Natural history

Reproduction

Most clupeiforms lay their eggs near shore, often close inshore or in fjords and bays. Few clupeiform species spawn far from shore or in the open sea, except, notably, the Atlantic herring (Clupea harengus harengus), which spawns on offshore banks. The majority of the spawning grounds are limited to shallow waters ranging from slightly below mean low-tide level to a depth of about 4 metres (about 13 feet). Some clupeiforms, however, such as the Atlantic herring, do spawn at depths of 40 to 200 metres (approximately 130 to 660 feet). The bottom of the spawning grounds, especially those of species with sticky eggs, tends to be clean, hard, and covered with gravel and sand. Spawning takes place above a soft muddy bottom only if there is a vegetative cover. The freshwater and anadromous clupeiform species spawn in currents of riverbeds with a low mineral content, in shallows of big lakes, and (less often) in river arms and riverine lakes.

The majority of clupeiform fishes have pelagic (free-floating) eggs, which float in the surface or bottom water layers. Egg position is maintained by the presence of a large swollen space between the egg itself and the outer membrane. Some forms (Clupea, Pomolobus) have sticky eggs with an adhesive secretion, so they stick to stones, gravel, or plants shortly after being released. Freshwater and anadromous clupeiforms usually have eggs slightly heavier than water. The slightest current and turbulence, resulting from wave action and convection of the water, constantly lift such eggs, which would normally sink to the bottom. In rivers they freely drift downstream above the bottom. Only a few freshwater forms, such as the freshwater sardine (Clupeonella abrau), have eggs that develop in the surface water.

The number of eggs produced varies greatly, but, in general, smaller species produce few eggs, larger species produce many. One of the smaller sprats (Sprattus sprattus phalericus), with a maximum size of 8 cm (about 3 inches), produces about 2,000 eggs; one of the biggest shads, Alosa kessleri kessleri, can produce more than 300,000 eggs; and menhaden (several species of Brevoortia) produce more than 500,000. Freshwater species usually have more eggs than marine species of comparable size, evidently an adaptation against the higher mortality in riverine conditions.

Those species of clupeiforms with adhesive eggs produce more eggs than do those with free-floating eggs. Apparently, eggs that develop while sticking to the bottom have a much higher mortality rate from predators than do eggs that develop while floating in the surface water. Of great importance in reducing mortality rates is “repeated portion” spawning. In the majority, if not in all, clupeiform fishes, the eggs in the gonads do not become ripe all at once but in two or more portions. As a result, more eggs develop in the limited space of the body cavity, and the chances that some survive are enhanced if the first are destroyed.

The many causes of spawn mortality range from those of a physical character, such as wave action and sudden temperature drops, to biological ones, such as predation by gulls and ducks. An important protective mechanism against destruction of the abundant schools is the remarkably early age at which they first breed; females begin frequently to spawn only a few months after hatching. This, coupled with high fecundity, gives the order a high reproductive potential.

The duration of egg development varies from a few hours to nearly two months. An important factor in the rate of development is the temperature of the surrounding water; the cold-water herrings have the longest developmental period. The egg development of the Atlantic herring takes as long as 47 to 50 days at a temperature of 0.1 °C (just above 32 °F) but only eight days at 19 °C (66 °F). Some shad eggs develop in about 75 hours at a temperature of 17 °C (63 °F); however, they require only 49 hours at 19 °C. The eggs of the Tanganyika sardine (Stolothrissa tanganicae), a species that spawns at the surface in open areas of freshwater environments, hatch in 24 to 36 hours. The eggs constantly sink from the surface to a depth of 75 to 150 metres (250 to 500 feet) at a temperature of 25 °C (77 °F).

Growth and mortality

The thin, threadlike, newly hatched larva has a shape characteristic of nearly all clupeiform fishes, but its behaviour varies greatly, depending on the habitat. In marine species such as the Pacific herring (Clupea pallasii), the larvae, shortly after hatching, tend to be concentrated near the surface and usually stay a long time in the area of the spawning ground. The larvae of the Atlantic herring at first tend to make short, upward movements from the spawning beds on the bottom and then sink back again. They start to make horizontal movements within two hours after hatching, and after six hours they start to form swarms. As their length increases, the vertical movements become more and more pronounced, particularly at night. Larvae have been found to be dispersed by currents at depths of from 1 to 600 metres (roughly 3 to 2,000 feet). Later, juveniles drift with the current on the surface, sometimes as far as 1,300 km (slightly more than 800 miles). The larvae of the Tanganyika sardine, less than 2 mm (0.08 inch) long, tend after hatching to come straight up by swimming movements of the tail, which is the only flexible part of the body; they sink, however, as soon as they stop movement. Such vertical movement is vital, because the larvae would not survive were they to sink below the level of oxygenated water (80–200 metres [roughly 260–650 feet]). As they grow, the larvae gradually move to the surface waters; when they are about 5 to 6 mm (0.2 to 0.24 inch) long, they move toward the shore. They form schools when about 10 mm (0.4 inch) in size. In the Atlantic menhaden (Brevoortia tyrannus), a species that spawns in riverine environments, the newly hatched pelagic larvae first drift downriver between fresh and brackish water and shoreward from spawning areas and into estuarine nursery areas. Later, pelagic juveniles tend to move upstream as far as 50 km (31 miles), emigrating into the sea only after nearly one year.

In the early stages of life, all clupeiforms are subject to a high mortality rate, by predation of larger fishes, birds, comb jellies (ctenophores), and arrowworms (chaetognathans) and by being carried out of sheltered bays into localities in which the proper food is lacking. Mortality has been estimated at well over 99 percent, but, because of the extremely high fecundity, the distribution, and the early maturity, the recruitment of new breeding individuals remains high. The age of first sexual maturity is seldom more than three years, and the length at maturity rarely exceeds more than 15 cm (approximately 6 inches). Late-spawning species are usually larger and move over long distances. The age at first breeding is broadly correlated with rate of growth of the individual and with maximum length attained by the species, but there are other determining factors, some of which are unknown. The Siberian shad (Alosa saposhnikovi), the Baltic sprat (Sprattus sprattus balticus), and the Clupeonella engrauliformis all mature at two to three years of age but at lengths of 160 to 200 mm (roughly 6.25 to 8 inches), 120 to 130 mm (4.75 to 5 inches), and 85 to 100 mm (3.3 to 4 inches), respectively. Different populations of a species may vary in their growth rates; the races of the Atlantic herring vary from two to seven years in age at maturity and from 100 to 185 mm (4 to 7.25 inches) in length at maturity. In anadromous populations of the alewife (A. pseudoharengus), sexual maturity occurs at three to four years of age and at 150 to 170 mm (roughly 6 to 6.75 inches) in length, but those of landlocked populations breed at one to two years of age and at 95 to 100 mm (3.75 to 4 inches).