The secret wildlife of golf courses

The cool morning air has strewn the lawn with small dewdrops. The green is bathed in flickering mist and shining dewdrops. Soon the green is filled with the sibilant sound of golf balls and walking golfers, but for a while, the course still belongs to someone else.

Water hazards of Hiekkaharju Golf in Vantaa (in the Helsinki metropolis area) provide suitable habitats for diverse species. Picture borrowed from

Keimola Golf, located in Vantaa (in the Helsinki metropolis area in Finland), is a true paradise for birds and amphibians. Whooper swan (Cygnus cygnus), common goldeneye (Bucephala clangula), and horned grebe (Podiceps auritus) pairs nest in the largest water hazard. In addition, the black woodpecker (Dryocopus martius) nests nearby. The number of horned grebes has declined worldwide, and the species is considered vulnerable in Finland. The Finnish population has decreased from 3000 to 6000 nesting pairs in the 1980s to the present 1200–1700 nesting pairs.

On the other hand, all Finnish amphibian species, except one, can be found living in one of the smallest water hazards of Keimola Golf. Only the Northern crested newt (Triturus cristatus) does not occur there. The Northern crested newt is critically endangered in Finland, and can only be found in a few places in eastern Finland. In spring, the common frog (Rana temporaria), the moor frog (Rana arvalis), and the common toad (Bufo bufo) croak vigorously. The smooth newt (Lissotriton vulgaris) does not croak, but mating males bring tropical colors into an otherwise brownish landscape.

Mating smooth newt males are springtime color spots in a wetland. ©Mia Vehkaoja

By Midsummer, golf courses are swarming. On dry land, golfers enjoy their sport in warm summer weather, while hatched ducklings and tadpoles are concurrently going through growth spurts around the water hazards. Golf courses provide lots of nutrition for ducklings and tadpoles. Water hazards, as most wetlands, are habitats for several invertebrates, such as mosquito (Culicidae), nematocera (Nematocera), and trichoptera larvae, as well as for phyto- and zooplankton. Amphibians prefer open and sunny wetlands because higher temperatures escalate tadpole development. Ducklings, on the other hand, prefer wetlands with luxuriant shoreline vegetation (for example club rushes and sedges). Vegetation provides cover against predation.

Luxuriant shoreline vegetation provides cover for ducklings against predation, whereas openness increases water temperature and escalates tadpole development. ©Mia Vehkaoja

Golf courses are oases for wetland-associated species, especially in urban environments, where most wetlands are isolated from each other. For numerous species, water hazards and golf greens offer nearly free access between wetlands and other habitats. Golf courses are currently not planned to consider nature and its needs. What if nature were taken into account during planning, with at least a 10% effort? Keimola Golf’s extraordinary biodiversity has arisen through chance. Waterfowl diversity is due to an island left in the middle of the largest water hazard. The island has some ten trees and bushes. The whooper swan and common goldeneye nest on this island.

Both national and international designers have planned Finnish golf courses. Keimola Golf was planned in Great Britain. More and more, architects plan golf courses by initially outlining the routes, after which the planning is continued on-site concurrently while the course is being constructed. This method enables taking nature into account during the planning process.

Architects could pay attention to small things that benefit animal and plant species when planning water hazards and groves. For example, bushes and shoreline vegetation could be left next to the shoreline that is not close to the green. This has been done at Keimola Golf. Paying attention to such small details does not even cause additional costs. Furthermore, most golfers enjoy the sport because they can be outside and “enjoy” nature. If nature were actually taken into account during planning, golfers could actually play their sport “in the wild”.


The beaver – our wetland rescuer

The beavers (Castor canadensis and Castor fiber) have recovered from near extinction, and come to the rescue of wetland biodiversity. Two major processes drive boreal wetland loss: the near extinction of beavers, and extensive draining (if we exclude the effects of the ever-expanding human population). Beaver dams have produced over 500 square kilometers of wetlands in Europe during the past 70 years.


The wetland creation of beavers begins with the flood. As floodwaters rise into the surrounding forest, soil and vegetation are washed into the water system. The amount of organic carbon increases in the wetland during the first three impoundment years, after which they gradually begin reverting back to initial levels. The increase in organic carbon facilitates the entire wetland food web in stages, beginning with plankton and invertebrates, and ending in frogs, birds and mammals.

The previous shoreline is very evident from an aerial photograph. Also the beaver flooded area shows clearly. © Antti Nykänen

The previous shoreline is very evident from an aerial photograph. Also the beaver flooded area shows clearly. © Antti Nykänen

Beaver-created wetlands truly become frog paradises. The wide shallow water area creates suitable spawning and rearing places. The shallow water warms up rapidly, and accelerates hatching and tadpole development. Beaver-created wetlands also ensure ample nutrition. The organic carbon increase raises the amounts of tadpole nutrition (plankton and protozoans) in the wetland, along with the nutriment of adult frogs (invertebrates). Furthermore, the abundant vegetation creates hiding places against predators for both tadpoles and adult frogs.

Beaver-created wetlands are perfect rearing places for frogs. The warm water accelerates hatching and the abundant aquatic vegetation gives cover against predators. © Mia Vehkaoja

Beaver-created wetlands are perfect rearing places for frogs. The warm water accelerates hatching and the abundant aquatic vegetation gives cover against predators. © Mia Vehkaoja

The flood and beaver foraging kill trees in the riparian zone. Deadwood is currently considered a vanishing resource. Finnish forests have an average 10 cubic meters of deadwood per hectare, whereas beavers produce over seven times more of the substrate into a landscape. Beaver-produced deadwood is additionally very versatile. Wind, fire and other natural disturbances mainly create two types of deadwood: coarse snags and downed logs. Beavers, on the other hand, produce both snags and downed logs of varying width, along with moderately rare deciduous deadwood. The more diverse the deadwood assortment is, the richer the deadwood-dependent species composition that develops in the landscape.

Beaver-created wetlands produce  especially standing deadwood. © Mia Vehkaoja

Beaver-created wetlands produce especially standing deadwood. © Mia Vehkaoja

Deadwood-dependent species are one of the most endangered species groups in the world. The group includes e.g. lichens, beetles and fungi. Currently there are 400 000 to a million deadwood-dependent species in the world. Over 7000 of these inhabit Finland. Pin lichens are lichens that often prefer snags as their living environment. Beaver actions produce large amounts of snags, which lead to diverse pin lichen communities. Snags standing in water provide suitable living conditions for pin lichens; a constant supply of water is available from the moist wood, and the supply of light is additionally limitless in the open and sunny beaver wetlands.


The return of beavers has helped the survival of many wetland and deadwood-associated species in Finland, Europe and North America. Only 1000 beavers inhabited Europe at the beginning of the 20th century. Now over a million beavers live in Europe. I argue that this increase has been a crucial factor benefitting the survival and recovery of wetland biodiversity. Finland and the other EU member states still have plenty of work to do to achieve the goals of the EU Water Framework Directive. Both the chemical conditions and the biodiversity of wetlands / inland waters affect the biological condition and quality of wetlands.


The whole research published here

Frogs love beaver-created wetlands

Amphibian and wetland loss is occurring globally at an increasing rate. Since the 1900s, approximately half of the world’s wetlands have been destroyed. During this time up to two-thirds of European wetlands were lost at a regional scale. This trend is reflected by the fact that 23% of Europe’s amphibians are threatened.

Wetlands in the boreal region are frequently constructed through the damming activities of an ecosystem engineer, the beaver (Castor sp.). They create and maintain special habitats by constructing dams. Beaver-created wetlands are open and sunny due to tree felling and flooding-induced tree mortality. They produce large quantities of woody debris and detritivorous invertebrates, e.g. chironomids and Asellus. Beaver ponds contain structurally heterogeneous vegetation.

Beaver-created wetlands contain structurally heterogeneous vegetation, as well as open water areas. © Mia Vehkaoja

Beaver-created wetlands contain structurally heterogeneous vegetation, as well as open water areas. © Mia Vehkaoja

According to our new study beaver-created wetlands increase frog species heterogeneity and abundance. There are only three native anuran species in Finland (the common frog, the moor frog and the common toad), and all of them were found in beaver ponds. The moor frog (Rana arvalis) was only found in beaver ponds, where the common frog (Rana temporaria) was also most abundant. The common toad (Bufo bufo) prefers deeper wetlands than the other two species, but because beaver ponds contain both shallower and deeper parts, it was also found from the beaver ponds.

Beaver wetlands offer high quality habitats for anurans and facilitate the occurrence of moor frogs. The shallow and warm water areas accelerate the hatching and metamorphosis of tadpoles. The rich aquatic vegetation provides attaching places for spawn and protection against predators. The abundant vegetal detritus, zooplankton and aquatic invertebrates offer nutrition for both larvae and adults. In addition, beaver-created wetlands create overwintering habitats that are less likely to freeze down to the bottom.

Both the moor frog (Rana arvalis) and the common frog (Rana temporaria) are the most abundant in beaver ponds. © Mia Vehkaoja

Both the moor frog (Rana arvalis) and the common frog (Rana temporaria) are the most abundant in beaver ponds. © Mia Vehkaoja

Beaver facilitation includes both habitat amelioration and resource enhancement. Frogs are not the only group that is benefitted by beaver activity. Other such groups are ducks, bats, woodpeckers and many invertebrates. These ecosystem engineers could be used in wetland restoration, and furthermore the beaver clearly promotes amphibian conservation.

Tadpole defence mechanisms

Predators are a major cause of mortality for amphibian larvae. Many insects, lizards and fish prey on tadpoles. Fish are particularly efficient predators of larval amphibians and often cause local extirpations of species. Luckily tadpoles have defensive mechanisms to avoid predation, including attaining a large body size, behavioural changes such as reducing activity or selecting a hiding place, the use of chemical signals, and toxicity and unpalatability.

Tadpoles choose their anti-predator defence depending on the aquatic conditions of wetlands. Tadpoles have just a few predators in temporary ponds that dry out periodically. It is more beneficial to grow rapidly in such habitats and reach the metamorphosis stage rather quickly.

On the other hand, tadpoles in more permanent wetlands must choose or exploit a much more diverse set of defensive mechanisms. Tadpoles detect predators through both direct and indirect chemical messages. Indirect chemical messages can be, for example chemical components secreted from the injured tadpoles. When a predator has been detected, most of the tadpoles reduce activity and hide among aquatic plants.

In the presence of predators, tadpoles hide between aquatic plants. © Mia Vehkaoja

In the presence of predators, tadpoles hide between aquatic plants. © Mia Vehkaoja

Reduced activity means reduced foraging, which leads to slower growth. A slower growth rate also indicates delayed metamorphosis for the common frog’s (Rana temporaria) tadpoles. Common and moor frog (Rana arvalis) tadpoles grow longer tails in the presence of predators. With longer tails tadpoles swim faster and are better at avoiding predation. The tadpoles of the common toad (Bufo bufo) also grow slower in the presence of predators, but the slower growth rate does not lead to delayed metamorphosis. The common toad is toxic. It has a toxin called bufotoxin, which makes it taste bad and causes several symptoms, such as irritation of the mucous membranes, and increased salivation and vomiting. Tadpoles of the common toad also have secreting glands.