in any other great region of the world.

(2.) SUB-PROVINCE: _Africa, South of lat. 30°._

Balanus tintinnabulum (1 to 5). Capensis* spongicola (1) and West Indies (?). amphitrite (1 to 5). crenatus (1) and West Indies. Acasta spongites (1). Tetraclita serrata* rosea (4). Chthamalus dentatus* also on West Coast of Africa. Octomeris angulosa* Scalpellum ornatum*

In this small region we have only 11 species, of which five are peculiar: _Balanus Capensis_ and _Tetraclita serrata_, seem to be representatives of _B. psittacus_ of S. America and of _T. porosa_ of that and several other regions.

(3.) SECOND PROVINCE: _West Coast of South and North America, from Tierra del Fuego to Behring's Straits._