Fish
Fish are by far the most important vertebrates inhabiting fresh waters. In Britain, they are more widely distributed and more abundant than other aquatic vertebrates and their value both in terms of human food and sporting interest runs into millions of pounds annually. Relatively few fresh waters in Britain are fishless though many isolated lakes must owe their present fish stocks to man-made introductions in the past. Fish are not found in Britain in lakes above 850 m in altitude and although attempts have been made to introduce fish to lakes above this level the stocks soon died out. Waters that frequently dry out in summer months are also fishless, as no British species is truly resistant to desiccation, although certain cyprinids, such as the tench
Tinea tinea, may survive for short periods in wet mud. Very acidic peat pools are also generally without fish, as pH values of less than about 4.5 are directly toxic to most fish species. Fish are now absent from the most grossly polluted stretches of rivers running through industrialised Britain, and from a small number of lakes where ancient mine workings cause unusually high concentrations of toxic heavy metals in the water.
The habitat preferences of the British freshwater fish are somewhat obscured by the uneven geographical distribution of several species. With the exception of those which can tolerate sea water and can enter the sea, the powers of natural dispersal of freshwater fish are very limited. Eels Anguilla anguilla, salmon Salmo salar, brown trout Salmo trutta (and its sea-going race the sea trout), flounder Platichthys flesus, three-spined stickleback Gasterosteus aculeatus, ten-spined stickleback Pungitius pungitius, marine lamprey Petromyzon marinus, and river lamprey Lampetra fluviatilis, frequently enter the sea and have become distributed in much of the British Isles in suitable waters. Apart from these, few fish have spread further north than the central lowlands of Scotland after their recolonisation of south-east England in the last interglacial, while Britain was still connected to the Continent. The two most successful colonists are the minnow Phoxinus phoxinus, which has been largely spread by man as fish bait, and the perch Perca fluviatilis, which lays long sticky ribbons of eggs which attach to wildfowl. Until man recently assisted their dispersion by canal-building and deliberate introductions, some species (e.g. barbel Barbus barbus, bleak Alburnus alburnus, ruff or pope Gymnocephalus cernua, white (or silver) bream Blicca bjoerkna and burbot Lota lota) were more or less restricted to those British rivers which flowed into the Rhine when the landbridge between Britain and the Continent was intact. Gudgeon Gobio gobio, dace Leuciscus leucisais, chub Squalius cephalus, rudd Scardimus erythro- plithalmus, tench Tinea tinea and bream Abramis brama, have extended as far north as the Scottish Border. Some species have crossed the barrier of the Southern Uplands of Scotland and have colonised lowland Scotland and some have just penetrated the Highlands. This group comprises grayling Thymallus thymallus, roach Rutilus rutilus, stone loach Noemacheilus barbatulus, and bullhead Coitus gobio. The pike f^sox Indus extends north as far as Ross but its natural distribution is obscured by its introduction as a food fish in the past. Other species such as the Coregonus spp., C. lavaretus and C. albula, and charr Salvelinus alpinus, are glacial relicts and are now restricted to generally deep waters in the north and west.
OLIGOTROPHIC   LAKES
In Britain such waters generally have a fish fauna dominated by salmonids. In those lakes containing fish, brown trout are invariably present and, where the outflow gives uninterrupted access to the sea, these are joined by salmon and sea trout. These species of salmonid require a clean gravel through which a current of water is flowing in which to spawn. Many such lakes, particularly those that are large and deep, contain charr, and a few in Wales, the Lake District and South Scotland contain white fish Coregonus lavaretus or vendace C. albula. Charr can seldom survive in competition with Coregonus spp. and few lakes in Britain contain both. Oligotrophic lakes also contain the ubiquitous eel and three-spined stickleback as well as one or more of the three lamprey species (Lampetra planeri, L. fluviatilis and Petromyson marinus). Minnows are not found in Scotland much further north than the Great Glen but are widespread in stony-shored lakes further south. Perch and pike are also found in many oligotrophic lakes but become more dominant in slightly richer waters. In lakes within the range of the cyprinid fishes (south of Loch Lomond) species such as roach may be found but only become abundant in more eutrophic waters. Stream-dwelling fish such as stone loach and bullhead may extend out from stream mouths into oligotrophic lakes.
MESOTROPHIC   LAKES
In mesotrophic lakes in England and Wales perch, pike and various cyprinid fishes, especially roach, may begin to replace the salmonid fishes. Thus, in Esthwaite Water with increasing nutrient enrichment, roach and rudd have greatly increased in numbers in recent years at the expense of the trout population. Small numbers of roach and rudd have been caught more recently in the south basin of Winder-mere. Esthwaite is the most evolved of the larger Lake District lakes and is the only one which does not contain either Coregonus spp. or Salvelinus alpinus. Charr are generally only found in oligotrophic lakes in Britain, though they were formerly found in Loch Leven (alkalinity = 35-70 p.p.m. CaCO3), Kinross, and occur in one marl lake, Loch Borralie (alkalinity = 140-150 p.p.m. CaCO3). The richest water in which Coregonus is now found in Britain is the Mill Loch, Lochmaben (alkalinity = 50-60 p.p.m. CaCO3), and they have disappeared from the neighbouring shallower and more enriched Castle Loch.
EUTROPHIC LAKES
Cyprinids are usually the most abundant fish in eutrophic lakes in England and Wales but, in Scotland, outside the range of this family the fish fauna may be very similar to that of oligotrophic waters. In eutrophic waters in Scotland self- maintaining populations of fast-growing brown trout may exist. On the other hand, the silted conditions of the inflows in many eutrophic waters in the south of Britain may prevent natural spawning and the maintenance of a trout fishery will depend on regular re-stocking. In such cases rainbow trout Salmo gairdneri are often stocked instead of the native brown trout. The murky phyto-plankton-laden waters of many eutrophic lakes militate against visual feeders such as the salmonids and favour fish such as the cyprinids which use their sense of smell to locate food. The latter also require weed-beds in which to spawn and these are generally more abundant in eutrophic lakes. Consequently, where there is natural competition between these two groups, the cyprinids are usually favoured in eutrophic waters and the salmonids often eliminated.
Of the few species of fish which only occur in abundance in enclosed standing waters, carp and tench are bottom feeders, and rudd feed at the surface. Many other species found in standing waters also occur in the slower stretches of lowland rivers (the so-called bream zone).
A few exotic species have been introduced into small lakes and ponds in southern England. These include the wels Silurus glanis, which came into Britain with the aquarium trade and has since been spread by anglers, the pike-perch Stizostedion lucioperca, and various American bass belonging to the Centrarchidae. The carp was originally introduced into Britain by monks in the sixteenth century and is now widely distributed in scattered localities throughout lowland Britain, but is abundant only in the southern half of England. The bitterling carp Rhodeus sericeus is now fairly widely distributed in canals and slow-flowing rivers in England, where it has been introduced by aquarists.