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K DO NOT SHELVE! THIS IS AN INTERLIBRARY LOAN REQUEST AUTHOR: Pucek, Zdzistaw; Smithsonian Institution; National Science Foundation (U.S.) TITLE: Keys to vertebrates of Poland, mammals IMPRINT: Warsaw, Poland: Translated and published for the Smithsonian Institution and the National Science Taylor fertebrates imals ski-Mammalia) ited from Polish viga Stachurska Zdzisfaw Pucek Due Date: 09/01/2021 ILL 208123974 AMERICAN MUS OF NATURAL HIST Interlibrary Loan (YAM) Borrowing library, return to: University of Minnesota DO NOT SHELVE! THIS IS AN INTERLIBRARY LOAN REQUEST ublishers - Warszawa
KEYS TO VERTEBRATES OF POLAND MAMMALS Translated from Polish by Magdalena Bibrich and Jadwiga Stachurska Collective Work edited by ZDZISLAW PUCEK Mammals Research Institute Polish Academy of Sciences Bialowieza, Poland PWN — Polish Scientific Publishers — Warszawa 1981 9
100 Systematic account from the posterodorsal part of the body to the front and down the sides to the belly. The spring molt proceeds in the opposite direction starting on the ventral surface of the head and running dorsoposteriorly. Males molt earlier than the females. Three pairs of nipples are present. The skull is more delicate and relatively longer in the rostral portion than in C. leucodon. The unicuspids are very diverse. Choanae do not narrow posteriorly and have a U or V-shaped cross-section when viewed from the rear. C. suaveolens inhabits fields, orchards and gardens. In autumn it moves into farm buildings where it remains till the spring. It is found on very rare occasions and not much is known about the biology of this shrew. Active during the day and night, these animals come out at more or less regular time intervals—once or more times per hour. Two peaks of activity have been observed—early in the morning and in the midafternoon. Total sum of activity is longer in summer than in winter. Males have larger home ranges than the females (averaging 50 and 27 m. in diameter respectively). Fig. 50. Distribution of lesser white-toothed shrew, Crocidura suaveolens (Pallas). After Pucek and Michalak, from Pucek and Raczyriski (1981) Bats — Chiroptera 101 The diet consists chiefly of small invertebrates, especially insects and slugs The animal eats its own weight in fresh food per twenty four hours. Lesser white-toothed shrews build their nests in grasses or m burrows abandoned by small rodents. The reproduction period lasts from spring ti early in autumn. The female produces 3-4 litters per year and the young can reach sexual maturity in the same year they were born in. The gestation period is about 28 days and each litter contains up to 10 young. Lactation lasts for about 3 weeks. The fur develops within 9 days after birth. On the ten y ofhfethe young open their eyes but they can be seen caravamng already 3t Thi^species^nhabitJsouthern and central Europe, northern Africa^andI the Palearctic regions of Asia. In eastern Europe it extends further to the nort than in the western parts of its range, reaching as far as Estonia and the regions °f "nd'fheTerser^Wte-toothed shrew is distributed in western and southern regions of the country. Its range includes thewestern part _of Pome™£ western and central parts of Wielkopolska-Kujawy Inland. Hills Lower and Upper Silesia, southern part of Kxakow-Wielun Uplan i , West' and East Sudetes, West and East Beskids J southern part of Maiopolska Upland, Sw.Stokrzysk,eMts. andtheSandom, Lowland (Fig. 50). In the southeastern region of Poland tofe ser white -toothed shrew occurs sympatrically with C. leucodon The tives of this species have been classified into the subspecies C. 5. suaveolens (Pallas, 1811), described from the Crimea. • Order: BATS—CHIROPTERA by KAZIMIERZ KOWALSKI and ANDRZEJ L. RUPRECHT All bats are adapted for active flight by the modification of the f°rel™bs and development tf the flight membrane. This order is dividedI into two subordersMegachiroptera and Microchiroptera comprint g . species In Europe thefe are only insectivorous bats, hence the below characteristics are those of the representatives of European, other In the forelimb, the first digit (thumb) is short and bears a Jaw. The other digits are elongated and have no claws. TheseThTflight carpal bones give support to the flight membrane J"™ membrane consists of two layers of skin with a network inbetween. Its major part, known as the wing membrane or plagiopatag.um
102 Systematic account is stretched between the side of the body and the fifth digit. Between the other digits there is the interdigital membrane, called dactylopatagium (= chiropatagium). A narrow strip of membrane, called the propatagium extends between the arm, forearm, neck and head. The tail and the hindlimbs are joined by the interfemoral membrane—uropatagium. The margins of the uropatagium are reinforced and stretched by the calcar, which develops on the calcaneum. In some genera, on the outer side of the calcar there is a small calcarial keel, the so called epiblema (Fig. 51). tragus uropatagium Fig. 51. Body structure of the bat, Eptesicus serotinus (Schreber) Along with the development of wings, there are other morphologica adaptations to flight: the sternum is well developed and has a crest, the shoulder girdle is stiffened. The hindlimbs are relatively poorly developed and are mainly used to grip the perch from which the bat is suspended. The weight of the hanging body fixes the claw in the curved position as the claw and the terminal phalanx curve automatically when loaded (the so-called pulley system). Hence, no energy is wasted in muscular effort. The skull of bats is shorter than that of Insectivora. The hard palate is much indented anteriorly and the incisura incisiva is present between the internal margin of the first upper incisors. From incisors to molars, the teeth form a continuous row without crevices. In some genera, the smaller premolars are suppressed and thus invisible from the side. The larger premolars (P3) and the molars are multicuspid (similar to a letter W) and have sharp edges. Bats — Chiroptera 103 There are rare instances of the retention of premolars (P2) or sporadic occurrence of extra teeth (incisors and premolars). Sexual dimorphism is inconspicuous, usually females are larger than males. The male is easily distinguishable from the female by its pendant penis. The baculum is present. The ears are large, sometimes enormous in size. In Vespertilionidae, the external auditory meatus bears a membraneous lobe of skin, the so-called tragus, which is absent in Rhinolophidae. The eyes are small. The fur is long and soft ranging in color from various shades of brown to almost black. Bats are mainly nocturnal although some of them emerge before sunset (as for instance the common pipistrelle and the common noctule) and are still active at dawn. The species in Poland feed on arthropods, chiefly insects, which are mostly caught when flying or snapped from leaves, tree trunks or from the ground. The orientation of bats in space is ensured by echolocation. The sight is of a minor importance. In winter, bats hibernate in caves, cellars, well-hidden hollow trees and the like. In summer, they often find shelter in attics, tree holes, birdboxes, less frequently in caves and slits in rocks. Bats have few predators. Sometimes they fall prey of tawny and barn owls. Those species which come out in search of food in the vanishing light of the day may be killed by hawks. The thermoregulation of bats is unique. They have the ability to lower body temperature (heterothermia) when roosting at daytime and, especially, during hibernation. Bats often form colonies. The colonies may consists of members of both sexes, or males and females may separate. In winter, they sometimes form groupings of several species. During summer, females congregate in nursing colonies and males form single-sex groups. The estrus and copulation usually take place in autumn, less frequently in winter or spring. The sperm remains active for a long period in the female reproductive tract. Fertilization is delayed and occurs in spring. The development of the fetus takes 2-3 months. Gestation varies in length depending upon the ambient temperature. Usually one to two young are born. Initially, the young cling strongly to the body of their mother, later, when the females go hunting, they are left in colonies. At the age of about six weeks, the voung are able to fly. Toward the end of summer, young bats born that year are distinguishable from the adults only by the grayish shade of their fur and epiphysal cartilages which form characteristic thickenings of the metacarpodigital and digital joints. The life span of bats is longer than that of other small mammals. The serotine is known to reach a maximum age of 19 years and 2 months (Hanak, 1976), the large mouse-eared bat 18 years and 7 months (Pieper, 1968), and Bechstein's bat—19 years (Henze, -1970).
104 Systematic account The upper age limit for the greater horseshoe bat is 23 years and 6 months (Roer, 1960). Some species fly over great distances between their summer and winter quarters. Bats are most abundant in the tropics. To the north, some eurotopic species, e.g. the northern bat, parti-colored bat and the whiskered bat, occur beyond the polar circle. They are also found high in the mountains, where they reach slightly above the upper limit of forest. In Poland there live 2 families including 21 species. All Polish bats are useful since they exterminate insects including many crop pests as well as mosquitoes, which are also harmful to man. All bats of Poland are under protection. Key to families by external features 1. Nose-leaf present, ear without tragus. . . Rhinolophidae, page 104 —. Nose-leaf absent, tragus present . . . Vespertilionidae, page 109 Key to families by skulls 1. Intermaxillaries loosely connected to skull; one incisor on each side (often damaged when prepared). Nasal bones spherically expanded. Two incisors on each side of mandible. . . Rhinolophidae, page 104 —. Intermaxillaries fused to skull; two incisors on each side. Nasal bones not spherically expanded. Three incisors on each side of mandible . . Vespertilionidae, page 109 Family: Horseshoe bats—Rhinolophidae Around the nostrils there is the nose-leaf formed by characteristic cutaneous outgrowths: the horseshoe, crest and lancet (Fig. 52). Between the horseshoe and the crest there is a sella. The ears are pointed and devoid of tragi. The wings are wide, the tail is thin and does not extend beyond the flight membrane. The intermaxillary bones are in the form of a bony plate loosely connected to the hard palate (Fig. 58). One incisor is present on each side of the premaxilla. In the mandible there are two incisors on each side. The number of teeth is 32. The dental formula is: 112 3 2 13 3 The deciduous teeth develop in the fetus and are absorbed before parturition. Bats — Chiroptera 105 Fig. 52. The nose-leaf in Rhinolophid bats; seen from the.front (a) and from the side (b). A — the greater horseshoe bat, Rliinolophus ferrumequinum (Schreber), B — the lesser horseshoe bat, Rhinolophus hipposideros (Bechstein); c — crest, h — horseshoe,! — lancet, s — sella. From Toschi and Lanza (1959), modified Echolocation is based on the emission of ultrasounds through the nose. These bats roost always alone hanging with the flying membrane wrapped around the body (Fig. 53). The family is widespread in the Old World, primarily in the tropical and subtropical regions. Only one genus lives in Europe. Genus: Rhinolophus Lacepede, 1799 In Poland there lives one species, the lesser horseshoe bat, Rhinolophus hipposideros (Bechstein, 1800). The other species, the greater horseshoe bat, Rhinolophus ferrumequinum (Schreber, 1774) has been recorded only once, so it probably occurs sporadically. Key to species by external features 1. Length of forearm less than 42 mm Rhinolophus hipposideros (Bechstein), page 107 —. Length of forearm more than 51 mm Rhinolophus ferrumequinum (Schreber), page 108
106 Systematic account Key to species by skulls 1. Cb. length of skull less than 16 mm.*, length of mandible 9.510.4 mm., height of mandibular ramus 1.9-2.3 mm Rhinolophus hipposideros (Bechstein), page 107 . Cb. length of skull more than 20 mm., length of mandible 15.116.3 mm., height of mandibular ramus 3.6-4.4 mm Rhinolophus ferrumequinum (Schreber), page 108 * The condylobasal length of the skull of Rhinolophidae is measured from the alveolus of the upper canine to the extreme point of the occipital condyle. Bats — Chiroptera 107 Lesser horseshoe bat—Rhinolophus hipposideros (Bechstein, 1800) Measurements of 93 specimens from Slovakia are as follows: head and body, 38-45 mm.; tail, 23-33 mm.; ear, 15-19 mm.; forearm, 37^42.5 mm. Body weight, 3-9 gms. Cb. length of skull, 13.4-14.5 mm. (FeriancovaMasarova and Hanak, 1965). Wingspan, 19.2-25.4 cm. (Gaffrey, 1961). Around nostrils, the following outgrowths can be distinguished: the horseshoe, which surrounds the nostrils with its two leaves and has a slightly corrugated edge, the crest, perpendicular to the former in the sagittal plane and evenly narrowing toward the end and, finally, the lancet, which lies flat on the forehead and has three depressions at the base. The ears are pointed and lack tragi. The tail, when not stretched, is arched upward and not downward as in vespertilionid bats. In the female, apart from thoracic nipples, there is an additional pair of rudimentary pubic nipples. These are the cutaneous folds to which the young attach themselves. They become conspicuous only in the females which are over two years of age, after the first young has been reared. On the dorsal side the fur is light brown with a suggestion of red, the underparts being lighter, somewhat grayish. The young, during the first year of life are distinctly grayish in color. The lesser horseshoe bat is primarily associated with rocky areas. In summer, small colonies of females can be found in caves and in attics. Solitary males may also sometimes be found there. Most males at that time shelter alone in rock crevices, caves and the like. In winter, colonies, containing both sexes, sometimes of up to several hundred individuals, occur in caves where the temperature is 5-7°C. In these concentrations there is no direct contact between bats, they hang separately, often not high above ground. Lesser horseshoe bats do not migrate very far inhabiting a small area during the whole year and flying from one cave to another even in winter time. They feed on insects and spiders catching their prey when flying or snapping it from rocks, fences and tree trunks. The flight is characterized by rapid movements of the wings and frequent changes of direction. The lesser horseshoe bat is distributed in Europe from Ireland, England, through central Federal Republic of Germany, southern Poland and Ukraine, in northern and eastern Africa and in western Asia (the Caucasus, Turkestan, Kashmir, Asia Minor, Arabia). In Poland its range covers the East Sudetes, the West and East Beskid Mountains, the Krakow-Wieluri Upland and the Sandomierz Lowland where it reaches its northeastern limit (in Nienowice) (Fig. 54). In the mountains B*
108 Systematic account Rhinolophus hipposideros Fig. 54. Distribution of the lesser horseshoe bat, Rhinolophus hipposideros (Bechstein). After Ruprecht, from Pucek and Raczynski (1981) it has been observed at altitudes of up to 1000 meters (the Trzy Kopce Cave in the Silesian Beskid Mountains) (Ruprecht, 1981). The Polish fauna contains the nominative subspecies R. h. hipposideros (Bechstein, 1800), for the first time described from France. * Greater horseshoe bat—Rhinolophus ferrumequinum (Schreber, 1774) Measurements of 45 specimens from Slovakia are as follows: head and body, 57-71 mm.; tail, 35-69 mm.; forearm, 54-60mm.; body weight, 19.5-29.5gms. Cb. length of skull, (19) 20-22 mm. (Feriancova-Masarovd a. Hanak, 1965). Length of ear, 20-26 mm.; wingspan, 33-40 cm. (Gaffrey, 1961)- f . The structure of the nose-leaf is similar to that of the lesser horseshoe bat but the edge of the leaf is not corrugated and the crest is narrow in the middle. The fur is also similar in color to that of the above described species. The first upper premolar (P1) and the second lower premolar (P2) are very small, suppressed and protruding from the tooth-row, sometimes they are absent. The biology and habits of the greater horseshoe bat resemble those of Rhinolophus hipposideros. Bats — Chiroptera 109 It is distributed in southern and western Europe, northern Africa, western, central and eastern Asia up to China and Japan. In Europe the range of this species is similar to that of the lesser horseshoe bat but does not reach as far to the north. It comes as close to the Polish territory as the southern part of Czechoslovakia, where it permanently occurs. The former reports of its occurrence in the Czechoslovak part of the Sudetes were probably erroneous. In Poland the greater horseshoe bat was recorded only once from the Krakow-Wielun Upland (The Nietoperzowa Cave near Lazy-Jawor) in the winter of 1962 (Harmata and Wojtusiak, 1963). In Czechoslovakia lives the nominative subspecies, R. f ferrumequinum (Schreber, 1774), for the first time described from France, and it is this subspecies that probably appears in Poland. The measurements of the female specimens found in the Nietoperzowa Cave (Dec. 16, 1962) are the following: head and body, 66 mm.; tail, 38 mm.; ear, 26 mm.; forearm, 58 mm.; body weight, 19 gms.; Cb. length of skull, 21.5 mm. Family: Common bats—Vespertilionidae The nose-leaf is absent. The ears always contain the tragus. The wings may be broad or narrow. The tail extends to the edge of the interfemoral membrane or slightly beyond it. Two incisors are present on each side of the intermaxillary bone and three on each side of the mandible. The nasal bones are not spherically extended. They fuse early with the other bones of the skull and the sutures disappear. Echolocation is based upon the emission of ultrasounds through open mouth. These bats are colonial and frequently hang close to each other or to a wall. Sometimes they are found hanging free. When resting, the flying membranes are neatly folded at the sides (Fig. 53), except in the common long-eared bat, which hibernates with the front part of the body covered by unspanned wings, thus resembling Rhinolophidae. Vespertilionidae are the most widely distributed family of bats, found in all continents and in many islands, both in the tropics and in the tempeiate zone including the northern limits of the order range and reaching highest in the mountains. Many species of the temperate climatic zone migrate seasonally between their summer and winter quarters. In Poland there are seven, and possibly eight, genera. Key to genera by external features 1. Ears joined at inner bases; nostrils open upward . . 2 —. Ears not joined; nostrils open forward 3 2. Ears large (more than 30 mm.), eliptic, equal to or less than length
110 Systematic account of forearm; tragus elongated, lanceolate; snout furred from nose to eyes; calcar without keel .... Plecotus Geoffroy, page 148 —. Ears small (up to 18 mm.), wide, less than half length of forearm; tragus cultrate, snout naked from nose to eyes, calcar with keel. . . Barbastella Gray, page 152 3. Calcar without keel (Fig. 55A), lower edge of ear does not reach eye 4 —. Calcar with keel (Fig. 55B), lower edge of ear reaches eye . . . 5 4. Ears eliptic, equal to or more than half length of forearm (Fig. 56); tragus cultrate. Second phalanx of third digit twice as long as first. . . Myotis Kaup, page 117 —. Ears triangular (11-13 mm.) less than half length of forearm; tragus crescent-shaped. Second phalanx of third digit three times as long as first. Terminal portion of unspanned wing curves inward (Fig. 57). . . Miniopterus Bonaparte, page 154 5. Wings narrow; ventral side of flight membrane along forearm covered with rusty hairs. Third metacarpal 8-11 mm. longer than fifth. Tragus much shortened, end widened and rounded (Fig. 56) . . . Nyctalus Bowdich, page 143 Eptesicus serotinus (Schreber) Wings wide; ventral side of flight membrane along forearm usually naked. Third metacarpal equal to or 1-4 mm. longer than fifth. Tragus more or less elongated, crescent-shaped, end never widened (Fig. 56) 6 Fig. 56. Ears of bats. 1 — large mouse-eared bat, Myotis myotis (Borkhausen), 2 lesser mouse- -eared bat, Myotis blythi (Tomes), 3 — Bechstein's bat, Myotis bechsteini (Kuhl), 4 — Geoffroy's bat, Myotis emarginatus (Geoffroy), 5 — pond bat, Myotis dasycneme (Boie), 6 — Daubenton s bat, Myotis daubentoni (Kuhl), 7 — whiskered bat, Myotis mystacinus (Kuhl), 8 — Natterer s bat, Myotis nattereri (Kuhl), 9 — common noctule, Nyctalus noctula (Schreber), 10 — lesser noctule, Nyctalus leisleri (Kuhl), 11 — common pipistrelle, Pipistrelluspipistrellus (Schreber), 12 —Nathusius' pipistrelle, Pipistrellus nathusii (Keyserling et Blasius), 13 — parti-colored bat, Vespertilio murinus Linnaeus, 14 — northern bat, Eptesicus nilssoni (Keyserling et Blasius), 15 — serotine, Eptesicus serotinus (Schreber), 16 — barbastelle, Barbastella barbastellus (Schreber). After Feriancova-Masarova and Han&k (1965)
112 Systematic account Fig. 57. The wing of Schreiber's bat, Miniopterus schreibersi (Kuhl). A — a scheme illustrating the way of taking measurements of first and second phalanges of the third digit. FromToschi and Lanza (1959), modified. B — curving the end of bat's wing 6. Forearm less than 35 mm Pipistrellus Kaup, page 139 —. Forearm more than 35 mm 7 7. Calcar with definite keel, reinforced by transverse cartilage. Conspicuous difference in coloration between dorsal and ventral sides, hairs of back tipped with silver Vespertilio Linnaeus, page 134 —. Calcar without definite keel, no transverse cartilage. No conspicuous difference in coloration between dorsal and ventral sides, hairs of back tipped with gold or tawny. . . Eptesicus Rafinesque, page 136 Key to genera by skulls 1. Three premolars in upper and in lower jaw; rostral portion of skull elongated, ten alveoli (Pi-Ma) Myotis Kaup, page 117 —. One or two premolars in maxilla, two or three in mandible; rostral portion of skull shortened 2
126 Systematic account Natterer's bat—Myotis nattereri (Kuhl, 1818) Measurements of 38 specimens from Lower Silesia (Haillinger, 1976) are as follows: head and body, 42.6-55.5 mm.; tail, 37.CM6.5 mm.; ear, 15 0-18.3 mm.; forearm, 35.0-12.3 mm. Body weight, 5.0-12.0 gms. Wmgspan, 22.0-26.0 (27.5 cm.) (Gaffrey, 1961). Cb. length of skull of 30 specimens from Poland, 14.0-15.6 mm. A distinctive feature of this species is that the edge of the ui opatagium between the ends of calcars and that of the tail are thickened, indented and covered with two rows of stiff, curved hairs (Fig. 55). The ear is rather long and broad. The length of the tragus equals 2/3 length of the ear. The wings are relatively broad. The plagiopatagium nearly reaches the base of the digits of the foot. The interfemoral membrane extends to the end of the tail. On the dorsal side, the fur is light brown. The hairs are darker at bases and lighter toward the ends. On the venter, the fur is grayish. Here, the hairs have dark bases and are tipped with white. The second upper (P2) and lower (P2) premolars are almost the same size as the first one. , . . In summer, colonies of Natterer's bat containing both males and females can be found in hollow trees, attics, crevices in walls, and in birdboxes. A very numerous summer colony of this bat occupies the roof space of the Cameldolites monastery at Bieniszew near Konin. Natterer's bats mainly spend winter in caves and cellars, where they form large, often mixed colonies. They fly low above ground among trees in search of flying prey but also snap insects and spiders from plants. _ This species occurs in Europe and in the temperate regions of Asia up to Japan in the east. In Poland its range covers almost all of the country except the central and western parts of the Baltic Coast, north-eastern regions of Masuria, Upper Silesia, NowyTarg Valley, the Tatra, Pieniny and Bieszczady Mountains, and Sandomierz Lowland. The West and East Beskid Mountains are its southernmost localities; to the north-east, it reaches as far as the Bialowieza primeval forest. In Poland lives the nominative subspecies, M. n. nattereri (Kuhl, 1818), originally described from Germany. Geoffroy's bat—Myotis emarginatus (Geoffroy, 1806) Measurements of 30 specimens from Czechoslovakia are as follows: head and body, 41-53 mm.; tail, 38-48 mm.; ear, 16-17 mm.; forearm, 36.0-40.5 mm. Cb. length of skull, 14.0-15.7 mm. Body weight, 5.9-9.0 gms. Bats — Chiroptera 127 (Feriancova-Masarova and Hanak, 1965). Wingspan, 22.0-24.0 cm. (Gaffrey, 1961). The ear has a conspicuous, deep indentation at one third of its length. The plagiopatagium reaches the base of the outer digit of the foot. The edge of the flying membrane between the ends of calcars and that of the tail is scarcely covered with soft, straight, thin hairs. The hair on the basal part of the interfemoral membrane is reddish and dense. The tail extends about 1 mm. beyond the flight membrane. The fur is dense, wavy and often contains woolly hairs. The dorsum is brown, varying in shades from reddish to chestnut. Individual hairs are tricolor (dark at the base followed by a band of lighter shade and tipped with brown). The second upper premolar (P2) is very small, protruding from the tooth-row and is inconspicuous from the buccal side. The second lower premolar (P2) is almost as large as the first one (Px). In summer, Geoffroy's bat can be found in attics, where females congregate in colonies. Single individuals or small colonies spend winter in caves, where they usually choose the places with a rather high temperature. The Myotis emarginatus Fig. 66. Distribution of Geoffroy's bat, Myotis emarginatus (Geoffroy). After Ruprecht, from Pucek and Raczynski (1981)
128 Systematic account winter sleep is long and never interrupted. In Poland, this species is associated with rocky areas and forests. Geoffroy's bat is distributed from western Europe to the Crimea, Caucasus, Turkestan, Iran and Palestine in the east. It is absent in the British Isles. In central Europe its northern range limit runs across Holland, southern Federal Republic of Germany and southern Poland. In Poland, this species occurs only in a few localities in the West Beskid Mountains and the Krakow-Wielun Upland (Fig. 66). It has also been recorded from the Karkonosze Mountains near the Polish frontier. The nominative subspecies, M. e. emarginatus (Geoffroy, 1806), originally described from France, also inhabits Poland. Whiskered bat—Myotis mystacinus (Kuhl, 1819) Measurements of 159 specimens from Czechoslovakia are as follows, head and body, 35.0-48.0 mm.; tail, 30.0-43.0 mm.; ear, 12.0-16.5 mm.; forearm, 31.0-37.0 mm. (Hanak, 1971). Cb. length of skull 12.0-13.2 mm. (Rybar, 1976). Body weight of specimens from Lower Silesia, 3.0-5.0 gms. (Haitlinger, 1976). Wingspan, 19.0-22.5 cm. (Gaffrey, 1961). This is the smallest Polish representative of the genus Myotis Kaup. The wings are rather narrow. The ears are shorter than those of Brandt s bat, with a slight indentation on the outer edge. When laid forward, the ear does not extend beyond the end of the nose. The snout is overgrown with long tactile hairs. The tip of the tail extends about 2 mm. beyond the flying membrane. The wing membrane is attached to the base of the foot digits. The "calcarial keel" is in the form of a rudimentary cutaneous fold, somewhat resembling an epiblema. The fur shows a great variety of colors. The specimens from Poland are usually dark chestnut on the back and lighter on the venter. Almost black and olive gray individuals are also found. The hairs are dense, long and silky. The color of the ears is dark, nearly black. The flying membrane is also dark. According to Hanak (1970), the whiskered bat's baculum, when compared with that of Brandt's bat, is much smaller, saddle-shaped and concave from side view. It is 0.43-0.62 mm. long and 0.22-0.29 mm. wide. The penis is thin all over its length and is not bulged at the end. The rostral portion of the skull is slender and the skull's profile is smooth. The small upper and lower premolars (P1 and Px) are definitely larger than P2 and P2, the P2 being usually suppressed between Px and P3. The large premolar (P3) has no conspicuous cingulum, a poorly developed protocone and Bats — Chiroptera 129 its two small buccal roots are at an obtuse angle to the axis of the palate (Fig. 67). Similarly as in the other representatives of the subgenus Selysius Bonaparte, 1841, the upper molars Mx-M3 of the whiskered bat have no protoconules (Fig. 63). The whiskered bat is the eurytopic species. In Poland it occurs sympatrically with Brandt's bat. It often hunts its prey over water. In summer it shelters in attics, hollow trees, birdboxes and may be found hanging behind blinds or beneath loose bark. At that time the females congregate in small colonies and the males remain solitary. In winter single individuals hide in caves, usually Fig. 67. Arrangement of the roots of left third upper premolar (P3) in the whiskered bat, Myotis mystacinus (Kuhl) (A) and in Brandt's bat, Myotis brandtii (Eversmann) (B). After Rybar (1976) in very damp places. The whiskered bat well resists adverse weather and remains active from early spring to late autumn. It is a fast and agile flier. It catches insects when flying but also captures them, as well as spiders, from plants or from the ground. The whiskered bat has a very wide range including all of Europe, a major part of Asia, up to Siberia and Kamchatka in the north, and reaching as far as India and Indonesia to the south. In Poland it inhabits all of the country being rather rare in the northern part (Ruprecht, 1974b). Its known localities are restricted in number due to the fact that, until recently, it was confused with Brandt's bat and hence the former data could not be taken into account. In the Tatra Mountains it has been observed as high as the Ciemniak Lodowa Cave (1715 m. above sea level), but probably reaches even higher. The Polish whiskered bat belongs to the nominative subspecies M. m. mystacinus (Kuhl, 1819), originally described from Germany.
130 Systematic account Brandt's bat—Myvtis brandtii (Eversmann, 1845) Measurements of 53 specimens from Czechoslovakia are as follows: head and body, 39.0-50.0 mm.; tail, 32.0-44.0 mm.; ear, 12.5-17.0 mm.; forearm, 33.2-37.7 mm. (Hanak, 1971). Cb. length of skull, 13.0-14.4 mm. (Rybar, 1976). Body weight of specimens from Lower Silesia, 4.0-7.0 gms. (Haitlinger, 1976). Wingspan, 19.0-22.5 cm. (Gaffrey, 1961). A characteristic feature of Brandt's bat are its rather long ears with a shallow indentation on the outer edge. When laid forward, the ear extends 1-3 mm. beyond the end of the nose. This is a small bat with relatively wide wings. The flight membrane is attached to the base of the feet digits. The calcar has no keel. The tail slightly extends beyond the uropatagium. The fur is various shades of brown, lighter on the ventral side, usually brownish white. The flying membrane is brownish or blackish. According to Hanak (1970), the baculum of Brandt's bat is much larger than that of the whiskered bat. It is saddle-shaped with a prominent protuberance on the proximal part of the dorsal surface. Its ventral side has a conspicuous depression, the sulcus urethalis. The baculum is 0.81-0.97 mm. long and 0.41-0.59 mm. wide. The penis is thick and bulged at the end. The skull is slender in the rostral portion and smooth from side view. There is some difference in the size of small premolars, the first upper and lower ones being slightly larger than corresponding second premolars (P] > P^). The large upper premolar (P3) has a well-developed protocone and its two small buccal roots are almost in the same line as the longitudinal axis of the palate (Fig. 67). Mx-M3 bear conspicuous protoconules, which are also characteristic for the other representatives of the subgenus Leuconoe Boie, 1830. Brandt's bat, being originally a central European species, is known from the Pleistocene of Hungary (the penultimate glaciation—Riss) (Topal, 1963) and also from the Holocene of the Polish Tatras (Woloszyn, 1970), where its remains are present in caves situated not too high in the mountains. Sometimes they are found together with those of the whiskered bat though the latter predominates in higher altitudes. It is the eurytopic species. In Poland, Brandt's bat occurs in woods, open lowlands and plateaus (Ruprecht, 1974 b). In summer this species can be seen behind blinds of wooden houses (Bialowieza primeval forest). In Zamosc region it was found to have summer shelters in wooden houses situated far in the forest (gamekeeper's lodge at Stoki in the Kosobudy forest inspectorate). In winter it hides in caves or in various underground passages that ecologically substitute the former. Brandt's bat inhabits the temperate zone of the Palearctic. Its range covers Bats — Chiroptera 131 Eurasia from the British Isles in the west to the Sakhalin, Primorski i Krai and Kamchatka in the east. According to Fedorov (1978), its northern range extends to a lattitude of 64° (near the town of Magadan). In the south, it reaches the southern part of Primorskii Krai (40° N) and goes further to eastern and central Mongolia. In Poland, Brandt's bat is known only from a few localities along the coastline and in Masuria, Pomerania, Wielkopolska-Kujawy Lowland, West Sudetes, Krakow-Wielun and Malopolska Uplands, Swi^tokrzyskie Mountains, Roztocze and the Bialowieza primeval forest (Ruprecht, 1981). The first Polish specimens of this species are those collected by I. K. Tarnani (Ruprecht, 1975) in the artificial cave at Pulawy. Hibernating bats were recorded in the Polnocna Cave at Wojcieszow Gorny (West Sudetes) (Haitlinger, 1976). Few specimens of the Brandt's bat spend winter near Mi?dzyrzecz on the Obra river in bunkers dating fiom World War II (E. Urbanczyk personal communication). Most of the known localities of Brandt's bat are those found in summer. In some of them this species co-occurs with the whiskered bat. Brandt's bat was originally described from Russia in the foreland of the Ural Mountains on the Sakmara river. So far no subspecies have been distinguished. Pond bat—Myotis dasycneme (Boie, 1825) Measurements of 9 specimens from Lower Silesia are as follows: head and body, 59.0-68.0 mm.; tail, 43.5-53.0 mm.; ear, 14.9-16.6 mm.; forearm, 44.1-49.2 mm. Body weight, 11.0-23.0 gms. (Haitlinger, 1976). Cb. length of skull of 10 specimens from Poland, (15.1) 15.7-17.4 mm. Wingspan, 20.0-30.0 cm. (Gaffrey, 1961). This is one of the Polish biggest bats. The wings are large and broad. The ventral side of the uropatagium along the shanks is covered with white hairs. The wing membrane reaches the heel. The feet are large and have long calcars which take two-thirds of the margin of uropatagium. The end of the tail extends about 3 mm. beyond the interfemoral membrane. The fur is not very long but it is very dense. There is a conspicuous difference in coloration of the dorsal and ventral sides of the body. The gray of the belly becomes mixed with brown on the back. All hairs have black bases; those on the back are tipped with brown whereas those on the belly have white ends. The skull is flat and wide and has a smooth profile. The first and second upper premolars (P1, P2) protrude lingually from the tooth-row. The protoconule is present on the proximal edge of the upper molars (Mx-M3).
132 Systematic account This is a lowland species associated with extensive water basins. In summer, single individuals or colonies can be found in attics and in hollow trees. The pond bat spends winter either solitarily or in groups, sometimes it joins the colonies of other bats. Late in the evening, it comes out to hunt over waters, flying low and catching insects in the air or snapping them from the surface of water. These bats apparently migrate from their summer quarters to the winter ones. . The pond bat inhabits central and eastern Europe ranging from Bohemia and Moravia (where it is rare) in the south to Belgium, Holland, Denmark and Sweden in the north. East of Europe it extends throughout West Siberia to the Yenisei. , x _T ^ In Poland this bat probably is present throughout the country. No records from Pomerania indicate lack of collecting rather than its actual absence there. Most of its known localities are in Masurian Lake Region, Wielkopolska- -Kujawy and Mazovian Lowlands, Krakow-Wielun and Matopolska Uplands, West Sudetes, Swi?tokrzyskie Mts., Pieniny, Tatras and Upper Silesia. The Polish fauna contains the nominative subspecies M. d. dasycneme (Boie, 1825), originally described from Germany. Daubenton's bat—Myotis daubentoni (Kuhl, 1819) Measurements of 78 specimens from Lower Silesia are as follows: head and body, 43.0-60.5 mm.; tail, 31.0-44.5 mm.; ear, 10.5-14.2 mm.; forearm 35.0-40.8 mm. Body weight, 4.0-8.0 gms. (Haitlinger, 1976). Cb. length of skull of 18 specimens from Poland, 13.2-14.6 mm. Wingspan, 21.0-25.0 cm. (Gaffrey, 1961). . , Daubenton's bat is one of the smaller Polish bats. The wing membrane reaches to the middle of the foot. The end of the tail extends about 3 mm. beyond the interfemoral membrane. The wings are rather broad. The feet are Iqi'00, The fur is dense and short. The dark brown back, often with a ieddish tinge, is well contrasted with the gray color of the belly. Individual hairs are bicolor. An albino Daubenton's bat has been recorded by Cerveny and Burger (1977); with partial albinism of tip of wings by Haensel (1968). The skull is flat and wide and has a smooth profile. It is likely to be confused with that of Brandt's bat, from which it differs by haying wider brain case, interorbital constriction and rostrum. In Daubenton s bat, the coronoid piocess of the mandible is a straight line, unlike that of Brandt s bat, which is parabolic in shape. In Daubenton's bat, the lower canine is wide and low, whereas that of Brandt's bat is narrower and higher (Fig. 68). Bats — Chiroptera 133 Fig. 68. Left mandibles. A — Daubenton's bat, Myotis daubentoni (Kuhl), B — Brandt's bat, Myotis brandtii (Eversmann), C — whiskered bat, Myotis mystacinus (Kuhl). Arrows indicate differences in the size of canines and second premolars The upper molars (Mx-M3) bear an additional cusp (protoconule) on the anterior edge of the crown (Fig. 62). The second upper premolar (P2) is half the size of the first one (P1). The second lower premolar is also much smaller than the first one (Pi). Daubenton's bat is associated with water but not as strictly as the pond bat. In summer, it takes up residence in hollow trees, attics, in crevices of rocks and walls, under bridge decks and in sand martins' nests. Its summer colonies contain members of both sexes and sometimes are mixed with those of other species. In winter, single individuals or colonies hide in caves and cellars. This bat h*s been observed to fly over long distances but not much is known about its regular migrations. It hunts both over waters and among trees, consuming its prey when flying. The flight is fast and irregular. Daubenton's bat is present almost throughout Europe. In the Palearctic
134 Systematic account part of Asia, it reaches up to a latitude of 60° N and extends as far as Japan in the east. In Poland it is rather evenly distributed all over the country. So far, it has not been recorded from the Baltic Coast, Podlasie, Sandomierz Lowland, Lublin Upland, Bieszczady, West Beskid, Tatra Mts., and the Nowy Targ Valley! The Polish fauna includes the nominative subspecies, M. d. daubentoni (Kuhl, 1819), originally described from Germany. Within the species Myotis daubentoni (Kuhl) a new form, Myotis nathalinae Tupinier, 1977, has been described from Spain. The lesser Daubenton's bat is a close relative of M. daubentoni, from which it differs by a smaller size (especially of the forearm, 34-35 mm. and 36-37 mm. respectively and of the hind foot, 8.4-9.0 and 9.4-10.1 mm.), and by the lack of the protocone on the third upper premolar (P3). It also has a different structure of the os penis, which m this species has a conspicuous protuberance on the dorsal side. The holotype and several paratype specimens were caught in a deserted mine in the province of Ciudad-Real south of Madrid. There were hundreds of individuals spread m small groups in which males predominated. They took up residence in flooded passages in the vicinity of greater horseshoe bats. Other habitats of this species were those occupied by Daubenton's bat at different times of the year. It has also been recorded from France and Switzerland, in the basins of the lower Loire and Rhone rivers. The lesser Daubenton's bat may occur m the Polish fauna since the skulls of M. daubentoni that were found in Poland contained a few morphotypes of P3 with structure analogous to those of M. nathalinae. These come from: Pomerania, Wielkopolska-Kujawy, lazovian Lowland and Malopolska Uplands, and Bialowieza primeval forest. Genus: Vespertilio Linnaeus, 1758 Polish fauna contains only one species. Parti-colored bat —Vespertilio miirinus Linnaeus, 1758 Measurements of 66 specimens from Poland are as follows: head and body, 47.j-64.0mm.; tail, 29.7-47.0mm.; ear, 12.9-18.8mm.; forearm, 39.6-48.2mm. Body weight, 10.7-16.4 gms. Cb. length of skull, 14.0-15.7 mm. Wingspan (acc. to Gaffrey, 1961), 26.5-30.0 cm. The ears and flying membranes are thick and dark. The ears are short. The tragus is short and slightly rounded at the end. The wings are rather narrow. The tip of the tail extends about 3.5-5.0 mm. beyond the interfemoral membrane. The calcarial keel is present. Bats — Chiroptera 135 The color of the fur much varies with the individual but dorsal hairs are always tipped with silver. The hair bases may be cherry red, dark brown or almost black. The fur on the venter is white or gray. The hairs on the face are dark all over their length. The belly is well contrasted with the back. The fur is short and dense. The number of teeth is 32. Dental formula: 2 113 3 12 3 The skull is massive, and the rostrum is quadrate in general outline. The incisura incisiva is deep and reaches the line across the posterior edges of the upper canines (Fig. 58). The second upper incisor (I2) is half as high as the first one (I1). The last upper molar (M3) is not suppressed in the anteroposterior diameter (Fig. 59). Little is known about biology of this species. Mountains and steppe regions are thought to be its chief habitats though it is often found in big towns of Europe. One thousand mandibles of parti-colored bats were recorded in pellets containing remains of 6000 vertebrates, produced by eagle owls in the Tatra Mts. (Czechoslovakia, Schaefer, 1974). In summer female colonies, single males and, occasionally, also small groups of males shelter in roof spaces, hollow trees or crevices in rock cliffs. Sometimes it forms mixed colonies with other species of bats, especially with Nathusius' pipistrelle and the common pipistrelle. -In winter it hides in caves, cellars and hollow trees. In autumn the parti-colored bat makes courtship flights during which it emits a characteristic squeek. It flies fast and high. This species has a very extensive range which includes all of central Europe reaching Denmark and southern Sweden to the north, Switzerland and Austria to the south, as well as eastern Europe and Palearctic part of Asia up to Japan in the east. Until recently, the parti-colored bat was believed to be rare in Poland. However, this opinion has proved to be erroneous since this species is quite common in several localities situated in: Masurian, Wielkopolska-Kujawy and Mazovian Lowlands, Podlasie, Bialowieza primeval forest, Krakow-Wielun, Malopolska and Lublin Uplands, Roztocze, Swi^tokrzyskie Mountains, Tatra, Pienines and West Beskid Mts. In western Poland it has only been recorded from the seaside, near the German frontier, and from Lower Silesia (Fig. 69). Further systematic study is needed to provide more information about the distribution of V. murinus, especially in the western part of the country. In Poland lives the nominative subspecies, V. m. murinus Linnaeus, 1758, originally described from Sweden.
136 Systematic account Genus: Eptesicus Rafinesque, 1820 It resembles the genus Vespertilio Linnaeus, from which it differs by having a much reduced M3 (Fig. 59), by the size of incisura incisiva, which does not reach the line across the posterior edges of the upper canines as it is in the former genus, and by the triangular outline of the rostrum (Fig. 58). The calcarial keel is poorly developed and lacks the reinforcing cartilage. The ears are short, triangle-shaped and black in color. The number of teeth is 32. Dental formula: 2 113 3 12 3 The height of the second upper incisor (I2) is more than half the height of the first one (I1). On the back, the fur has no silver tinge. It is poorly contrasted with that of the belly, the latter being slightly lighter. The wings are rather wide and bluntly ended. These bats are associated with forests, mountains and human settlements. Two species of this genus occur in Poland. Bats — Chiroptera 137 Key to species by external features 1. Hairs on back usually tipped with gold. Length of forearm less than 44 mm Eptesicus nilssoni (Keyserling et Blasius), page 137 —. Hairs on back usually tipped with buff. Length of forearm more than 48 mm Eptesicus serotinus (Schreber), page 139 Key to species by skulls 1. Height of second upper incisor (I2) equals 2/3 that of first one (I1). Cb. length of skull less than 16 mm.; length of mandible, 10.511.6 mm.; height of mandibular ramus,^2.9-3.9 mm Eptesicus nilssoni (Keyserling et Blasius), page 137 —. Height of second upper incisor (I2) less than 1/2 that of first one (I1). Cb. length of skull more than 18 mm.; length of mandible, 14.215.8 mm.; height of mandibular ramus, 5.1-6.0 mm Eptesicus serotinus (Schreber), page 139 Northern bat—Eptesicus nilssoni (Keyserling et Blasius, 1839) Measurements of 20 specimens from Czechoslovakia are as follows: head and body, 45-64 mm.; tail, 37-48 mm.; ear, 11.5-17.0 mm.; forearm, 37.0^43.5 mm. Cb. length of skull, 14.0-15.6 mm. Body weight, 8.5-12.5 gms. (Feriancova-Masarova and Hanak, 1965). Wingspan, 24.0-27.0 cm. (Gaffrey, 1961). The ears are short and wide. The tragus is blunt and short, with the middle portion most expanded. The wing membrane reaches the base of the foot digits. The tip of the tail extends 2-3 mm. beyond the interfemoral membrane. The hairs are very long, dark chestnut, and tipped with gold on the back. The ventral side of the body is well contrasted in color since the hairs are light brown on ends. The ear and flying membrane are of a darker color. The first upper incisor (I1) is slightly larger and longer than the second (I2). The Northern bat inhabits forests and mountains. In summer, it takes refuge in tree hollows, attics and in rock slits, where single males or female colonies can be found. In winter, it hides in caves choosing dry or cool places with a temperature slightly below 0°C. The flight is fast with frequent turns. 10
138 Systematic account The range of this species covers central Europe, northern and eastern regions of Asia up to Mongolia, Tibet, Turkestan and probably also xManchuria and Korea. In Europe, it reaches beyond the polar circle (Norway, Kola Peninsula). In the south, it extends to the Alps, Sudetes, Carpathians and Himalayas. As presently known, the Northern bat is distributed in most regions of Poland. So far it has not been recorded from the Baltic Coast, Pomerania, Lower and Upper Silesia, Trzebnickie Hills, East Sudetes, Bieszczady Mountains, Roztocze and Lublin Upland (Fig. 70). Similarly as in the case Fig. 70. Distribution of the northern bat, Eptesicus mlssoni (Keyserling et Blasius). After Ruprecht, from Pucek and Raczyriski (1981) of the parti-colored bat, the absence of the Northern bat in western Poland seems to be conspicuous enough to give grounds to further fauna research In Poland lives the nominative subspecies, E. n. nilssoni (Keyserling et Blasius, 1839), for the first time described from the mountains of Scandinavia (Sweden). Bats — Chiroptera 139 Serotine—Eptesicus serotinus (Schreber, 1774) Measurements of 457 specimens from Poland are as follows: head and body, 62.6-82.0 mm.; tail, 34.0-66.5 mm.; ear, 12.0-21.0 mm.; forearm, 48.0-56.0 mm. Cb. length of skull, 18.1-21.2 mm. Body weight, 14.4-33.5 gms. Wingspan (acc. to Gaffrey, 1961), 31.5-38.1 cm. The serotine is one of the largest bats in Poland. The ears are long. The tragus is elongated and widest at one third of its height. The wings are wide and blunt. The wing membrane reaches the base of the foot digits. The end of the tail extends 7-8 mm. beyond the interfemoral membrane (Fig. 55). The fur is dark, on the back the hairs are chestnut with lighter ends. The ventral side of the body is similar in color but is a bit lighter and has a tinge of gray. The serotine is primarily associated with human settlements. In summer, nursing colonies and individual specimens can be found in the roof spaces of houses both in the town and forest. In winter, it finds shelter in cellars, attics and caves. The serotine hunts in the gardens, parks and on the edge of forests. It is a slow flier and often makes circles o\er one spot, such as a clearing for instance. ' This bat is distributed almost throughout Europe, northern Africa and Asia up to China and Korea. The serotine belongs to the most common species of Polish bats inhabiting almost the entire country. It is rarer in the southwestern parts of Poland. So far it has not been recorded in the Trzebnickie Hills, East Sudetes and in the Nowy Targ Valley. In Poland there occurs the nominative subspecies, E. s. serotinus (Schreber, 1774), originally described from France. Genus: Pipistrellus Kaup, 1829 - These are the smallest bats of Poland. The wings are narrow. The calcarial keel is well developed and reinforced with a transverse cartilage. The ears are small and triangular. The flying membrane is thick, somewhat skin-like and blackish in color. These bats live in forests. The number of teeth is 34. Dental formula: 2 12 3 3 12 3 The fauna of Poland includes two species. Occurrence of the third one was claimed erroneously since it is very doubtful that it makes even occasional trips to the Polish territory. \
140 Systematic account Key to species by external features 1. Last coccygeal vertebra extends about 4 mm. beyond interfemoral membrane. Hairs on back tipped with yellowish gold Pipistrellus savii (Bonaparte), page 141 —. Interfemoral membrane includes only half of last vertebra which extends about 1 mm. beyond it. Hairs on back not tipped with yellowish gold 2 2. Ear, when laid forward, does not extend beyond end of nose. When wing folded, length of thumb with claw less than width of carpus (Fig. 71). Length of forearm usually less than 33 mm Pipistrellus pipistrellus (Schreber), page 141 . Ear, when laid forward, extends beyond end of nose. When wing folded, length of thumb with claw equal to or slightly more than width of carpus (Fig. 71). Length of forearm usually more than 33 mm. . . . Pipistrellus nathusii (Keyserling et Blasius), page 142 Fig. 71. The length of the thumb in relation to the width of the wrist. A the common pipistrelle, Pipistrellus pipistrellus (Schreber), B — Nathusius' pipistrelle, Pipistrellus nathusii (Keyserling et Blasius). From Toschi and Lanza (1959), modified Key to species by skulls 1. First upper incisor (I1) unicuspid. First upper premolar (P1) absent or very small, not visible from buccal side. Length of mandible 9.3-10.2 mm.; height of mandibular ramus more than 2.8 mm. (2.8-3.3 mm.) Pipistrellus savii (Bonaparte), page 141 —. First upper incisor (I1) bicuspid. First upper premolar (P1) visible from buccal side. Height of mandibular ramus less than 2.8 mm. ... 2 Bats — Chiroptera 141 2. Third lower incisor (I3)set close to I2. First upper premolar suppressed between canine and large premolar (P2). Cb. length of skull, 10.012.3 mm.; length of mandible, 7.7-8.6 mm.; height of mandibular ramus, 2.1-2.6 mm • Pipistrellus pipistrellus (Schreber), page 141 —. Third lower incisor (I3) spaced from I2. First upper premolar not concealed behind canine. Cb. length of skull, 12.2-13.2 mm.; length of mandible, 8.8-9.7 mm.; height of mandibular ramus, 2.22.7 mm. . . . Pipistrellus nathusii (Keyserling et Blasius), page 142 **Savi's pipistrelle—Pipistrellus savii (Bonaparte, 1837) Measurements of south European specimens are as follows: head and body, 43-52 mm.; tail, 34-40mm.; ear, 10-17mm.; forearm, 31-38 mm. Cb. length of skull, 12.8-14.0mm. Body weight, 6-10 gms. (Konig, 1969). Wingspan, 22.0-22.5 cm. (Gaffrey, 1961). It is a small bat but larger than the other species of the genus Pipistrellus Kaup. Two indentations are present at the base of the outer edge of the tragus. The tail extends about 4 mm. beyond the interfemoral membrane. Near the base, the penis is bent at a right angle forming a reversed letter L. The fur consists of dense and long hairs, which are dark at bases and lighter at ends. Hairs on the back are tipped with yellowish geld. The flying membrane is dark in color. The first upper premolar is very small and invisible from the buccal side. In the upper jaw, the carjine touches the large premolar. Sometimes P1 may be retained. Savi's pipistrelle inhabits both lowlands and mountains, where it reaches high altitudes. In summer, colonies of females occupy the roof spaces of houses and single males are found in various crevices of buildings. They also take up residence in hollow trees. This bat is distributed in the European part of the Mediterranean region reaching the Alps. It also lives in the Canary Islands, Crimea, Caucasus and in central-eastern Asia up to Vladivostok in the north. Individual specimens of Savi's pipistrelle were reported to have been caught in Leipzig and Wroclaw in the years 1931 and 1939. However, further study proved that these specimens belonged to Pipistrellus nathusii (Keyserling et Blasius). It is very unlikely that Savi's pipistrelle will ever occur in Poland. In southern Europe lives the nominative subspecies P. s. savii (Bonaparte, 1837) described from Italy. Common pipistrelle—Pipistrellus pipistrellus (Schreber, 1774) Measurements of 135 specimens from Poland are as follows: head and body, 32.0-48.7 mm.; tail, 23.0-35.0 mm.; ear, 7.0-12.0 mm.; forearm, 26.0-33.2 mm. Body weight, 3.5-9.2 gms. Cb. length of skull, 10.0-12.3 mm. Wingspan (acc. to Gaffrey, 1961), 18.0-23.2 cm.
142 Systematic account This is the smallest Polish bat. The wings are rather narrow. The ears are short. The outer edge of the tragus is indented at the base. The end of the tail extends about 1 mm. beyond the interfemoral membrane. The thumb is short; when the wing is folded, it is shorter than the width of the carpus (Fig. 71). The dorsal surface of.the interfemoral membrane is scarcely haired for the basal third, being naked in the distal half of the shank. The fifth digit of the hand is about 40 mm. long. The fur on the back is brown with a slight golden tinge, that of the venter is similar but lighter in color. The hairs are short but dense. / The first upper premolar is visible from the buccal side though it is small and protrudes lingually from the tooth-row. The lower canine is wide and short. The common pipistrelle lives in forests and near human settlements. In summer, the bats take up their abode in attics, hollow trees, nest-boxes, crevices in buildings and beneath loose bark. At that time the females congregate in colonies which occasionally contain single males. Most males, however, shelter alone. Sometimes the common pipistrelle forms colonies mixed with other species of bats, e.g. Nathusius' pipistrelle and the particolored bat. In winter,, farge groups hide in various crevices in houses, hollow trees and, on rare occasions, in caves. These bats are rather early fliers; they appear at dusk and sometimes even before sunset. Gardens, parks and the edges of forests are their hunting areas. The flight is characterized by frequent changes of direction. The common pipistrelle is spread throughout Europe, northern Africa, and western and central Asia up to Turkestan and Kashmir. This bat is common in most parts of Poland. In southwestern regions of the country it is rare. So far it has not been reported from the Trzebnickie Hills, Krakow-Wielun Upland, West and East Sudetes, Tatras, Nowy Targ Valley, Bieszczady Mts. and in the Sandomierz Lowland. In Poland lives the nominative subspecies, P. p. pipistrellus (Schreber, 1774), originally described from France. Nathusius' pipistrelle—Pipistrellus nathusii (Keyserling et Blasius, 1839) Measurements of 69 specimens from Poland are as follows: head and body, (42.2) 46.1-55.8 mm.; tail, 30.1-44.6 mm.; ear, 10.0-13.5 mm.; forearm, (30.8) 31.5-37.0 mm. Body weight, 6.0-9.0 gms.; Cb. length of skull, 12.113.2 mm. Wingspan (acc. to Gaffrey, 1961), 23.0-24.5 cm. In general appearance, Nathusius' pipistrelle is similar to the common pipiBats — Chiroptera 143 strelle but it is slightly larger, and has bigger ears and a little broader wings. When the wing is folded, the length of the thumb equals or exceeds the width of the carpus (Fig. 71). In the hand, the fifth metacarpal with its corresponding digit are 46 mm. long. Dorsal surface of the interfemoral membrane is thickly haired for basal half and along the shank. The penis is not bent at the base. The fur is longer and more dense than in the common pipistrelle. The hairs have black or dark brown bases. The fur is dark brown on the dorsum and lighter on the venter. The first upper premolar (P1) is well developed and slightly shifted lingually. The lower canine is high and narrow. Nathusius' pipistrelle inhabits wooded areas and human settlements. During summer, it can be found in hollow trees, birdboxes and in attics. Females congregate in colonies where single males can also be observed. Sometimes Nathusius' pipistrelle forms mixed colonies with other species. In winter, it hides about buildings and in hollow trees. It hunts in parks and at the edges of forests. This species is distributed nearly all over Europe; to the north its range extends to southern Sweden and Leningrad (60°) and to the east it reaches the Caucasus, Persia and Asia Minor. In Poland, Nathusius' pipistrelle primarily occurs in the lowland regions of the country. At present it is mostly located on the Baltic Coast, in the Masurian and Pomeranian Lake Regions, in Wielkopolska-Kujawy, Mazovian and Sandomierz Lowlands, as well as in Podlasie and in the Bialowieza primeval forest. It has also been recorded in Upper and Lower Silesia and in the Malopolska and Lublin Uplands (Ruprecht, 1976 b). No subspecies have been distinguished. Genus: Nyctalus Bowdich, 1825 The wings are narrow, the fifth digit shortened. The flying membranes are blackish in color. The ears are short and triangular, reaching the eye with the lower edge. The tragus is shortened and expanded in the upper portion. The calcar bears a large keel reinforced with a transverse cartilage. The number of teeth is 34. Dental formula: 2 12 3 3 12 3
144 Systematic account The first upper premolar (P1) is usually very small. It is invisible from the buccal side and bften inapparent from the lingual side. The skulls of this genus are characterized by a nearly squarish outline and a very large incisura incisiva whose distal edge reaches the line running across the middles of the large premolars (P2). These bats are associated with wooded areas and are good fliers. Key to species by external features 1. Length of forearm less than 46 mm. Hair bases on dorsal and ventral sides darker than ends .... Nyctalus leisleri (Kuhl), page 146 —. Length of forearm more than 46 mm. Hairs usually unicolor; if with inconspicuous bicoloration on dorsal side, then lighter at bases than on ends 2 2. Length of forearm less than 58 mm Nyctalus noctula (Schreber), page 145 —. Length of forearm more than 62 mm Nyctalus lasiopterus (Schreber), page 145 Key to species by skulls 1. Back of skull characteristically "upturned" from side view. Cb. length of skull more than 20 mm. Length of maxillary tooth-row more than 8.5 mm. Length of mandible more than 17.0 mm.; height of mandibular ramus more than 5.0 mm Nyctalus lasiopterus (Schreber), page 145 —. Back of skull not "upturned" from side view. Cb. length of skull less than 20 mm.; length of maxillary tooth-row less than 8.5 mm.; length of mandible less than 17.0 mm.; height of mandibular ramus less than 5.0 mm 2 2. First upper premolar (P1) invisible from buccal side. Cb. length of skull more than 17.4 mm.; length of maxillary tooth-row more than 6.7 mm.; length of mandible, 13.4-15.0 mm.; height of mandibular ramus, 4.3^1.8 mm Nyctalus noctula (Schreber), page 145 —. First upper premolar (P1) partly visible from buccal side. Cb. length of skull less than 16.5 mm.; length of maxillary tooth-row less than 6.3 mm.; length of mandible, 10.9-11.9 mm.; height of mandibular ramus, 2.9-3.6 mm Nyctalus leisleri (Kuhl), page 146 Bats — Chiroptera 145 *Giant noctule—Nyctalus lasiopterus (Schreber, 1780) Measurements, according to Hanak and Danko (1975), are as follows: head and body, 84.0— 104.0 mm.; tail, 55.0—65.0 mm.; ear, 21.0—26.0 mm.; forearm, 63.0—69.0 mm.; wingspan, 41.0—45.0 cm. Cb. length Uf skull, 21.9—23.6 mm. Body weight, 41—76 gms. This is the largest European bat, similar to the common noctule but larger. The ear is distinctly wide, much wider than that of N. noctula, and its lower margin reaches the eye in the form of a thickened open tube. On the ventral side of the propatagium there is a characteristic band of rusty colored hairs. The back is yellowish or brownish red and the belly is slightly lighter. A well-developed calcarial keel is reinforced with a transverse cartilage. The crista lambdoidea is extremely well developed and much upturned toward the front of the skull where it passes into the external occipital protuberance. The teeth resemble those of N. noctula but are relatively larger. The first upper premolar (P1) is so small that it is invisible without the aid of a lens. The giant noctule primarily inhabits hollow pine, lime and oak trees and is very difficult to find. It was observed in mixed colonies with the common noctule but it mainly forms small singlespecies colonies. This bat feeds on large insects, which it takes during flight, high above ground. Two young are born in June. The range of this species covers the European zone of deciduous forests from Spain through France, Switzerland, Italy, Yugoslavia, Greece, Bulgaria, Turkey, Czechoslovakia, Poland, Hungary to Byelorussia and Ukraine. In the U.S.S.R., according to Strelkov (1963), the giant noctule ranges from Moscow district and Gorki in the north to the southern Crimea and southwestern Transcaucasia. In Poland, N. lasiopterus was found only once in pellets of the barn owl at Krolikow in the Wielkopolska-Kujawy Lowland (Ruprecht, 1970). The fact that the giant noctule's locality is situated in the central part of the country and that its remains have been found in the pellets of this particular species of the owl (which has a very small home range) indicate that this bat is the native element of the Polish fauna. In Europe there lives its nominative subspecies N. I. lasiopterus (Schreber, 1780) described from Italy. Common noctule—Nyctalus noctula (Schreber, 1774) Measurements of 207 specimens from Poland are as follows: head and body, 60.0-84.8 mm.; tail, 41.0-60.6 mm.; ear, 10.5-21.2 mm.; forearm, 50.058.5 mm. Cb. length of skull, (16.9) 17.4-19.9 mm. Body weight, 16.7-44.2 gms. Wingspan, acc. to Hanak and Danko (1975), 32.0-40.0 cm. This is one of the largest bats of Poland. The wings are narrow and long. The fifth digit is shortened. A well-developed calcarial keel is reinforced with a transverse cartilage. The wing membrane comes to the heel. The tail extends 2-3 mm. beyond the interfemoral membrane. The ears are short, rounded and thickened at the edge. The tragus is short with a flat and expanded end. The dense fur is made up by short hairs which are either unicolor or lighter