Three-spined stickleback
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[Photo] Male Three-spined Stickleback (Gasterosteus aculeatus) with red throat and shiny blue eye. Date April 9. 2007. Author Piet Spaans (http://commons.wikimedia.org/wiki/User:Viridiflavus)

Copyright (C) 2007 Piet Spaans Permission is granted to copy, distribute and/or modify this document under the terms of the GNU Free Documentation License, Version 1.2 or any later version published by the Free Software Foundation; with no Invariant Sections, no Front-Cover Texts, and no Back-Cover Texts. A copy of the license is included in the section entitled "GNU Free Documentation License".

Distribution and Morphological Variation
The three-spined stickleback, Gasterosteus aculeatus, is a fish native to much of northern Europe, northern Asia and North America. It has been introduced into parts of southern and central Europe.

Three subspecies are currently recognised by the IUCN:

Gasterosteus aculeatus aculeatus is found in most of the species range, and is the subspecies most strictly termed the three-spined stickleback; its common name in England is the tiddler, although "tittlebat" is also sometimes used.
G. a. williamsoni, the unarmored threespine stickleback, is found only in North America; its recognised range is southern California, though there are isolated reports of it occurring in British Columbia and Mexico;
G. a. santaeannae, the Santa Ana stickleback, is also restricted to North America.

These subspecies actually represent three examples from the enormous range of morphological variation present within three-spined sticklebacks. These fall into two rough categories, the anadromous and the freshwater forms.

The anadramous form spends most of its adult life eating plankton in the sea, and returns to freshwater to breed. The adult fish are typically between 6 and 10 cm long, and have 30 to 40 lateral armour plates along their sides. They also have long dorsal and pelvic spines. The anadromous form is morphologically similar all around the Northern Hemisphere, such that anadromous fish from the Baltic, the Atlantic and the Pacific all resemble each other quite closely.

Three-spined stickleback populations are also found in freshwater lakes and streams. These populations were probably formed when anadromous fish started spending their entire life cycle in freshwater, and thus evolved to live there all year round. Freshwater populations are extremely morphologically diverse, to the extent that many observers (and some taxonomists) would describe a new subspecies of three-spined stickleback in almost every lake in the Northern Hemisphere. One consistent difference between freshwater populations and their anadromous ancestors is the amount of body armour, as the majority of freshwater fish only have between 0 and 12 lateral armour plates, and shorter dorsal and pelvic spines. However, there are also large morphological differences between lakes. One major axis of variation is between populations found in deep, steep sided lakes and those in small, shallow lakes. The fish in the deep lakes typically feed in the surface waters on plankton, and often have large eyes, with short, slim bodies and an upturned jaw. Some researchers refer to this as the limnetic form. Fish from shallow lakes feed mainly on the lake bed, and are often long and heavy bodied with a relatively horizontal jaw and a small eye. These populations are referred to as the benthic form.

Since each watershed was probably colonised separately by anadromous sticklebacks, it is widely believed that morphologically similar populations in different watersheds or on different continents evolved independently. There is a unique population in the meromictic Pink Lake in Gatineau Park, Quebec.

One fascinating aspect of this morphological variation is that a number of lakes contain both a limnetic and a benthic type, and these do not interbreed with each other. Evolutionary biologists often define species as populations that do not interbreed with each other (the Biological Species Concept), and thus the benthics and limnetics within each lake would constitute separate species. These species pairs are an excellent example of how adaptation to different environments (in this case feeding in the surface waters or on the lake bed) can generate new species. This process has come to be termed ecological speciation. This type of species pairs is found in British Columbia in Western Canada. The lakes themselves only contain three-spined sticklebacks and cutthroat trout, and all are on islands. Tragically, the pair in Hadley Lake on Lasqueti Island was destroyed in the mid 1980s by the introduction of a predatory catfish, and the pair in Enos Lake on Vancouver Island has started to interbreed and are no longer two distinct species. The two remaining pairs are on Texada Island, in Paxton Lake and Priest Lake, and they are listed as Endangered in the Canadian Species At Risk Act.

Other species pairs which consist of a well-armored marine form and a smaller, unarmored fresh water form are being studied in ponds and lakes in Southcentral Alaska that were once marine habitats such as those uplifted during the 1964 Alaskan Earthquake. The evolutionary dynamics of these species pairs are providing a model for the processes of speciation which has taken place in less than 20 years in at least one lake. In 1982, a chemical eradication program at Loberg Lake, Alaska, killed the resident freshwater populations of sticklebacks. Oceanic sticklebacks were introduced and colonized the lake. In just 12 years beginning in 1990, the frequency of the oceanic form dropped steadily, from 100% to 11%, while a variety with fewer plates increased to 75% of the population, with various intermediate forms making up another small fraction (Carroll 2006). This rapid evolution is thought to be possible through genetic variations that confer competitive advantages for survival in freshwater when conditions shift rapidly from salt to freshwater. However, the actual molecular basis of this evolution still remains unknown.

Although sticklebacks are found all round the coast of the Northern Hemisphere and are thus viewed by the IUCN as species of least concern, the unique evolutionary history encapsulated in many freshwater populations means they deserve more legal protection than they currently receive.

Life History
Many populations take 2 years to mature and experience only one breeding season before dying and some can take up to 3 years to reach maturity. However, some freshwater populations and populations at extreme latitudes can reach maturity in only 1 year. In spring, males defend territories where they build nests on the bottom of the pond or other body of water; the sequence of territorial, courtship and mating behaviours was described in detail by Niko Tinbergen in a landmark early study in ethology. Territorial males develop a red chin and belly colouration, and Tinbergen showed that the red colour acted as a simple sign stimulus, releasing aggression in other males and the first steps in the courtship sequence from gravid females. Red colouration is produced from carotenoids found in the diet of the fish. As carotenoids cannot be synthesised de novo, the degree of colouration gives an indication of male quality, with higher quality males showing more intense colouration. However, it is noteworthy that the response to red is not universal across the entire species complex, with black throated populations often found in peat-stained waters. Males also develop blue irises on maturation. Only the males care for the eggs once they are fertilised. Parental care is intense, involving nest maintenance and fanning of the eggs to ensure a fresh water supply. Males build the nests from vegetation, sand, pebbles and other debris, adhering the nest together with spiggin, a proteinaceous glue-like substance secreted from the kidneys. Sticklebacks have four colour photoreceptors in their retina, making them tetrachromatic. They are capable of perceiving ultraviolet wavelengths of light invisible to the human eye and use such wavelengths in their normal behavioural repertoire.

Genetics
Three-spined sticklebacks have recently become a major research organism for evolutionary biologists trying to understand the genetic changes involved in adapting to new environments. The entire genome of a female fish from Bear Paw Lake in Alaska was recently sequenced by the Broad Institute and many other genetic resources are available. This population is under risk by the presence of introduced northern pike in a nearby lake.

Trivia
The three-spined stickleback was featured on a 14-franc postage stamp issued by Belgium in 1990.

It is also referred to in the very beginning of Dickens' "Pickwick Papers", in the title of Mr. Pickwicks illustrious work ??? "Speculations on the Source of the Hampstead Ponds, with some Observations on the Theory of Tittlebats".

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