Bats of Britain and Europe

Bats – the mystery beings

With over 1,200 species, bats are a very successful and species-rich order among the mammals. They are extremely varied and have inhabited all continents and almost all habitats of the Earth with the exception of Antarctica. As a typical mammal, they have fur, external ears and a mammalian dentition, and they are homothermic, and they give birth to live young and suckle them. They also have some special features that make them unique among the mammals.

They can actively fly

Bats are the only group among the mammals that is capable of active flight, which distinguishes them from gliders such as flying squirrels, possums and colugos. Their extended fingers and metacarpals are connected to the body by flight membranes, which is why they got the name Chiroptera – ‘hand wing’.

They can reach a very old age

Bats can get very old in relation to their generally small body size (Box 1). Some species reach over 30 years. They probably reach this great age due to low predator pressure (at night there are few enemies) and with that a relatively high survival rate. It is also believed that bats are better at reducing cell toxins that are responsible for many ageing processes. A link between hibernation and old age is assumed.

Distribution of bat species in the New World within 100 x 100km grid squares depending on the geographical latitude. The size of the circles is proportional to the number of cells with the same number of species at the geographical longitude. An increase in biodiversity in the direction of the tropics is clearly apparent. Graph: W. Lang, modified after Ramos Pereira & Palmeirim 2013.

Emergence of a colony of several million Wrinkle-lipped Free-tailed Bats (Chaerephon plicata) and Black-bearded Tomb Bats (Taphozous melanopogon) from a Vietnamese cave.

Their sperm has a long viability

In the temperate zones bats hibernate and additionally have developed a mechanism to store sperm alive for several months. Thus, bats can mate in late summer and autumn, but the actual fertilisation only happens in the course of the next spring, when the optimal conditions for pregnancy and birth are available. Other winter hibernators, such as the bent-winged bats, have developed a different strategy. They delay the development of the fertilised egg. In the tropics bats adapt the time of birth to the optimal period of favourable food availability for the rearing of the young.

They are very diverse

No other mammal group has conquered its habitats so widely and occupied such a variety of ecological niches. Besides the typical and original insectivorous feeding, there are carnivorous and frugivorous species; some bats are specialised in fishing, others feed on blood or, hummingbird-like, collect nectar and pollen from flowering tropical plants. These bats often play a core role as ‘pest controllers’, seed distributors and pollinators in their respective habitats.

They use echolocation to find their way

Last but not least bats (with the exception of flying foxes) have developed a unique orientation system among land animals, namely ultrasonic echolocation. We, as humans, are animals of sight, and with our passive sense of sight we depend on sunlight. With their active orientation system bats could become independent of daylight and conquer the night as a niche. The discovery of echolocation by Donald Griffin in the 1940s explained what for us had long been a mystery of a sixth sense connected with the flight manoeuvres of bats. This was probably one of the reasons why bats in our regions had a bad reputation for a long time; if we can’t explain something, we are either afraid or we adore it. Therefore, it is not surprising that in other cultures, like the Maya, bats were worshipped as deities, while in China they are equated with luck. The author of Dracula, Bram Stoker, shares responsibility for the bad reputation of bats; he made, for the first time and very successfully, a connection between bats and vampires. Today bats have a better reputation and any disgust has mostly given way to fascination.

The Indian False Vampire Bat (Megaderma lyra) is widespread in South Asia and belongs to the carnivorous species that hunt small vertebrates.

A spectacular symbiosis is the coexistence of Hardwicke’s Woolly Bat (Kerivoula hardwickei) and pitcher plants: the plants offer a roost and the droppings of the bats provides valuable mineral nutrients, especially nitrogen, to the plants. Photo: A. Seibert.

Box 1: Records in bats

Body structure of bats

Built to fly

The body of a bat is built for lightness and energy efficiency. In spite of many special adaptations to flight the five-digit construction of the mammal limbs can still be seen in the wing. In particular, the metacarpals and phalanges are greatly elongated and thin, and the similarly long forearm consists of the radius, while the ulna is absent. The second finger has only one phalange, the third has three phalanges, and both the fourth and fifth fingers have two phalanges. Only the thumb (in the fruit bats also the second digit) has a claw that is used for climbing. A flight membrane is formed between the extremities, the body and the tail. Among the European bats, the tail is largely free only in the European Free-tailed Bat; in all other species the tail is integrated into the flight membrane. The flight membrane is criss-crossed with blood vessels, nerves, sensory cells, muscles and elastic fibres. Some veins in the wings have contractile areas that can be described as minor hearts. They ensure that an adequate blood pressure can be maintained and also have a function for heat balance. Merkel cells on the flight membranes, sensory papillae associated with a hair, measure the airflow and provide a stable flight. Because the flight membrane must not dry out, much time is spent by bats rubbing on a secretion from the mouth to keep the membrane elastic.

Body plan and naming of individual parts. Diagram: R. Roesler.

A cartilaginous spur, the calcar, extends from the heel towards the tail and helps to support the margin of the tail membrane and sometimes has a membranous lobe, the post-calcarial lobe, or epiblema.

The comparatively low weight of bats is an adaptation to travel by flying. Even the largest species, fruit bats of the genus Pteropus, are, for a mammal with a 1.7m wingspan and weight of about 1.5kg, no giants. The majority of bats are much smaller and in the Bumblebee Bat (Craseonycteris thonglongyai) evolution has probably produced the physiologically smallest possible mammal. Small animals have a relatively large surface area and lose more energy. Hair can reduce this energy loss, so the body of bats is densely haired. Only a few desert bats have short fur and even naked skin. Parts of the face, ears and membranes are glabrous. The fur is usually moulted once a year, from late summer.

Left Bourret’s Horseshoe Bat (Rhinolophus paradoxolophus) has bizarre-looking nose adornments representing an extreme adaptation for emitting echolocation calls.

Right Andersen’s Leaf-nosed Bat (Hipposideros pomona) has very large ears and hunts with an agile flight for small flying insects within tropical vegetation.

Upside down as a success principle

Bats hang with the head down, which allows, among other things, a rapid take off by simply dropping. Many bats can also rest horizontally (for example in rubble in the winter), and for hunting – as in the Greater Mouse-eared Bat – can quickly and nimbly walk on the ground and take off with a jump into the air. With the hanging way of life, the feet are twisted backwards. A rotation of the leg at the knee joint upwards and outwards is unique among mammals. The claws of the feet enable safe hanging, and only grip from the weight of the bat using a special locking mechanism. Thus, a bat can save energy in the winter and even remain hanging when dead. The legs stretch or fold the tail membrane, in horseshoe bats onto the back, in plain-nosed bats to the belly. The mammary glands of female bats lie in the pectoral region. The two nipples (the Parti-coloured Bat has two pairs of nipples) are clearly visible in lactating females. Horseshoe bats have a pair of false nipples in the pelvic region. The penis of a male bat has a small, species-specific penis bone (baculum). The testes and epididymides are clearly visible, at least during the mating period. Bats have a relatively large stomach, and a short intestine that allows rapid digestion.

Strange faces and a strong set of teeth

The reason for the big eyes and small ears of the flying foxes is obvious. They are visual animals like humans, and are mainly visually oriented. In that, they differ from all the other echolocating bats, with their small eyes and mostly large ears.

‘No other mammal group can surpass the diversity and curiosity of the face design of bats.’ Martin Eisentraut 1968.

In the course of evolution and colonisation of new habitats and food resources, a number of differences in the structure of the ears, the size of the eyes, the form of the face, often characterised by bizarre nose appendages, have been developed. The ears, but also the mouth and nose region, have been developed as receiver (ears) or transmitter (mouth, nose) of sounds for echolocation. The ears are very agile and are constantly aligned in horseshoe bats. Long ears can be relaxed by transverse folds, and in long-eared bats can be completely folded. Plain-nosed bats have a diverse shaped tragus inside the auricle that is absent in the horseshoe bats. The nose and nostrils, as in most plain-nosed bats, can be small and inconspicuous, but they can also be extremely enlarged and be fitted with various appendages. Obvious large nostrils, as in the barbastelle and long-eared bats, are also evidence in plain-nosed bats that, depending on the situation, these species can emit echolocation calls through the nose and not only, as with many other species, through the open mouth.

Massive muscle packages are found on the skull. The neck muscles support the head during flight and the jaw muscles allow a powerful bite, which leaves little chance for even stubborn prey. A specialised set of teeth, resembling at first glance a predator dentition, is enhanced by the large canines. The diet determines the number and type of teeth. Fruit-eating species have only flat cusps on the teeth, an adaptation to grinding softer food. Nectar-eating bats have greatly reduced dentition and often long and delicate jaws. Their long and narrow faces are adapted, with a long tongue for visiting deep calyces with hummingbird-like hovering. Carnivorous and insectivorous bats have a more predator-like dentition, with strong canines, many cusps and cutting edges. The basic blueprint of the dentition of the bats with 38 teeth is found repeatedly in the indigenous and globally distributed genus Myotis (Box 2). In many species, however, the number of teeth is reduced. There is a high selection pressure on the shape and size of the teeth. The length of the teeth, the relative heights of individual teeth and the shapes of cusps can thus be used to distinguish between even closely related species.

And vision?

Bats see the world around them in black and white and some species of flower-visiting bats also see ultraviolet light as it is reflected by some flowers and thus serves as a marker to fly to. Bats also have a magnetic sense. With this they can follow the Earth’s magnetic field lines during long-distance flights, similar to the way birds do.

The Long-tailed Fruit Bat (Notopteris macdonaldi) feeds on nectar; flower bats with much longer muzzles occur in the New World.

Box 2: Dental formula of the genus Myotis

A long success story

Preserved for a long time

The oldest fossil bat species from the early Eocene (about 52 million years ago) corresponds closely in body structure to today’s bats. Only Onychonycteris finneyi, described as recently as 2008, has original characteristics: thus it has a claw on all five fingers, relatively short wings and long legs. It was, nevertheless, able to fly over long distances. The approximately 47 million-year-old bat species of the genera Paleochiropteryx, Archaeonycteris, and Hassianycteris from the Messel quarry (Germany) could orientate themselves using echolocation. Their different wing shapes occupy similar niches to those of the recent bats.

Bats and dinosaurs?

Molecular methods assume that the first bats were formed 65 million years ago, shortly after the mass extinction of the dinosaurs and during the transition of the Cretaceous/Tertiary. It is, therefore, likely that primeval bats already existed on the Earth during the time of the dinosaurs. In the Eocene, a rapid radiation of the bat families occurred, as happened with many mammal groups.

Everything turned upside down

Molecular analysis in the last two decades provided even more surprising results: the classic divisions based on morphological features of the Microchiroptera and the Megachiroptera turned out to be artificial. Some representatives of the horseshoe bats (Rhinolophoidea) are more closely related to the fruit bats than to all other bats. At the same time, the Nycteridae, a family of this same group clearly belonged to the other bats. The modern family tree of bats shows two new groups, the Pteropodiformes (or Yinpterochiroptera) and Vespertilioniformes (or Yangochiroptera).

Paleochiropteryx tupaiodon, an Eocene bat from the Messel quarry, Germany. Photo: D. Nill.

Family tree of bats based on molecular genetic investigation, after Simmons (2005). Diagram: S. Wöhl.

The new classification leads to a new debate: did echolocation develop twice independently, or have the flying foxes lost their echolocation ability? And how can it be explained that some representatives of the fruit bat genus Rousettus have developed a simple form of echolocation by tongue clicks and can thereby also use caves and other dark places as roosts?

The other findings of molecular biology, relating to the question of which animal is the closest relative of the bats, are no less amazing. Bats probably originate from the group of the Laurasiatheria, which was formed from insectivores, carnivores, whales and the odd- and even-toed ungulates.

The flight of bats

He who can fly has a clear advantage

Pterosaurs, birds and bats are distinguished from other vertebrate animals by their power of flight. All three use their forelimbs as wings, but the specific modification of individual bones is different in each case. Pterosaurs had flight membranes that were attached to the fourth finger. Birds use feathers as a ‘replacement’ for the reduced finger and metacarpal bones. Bats stretch their flight membrane between four fingers, the legs and the tail.

Flying is an energy-intensive form of locomotion per unit time, but per travelled distance it is efficient. And it is the most efficient way for bats to travel, because they need to fly repeatedly from their roosts, for example in caves, to their distant hunting territories. Bat flight is often very manoeuvrable through the flexibility of usable membranes. Bats can quickly reduce speed, fly the tightest curves and even catch prey with their wing membranes. Some, such as the long-eared bats, can hover in the air and search for prey from leaf to leaf. They consume consistently less energy in their flight than similarly sized birds. Even while hovering, flower-visiting bats show more efficiency than hawkmoths and hummingbirds.

Left A Brown Long-eared Bat in flight. Long-eared bats can take off well from the ground with their short, broad wings and do not have to jump up as do many other species. Photo: R. Klenk.

Right Greater Horseshoe Bats usually catch their prey with the wing, then transfer it to the mouth. Photo: D. Nill.

Wing loading (ratio of weight to wing area; wing loading [Nm-2]) and wing aspect ratio (ratio of wing length to width of wing; aspect ratio) for European bat species. Graph: S. Wöhl.

Slow or fast? – the optimal adaptation brings success

Most bats appear relatively slow with respect to their airspeed. This also applies to most small species that travel at 5–8m/s (= 18–29km/h) while in flight. Species such as the European Free-tailed Bat and Noctule Bat often reach speeds of over 50km/h, which are also significantly exceeded in a dive. Looking at the wing shapes, the fast-flying species can be immediately identified by the long, narrow wings while slow manoeuvrable species have rather shorter and broader wings. There are many variations of this basic pattern, even supplemented by the shape and size of the tail membrane, the roundness of the wing-tip and a lot more. The variety of forms is a reflection of the niches occupied by bats and of particular hunting techniques or habitat requirements. Even a species’ ability to migrate can be interpreted from the wings in the form of an ideal combination of wing loading and long, narrow wings. It is also interesting that flight, echolocation and breathing are linked to each other. When the pectoral muscles are most contracted a call will be emitted at the same time; a call is thus emitted with the maximum energy at the most optimal time.

Bats are ‘high-performance machines’: the heart of a bat is about three times as large as in an equivalent-sized small mammal. They have ultrafast muscles (for echolocation), many red blood cells and a high content of haemoglobin, and the lungs can extract more oxygen from the air than those of any other mammal.

Habitats of bats

Hiding and oversleeping

Bats do not build nests. Only a few tropical bat species build their own refuges by biting through the mid-ribs of leaves of bananas, through which the leaf blades fold downwards like a tent. Temperate bats are strictly dependent on already existing hiding places. In such roosts they find protection against wind, weather and enemies, and can fall into torpor, raise their young, mate or hibernate.

The roosts of Grey and Brown Long-eared, Common Pipistrelle and Whiskered Bats occur in the settlement area of this village. Using connecting structures, such as hedges or tree lines, the bats get to their specific foraging areas.

Forests, meadows and mountains

Bats can be found everywhere in Europe – in the coniferous forests of northern Europe and the hardwood forests in central Europe, heaths, meadows, moors, villages, cities, the maquis of the Mediterranean, along the coasts and in the mountains. Some habitats offer few roosting places but are insect rich and thus well suited as hunting grounds. There are species that have adapted to inhospitable living conditions for bats, such as the deserts of North Africa.

Tree hollows, caves and crevice hiding places

European bats can be classified on the basis of three types of roosts: as cave and tree dwellers, and as crevice dwellers. As the requirements for the roost change in the course of the year, so a change of the basic roost types becomes possible, for example from tree hollows in summer to caves in winter. In the mainly forested central Europe, tree holes were the most important summer roosts for a long time, since here caves are either rare or too cold for bats’ offspring. But tree holes are small, and offer space for only small groups. They are also not as permanent as caves. Therefore tree-dwelling bats use many roosts and change them very often. Moreover, the colonies split from time to time and new groups are formed. Many dwellers of crevices behave in a very similar way because their refuges, such as peeling bark on tree trunks, are usually not permanent.

Through human construction activity, cave-dwelling bats could spread from the Mediterranean to the north. Even today they show their typical close fidelity to their roosts, which they use not only continuously in a summer, but also across generations. Within caves and attics large colonies can find the appropriate temperature regime, then choose the roosting sites. Tree-hollow and crevice dwellers have to move on to another roost if the temperature no longer suits them. Many former forest-living bats, either tree-hollow or crevice dwellers, have adapted to new roost possibilities in our buildings, and have thus moved into our settlements.

Left Traditional orchards are used by many bat species in central Europe, because they combine features of both forest and open landscape, thus providing for most species’ hunting habitats and prey.

Right Real forests have become very rare in the Mediterranean, so that specialist forest bats are mainly to be found in mountain regions, while the open-habitat species dominate in the lower regions.

Forest- and building-dwelling bats

Real forest bats, such as Bechstein’s Bat, find their roosts as well as their hunting grounds in forests. Real building bats, such as the Serotine Bat and Grey Long-eared Bat, roost in buildings and prefer to use open landscapes as hunting ground. But there are also many species that need both habitats and switch between them. Species such as Natterer’s Bat and the Brown Long-eared Bat are more flexible in their roost choice and use tree hollows and also small holes and crevices in buildings. The hunting grounds of these species can be found in forests, but also in open landscape. Greater Mouse-eared Bats use lofts for raising their offspring and forests mainly as hunting grounds. Single males of the Greater Mouse-eared Bat are often found in tree hollows. A more typical building-dwelling bat, the Common Pipistrelle Bat, hunts preferably and extensively in forests, but can also find its food in villages and towns.

Left Spruce monocultures offer hardly any natural roosts and a limited range of insect prey.

Right Deciduous forests have a variety of insects and there are more possibilities for natural roosts.

Forests and ‘forest bats’

Many bat species were originally forest animals, yet despite this the importance of forests for bats has been underestimated for a long time. Clearing forests away to create open cultivated landscape, and the local use of insecticides probably resulted in forests becoming increasingly important as refuges. At the same time an increasing absence of exploitation of forests led to a positive development of important structures for the conservation of species. This has again dramatically changed in the last decades, in particular through the increased demand for renewable raw materials. Despite this, above all European habitats, even today the highest number of species can still be found in forests, because they equally offer the two key resources of roosts and hunting grounds. Tree hollows or crevice-like hiding places can be found in great numbers in natural forests. Prey develops in many places, such as pools, dead wood, moist soil and forest edges and clearings, and they therefore offer food in abundance. Through the frequent, often daily, changing of the roosts, it also becomes clear why bats need a variety of roosting options. A single bat hole is not sufficient for the survival of a population because of competition with many other animals and many bat species. Core hunting grounds are often used by individual species over years. There is competition among the bats species on the hunting grounds. A way to avoid this is for bats to use different microhabitats, such as tree crowns, forest land, edge structures, single trees and clearings. This requires specific hunting strategies, especially in terms of echolocation and hunting techniques. In the past, these adaptations could then be refined for other habitats and were the basis of the colonisation of new habitats.

Forest types

Not all forest types are equally suitable for bats. Diverse structure and near-natural forests with a high proportion of deciduous trees are preferred, but coniferous forests, particularly pine, are also used. Monotone coniferous cultures are of little interest, but they are not empty of bats. Riparian forests are characterised by the dynamics of water, a wide roost choice and structure diversity, and by an outstanding food supply. They are thus of prime importance, not only in summer but also during the migration times of Noctule, Leisler’s and Nathusius’s Pipistrelle Bats. This could also influence their focus in migration along the rivers. In mountain forests in Germany, 19 species were identified, of which seven species also breed there, and in the Mediterranean, mountain forests are the home of rare species, such as the Greater Noctule Bat.

Left The Brown Long-eared Bat can be common in coniferous forests. Photo: M. Koch.

Right Bechstein’s Bat is a specialist of old beech and oak forests. Photo: R. Klenk.

Forests should be connected

There are only a few real forest bats, such as the Alcathoe Whiskered Bat and Bechstein’s Bat, but almost all bat species use forests. When forests are connected by linear structures, such as hedges or tree lines, they can also be reached from settlements by linear structure-bound flying species. These bat species avoid flying across open areas. In the absence of connecting structures, the way to and from forest areas is cut off. Roads can also be an obstacle.

What is a good bat forest?

A richly diverse forest should contain all natural forest development phases (initial, growth, peak, and decay phases). When these phases form a mosaic, a wide variety of hunting and roosting habitats can be found, from young-growth areas, open tree stands and open spaces, up to old stands. Today’s economic forestry does not offer this, missing the decay phase. The trees often do not even reach the age at which storm damage, rotting, woodpecker holes, lightning cavities and trunk cracks are formed. Young trees that show signs of damage, from which roost opportunities can develop are removed for use as firewood or because they hinder the future trees. The economic pressure is high. When ‘forest bats’ have to survive, individual trees with roosting options (at least 10 trees/ha) should be protected, or better, still the whole area should not be used for forestry. Bat boxes are certainly readily accepted by some species, but are not a good long-term means of conservation.

Threats to bats

Pesticides have multiple effects

From the middle of the 1950s, the European bat populations went through a deep collapse, that led to large-scale local extinction of horseshoe bat populations in the middle of the 1970s. Today, some species have not yet recovered from it. Since bats are predators and are at the top of the food chain, changes in their habitats have cumulative effect on their populations. It is not only the mechanisation of agriculture but also the general change of habitats, which leads to the mass reduction of insects, that affects bats. The use of pollutants has affected bats in multiple ways. Insecticides used in agriculture and forestry, also in the private sector, have led to a general decrease of the available insect biomass and thus the food of bats. Some poisons (like DDT) were accumulated in fatty tissue and in mother’s milk. These substances were released over the winter and poisoned the young animals. In some species, they have led to sterility. In actions to combat wood pests, building-dwelling bats were killed, often directly or indirectly. Other chemicals, such as tetraethyllead mixed in car gasoline, can increase the level of lead in bats’ tissues and result in the impairment both of the sense of hearing and the nervous system.

New poisons

More recently, anti-parasitic drugs to protect grazing animals have become a threat: dung-eating insects that play a key role for many bat species, such as the Greater Horseshoe Bat, are eliminated and other insects can also be affected. In some areas, large-scale campaigns against alleged pests, such as maybugs, are performed and lead to a general reduction of the insect biomass and therefore to food shortages for bats. The massive pesticide use in agriculture is ensuring that the diversity of insects is decreasing drastically, and no improvement is foreseen.

Left Many building-dwelling bats, such as the Grey Long-eared Bat, were poisoned by toxic wood preservatives in the summer roosts. Even five decades after such treatment one can find non-decomposed animals – they are preserved chemically.

Right During hibernation, bats like this Geoffroy’s Bat need a quiet environment in which to survive the winter on their fat reserve. Disturbances are increased by leisure activities such as geocaching, mineral collecting and cave tourism.

Destruction and disturbance of roosts

Another important risk factor is roost destruction and the subsequent loss of roosts. In forests, an increased need for timber and firewood leads to an over exploitation of forest stands. Forests