Fig. 297.—Oregon Lamprey, Entosphenus tridentatus, ascending a brook. (Modified from a photograph by Dr. H. M. Smith. Published by Prof. H. A. Surface.) Willamette River, Oregon.
"In 'running' they move almost entirely at night, and if they do not reach a suitable spawning site by daylight, they will cling to roots or stones during the day and complete their journey the next night. This has been proven by the positive observation of individuals. Of the specimens that run up early in the season, about four-fifths are males. Thus the males do not exactly precede the females, because we have found the latter sex represented in the stream as early in the season as the former, but in the earlier part of the season the number of the males certainly greatly predominates. This proportion of males gradually decreases, until in the middle of the spawning season the sexes are about equally represented, and toward the latter part of the season the females continue to come until they in turn show the greater numbers. Thus it appears very evident in general that the reproductive instinct impels the most of the males to seek the spawning ground before the most of the females do. However, it should be said that neither the males nor the females show all of the entirely sexually mature features when they first run up-stream in the beginning of the season, but later they are perfectly mature and 'ripe' in every regard when they first appear in the stream. When they migrate, they stop at the site that seems to suit their fancy, many stopping near the lake, others pushing on four or five miles farther up-stream. We have noted, however, that later in the season the lower courses become more crowded, showing that the late comers do not attempt to push up-stream as far as those that came earlier. Also it thus follows, from what was just said about late-running females, that in the latter part of the season the lower spawning beds are especially crowded with females. In fact, during the early part of the month of June we have found, not more than half a mile above the lowest spawning bed, as many as five females on a spawning nest with but one male; and in that immediate vicinity many nests indeed were found at that time with two or three females and but one male.
"Having arrived at a shoal which seems to present suitable conditions for a spawning nest, the individual or pair commences at once to move stones with its mouth from the centre to the margin of an area one or two feet in diameter. When many stones are thus placed, especially at the upper edge, and they are cleaned quite free of sediment and algæ, both by being moved and by being fanned with the tail, and when the proper condition of sand is found in the bottom of the basin thus formed, it is ready to be used as a spawning bed or nest. A great many nests are commenced and deserted. This has been left as a mystery in publications on the subject, but we are well convinced that it is because the lampreys do not find the requisites or proper conditions of bottom (rocks, sand, etc., as given below) to supply all their needs and fulfill all conditions for ideal sites. This desertion of half-constructed nests is just what would be expected and anticipated in connection with the explanation of 'Requisite Conditions for Spawning,' given below, because some shallows contain more sand and fewer stones, and others contain many larger stones but no sand, while others contain pebbles lying over either rocks or sand. The lampreys remove some of the material, and if they do not find all the essentials for a spawning nest, the site is deserted and the creatures move on."
Requisite Conditions for Spawning with Lampreys.—"For a spawning site two conditions are immediately essential—proper conditions of water and suitable stream bed or bottom. Of course with these it is essential that no impassable barriers (dam or falls) exist between the lake and the spawning sites to prevent migration at the proper 'running' season. Lampreys will not spawn where there is no sand lying on the bottom between the rocks, as sand is essential in covering the eggs (see remarks on the 'Spawning Process'); neither will they spawn where the bottom is all sand and small gravel, as they cannot take hold of this material with their mouths to construct nests or to hold themselves in the current, and they would not find here pebbles and stones to carry over the nest while spawning, as described elsewhere. It can thus be seen that, as suggested above, the reason they do not spawn in Fall Creek and Cascadilla Creek, between the lake and the falls, is that the beds of these streams are very rocky, being covered only with large stones and no sand. There is no doubt that the lampreys find here suitable conditions of water, but they do not remain to spawn on account of the absence of the proper conditions of stream bed. Again, they do not spawn in the lower course of the inlet for a distance of nearly two miles from the lake, because near the lake the bed of the stream is composed of silt, while for some distance above this (up-stream) there is nothing but sand. Farther up-stream are found pebbles and stones commingled with sand, which combination satisfies the demands of the lampreys for material in constructing nests and covering eggs. The accessibility of these sites, together with their suitable conditions, render the inlet the great and perhaps the only spawning stream of the lake; and, doubtless, all the mature lampreys come here to spawn, excepting a few which spawn in the lower part of Six-mile Creek, a tributary of the inlet.
"As the course of the stream where the beds abound is divided into pools, separated by stony ripples or shallows, the nests must be made at the ends of the pools. Of the spawning beds personally observed during several seasons, nine-tenths of the entire number were formed just above the shallows at the lower ends of the pools, while only a few were placed below them. An advantage in forming the nest above the shoals rather than below it is that in the former place the water runs more swiftly over the lower and middle parts of such a bed than at its upper margin, since the velocity decreases in either direction from the steeper part of the shallows; and any organic material or sediment that would wash over the upper edge of the nest is thus carried on rather than left as a deposit. When formed below the shallows, owing to the decreased velocity at the lower part of the nest compared with that at the upper, the sediment is likely to settle in the hollow of the nest, and, through the process of decay of the organic material, prove disastrous or unfavorable for the developing embryos.
"The necessity of sand in the spawning bed indicates the explanation of why we see so many shallows which have no spawning lampreys upon them, while there are others in the same vicinity that are crowded. There will be no nests formed if there is too little or too much sand, not enough or too many stones, or stones that are all too small or all too large. The stones must vary from the size of an egg to the size of a man's hand, and must be intermingled with sand without mud or rubbish.
"The lampreys choose to make their spawning nests just where the water flows so swiftly that it will carry the sand a short distance, but will not sweep it out of the nest. This condition furnishes not only force to wash the sand over the eggs when laid, but also keeps the adult lampreys supplied with an abundance of fresh water containing the dissolved air needed for their very rapid respiration. Of course in such rapid water the eggs are likely to be carried away down-stream, but Nature provides against this by the fact that they are adhesive, and the mating lampreys stir up the sand with their tails, thus weighing down the freshly laid eggs and holding them in the nest. Hence the necessity of an abundance of sand at the spawning site."