Observations of a Naturalist in the Pacific Between 1896 and 1899, Volume 2 Plant-Dispersal
CHAPTER XXXII
THE WEST COAST OF SOUTH AMERICA
The littoral floras of the West Coast of South America.—The Convolvulus soldanella zone of Southern Chile.—The plantless or desert zone of Northern Chile.—The Sesuvium zone of Peru.—The Mangrove zone of Ecuador and Colombia.—The two varieties of Rhizophora mangle, the “mangle chico” and the “mangle grande.”—The floating vegetable drift of the Guayaquil River.—The Humboldt current and the climate of the West Coast of South America.—The advance northward of the arid climatic conditions of the Peruvian sea-border.—The retreat of the mangroves.—Evidence of ancient coral reefs on the coast of Peru.—The shore plants and stranded seed-drift of the Panama Isthmus.—Summary.
MY acquaintance with the strand-flora of the west coast of South America began at Corral, the port of Valdivia, in Southern Chile in lat. 40° S., and terminated at the mouth of the Guayaquil River, in Ecuador, about 2° south of the equator. During the period December 23, 1903, to March 17, 1904, I examined the coast plants at sixteen localities in this region, which covers 38 degrees of latitude and thus measures about 2,300 miles. Travelling in a steamer to Callao that was trading on the coast I had opportunities of staying for periods ranging from half a day to a couple of days at a considerable number of places; and a week spent at Valparaiso gave me a good opportunity of examining the beaches north and south of it. At Lima I spent some weeks, and from that centre examined the shore-plants at Callao, Ancon, and Chancay to the northward. North of this I had not the same opportunities, until we passed the Peruvian and Ecuadorian boundary; but from a visit to the shore at Paita, from the general look of the country in places as we coasted along, and from information derived from other sources, I was able to obtain a fair general idea of the prevailing character of the beach plants. After my previous experience to the southward, one could fairly gauge the character of the beach-flora from the appearance of the land behind. In the Gulf of Guayaquil and in the vicinity of the city of that name I spent about three weeks in the investigation of the coast flora.
If it were not for the interposition of the great rainless deserts of Northern Chile and for the scantily vegetated, scantily watered and semi-sterile condition of almost the whole coast of Peru, the botanist would be presented with a splendid opportunity of studying the distribution of shore-plants along a meridian stretching through some fifty degrees of latitude from Patagonia to Ecuador. As it is, drought and sterility in one form and another reign over about half of this great stretch of continental coast. This is reflected in the beach-flora; and though the observer will often have his interest attracted by the wonderful climatic anomalies arising from the presence on the coast of the cold Humboldt current, to which the sea-border of North Chile owes its desolation and the coast of Peru its semi-sterility, yet for a long time he will feel as if Nature had hardly dealt fairly with him.
Along the sea-border corresponding to the deserts of North Chile there would seem to be practically no plants growing on the beaches, except here and there where some stray plant from the saline districts inland intrudes on the coast. Along the whole sea-border of Peru from Arica north to Tumbez on the borders of Ecuador, the coast-districts, though more or less rainless, receive the benefit of the drizzly garuas and sea-fogs, and the sterility of the land immediately backing the beaches is much less pronounced than with the sea-border corresponding to the deserts of Northern Chile. This difference shows itself in a peculiar type of littoral vegetation, a strand-flora that is very scanty but one where on the beaches Sesuvium prevails. North of Tumbez the mangrove-formation predominates along the sea-borders of Ecuador and Colombia to Panama, excepting on a stretch of sterile coast extending north from the Gulf of Guayaquil to the equator.
Though in one sense the botanical observer will be disappointed with the littoral floras of the west coast of South America, in another sense when he remarks the manner in which the coast-vegetation reflects the abrupt changes in the prevailing climatic conditions he will be fascinated by the interesting problems presented to him. We are accustomed to connect a tropical coast with mangroves, coral-reefs, and beaches of calcareous sand supporting a luxuriant littoral flora. Climatic conditions banish all these from the tropical west coast of South America until within four degrees of the equator, and then with startling suddenness the dominion of the mangrove begins, the neighbouring hills commence to be clothed with tropical jungle, and the climate is completely changed. Mr. John Ball, who sailed along this coast about twenty years ago, referring to this remarkable phenomenon on the borders of Peru and Ecuador, remarks that no such abrupt and complete change both in climate and vegetation is known elsewhere in the world, and he adds that few parts of the American coast better deserve careful examination (_Naturalist in South America_). This subject has since been discussed at length by Dr. Wolff in his “Geografia y Geologica del Ecuador,” and by Baron von Eggers in a paper to be subsequently quoted, two very competent observers, but the latter considers that the subject still requires a systematic investigation, and suggests that an observing station should be established on this coast by the combined meteorological societies of Europe. A sojourn of more than a week in the swamps at Puerto Bolivar, a few miles from Tumbez, enables me to appreciate the nature of the problem, and to throw a little light on the line of investigation required.
But to return to the general subject of the littoral floras of the west coast of South America, I may say that beginning with the island of Chiloe in lat. 42° S., this coast may be divided into four zones.
(1) The Convolvulus soldanella zone of Southern Chile, which extends as far north as Coquimbo about 30° S. lat.
(2) The Plantless or Desert zone stretching north to the vicinity of Arica in lat. 18° 30ʹ, and corresponding to the coast of Northern Chile.
(3) The Sesuvium zone, extending north from Arica to the 4th parallel of south latitude in the vicinity of Tumbez, a sea-border of semi-sterility that comprises the entire coast of Peru.
(4) The Mangrove zone, stretching from Tumbez, on the frontiers of Ecuador, to the equator and on to Central America, but interrupted at first by a strip of sterility on the coast extending from the Gulf of Guayaquil to the borders of Colombia, or, strictly speaking, to the equator.
THE CONVOLVULUS SOLDANELLA ZONE (SOUTHERN CHILE).
This zone, which answers to the coast of Southern Chile, from Chiloe as far north as Coquimbo, corresponds to watered and vegetated inland regions, in which, however, the amount of rain and the degree of fertility decreases from south to north, that is to say, as we approach the desert regions. Here we find none of the dry beaches that prevail for twenty-five degrees of latitude north of Coquimbo. When we scoop with our hands to a depth of three or four inches in the sand we find it relatively cool and more or less moist, as in an English beach. In a hot summer’s day on a Valparaiso beach we should find that the temperature of the sand at the surface (half-inch deep) was about 112° F., and at a depth of four inches about 80°. This would be above the average for the zone, which would be probably near the typical summer-temperature of an English beach, namely, 102° at the surface and 77° four inches down. This subject of beach temperature is discussed in Note 70.
Plants typical of the beaches of this zone, and evidently occurring over the length of it, are Convolvulus soldanella, Nolana (paradoxa?); Polygonum maritimum, Salsola Kali, and Selliera radicans. Nolana is a Chilian and Peruvian genus. This beach plant, which is especially abundant on the beaches near Coronel and at Bahia San Vincente, has the creeping habit of its associate, the Convolvulus. However, it possesses seeds, or rather seedvessels, of more limited buoyancy; and it is shown in Note 71 that prolonged drying is needed for effective dispersal by currents over great distances. This beach species of Nolana has narrowly escaped being a widely-spread littoral plant; whereas it is now restricted to the Chilian beach flora. Selliera radicans, a little creeping Lobeliaceous plant, growing under the shade of tall clumps of Juncus at the edge of the beach or in wet places where springs ooze out in the sand, is a very interesting species that occurs also on the other side of the Pacific in Australasia. Of the mode of dispersal of its small seeds I know nothing, as the fruits were not ripe at the time of my visit; but I would suggest that some resident botanist should investigate this important point. I found it at Corral and at Coquimbo; and Gay speaks of it as growing on wet coast places from Chiloe to Coquimbo, a range of 12-1/2° of latitude.
It is probable that all the shore-plants of this zone extend south to Chiloe in latitude 42° S.; and it is likely that some of them reach towards the Straits of Magellan. I did not find any of them within the Straits on the beaches in the vicinity of Punta Arenas, where, however, I noticed the three plants recorded by Ball, namely, Armeria maritima, var. andina; Senecio candidans, also found in the Falkland Islands; and Plantago maritima; besides a Chenopodiaceous plant not in fruit. The Plantago has no capacity for dispersal by currents, and probably none of the other plants are thus dispersed. I formed the opinion when in the Straits that the beach plants on the Pacific and Atlantic coasts of Patagonia could have but little communication by the currents, and that they are in this respect quite cut off from each other. A botanist who investigates the strand-flora of Patagonia and Tierra del Fuego in connection with the littoral plants of the opposite coasts ought, if he has not already done so, to obtain some very interesting results from the standpoint of plant-dispersal.
The northern limit of the plants of this zone near Coquimbo, in lat. 30° S., is not determined by the change in climatic conditions that goes normally with decrease in latitude, but by the vicinity of the great deserts of Northern Chile, the aridity extending to the beaches.
Amongst the other plants occurring generally in the Convolvulus soldanella zone of Southern Chile, species of Salicornia and Samolus are to be observed in wet places. On the beaches near Valparaiso and in the vicinity of Talcahuano there thrives a species of Franseria, a Composite plant possessing prickly fruits well suited for conveyance in bird’s plumage, but not adapted, as shown in Note 71, for dispersal by currents. Mesembryanthemum is a typical beach-plant at Coquimbo, and an intruder from the adjoining hill-slopes at Valparaiso. Raphanus, seemingly R. maritimus, occurs in places, but apparently only as an intruder from the cultivated districts behind the beaches. One or two species of Euphorbia are not uncommon. A few small trees or bushes of Acacia farnesiana grow typically on the beach at Coronel and in neighbouring sandy tracts at Talcahuano, though the plant, as Gay observes, has been introduced. Sophora tetraptera, found also in New Zealand, and one of the most interesting plants of the Antarctic flora, thrives as a small tree on the hill slopes overlooking the harbour of Corral, becoming bushy where in places it intrudes on the beaches, and fruiting there as freely as on the slopes above. It was by testing the buoyancy of the seeds of this plant that I was led to the discovery of its mode of dispersal by the currents (I am indebted to Mr. Holland for the specific determination of the fruits sent by me to the Kew Museum). Other shore-plants, of course, occur in this zone; but I have gone far enough to illustrate the subject. Of the numerous occasional intruders from the neighbouring inland districts, frequently Compositæ, I say nothing. The results of my observations on the floating power of the seeds and seed-vessels of some of the shore-plants of this zone are given in Note 71.
Stranded seeds and fruits that belong to the proper beach-drift are not easily found on the beaches of Southern Chile, as they are often buried in rubbish. Those most characteristic are seeds of Convolvulus soldanella and drupes of Nolana (paradoxa?), both typical beach-plants of the zone. Portions of Salsola Kali bearing mature fruits, as described in Note 17, are also frequent. Seeds of Sophora tetraptera were found on the beach of Bahia San Vincente, whither they must have been brought by the Humboldt Current from the south, as I did not observe the tree in the vicinity. On this beach, as well as at Valparaiso, the prickly fruits of Franseria were abundant in the drift, doubtless derived from the plants growing on the same beaches. In addition we get as frequent components of the beach-drift materials that mark the white man’s presence over much of the globe. Corks are widely distributed over the beaches of the world; but on no coast have I found them more numerous than on the Valparaiso beaches. Here we find Medicago fruits, the empty stones of the cherry, the plum, and the peach, empty filberts and other materials, all of which I have gathered on the shores of the Straits of Messina and on English beaches. Amongst this medley we find also Casuarina cones and fruits of Eucalyptus. Then we find special indications of the New World in the pea-nut (Arachis hypogæa) and in the abundant seeds of a huge pumpkin (Cucurbita), which is a favourite food with the Chilian indigenes. These seeds are cited as an example of futile buoyancy in Chapter XIII.
THE PLANTLESS OR DESERT ZONE (NORTHERN CHILE).
This zone of the coast, which stretches north for some 700 miles from Coquimbo to near Arica (30°-18°30ʹ S. lat.), corresponds to the great desert region of North Chile. On the beaches of Antofagasta, Tocopilla, and Iquique, which are situated in the midst of this zone, I found no plants. This rainless sea border of barren mountains, presenting to the eye of the traveller from the deck of a passing steamer nothing but rock and sand, must be one of the most desolate coasts on our globe. It is therefore not a matter for surprise that the beaches are of dry loose sand in which the hand fails to find on scooping below the surface that refreshing coolness which is the character of beaches in all latitudes where the land is vegetated and a subsoil drainage seaward exists. Under ordinary conditions the sensation of moisture in the sand a few inches down is not produced by the mere proximity of the sea. On the Antofagasta and Iquique beaches the temperature in the heat of the day of the surface half inch ranged from 120° to 130° F., whilst four inches down it was 95° to 100°, and no moisture was found by scooping five or six inches down. On the Taltal beach, which lies towards the southern end of the desert region, I noticed, besides a few plants of Suæda fruticosa, two other species of the orders Santalaceæ and Nolanaceæ, evidently intruders from the inland regions. Where the zone of extreme aridity terminates at the north between Pisagua and Arica a few bushes are to be seen on the hill-slopes behind the beaches.
Very little seed-drift came under my notice on the beaches of the desert zone. Here and there I found a few Medicago pods and some seeds of the large pumpkin above noticed, but that was all. This is due as a rule to the seed-drift being masked by an enormous amount of rubbish, mostly brought from the south by the Humboldt Current. My walk for five miles along the beaches immediately north of Antofagasta gave me an experience in the way of stranded drift such as I have never met with on the beaches of any other region. All the dead bodies of the Chilian coast to the southward seem to have been stranded in the bend of Moreno Bay, on the shore of which Antofagasta lies; and the air was tainted with decaying flesh, the past being mixed up with the present in a most unrefreshing fashion. Besides carcases of sea-lions, six feet in length, sharks, dog-fish, and fish of many sorts, some of them dried up, others in a state of putrefaction, there were dead penguins, dead pelicans, dead sea-birds of other kinds, the bodies of horses, cattle, dogs, &c., all preyed upon by the numerous vultures and skuas, and in some localities by hungry-looking dogs of large size that took no notice of me as they slunk along. The past was represented by great quantities of bones that lay bleaching on the sand, with here and there a vertebra of a whale, making in all quite a varied osteological collection. But this was not all. Carcases of all sorts were drifting towards the beach. Here a vulture, there a skua, there again a dog stood just beyond the tide-wash looking keenly seaward; and by following the direction of their gaze one could see that each had marked down a carcase slowly drifting in. Now and then they would make a dash, scarcely waiting for the new arrival to be washed up by the waves. But there was no competition, since there was enough for all.
Under such conditions my investigation into the seed-drift was out of the question; but I saw what would be considered by some as more interesting, namely, the dead of many latitudes piled up on the beach by the Humboldt Current.
THE SESUVIUM ZONE (THE PERUVIAN COAST).
This zone, which comprises the whole Peruvian sea-border from Arica in 18°30ʹ S. to the vicinity of Tumbez in about 3°30ʹ S., usually possesses in its scanty littoral flora one or two species of Sesuvium, and in some places Sesuvium alone occurs on the beach. The beaches here do not line a region of almost complete aridity, as in the coast corresponding to the great desert region of North Chile. Though here also scarcely any rain falls, the sea-border receives the benefit of the “garuas” or drizzling sea-fogs; whilst the region immediately behind the coast may either be desert or semi-sterile during much of the year, or may be scantily vegetated, or, as along the river-valleys, may display a vegetation more fitting to the latitude. The general aspect, however, of the coast of Peru is one of aridity; but there are probably few beaches where a certain amount of subsoil drainage from the land sea-ward does not exist. This is well exhibited at Ancon, north of Callao, where in the most unlikely situations water is reached by digging wells; but in spite of this the Sesuvium alone grows on the beach. The beaches examined by me in the heat of the day in February, as at Mollendo and Ancon, had much the same surface-temperature noticed in the preceding month on the beaches of North Chile, namely, 120° to 130° F., and in one place 135°; whilst at a depth of four inches the sand was rather cooler, and instead of being between 95° and 100°, as on the Antofagasta and Iquique beaches, it was here usually only about 90°. But it was only occasionally that the sand felt at all moist at a depth of five or six inches; and in this zone, therefore, only a few shore plants of a peculiar type could be expected to find a station on the beaches, excepting, of course, those localities where low marshy districts or lagoons lie behind the beach.
The beach plants of the coast of Peru as observed by me though usually scanty, presented two types according to the character of the district bordering the beach. I make no mention here of those local plants, often belonging to the Compositæ, that as at Callao and Arica descend the valleys to the beaches, or to those numerous introduced plants that accompany cultivation, such as we find at Arica. In those coast localities, as at Arica, Callao, and Chancay, where salt-water pools or brackish lagoons lie behind the beach, or where a stream or a river empties into the sea, Sesuvium portulacastrum, Heliotropium curassavicum, and a Salicornia are to be generally noticed, and, as at Callao, Batis maritima may also abound. On the Chancay coast, about 30 miles north of Callao, there lies inside the shingle-beach a large shallow lagoon of brackish water (spec. gr. 1·012) with extensive muddy marginal flats, the temperature of the water at the edge being at mid-day on Feb. 3rd, 90° F. In the water flourished Ruppia maritima, which was also exposed in dead, dry, matted masses on the bordering mud-flats. On these mud-flats grew Sesuvium portulacastrum, which near the water’s edge was associated with a small species of Salicornia, whilst further away from the water it was accompanied by Heliotropium curassavicum.
But the most typical beach-flora of the Peruvian coast is such as we find on the dry beaches skirting the base of sand-covered or barren hill-slopes such as occur at Mollendo, Ancon, and Paita. As at Ancon, sand-covered hills and plains may extend miles inland, displaying here and there lines of shifting sand-mounds or “medanos.” On such beaches we may often find only a solitary plant, a species of Sesuvium which seems to differ only in its larger flowers, its much larger leaves (2 inches long), and its stout stems, of the thickness of the little finger, from the ordinary Sesuvium portulacastrum. This seems to be the only plant that can make its home on such beaches. At Mollendo, where there are signs of desiccated pools behind the beach which are occasionally filled with sea-water, the vegetation was of an intermediate character and more abundant; and here grew Sesuvium portulacastrum, a tall Salicornia, and Suæda fruticosa; whilst the commonest plant was a prostrate Nolanaceous species with a handsome purplish flower.
Excepting with the fruits of Batis maritima, and perhaps the buoyant joints of Salicornia, scarcely any of the prevailing shore-plants of the coast of Peru possess a capacity for dispersal by currents. In this zone I rarely found any seed-drift on the beaches. Much rubbish, such as roots of bamboos, however, may be brought down by the rivers; and where the Humboldt Current strikes a bend in the coast we get a repetition, on a smaller scale, of the scenes on the Antofagasta beaches. Ancon Bay, for instance, receives much of the floating offal of the south.
THE MANGROVE ZONE (THE COASTS OF ECUADOR AND COLOMBIA)
We come now to the mangrove zone which comprises, with the remarkable exception of a long stretch of arid sea-border to the north of the Gulf of Guayaquil, the whole remaining western sea-border of South America, namely, the Ecuadorian and Colombian coasts. My own acquaintance with this region is limited to the estuary of the Guayas or the Guayaquil River and to the southern shore of the Gulf of Guayaquil; but I am able to avail myself of the researches of Baron von Eggers, which cover the entire Ecuadorian coast; and with Ecuador, therefore, I will bring this brief sketch of the littoral flora of one side of a large continent to a conclusion.
The Ecuadorian coast, lying, as Baron von Eggers observes, between the rainless and desert coasts of Peru and the “ewig grüne” coasts of Colombia, may be regarded as a transition-area presenting very varied and complicated conditions. With the cause of the remarkable contrasts exhibited by the strand-flora, not only on the coast of Ecuador, but along the whole west coast of South America through some forty-five degrees of latitude from Patagonia to Colombia, I will presently deal. Here it may be remarked in passing that the Humboldt Current has played the determining part in producing the abnormal climatic conditions to which these remarkable contrasts in the strand-flora of this coast of the continent are mainly due.
The mangrove zone, marking a more or less abrupt transition from a region of drought and semi-sterility to one of humidity and rank tropical vegetation, begins about lat. 3° 30ʹ S., that is, in the vicinity of Tumbez, or perhaps nearer the boundary-line between Ecuador and Peru in lat. 3° 20ʹ (see Note 72). Occupying the southern shore of the Gulf of Guayaquil it extends up the Guayas estuary to Guayaquil and rather beyond. But when we follow the coast of Ecuador northward from the island of Puna towards Santa Elena Point, we come upon one of the most remarkable phenomena presented on the west coast of South America. The dry region begins again and the mangroves disappear; and these conditions continue through about 2-1/2 degrees of latitude until we reach the equator, when the mangrove zone soon recommences, and, as I infer, continues northward without a break to the coast of Central America.
Dealing first with the mangrove districts of the south side of the Gulf of Guayaquil and of the Guayas or Guayaquil estuary, we may observe that probably in few localities of the globe have the forces of nature worked more in unison to produce the conditions favouring the growth of the mangrove. The reason why this particular locality has been thus favoured will be discussed later on in this chapter. I may here observe that Baron von Eggers was so struck with the exceptional features of the mangrove-growth in this region that he was inclined to look for the American centre of the genus Rhizophora, the prevailing mangrove, in the estuary of the Guayas River.
I will not enter into a detailed description of the mangrove-formation of this coast, which has indeed been given by the German botanist; but I will merely refer to the leading features such as they presented themselves to me. In the first place, reference will be made to the sea-border of the province of Eloro, where I spent nine or ten days, making Puerto Bolivar, the port of Machala, my headquarters—a locality about thirty miles east of Tumbez. Except in the Guayas estuary I have never seen such a magnificent growth of mangrove.
By following the line of light railway that runs about six kilometres inland from Puerto Bolivar to Machala, the capital of the province, we obtain a good section of the mangrove-belt, which may be thus described. The mangrove-swamp proper extends about three kilometres inland. Whilst the small variety of Rhizophora mangle (mangle chico) immediately fronts the sea, Laguncularia grows on the islets close to the seaward margin of the swamp. When we enter one of the numerous broad creeks that intersect the border of the mangrove-belt we soon find ourselves in the true mangrove forest, where prevail tall trees of Rhizophora mangle (mangle grande) that rise to a height of 70 or 80 feet or more. Gloomy as the depths of the swamp are, they acquire quite a funereal aspect, the branches of the trees being draped with pendent Tillandsias. These long, hair-like, tangled growths hang vertically from the branches of the trees and may be 20 or 30 feet in length. In the rear of the zone of tall mangroves we come upon a more open district of the swamp. The forest proper gives place to a tract occupied by small trees of Rhizophora, Laguncularia, and Avicennia, with here and there whole acres occupied only by the shrubby Salicornia peruviana which attains the height of a man.
[_To face page 484._
(The main stream of the Humboldt current, as indicated by the arrows, turns off to the north-west at Cape Blanco; whilst a small branch crosses the mouth of the Gulf of Guayaquil and flows along the Ecuador coast north of Santa Elena Point.)
Here terminates the mangrove-swamp proper, and about three kilometres from the sea it passes gradually into a region of extensive bare mud-flats which are penetrated by salt-water creeks, two or three yards across and a foot or two in depth, that are bordered by low and shrubby Avicennias, the Salicornia bushes above noted, and dwarfed trees of Rhizophora mangle only four or five feet high. These flats, which are evidently only overflowed by the sea at the higher spring tides, were at the time of my visits much sun-cracked and in some parts incrusted with salt; but the mud was rather soft, and in places Sesuvium portulacastrum and Batis maritima flourished in quantity on it. These mud-flats, about two kilometres across, pass by degrees into the low-lying level district known as the Machala plains, on which the capital of the province is built. Here the soil is dryish, and, notwithstanding that it displays on its surface when exposed to the sun a white saline efflorescence, a dry jungle type of vegetation of the xerophilous character here thrives. I noticed casually the Algaroba (Prosopis), a yellow-flowered Cordia, cacti of the Opuntia and Cereus kinds, besides several small trees and shrubs often thorny.
These Machala plains, on account of the fine saline incrustation above mentioned, are of much interest, since at a distance of six kilometres from the coast they thus display on their surface the effect of sea-water infiltration, their level above the sea being only a few feet. We have seen the three stages of this infiltration landward of sea-water: first, the mangrove-swamps daily overflowed by the tide; second, the mud-flats behind them which are only overflowed by the fortnightly spring-tides; third, the vegetated plains behind all, which are sufficiently raised to be above the reach of the tides, but which are nevertheless soaked with sea-water that displays its presence in the salt left by evaporation on the surface of the soil.
But another interesting point is here raised. At the back of the mangrove-belt, in most parts of the world, we usually find a particularly rank and luxuriant vegetation where the Scitamineæ often take a leading part; whereas on the sea-border of this part of Ecuador the mangrove-swamps pass gradually into arid saliniferous plains. With this singular fact is to be associated the circumstance that we see here in operation, some four or five miles from the coast, a process by which great quantities of sea-salts are accumulating below the surface. This may possibly be concerned with the origin of the great saline deposits of Northern Chile. However this may be, there is some reason for believing—and I understand that this is the opinion of Dr. Wolff, the historio-geographer of Ecuador—that in the course of ages the tendency will be towards an extension of the dry, sterile regions of Northern Peru into Ecuador. This subject is referred to again in a later page of this chapter.
Whilst in this neighbourhood I made the ascent for some fifteen miles of the Santa Rosa River, which opens into the sea near Puerto Bolivar. It is a tidal estuary that has no proportion in size to the small river that enters it. In its lower third we passed at first between long mangrove-islands formed almost entirely, as viewed from the boat, of the tall Rhizophora trees draped with Tillandsias, and presenting really a magnificent spectacle. In the middle third we were penetrating into the rear of the mangrove-belt. The giant swamp-fern (Chrysodium aureum) abounded, and here and there we passed by a patch entirely held by the large shrubs of Salicornia peruviana. The tall Rhizophora trees were replaced by the short variety, the “mangle chico,” which ceased altogether about ten miles from the mouth of the estuary, but probably only about five miles from the nearest part of the coast. The water at the place where the Rhizophora trees ceased was evidently quite fresh during nine out of the twelve hours, being only salt in the latter part of the rising tide. Above the mangroves, in the upper third of the ascent, Hibiscus tiliaceus, with Chrysodium aureum, flourished on the banks. The shallows at the margins were occupied by a considerable variety of semi-aquatic and other plants, such as Pontederia (two species); one of the Alismaceæ, with the flower and fruit of Sagittaria and the leaves of Alisma; Typha, Polygonum, and an Amaryllid like Crinum. Plants of Pistia and Pontederia floated in the stream.
I have said enough to give a general idea of the composition of the mangrove-belt of the Ecuador littoral, and will refer but briefly to the mangroves and other river-side plants in the neighbourhood of the city of Guayaquil, some forty miles up the Guayas estuary. As I have remarked in Note 38, the water of the river off the city is usually quite fresh except at high water; but the sea has much freer access to the channels at the back of Guayaquil, where at high water the density was 1·014. In these channels are displayed the typical mangrove formation, trees of Rhizophora mangle bordering the water, whilst behind they are mingled with Avicennia tomentosa and Laguncularia. On the banks of the main river, where they are overflowed at high water, Anona paludosa was the most frequent tree, being associated with the Rhizophora, Hibiscus tiliaceus, and other trees. Above the city, Polygonum glabrum was growing in dense masses at the river’s edge, whilst Pontederia and Pistia flourished on the low muddy banks and floated in quantities in the river.
Before quitting the subject of the mangrove-formation of Ecuador, I will refer shortly to the two varieties of Rhizophora mangle that here occur. Baron von Eggers received the impression that the common type of this species, a low tree bordering the coast, did not exist in Ecuador, such a type as he says is characteristic of the West Indies and of Central America, and, I may add, also of Fiji. The species he regards as acquiring a new facies in Ecuador, where it exists as tall forest-trees, branchless for half their height, and exhibiting other divergent characters. However, I found that the common type of the species occurs normally on the coast in the vicinity of Puerto Bolivar, thirty miles east of Tumbez, a district above described.
There are two distinct forms of Rhizophora mangle exhibited in the mangrove-belt of the coasts around Puerto Bolivar. One of them, which the indigenes name “mangle chico,” is a small tree, 10 to 15 feet high, with useless timber, that immediately borders the sea, and, in fact, largely forms the margin of the swamp, not only on its seaward side, but also on the land side, where it passes into drier ground. The other, the “mangle grande,” a tall tree reaching to 60 or 80 and sometimes perhaps to 100 feet in height, composes the interior, and indeed the bulk, of the mangrove-belt, and possesses a hard and durable timber much employed in the district.
Distinct as these two types are, it is not difficult to find intermediate forms, and, in truth, in some localities they prevail. But the interesting point is that this peculiar Ecuadorian type of the species, a type that attracted the attention of the eminent German botanist, comes near the “Selala,” the mysterious seedless Rhizophora of the Fijian swamps—a subject fully discussed in Chapter XXX., where I have compared the Fijian and Ecuadorian Rhizophoras. Both the “Selala” of Fiji and the “mangle grande” of Ecuador are intermediate between the American Rhizophora mangle and the Asiatic R. mucronata, resembling the last in their inflorescence, but in other points approaching the American species. The “Selala,” however, comes nearer to the Asiatic tree, whilst the “mangle grande” comes nearer to the American tree. Unlike the Fijian tree, that of Ecuador is not sterile, but matures its fruit; and it displays no evidence of the vegetative reproduction so characteristic of the “Selala.”
Sandy beaches are not common on the mangrove-fronted shores of the south side of the Gulf of Guayaquil. However, on the seaward side of the long low mangrove island of Jambeli, on which the lighthouse is placed off Puerto Bolivar, there is a long stretch of beach of whitish, mainly non-calcareous, sand. The Coco palms behind the beach give the coast quite the aspect of a Pacific island strand. Ipomœa pes capræ flourishes on the sand nearest to the sea; and immediately behind, the beach is more or less occupied by a Cyperus 2 to 3 feet high, and by Canavalia obtusifolia. Further back grows a small Acacia tree, and behind it the yellow-flowered Cordia tree of the district; and in the rear of all lie extensive mud-flats, partly occupied by stunted bushes of Avicennia tomentosa and by Sesuvium portulacastrum, which in their turn pass into the mangrove-swamps.
On account of the enormous amount of drift of all descriptions that is carried to the sea by the Guayas or Guayaquil River, floating vegetable materials are abundant in the Gulf of Guayaquil, and are thrown up in quantity on the coasts of Ecuador. One of the most interesting spectacles at Guayaquil is presented by the floating river-drift. Huge tree-trunks and floating islets, the last-named ranging from 3 or 4 to 30 or 40 feet or more across, were, at the time of my visit in February, being carried to and fro unceasingly in front of the city by the tide, gradually making their way down the river, and ultimately reaching the open waters of the gulf. Floating plants of Pistia were in abundance; and their fate when they reached the sea must have been tragical. The islets were exceedingly interesting; they were evidently formed of materials lifted up bodily from the shallows at the margin of the river, and then carried off in the stream. They were mainly composed of two species of Pontederia and of Polygonum glabrum in the position of growth; the first often in flower. Pistia and a variety of smaller plants nestled among them, such as Salvinia, portions of Azolla, Lemna, &c.; and in one islet I noticed, oddly enough, the growing rhizome of a sensitive plant (Mimosa pudica). A great quantity of floating seeds collect amongst the roots and stems of the plants composing the islets, and here I obtained much of the smaller seed-drift.
Most frequent in the floating drift of the river at Guayaquil were the seeds of Anona paludosa, often in a germinating condition. The seeds are liberated by the decay of the floating fruit, which was also common in the drift. Amongst the larger materials were the seeds of Entada scandens and of Mucuna; the empty seeds of the vegetable-ivory palm (Phytelephas macrocarpa), the sound seed possessing no floating power; the “stones” of Spondias lutea, L., as identified by Mr. Holland, of the Kew Museum; the empty small nuts of several palms, including, apparently, Oreodoxa, &c. Amongst miscellaneous materials were small gourds, which are referred to in Note 47, and an occasional empty cacao fruit. Smaller seeds were also abundant, and included those of Hibiscus tiliaceus, Erythrina, Vigna, Ipomœa, and others. Carried into the river from the neighbouring mangrove-creeks, where they abound, there were floating seedlings of Rhizophora and Avicennia, fruits of Laguncularia often germinating, and the seeded joints of Salicornia peruviana.
There was of course, in addition, much that was strange in the floating drift of the Guayas River, which received its contributions not only from the river-side vegetation and the neighbouring mangrove-swamps, but also from the interior mountain ranges culminating in Chimborazo, the slopes of which are drained by its tributaries. I had several opportunities of meeting the drift of the Guayas River in the open waters of the Gulf of Guayaquil. Much of it is carried along the south side of the gulf; and I picked up at sea, ten to twenty miles from the mouth of the estuary, many of the things above enumerated, such as Erythrina and Mucuna seeds, seeds of Hibiscus tiliaceus, the empty vegetable-ivory seeds, the seedlings of Rhizophora and Avicennia, and the germinating fruits of Laguncularia and Salicornia peruviana. Much of these materials mingled with local drift is stranded on the long beach of Jambeli Island, thirty miles from the mouth of the estuary. Here, besides the seeds of Canavalia obtusifolia and Ipomœa pes capræ derived from the locality, I found the seedlings of Rhizophora and Avicennia, and the fruits of Laguncularia and Salicornia peruviana, that might have been in part derived from the adjacent swamps, as well as much of the drift of the Guayas River, such as the seeds of Anona paludosa, Entada scandens, Erythrina, and Mucuna, the small gourds, the same small palm-nuts, the empty seeds of Phytelephas, the “stones” of Spondias lutea, and much other material previously familiar to me, but nowhere a sign of the floating Pistias and of the flowering Pontederia islets of that estuary.
_The Stretch of Dry Coast from the Vicinity of Puna Island to the Equator._— This remarkable piece of sea-border, covering nearly three degrees of latitude, and in its aridity and general character recalling the sterile sea-coast of Peru, is placed between the humid mangrove-fronted coast of the Guayas estuary and the similarly humid and mangrove-fronted coasts of Northern Ecuador and Colombia. The mangrove seems to be almost absent from this stretch of dry coast. Mr. F. P. Walker, of the Santa Elena Cable Station, tells me that some time ago a little mangrove-growth existed near the Point, but that it has disappeared; and Baron von Eggers implies the absence of mangroves from the whole coast. The first-named speaks of the dry character of the coast district from Santa Elena Point to within half a degree of the equator; and the last-named, in his description of the coast, mentions cacti and thorny plants as typical of the vegetation. Since this region represents a typical locality where the direct influence of the Humboldt current on the climate of almost the whole west coast of South America can be put to the proof, I will refer to its peculiar climatic conditions below in my discussion of the general question, and will here content myself with saying that on this dry portion of the coast of Ecuador we have reproduced, but in a less pronounced degree, the climatic conditions of the coast of Peru.
_The Humboldt or Peruvian Current and the Climate of the West Coast of South America._—The question we will now briefly consider is one that is concerned with the determining causes of the singular distribution of coast-plants on the west coast of South America. The reader will have already seen that the matter is an affair of climate; but it is an affair of climate in which (although it affects forty or more degrees of latitude), latitude, in a general sense, scarcely counts. All the naturalists, from Humboldt onward, who have sojourned in this region of the globe have displayed a deep interest in this subject; and I suppose there can be no region of the globe where there are so many climatic anomalies as interesting to the meteorologist. Here, for instance, might be obtained materials for solving the irritating mystery of a London fog; and if the suggestion of Baron von Eggers, before alluded to, is carried out, and a station is established by the Meteorological Societies of Europe and America at some suitable locality like Santa Elena on the coast of Ecuador, we might obtain, among other results, another line of investigating the causes of the fogs of our metropolis, a subject about which Captain Carpenter has recently made an important preliminary inquiry.
I will assume that my readers are already acquainted with the nature of the problem to be discussed relating to the climate of the west coast of South America, and that they are familiar with the view generally held that the aridity of this extensive coast region, stretching from the thirtieth parallel of south latitude to the equator, arises from the loss by the trade-winds of all their moisture in the interior of the continent before reaching the western countries of Chile and Peru. Mr. Ball, in his book on South America, opposed this view, though from reasons only partially valid, since he instanced the Ecuador coast as being, contrary to the theory, a wet coast, whereas we know that a large stretch of it is arid and not unlike Peru. The parting of the ways in this discussion lies in the answer to the query, Why should the south-east trade carry so much moisture to the east side of South America, whilst the south-west winds, that are equally prevalent on the west coast of the continent, are drying winds which convert the sea-border into a desert, as in Northern Chile, or into a region of semi-sterility, as in the instance of Peru? Other things being equal, we should expect both sea-borders of the continent in these latitudes to be well watered. In the answer to the question why the south-east trade should be a wet wind and the south-west wind a dry one lies a fatal objection to the prevailing view.
When Professor Davis, in his article on North America in the _Encyclopædia Britannica_ (vol. 25), observes in connection with the arid coast regions on the west side of the continent that the southerly flow of the winds along the Pacific coast gives them a drying quality, thus causing the extension to the coast in South California and in North Mexico of the arid regions of the interior, he seems to imply that these winds acquire their drying capacity in flowing from cooler to warmer latitudes. On this view all trade-winds should be drying winds, whereas the reverse would appear to be the case.
There is some condition, present on one coast of the South American continent and absent on the other, which determines why a southerly wind, blowing landward, is in the one case moist and in the other dry. According to my own view the winds of the arid coast regions of western North America cross the cool waters of the Californian current, and thus acquire their drying quality on striking a sea-border more highly heated than the winds themselves. On the tropical west coast of South America the winds also become drying winds by passing over the cold waters of the Peruvian or Humboldt current, where mists are in consequence of frequent occurrence; and on striking the more highly heated land-surface at the sea-border the moving air does not part with any more moisture until an altitude of some thousands of feet above the sea is reached, when the cloud-belt forms. On the mountains bordering the coast of the Antofagasta province, in January, the clouds gathered at an elevation of 4,000 to 5,000 feet. Perhaps the best way to contrast the east and west coasts of tropical South America in this respect would be to say that whilst the wind blows landward in both regions, the land is the condenser on the east side, and the sea, owing to the interposition of the cold Humboldt current, is the condenser on the west side.
During a fortnight spent in February at Ancon, about twenty miles north of Callao, I noticed that with the prevailing cool south-westerly wind the coast was clear, but it was misty at sea. On the few days when there were warm westerly and north-westerly breezes, the weather was thick at sea; and if this condition was pronounced, the whole coast was enveloped in mist; but more usually the coast-line was fairly clear except at the promontories, along the sides of which clouds blown in from the sea rolled in lines inland, not generally attaining an elevation over 300 or 400 feet, but sometimes reaching 900 or 1,000 feet, and gradually disappearing a mile or two inside the coast-line. These sea-born clouds thus vanished as they traversed the more highly heated land-surface; and the air-current continuing its inland course mounted the slopes of the adjacent mountain ranges of the Andes, some three or four miles from the coast, until at an altitude of some 5,000 or 6,000 feet condensation again occurred and the cloud-belt was formed at those cooler levels.
From the summit of a range rising to a height of about 2,500 feet to the north-west of Lima I had presented to me a splendid spectacle, on February 12th, in the formation of the coast-belt of clouds. The forenoon was clear, but about 2 p.m. the sea-born clouds began to roll inland, concealing the lower two-thirds of the island of San Lorenzo, which has an elevation of almost 1,400 feet, and completely covering up Callao and the low country bordering the sea, but extending only a mile or two from the coast-line. The dense cloud that covered Callao appeared, as I looked down upon it from my mountain-peak, like a billowy field of snow sparkling in the sun, with the summit of San Lorenzo standing out like some bare alpine summit from amidst the snows. Yet beneath that dazzling covering Callao lay all in gloom; whilst only six miles up the broad valley of the Rimac the city of Lima stood in a blaze of sunlight, its domes and towers reflecting back the light as I looked at the strange contrast it presented with the buried city of the coast. The mystery of a London fog seemed to lie unfolded at my feet, ready for the man who can read the signs aright.
That the mere presence of a cold current on a coast with the winds blowing off the land (as in the case of the Labrador current, which extends down the Atlantic coast of North America to Cape Hatteras and beyond) produces no sterilising effect on the vegetation of the sea-border of a continent is well brought out in the beautifully executed maps in Prof. Russell’s recent work on North America. The essential condition for producing sterility on the sea-border of a continent is not only that the waters of a cold current should wash its coasts, but that the regular winds should blow landward across its cool surface. These are what we find on the west coast of South America.
Not with the hope of adding anything new to our knowledge of the climatology of this region, but with the purpose of becoming personally acquainted with the problem involved, I paid considerable attention to this subject during the three months passed on the west coast of South America between Port Valdivia and Guayaquil. It was not until I had dropped my thermometer into the cool water of the Humboldt current and had watched the formation of the fogs on the sterile coast of Peru that the real nature of the problem presented itself. From the pages of a work like Tschudi’s _Travels in Peru_ one acquires an excellent idea of the extraordinary climatic conditions of this region, and the same may be said of the narratives of Darwin and other travellers; but it is necessary to be brought into personal contact with these conditions before one can appreciate their significance.
As is well known, says Baron von Eggers, the Humboldt current explains the anomalous climate of the coast of Peru, and one may add of North Chile and Ecuador. The current, which represents the extension northwards of the west wind-drift of the Roaring Forties (see Dickson in _Encycl. Brit._, xxxi. 404; and Admiralty Current Charts of the Pacific), begins on the coast between the 33rd and 40th parallels of south latitude, according to the season. North of Valdivia, as we approach Valparaiso, in lat. 33°S., the effect of its presence is at once seen in the increasing dryness of the climate and in the alteration in the character of the vegetation. It has, however, been shown that the current needs the co-operation of the prevailing southerly and westerly winds as they blow landward over its cool waters. On the coast of Peru these moist winds often generate fog and mist as they cross the current. They reach the coast as drying winds, having a temperature much cooler than the lower coast regions; and the air-currents do not precipitate any moisture on the land until an elevation of 4,000 to 6,000 feet is attained where the cloud-belt is formed.
In order to establish this theory it is, however, necessary to show that when the Humboldt current leaves the coast normal conditions of humidity occur, to which the vegetation responds, and that when the current strikes the coast again the conditions of aridity reappear. In its course northward the current divides off Cape Blanco, the principal mass of its waters making towards the Galapagos Group, whilst the remainder, after crossing the Gulf of Guayaquil, flow along the coast of Ecuador between Santa Elena and the equator. Now, it is along this stretch of the Ecuador coast that the conditions of aridity reappear and that the climate of the Peruvian sea-border is in a modified form reproduced. In the interior of the Gulf of Guayaquil, on the other hand, where the sea-border is no longer subjected to the influence of the cold waters of the Humboldt current, the genius of the tropics, repressed through so many degrees of latitude, bursts its bonds, and presents us with a spectacle of littoral vegetation that, so far as mangrove-growth is concerned, is probably unrivalled on our globe.
This contrast is well shown in the mean annual temperatures on the opposite sides of the Gulf of Guayaquil. Baron von Eggers, quoting Dr. Wolff, states that whilst the mean for the year at Puna is about 75° F., and at Santa Elena about 73°, on the south side of the gulf at Balao it is several degrees warmer and is evidently not under 80°. The mean temperature for the second week of March during my sojourn at Puerto Bolivar, which is near the beginning of the mangrove region on the south side of the gulf, was 79°, the mean daily range being 74° to 83·5°. This stretch of dry coast reaching north from Puna to the equator is evidently regarded by Baron von Eggers and others who have studied the climatology of Ecuador as the critical area required to confirm the theory connecting the aridity of the west coast of South America with the Humboldt current. Here the sea for the greater part of the year has a temperature (according to the British Admiralty chart of surface-temperatures) of 70° to 75°; the mean temperature of the air is 73° to 75°; the rainy season, instead of covering a period of six months and over, as in the humid regions north and south of this coast, has a duration of only two or three months; the prevailing wind is south-west; whilst the direct influence of the cool waters of the current is shown in the general cloudiness that prevails during the last half of the year and in the drizzling mists that are frequent from June to October. Reference has already been made to the manner in which the vegetation on this dry coast of Ecuador responds to the arid conditions, as, for instance, in the absence of mangroves and in the prevailing character of the plants of the sea-border, cacti, thorny plants, and such like. For my information on this exceedingly interesting tract of coast, which is the test-ground of the Humboldt current theory, I am indebted to the papers of Baron von Eggers (see end of this volume) and to Mr. F. P. Walker, of the Central and South American Telegraph Company’s Station at Santa Elena, who very kindly communicated with me by letter. Some additional remarks are given in Note 73, and my own observations on the temperature of the Humboldt current from Antofagasta northward are summarised in Note 74.
Before quitting the Ecuador coast a word may be said relating to the prediction of Villavicencio that the climate of this sea-border will assimilate itself to that of the rainless coasts of Peru. This is, I believe, also the opinion expressed by Dr. Wolff in his _Geografia y Geologia del Ecuador_ (Leipzig, 1892); and it is referred to by Mr. Webster in his article on Ecuador in the seventh volume of the _Encyclopædia Britannica_. There is a prevailing impression amongst the more observing residents that I met in the Ecuadorian province of Eloro, on the Peruvian border, that the country is drying up. A few pages back I have described how in the Machala district of this province the mangrove-belt passes landward into an arid region suggestive of the sea-border of Peru. This transition is startling to one who expects to find behind the mangrove-belt, as he would find in most parts of the world, a humid region where Nature revels in the rank luxuriance of plant-growth. This is, however, not always the case, since on the lee or dry sides of the large islands of Fiji the mangrove-belt is backed by extensive arid plains, for an explanation of which, as I have shown in Note 22, we have to appeal rather to the hygrometer than to the rain-gauge. This is true also of Ecuador; but whilst the reason is intelligible enough in Fiji, it only carries us a step farther back in the case of the Machala plains in Ecuador. These plains are continuous with similar districts across the Peruvian border where they reach the coast; and if the reader will refer again to my description of the section of the mangrove-belt and the plains in its rear from Puerto Bolivar to Machala, he will incline to the view that the desiccation of the sea-border of Ecuador is now in progress.
Evidence of a more direct nature could doubtless be supplied by those who have long resided on the coast of Ecuador, and in illustration I will give an extract from one of Mr. Walker’s letters dated May, 1904, from Santa Elena.—“The rainfall here might for the last ten years be put down at two showers per year. It is said that the last good rainy season was in 1891. The inhabitants say that formerly it always rained enough to make the grass grow every year, but during the eleven years I have been here there appears to be a marked falling off of the rainfall.”
It has been only possible to touch the fringe of this interesting question here; but from the standpoint of the study of the littoral flora of the west coast of South America it is of some importance. Immediately behind the epoch of the present marine molluscan fauna of this coast there lies an age when, as we learn from Philippi, the shells of Chile were more akin to those of the Atlantic and Mediterranean faunas than to those now found on the Chilian coasts. The transition is a sudden one; and amongst other explanations of this strange transformation Suess suggests the sealing up of a communication through the Panama isthmus by volcanic eruptions and the appearance of the Humboldt current (_Das Antlitz der Erde_, French edit. by Margerie, ii. 825). May it not be, my readers may ask, that the west coast of South America is still in the age of progressive sterility; and that before this age began Peru possessed a normal tropical strand-flora? It has been remarked in Chapter VIII. that the same species of mangroves occur on both the Atlantic and Pacific coasts of America, and that at all events their present distribution belongs to an age when the Gulf of Mexico was in communication with the Pacific Ocean. May we not, again, suppose that in that age the mangroves extended far south on the coast of Peru, just as they do now on the coast of Brazil?
Coral reefs are stated not to exist in tropical latitudes on the west coast of South America in our own day; but we might almost expect that at the close of the Tertiary period, and perhaps before the appearance of the Humboldt current, they existed with the mangroves on the coast of Peru. As bearing on the subject of a change of climate on that coast in times geologically not remote, I may allude to the circumstance, which is discussed more in detail in Note 75, that I found, sometimes in fair quantity, blocks of massive coral, long since dead, much pierced by boring shells, and in places undergoing a chemical change, at Arica (lat. 18° 25ʹ S.), at Callao (12° 3ʹ S.), and at Ancon (11° 45ʹ S.) on the coast of Peru.
These masses, which varied from a few inches to two or three feet in size, gave me the impression of having been torn off the bottom, in some cases in recent times, in others perhaps centuries ago, by the huge sea-waves that from time to time overwhelm this coast. At Ancon, where they were sufficiently abundant to be used for bordering the flower beds in the hotel garden, they were most numerous in the vicinity of a rocky spur of andesite that protruded from the beach between the tide levels and was more or less covered at high water. A few paces inland from the beach some of these coral masses, evidently stranded long ago, were undergoing that queer process of disintegration which everything calcareous seems to undergo on the beaches and plains of this almost rainless coast. Like the bones of the Incas lying bleaching on the neighbouring plains, like the sea-shells and bones of bird and beast cast up long ago on the beach, they were falling to powder where they lay, and the coral fragment lay often in the midst of its own _débris_. The blocks on the beach proper were for the most part still hard and compact, and the same may be said of those observed on the beaches of Callao and Arica.
The corals were quite different from those with which I was familiar in the reefs of the Pacific islands, and, bearing in mind the known distribution of coral reefs, I was a little dubious about them. Accordingly I sent some specimens to the British Museum, and Mr. Jeffrey Bell has kindly informed me that they seem to be decayed and much injured perforated examples of Porites. When powdered they effervesce in an acid, but the bulk of the material remains undissolved.
No more eloquent testimony could be afforded of the rainless climate than these corals crumbling on the Ancon plains when washed a few paces inland from the beach. They could be noticed in all stages of disintegration from the block surrounded by a little line of disintegrated material, representing the initial products of its own decay, to the crumbling mass, almost friable in the fingers, that was lying in the midst of its own dust and loose polyp-tubes, and finally to the little mound of _débris_ that alone remained. Mr. Darwin, in his _Journal of Researches_ (chap. xvi.), refers to a similar process of decay in the elevated shell-beds of San Lorenzo, off the coast of Callao. On the higher terraces a layer of saline powder, consisting of sulphates and muriates of lime and soda but with very little carbonate of lime, was the sole indication of the shell-beds. Dry climatic conditions at the sea-border evidently favour, as he observes, the early decay of exposed calcareous remains.
_The Shore-plants and Stranded Seed-drift of the Panama Isthmus._
I spent two days at Panama and two days at Colon in examining the neighbouring beaches and estuaries of the Pacific and Atlantic coasts of the isthmus. On the Panama side the mangrove-belt was formed on the seaward border of “mangle chico” (the small prevailing type of Rhizophora mangle), Laguncularia, and Avicennia; whilst behind it passed into extensive swampy tracts occupied by the Swamp Fern (Chrysodium aureum), Hibiscus tiliaceus, and other plants. On the Colon or Atlantic side the mangrove-belt had precisely the same composition and presented the same species, Rhizophora and Avicennia usually forming the outposts on the reef-flat, whilst Laguncularia was abundant in the rear. In the estuary of the Rio Chagres, Rhizophora and Laguncularia were abundant near the mouth, and Chrysodium aureum and Hibiscus tiliaceus by the waterside higher up. Dr. Seemann, in his volume on the botany of the voyage of H.M.S. _Herald_, observes that the species of Laguncularia common on both the Atlantic and Pacific coasts of the Panama isthmus is L. racemosa. This species differs in the form of its fruit from the Ecuador tree. Laguncularia racemosa, Rhizophora mangle, and I may add Anona paludosa and Conocarpus erecta, are all plants of the mangrove-formation that occur not only on the Pacific and Atlantic coasts of America but also on the west coast of Africa. It is likely, I may add, that the “mangle grande,” the Ecuadorian type of Rhizophora mangle, exists in the Panama isthmus, since in the higher part of the estuary of the Chagres I found trees approaching it in characters.
Amongst the plants growing on the Panama beaches I noticed Canavalia obtusifolia, Hibiscus tiliaceus, and Ipomœa pes capræ, all of which occur also on the Atlantic side of the isthmus. The Manchineel (Hippomane mancinella), found also on the Atlantic side of the continent, grows on the Panama beaches. Its fruits, which look like crab-apples, lose their outer fleshy covering when drying on the sand. Not being familiar with this poisonous tree, I allowed some of the milky sap of the fruits to touch the skin, and suffered great pain for five or six hours. The fruit possesses an inner coat of air-bearing cork-like tissue; and the stone, if I may so term it, thus acquires great floating power. I kept some afloat in sea-water for five weeks, and no doubt they will float for months.
The seed-drift to be observed stranded on the beaches and floating in the estuaries on both sides of the isthmus is, generally speaking, the same—a circumstance of great importance in plant-distribution, since we can here see rivers bringing down the same seeds from the same “divide” to the shores of the Pacific and Atlantic oceans. In the case of a plant like Entada scandens, which grows in the interior, this is a matter of much interest, as it thus possesses here a centre of dispersal from which its seeds can be carried by the currents eastward to the West African coast and westward across the Pacific to Malaya and (given time) around the shores of the Indian Ocean to the East African coast. In describing the possible routes of dispersion from this centre I have described the distribution of the species.
I am indebted to Mr. Holland, of the Kew Museum, for the identification of some of the drift-seeds and fruits collected by me on the isthmus, those identified by him being followed by the letter H. On the beaches and floating in the estuaries on both sides of the isthmus I found Rhizophora seedlings; seeds of Entada scandens and Mucuna urens (medic.), H.; seedvessels of Spondias lutea (Linn.), H.; Prioria copaifera (Griseb.), H., with decayed seed; and the empty nuts, 1-1/2 to 2 inches in size, of more than one species of Astrocaryum, H. Although in the case of the two last-named genera the seedvessels were useless for dispersal, being evidently brought down from the interior by the rivers, they serve to illustrate the important principle that the rivers bring down the same seed-drift on both the Atlantic and Pacific coasts of Central America. Mr. Hemsley includes amongst the seed-drift stranded on the coast of Jamaica the seedvessels of Spondias (probably S. lutea) and of Astrocaryum (_Bot. Chall. Exped._, iv. 299, 304).
Those of Spondias lutea were found by me floating in the Guayaquil River and stranded on the beaches of Ecuador and of the Pacific and Atlantic coasts of the Panama isthmus. This is the Hog-plum, which in tropical America and the West Indies is both wild and cultivated. Its buoyant “stone” has a covering of cork-like air-bearing tissue. This is a remarkable case of non-adaptation in the matter of buoyancy. The seedvessels cut across contained sound seeds; and they are provided with the essential qualities of “long floaters.”
_Summary._
(1) The strand-district of the west coast of South America is divided into four zones:—
(_a_) The Convolvulus soldanella zone of Southern Chile.
(_b_) The Desert or Plantless zone of Northern Chile.
(_c_) The Sesuvium zone of Peru.
(_d_) The Mangrove zone of Ecuador and Colombia.
(2) The mangroves do not extend south of Ecuador or, more strictly, south of Tumbez (3° 30ʹ S.).
(3) The absence of mangroves on the tropical coasts of Chile and Peru is attributed to the Humboldt current, which has so influenced the climate that it has converted the sea-border of North Chile into a desert and that of Peru into a region of semi-sterility.
(4) It is considered that this has been effected through the prevailing winds acquiring drying qualities on crossing the cold waters of the current in tropical latitudes.
(5) To establish this it is shown that when the Humboldt current leaves the coast at Cape Blanco mangroves thrive in the Gulf of Guayaquil, and that when it strikes the coast again near Santa Elena Point and courses along that seaboard to the equator we find the Peruvian conditions of semi-sterility reproduced.
(6) The probability that the arid climate of Peru is in our own time extending northward into Ecuador is pointed out; and from the presence of old coral blocks on the Peruvian beaches it is considered likely that when these corals throve the mangroves extended far down the coast of Peru.
(7) It is shown from the presence of the same species of mangroves on the Pacific and Atlantic coasts of America that there must have been, not long ago, a communication between these two oceans across Central America; but it is at the same time observed that this could not be inferred from shore-plants with buoyant seeds that, like Entada scandens, occur inland, since, although they occur on both sides of the continent, their distribution can be explained by the transport of their seeds by rivers to the Atlantic and Pacific coasts, such as we see in operation on the Panama isthmus in our own day.
(8) Stress is laid on the great development of mangroves in the Gulf of Guayaquil and in the Guayas estuary; and it is pointed out that there are in this locality two varieties of Rhizophora mangle, a large and a small variety, the first approaching in some of its characters the Asiatic species, R. mucronata, and being akin also to the seedless intermediate form of Fiji.
(9) Amongst other matters dealt with in this chapter are the floating seed-drift of the Guayas or Guayaquil River and the shore plants and stranded seed-drift of the Panama isthmus. From the locality last named we learn that rivers bring down from the interior to the Atlantic and Pacific coasts much the same seed-drift, and that from this centre littoral plants with buoyant seeds can be distributed over the whole tropical zone.