A contribution to the population ecology of Nyctalus noctula (Mammalia: Chiroptera).

PRfRODOVEDNfi PRACE DSTAVtl CESKOSLOVENSKE AKADEMIE VED V BRNE Acta scientiarum naturalium Academiae scientiarum bohemoslovacae - Brno Abbreviation: Acta Sc. Nat. Brno Tomus XIII, nova series - 1979 Editor-in-Chief: Academician Josef Kratochvil Assistant Editor: Juliana Hobstova Editorial Board: Ing. Vlastimil Barus, DrSc., Corresponding Member of the CSAV, Director of the Institute of Vertebrate Zoology of the CSAV, Brno, Doc. RNDr. Jaromir D e m e k. DrSc., Brno, RNDr. Milena Rychnovska, CSc., Head of the Ecological Department of the Institute of Botany of the CSAV. Brno, Prof. RNDr. Josef S e k a n i n a, DrSc., Corresponding Member of the CSAV Linguistic collaboration: Dr. Ing. R. Obrtel, CSc. (English), Dr. J. Gruna (German), Ing. E. BezdiCkova (Russian), Doc. Dr. V. S t u p k a, CSc. (French) © by ACADEMIA PRAHA Pfirodovidni prdce ustavi Ceskoslovenski akademie vid v Brni Acta scientiarum naturalium Academiae scientiarum bohemoslovacae — Brno Abbreviation: Acta Sc. Nat. Brno Pfirodovidni price ustavi Ceskoslovenske akademie vid v Brni (Acta scientiarum naturalium Academiae scientiarum bohemoslovacae - Brno) is a continuation of the periodicals Price Moravski pfirodovidecke spoleinosti (Acta Societatis scientiarum naturalium mora— vicae) I—XXVI, 1924—1954, and Price Brninski zikladny Ceskoslovenske akademie vid (Acta Academiae scientiarum ceclioslovenicae Basis brunensis) XXVII—XXXIV, 1955-1962. Published by the Czechoslovak Academy of Sciences in the Publishing House ACADEMIA., Vodifikova 40, 112 29 Prague, Czechoslovakia. Address of the Editor's Office (manuscripts, proofs, etc.): Institute of Vertebrate Zoology of the CSAV, Kv§tn& 8, 603 65 Brno, Czechoslovakia. Editor-in-Chief: Academician J o s e f Kratochvil, Brno Assistant Editor: Juliana Hobstovd Editorial Board: Ing. Vlastimil B a r u §, DrSc., Corresponding Member of the CSAV, Director of the Institute of Vertebrate Zoology of the CSAV, Brno Doc. RNDr. Jaromir Demek, DrSc., Brno RNDr. Milena Rychnovski, CSc., Head of the Ecological Department of the Botanical Institute of the CSAV, Brno Prof. RNDr. Josef Sekanina, DrSc., Corresponding Member of the CSAV, Brno Linguistic collaboration: Dr. Ing. R. Obrtel, CSc. (English), Dr. J. Gruna (German), Ing. E. Bezdidkovfi (Russian), Doc. Dr. V. Stupka, CSc. (French) Pri'rodovedne prdce ustavu Ceskoslovenske akademie ved v Brne Acta Sc. Nat. Brno, 13 (1): 1-38 Jin GaisJer, VSadimi'r Hanak, Jan Dungei A CONTRIBUTION TO THE POPULATION ECOLOGY OF NYCTALUS NOCTULA (MAMMALIA: CHIROPTERA) ACADEMIA Nakladatelstvi Ceskoslovenske akademie ved Praha 1979 Tomus XIII, nova series - Fasc. 1 Received: November 24, 1977 Scientific Editor: Academician Prof. Dr. J. Kratochvil, DrSc. Critical Revision by Ing. VI. Barus, DrSc., Corresponding Member of the CSAV Abstract From March to October colonies of Nyctalus noctula were found in hollow trees where also the reproduction takes place; winter colonies in trees are evidenced by oldish material. In buildings these bats were found throughout the year with the exception of June and July, and they do not reproduce there. One winter colony was found in a crack in a rock. According to the authors' material and according to data in literature the relation of different components of populations to different types of shelters is evaluated as well as the behaviour of N. n. in their hunting grounds. In Central Europe all females and most of the males mature at the age of about 3 months, i. e., immediately after finishing their growth. Most females older than 1 year have 2 young and the average number of young per one grown-up female is 1.8. The sex ratio at birth is 1:1 and is probably also maintained later on. The young are born from mid-June till early July and nursery colonies disintegrate in August. Old females as well as those born in that year move to mating quarters to territorial old males. The males of that year do not mate and in juvenile colonies they gradually exceed the females in number. In October at the latest the summer shelters are left till March or April. Males prevail in all winter samples from Western and Central Europe; the possible causes of this phenomenon are being discussed. The average population density in the summer season, found in two areas of the optimum habitat, is 0.32 individuals per ha. Although originally a forest species, N. noctula has managed to adapt to the civilization pressure and in places it exhibits a trend towards a hemisynanthropic way of life. Contents Introduction Material and methods Results Shelters Trees Buildings Other shelters Flight activity, hunting grounds Reproduction H Description of reproductive organs 11 Age and sexual maturity 13 Male sexual cycle 13 Female sexual cycle 13 Population 13 Sex ratio 18 Structure of the summer population 13 Density of the summer population 21 Discussion 25 Shelters 25 Hunting activity 28 Reproduction 23 Population 31 Summary 35 Peaio' e 36 Literature 35 Introduction Nyctalus noctula (Schreber, 1774) (hereinafter, N. n.) occurs in Europe south of approximately the 60th parallel, being frequent in woodland or park landscapes of lowland regions. It is a species with considerable migratory activity, 2 covering distances up to 1,500 km (Strelkov 1969, R o e r 19/1). Its ecology has been dealt with by a number of authors, particularly Lohrl (1936), Ryberg (1947), Kuz'akin (1950), Meise (1951), Blackmore (1963), Cranbrook & Barret (1965), Sluiter & Heerdt (1966), Stratmann (1968), Pan'utin (1963, 1970) and Sluiter, Voute, Heerdt (1973). Despite the fact that the number of papers devoted to this species is considerable, many aspects of its ecology have remained obscure. It is above all the fact that there are few quantitative data concerning the populations; also the yearly life cycle and reproduction have been.studied only in part. In the present paper we therefore summarize our observations carried out in the western part of Czechoslovakia (Bohemia and Moravia) in the course of the last 20 years. The main objective of the paper is the analysis of the occupation of shelters all the year round, the evaluation of important aspects of the population, such as the reproduction process, and the estimation of the structure and density of the summer population in the optimum habitat. The share of the individual authors is as follows: The third author dealt with reproduction. The second author obtained most of the material from Bohemia and elaborated the chapter on population. The first author obtained most of the material from Moravia and elaborated the remaining chapters, preparing the paper for print. For the assistance in obtaining our material we wish to thank numerous persons, above all, our students and colleagues; extraordinary help was given by Dr. I. Horacek and Mr. V. Bejcek (both from Prague). We also thank Dr. I. Flasar from the Museum at Teplice for supplying us with material of a great thanatocolony of the species studied. Dr. V. HrabS, CSc., from the Institute of Vertebrate Zoology, CSAV (Brno), has our thanks for carrying out the cross-sections through the teeth of the noctules. And, last but not least, we wish to thank the heads of our workplaces for enabling us to carry out the research, and the reviewers for valuable comments. Material and methods The region of the research is situated between 50°15' and 48°40' northern latitude and 13°10' and 18°15' eastern longitude. The period of the research lasted from 17 Sept., 1955, to 15 July, 1977. Altogether, 1,383 individuals were caught, out of which there were 963 grown-up and 420 juvenile individuals. Most of those individuals were banded and released. For studying the reproduction 18 grown-up males and 24 grown-up females were killed. Besides this material about 400 skeletons or mummies were found. In the above sample : those individuals flying away before their sex could be established and those observed in flight in the hunting grounds are not included. Owing to the fact that N. n. is one of the most frequent bats in Czechoslovakia (Hanak 1967) and in Central Europe in general, we do not give a list of collecting stations. The main regions of collection were: the south Bohemian . pond basin (about 50 °/o of the sample), central Bohemia, including the City of ' Prague, and southern Moravia including the southern part of the Ceskomoravska vrchovina Highland and the City of Brno. A detailed description of the main region under investigation in southern Bohemia is given in the chapter dealing with the density of the summer population. The stations studied are situated at elevations from 150 to '500 m. ! . For catching N. n. from hollow trees we most often used cages according to ,B e 1 s (1952). which, in the course of time, were variously modified. We found iA out that the probability of the whole colony flying out was increased by decreasing the compactness of the cage walls, particularly that opposite the opening of the cavity. It is not even necessary that the cage should be completely closed, as far as the wall opposite the opening safeguards knocking off or sliding of the outflying bats into a plastic bag fastened instead of the bottom of the cage. The type used at present, suggested by the second author, has a frame made of a strong wire and only a slanting roof consisting of silon fibres stretched at the distance of 1 cm from each other (PI. II, left). The cage can be folded and the catching bag removed, so that a large number of such devices can be transported. Another catching device are telescopic dural rods 8 m in maximum possible length in which the cage can be put and lifted to hollows situated at a major height. In the systematically followed region, next to the known holes there are permanently hammered hooks on which the catching cages are hung up. The catching device was placed before the entrance holes about one hour before the sunset and removed after finishing the catching. When working with cages of the older type (all walls made of wire netting and a fabric bag instead of the bottom) it sometimes happened that only some of the individuals flew out, but others stayed in the cavity from where they uttered sounds. If they were grown-up individuals (the young can be distinguished according to the sounds), the cage was left in place for several hours, twice even for the whole night. In spite of this it happened several times that part of the colony evidently remained in the hollow; those cases are excluded from the calculation of the size of the colonies. If the individuals flew quickly one after another and no sounds were heard after the last one had left the hollow, the catch was considered quantitative. From hiding places in buildings the bats were taken out with hand or with a long forceps or caught with a hand net. Such catches could not often be quantitative. A small sample described in the chapter on hunting grounds was mist-netted. At the beginning of the research several individuals were shot dead in their hunting grounds (they could not, however, be included in the material for studying the reproduction); besides we were several times given individuals caught in various environments, often unable to fly. Skeleton remains or mummies were, as a rule, found in building adaptations of houses. The above sample of 1,383 individuals was classified according to sex and further into juvenile and grown-up individuals. As for the grown-ups we tried to classify them into yearlings and old ones, but the identification of yearlings is not quite reliable, as described below. We should like to stress the fact that yearlings cannot be synonymized with subadult individuals, as our study of the reproduction has shown that the existence of the subadult stage in N. n. is questionable. The terms used in the paper are to be understood as follows: juveniles = young individuals at the stage of rapid growth, from the end of June to the end of September; grown-ups = all individuals older that juveniles; yearlings = individuals older than the young and younger than one year, from October to May, sexually mature as a rule; the old = all individuals older than the yearlings, from June of the year following the year of birth, sexually mature (adult). In the material of 42 specimens used for the study of the reproduction the following were determined: body mass, coloration of pelage, the stage of upper limb metacarpal and finger ossification, the dry mass of the eye, and the degi ee of dentition abrasion. Genital organs were dissected and preserved. In 5 males the right testis, the right epididymis and the vesicular gland were weighed; smears and histological sections of testes and epididymides were carried out. In females the size of nipples was estimated, the width of vaginal fissure measured as well as the length of uterus and the presence of embryos or sperms in the uterus established - the latter in the smear of the cut right uterine horn. Further, histological sections of ovaries, uteri and vaginae were made. In randomly selected specimens (7 males, 11 females) transversal sections through the right lower canine were made. . The ossification of long bones of the wing is well visible macroscopically in juvenile individuals when the light is passed through the wing; this has been known in bats for a long time (Barrett-Hamilton llf^h/newlv ginous zone of cartilago epiphysaria appears white in passing light, the ne y formed primary bone is reddish brown and the permanent secondary bone is gray. The growth zone is more or less macroscopically visible in yearlings m October and under a dissection microscope also in November. The later distinction of yearlings from older individuals was carried out according to the dentition with fine sharp points and darker hairs. In the material used foi the s y of reproduction where further criteria were available the distinction of yearlings is reliable, but in the sample under investigation of living animals the field the distinction of yearlings is rendered difficult by subjective erio taken into account in the evaluation. The dry mass of the eye was ascertained instead of the usual dry mass 01 lens (cf., Giles 1971) on the assumption that with progressing age the dry mass of'different eye parts increases, such as of sclera and the lens. The right eye was fixed for 7 days in 10% formaldehyde, then dried at 80 C. for 36 hours and immediately weighed. On the canine sections the increment zones of dentin were followed by means of the Klevezal & Klejnenberg (1967) method. The teeth were decalcified in 7 % nitric acid for 6 hours, rinsed in running water for 5 minutes, cut on a freezing microtome and stained with the Mayer haematoxyline. Histological sections of the genital organs were processed by current technique; they are not reproduced in the paper, as they agree with published materials on bats with a similar reproduction cycle. Results Shelters As in other countries of Central Europe, N. n. in Czechoslovakia was most often found in trees and, from time to time, in buildings (Gaisler & Hanak 1969, Hurka 1973). In our material (n= 133 finds of living individuals or colonies) 72.8% fall to tree cavities, 21.8% to buildings, and 5.4 /0 to remaining finds. Trees. As shown in Fig. 1, N. n. was found in tree cavities from Maich to October The average abundance in one shelter was the highest in July. In hollows we found individuals, pairs as well as colonies (for the definition of a bat colony, cf. Gaisler 1966). In Tab. 1 we distinguish o types of colonies according to their composition. Their number varies from 3 to 53 individuals, on the average the strongest colonies were those of grown-up females, found from March to June. The data in the table are distorted by the fact that they do not include the young incapable of flight which remained in the cavities 6 n.3 - t BUILDINGS J FMAMJ J.A SO J F MAM J J ASO N O Fig. 1. Composition of samples obtained in the two main types of shelter in the course of the year. Explanations: ordinate, the average number of individuals per shelter per month; abscissa, months; 1, grown-up males; 2, grown-up females; 3, juveniles; 4, sex and age not determined; n, number of finds per month. Tab. 1. The size of colonies in tree holes Type of aggregation Period Number of individuals in a colony Type of aggregation Period 3-10 11-20 21-30 31-40 41-50 51-60 X grown-up males + females May—Oct. 9 5 3 —• — — 11.7 grown-up males May—Sept. 7 — — — — — 6.9 grown-up females March—June 6 2 3 — 1 1 19.5 .nursery July—Sept. 4 4 4 1 1 — 18.6 juvenile Aug.—Sept. 11 2 — — — — „• 6.5 not stated May—Sept. 2 1 3 — — — t 16.5 total March—Oct. 39 14 13 1 2 1 13.4 and thus could not be caught. According to sound utterances and individual finds of quite small young the births of N. n. take place from mid-June to early July in the region under investigation. The first flying young were found ort 12 July. The number of members of nursery colonies is — particularly in the first half of July — substantially higher than that stated in Tab. 1, these colonies being in fact numerically strongest aggregations of the species studied in the summer season. Later on, their numbers drop because, starting from mid-August, the nursery colonies begin to disintegrate, as will be described in the chapter on population. By the number of members the smallest are the colonies of males and the juvenile colonies, the latter type being more frequent. A small number of grown-up males can be found in female and nursery colonies. Besides the data summarized in Tab. 1, in the cavities of trees individual males were found (3X), a pair of males (IX), and a male and a female (2X). The colonies of N. n. were found in trees from March to October, individuals or pairs from June to October. There are no finds in trees during the winter season in our material. In the collections of the National Museum in Prague there are specimens labelled as found in hollow trees in Prague in January and in "winter"; as far as could be established, the respective colonies were discovered in the cavities of a mighty lime-tree and a beech; their number, 7 Tab. 2. Species of trees and location of holes occupied by N. noctula Tree Height of the entrance opening above ground, m Individuals or colonies species n I"2 3-4 5-6 7-8 9-10 11-12 13-14 15-16 X Individuals or colonies oak 27 3 8 5 6 2 1 2 6.2 both linden 8 3 3 2 — — — — _ 3.2 both alder 4 2 2 2.5 both willow 3 1 1 — — 1 — — — 4.8 both aspen 2 — 1 — 1 — — — — 5.5 colonies pine 2 — 1 — 1 — — — — 5.5 colonies ash 1 — — 1 — — — — — 5.5 colony elm 1 — — — — 1 — — — 9.0 colony apple 1 1 2.0 colony acacia 1 — — 1 — — — — — 5.5 colony maple 1 — — 1 — — — — — 5.0 colony walnut 1 1 2.0 colony spruce 1 1 1.5 ind. male total 53 12 16 10 8 4 1 — 2 5.1 both however, is not exactly known (G a i s 1 e r 1956, Hanak, Gaisler, Figala 1962). Hurka (1973) published a find of 1 male behind the bark of a big oak in Plzen on 4 Dec. There are some more reports about winter finds of N. n. in hollow trees by wood workers which, however, cannot be verified. Tab. 2 summarizes those finds in which the species of tree is known and the height of the entrance measured or estimated. Those bats were found in 13 species of trees, most frequent of them being the oak (50.9% of cases). In the table each hollow is counted only once; the average frequency of oak would be still increased if repeated finds in the same hollows were considered. The height of the entrance varied between 1 and 16 m, the accuracy of the estimate dropping with increasing height. The average height of the aperture is 5.1 m, but more than a half (53.8 %) of the occupied hollows was at the height of 1 to 4 m. As to a possible objection that the occupation of higher situated cavities can be registered less easily than in cavities situated lower, we should like to note that the summer colonies of N. n. utter loud sounds during daytime (c/., Stratmann 1968), and accarding to our measurements those utterances can be well heard up to the distance of 20 m. Thus, the probability of discovering higher situated hollows will not be substantially lower than in the hollows near ground. Exceptions are cold days particularly at the beginning and at the end of the period of occurrence of N. n. in trees; it is, however, improbable that at that very time the bats should seek higher situated hollows. Only single individuals can be missed, because t'hey utter sounds only under special circumstances. This methodological error, pertaining to many analogical population studies on bats, cannot be eliminated so far. Besides cavities in trees, N. n. was found in bat boxes hung up on trees in two stations: the Velky Tisy Pond, southern Bohemia, and the reserve Pod Trlinou, northern Moravia. In one case they were 4 males; in another case, 1 female; and in two cases 1 male, all of them being old individuals. The last mentioned case was a bat hidden between the back wall of the box and the tree trunk. In other tree shelters we did not find these bats, but Dr. Bal&t (Brno) informed us on his observation of 4 individuals behind loose back of a pine tree. Buildings. Finds in buildings fall into the period from August to May 8 (Fig. 1) so that, to a certain extent, they alternate with the finds in trees as far as time is concerned. The average number of individuals in one shelter was highest in December and January. Owing to the fact that shelters in buildings are often difficult to find, the data on the numbers and especially on the missing of N. n. must be judged carefully. For the time being, there is no evidence of the occurrence of suckling females and non-flying young, so that nursery colonies do not seem to occur in buildings. The size of colonies in buildings varies from 3 to 54 individuals, the average being 10.7 individuals per shelter, which can be lower than the actual value. The colonies found are of only two types: mixed and male. Mixed colonies (tt=13) were found from August to March, including all components of the population with the exception of non-flying juveniles. Male colonies (n = 10) are known only from the autumn season, from September to November. In further 6 colonies it was possible to find out neither the composition nor ftie number. Besides, the following were found in buildings: 1 male in three cases, 2 males in 4 cases and once 1 female; the finds are from April, May, September, October, and November, respectively. The highest number of finds (n = 29) is from prefabricated houses of a new housing estate in Prague, Zahradni Mesto (= Garden City). The flats there are equipped with an inbuilt recess in the kitchen connected with the outside through a narrow ventilation shaft, partitioned from inside by removable metal sheet. The bats hide most frequently in that shaft which is heated in winter. The occurrence was evidenced from the first to the ninth floor, most often on the third and on the sixth foors of houses. The total situation is shown in PI. I. Further evidence is from a family house at Roztoky near Prague where a colony of N. n. was hiding in the space between the ceiling of an unoccupied room and a flat roof with an entrance slot over the window. We got to the place only after the owner had closed the aperture; we obtained 2 emaciated males, the other individuals had probably perished. There is some evidence from the City of Brno. According to sound utterances a winter colony was hiding in a bay under the roof of a four-floored building of the Medical Faculty, at least from October to January for five successive winters. Another supposed colony inhabited a space above the ceiling of a room on the first floor of a house, according to unverified statements of local people "for the whole year". Evidence to this are droppings fallen out of the entrance and, besides, a male N. n. which flew into the room below the shelter in August. Besides, three times there were found individual males creeping on the ground in Brno: In October, January, and April. Another find of a winter colony was reported by Z. Rumler (Olomouc). The shelter was a narrow fissure between the wooden panelling and the wall of a window recess on the second floor of the castle building at Sternberk (northern Moravia). On 11 Feb., 1976, 413 individuals of Pipistrellus pipistrellus were found there, 1 female Vespertilio murinus which, in captivity, bore 2 young (for details c/., Rumler 1977) and 22 male and 10 female N. n. And, finally, we were supplied with two individuals of N. n. from villages: a dead male found in May in the hall of a farmer's house at Pozcfatin (the Ceskomoravska vrchovina Highland) and a living female which, in October, flew out of the stove of a recreation cottage at Celadna (the Beskydy Mountains). Large series of skeletal remains (thanatocolonies) were found in Prague and at Teplice (northern Bohemia). In building adaptations of the former abbey Na Slovanech in Prague, more than 200 skeletons of N. n. were found in 9 a bricked-up recess high in the attic turned towards the Vltava River. In the castle at Teplice, 166 skeletons of N. n. were discovered behind a metal sheet cover of the roof cornice. Both cases were most probably winter colonies. Besides we found or were given mummies of individual specimens which had lain in lofts or towers of various buildings. Other shelters. The most remarkable shelter, evidencing the occurrence of N n., is a crack in a vertical rock wall near Srbsko (central Bohemia), about 50 m above the water level of the Berounka River. According to sound utterances, a colony stays there at least from November to January, probably during the whole period of hibernation. The crack is not normally accessible, with the kind assistance of mountaineers, 2 females of N. n. were caught there in November. The situation is shown in PI. II (right). The last find was made by dr. K. Hudec (Brno) in a reed stand of Prostredni rybnik Pond (southern Moravia). In May, he found there a pregnant female hung up at a cluster of Typha stalks 1 m above the water level. The individual, which we could examine, did not show any symptoms of disease and was able to fly. Flight activity, hunting grounds N. n. belongs to bats having a very characteristic outline (G a i s 1 e r 1959) and way of flying. The characters distinguishing this species in the field were very instructively described by Klawitter & Vierhaus (1975). In this chapter we only summarize those observations when the determination of the species is beyond doubt; in several instances it was verified by shooting, catching from cavities or netting. Flying (hunting) individuals were observed from April to October. From April to September, we noted the beginning of the flight activity 16 times according to the moment of leaving the shelter, mostly a tree cavity. In all cases the bats flew out only after the astronomical sunset; the difference with respect to the sunset was + 2 to + 39 min., on the average + 19 min. In culminating summer, the bats seem to leave the shelter relatively later, but due to a small number of observations we do not consider this observation conclusive. In five cases, we observed hunting individuals of N. n. at daytime in full light: once in April (10 Apr., 1973), once in September (8 Sept., 1972) and three times in October (14 Oct., 1973, 13 and 19 Oct., 1974). In the last two cases mentioned uninterrupted observations were carried out from 12 to 14 h, using a 7X50 fieldglass. Throughout the observation, 10 to 12 individuals of N. n. were flying over the station (ponds and surrounding small woods near Lednice, southern Moravia). The height of the flight was estimated to be 20 m. During the night of 12/13 Oct. there had been a heavy rain; during the night of 18/19 Oct., the temperature was as low as —2 °C. During the observations there was sunny weather with temperatures of 4 to 8 °C. Due to the fact that the station is regularly visited by student excursions at the beginning of October, observations of this kind were also carried out in subsequent years but in spite of using stronger telescopes, the result was negative. Hunting individuals were observed in 35 stations. Most of them (62.9 %) fall to ponds and their immediate surroundings; they are both ponds situated in woods as well as those between fields and woods, in castle parks and at the outskirts of towns (Prague, Zabehlice — Krc). The majority of such observations can also be the result of the orientation of our research. We further observed N. 7i. flying over the edges of woods, wood clearings and roads, old lanes and 10 fields. Inside the built-up areas of the towns hunting N. n. have not been observed so far, although a systematic research of urban chiropterofaunae is in progress at present. In our study area, we metted bats in 9 stations (altogether 92 net-nights), but in only one station did we also get hold of N. n. The station is the pond Hlad near Studenec (the Ceskomoravska vrchovina Highland), where nets were put on the shores and in the surroundings of the pond. In 1968 and 1969, altogether 17 males and 2 females of N. n. were netted, always in the same net placed across the pond shore in such a way that 1 m of the net was on shore and 5 m above the water. In close surroundings there was a willow-tree with a hollow in which 3 times a male colony of N. n. was found, counting 6 to 9 individuals. At the time of successful netting, the cavity was mostly empty but banding proved that 3 males that had been hiding in the hollow were netted later on. Nettings and indirect observations showed that round the willow-tree there was a regular flight route at the height of 1 to 3 m above the water surface of the pound. In all other stations, hunting N. n. were observed at least 4 m above ground, as a rule 10 to 20 m high. Fig. 2. Reproductive organs of the adult male in June, dorsal side. Explanations: T, testis; E, epididymis; DD, ductus deferens; VS. vesicular gland; GA, ampullary gland; VU, vesica urinaria; SU, sinus urogenitalis; GB, bulbourethral gland; P, penis. Reproduction Description of reproductive organs. Since we did not find detailed descriptions of the morphology of genital organs of N. n. in literature, VU 11 we give their description here (Figs. 2, 3). The penis of the male is covered with hairs up to the tip, it is slightly flattened and pointed at the end, without a preputium. Inside the big glans penis there is a rod-like os penis. The scrotum is formed only in the functional period in dependence of the descent of the abdominally positioned testes and the increase in the volume of epididymides. Both halves of a sac-like scrotum are symmetrically adjacent to the root of the tail, resting firmly in the uropatagium, where they are easy to see ventrally. In the non-functional period only the caudae epididymidis are situated in that position. The testis is of the usual bean shape and the caput epididymidis is attached to its cranial part. The ductus epididymidis penetrates in numerous loops both the caput and the considerably long cauda epididymidis, which is attached to the caudal part of the testis. Ductus deferens is long and conspicuous when full of sperm in the functional period, similarly as cauda epididymidis. Before opening into the sinus urogenitalis, the former forms a pars glandularis ductus deferentis (ampulla ductus deferentis). In this part of the male reproductive tract, three anatomically and histologically distinct glands are situated dorsal of the bladder. The most conspicuous paired accessory glands are considered to be the vesicular glands; the smaller paired glands are considered to be the ampullary glands. The gland situated medially at the neck of the bladder, thus not seen in Fig. 2, is the prostate. Glandulae bulbourethrales (Cowper's glands) are situated more distally at the root of the penis, at the boundary of the pars pelvina and the pars penis. Fig. 3. Reproductive organs of the adult female after copulation in September, ventral side. Explanations: O, ovary; OV, oviductus; CU, cornus uteri; U, uterus; VU, vesica urinaria; V, vagina; VV, vulva and vaginal fissure. 12 The female has a transversal opening of the vulva, a shortened and flattened clitoris, an inconspicuous mons veneris; the labia are missing and the postand circumanal fold is indistinct. In the width of the vaginal fissure there is a clear difference between nulliparous and multiparous females; the border between the two groups is 1.5 mm. The vagina is an elongated thick-walled organ into whose lumen protrudes the cervix uteri. The uterus is of the bicornis type, formed by the corpus uteri and the cornua uterina. Both horns are morphologically as well as functionally equivalent and the length of either of them in non-pregnant adult females is 6.0 to 7.0 mm. The left and the right ovary are of the same size, in adult females they measure 1.5 mm on the average. The oviducts are short and coiled. The sperm is stored in uterine horns and the oviducts. An important additional criterion for judging whether the females take part in the reproduction process are the nipples, papillae mammae, placed near the lateral edge of the superficial part of musculus pectoralis major. In multiparous females the nipples are much cornified on the surface, elongated and hairless. In nulliparous females, both juvenile and adult, the nipples resemble those of males, with no signs of cornification, they are point-like and covered with fine hairs. Their length is difficult to measure, but the difference is maeroscopically wel perceptible. Age and sexual maturity. The criteria used for the determination of age are sumarized in Tabs. 3 and 4. The sexual activity of males is considered to be evidenced by the presence of sperm in the cauda epididymidis; of females, by the presence of sperm in the uterus, by pregnancy, or lactation. The individuals showing contrary signs are evaluated as sexually inactive. In calculating the age with the precision of months, one starts from the fact that the young are born within a short time interval at the break of June and July. The age Tab. 3. Age determination of males. Explanations: colour (of pelage) — 1, dark ochreous; 2, glossy ochreous; 3, light ochreous; ossification — 1, a broad zone of cartilago epiphysaria; 2, a narrow zone of c. epiphysaria; 3, ossification completed; eye = dry mass of right eye; abrasion of teeth — 1, none; 2, medium; 3, considerably worn; dentin layers = number of dentin layers on cross-section through right upper canine; (1), developing first layer Date Weight (g> Colour Sexual OssifiEye Abrasion of Dentin Age Weight (g> Colour activ. cation (mg) teeth layers years months 13 Jan. 24.0 2 + 3 1.28 1 (1) 0 6 13 Jan. 21.0 2 + 3 1.32 1 (1) 0 6 12 June 23.5 3 — 3 1.66 3 2 11 12 June 25.0 3 — 3 1.52 3 2 11 10 July 27.0 3 — 3 1.70 3 5 5 0 6 Aug. 22.0 1 — 1 1.14 1 0 1 6 Aug. 22.8 1 — 1 1.18 1 0 1 9 Aug. 22.5 1 — 1 1.17 1 0 1 10 Sept. 31.2 3 + 3 1.42 2 2 2 2 10 Oct. 27.0 3 + 3 1.68 3 3 3 25 Oct. 28.0 2 + 2 1.11 1 0 4 25 Oct. 26.0 2 + 2 1.20 1 8 4 25 Oct. 27.0 2 + 2 1.10 1 0 4 12 Nov. 26.0 2 + 2 1.18 1 0 4 12 Nov. 23.5 2 + 2 1.13 1 0 1 0 4 12 Nov. 24.0 2 1 + 2 1.20 1 0 0 4 18 Dec. 22.0 2 + 3 1.12 1 0 0 5 18 Dec. 19.0 2 3 1.09 1 a, 0 5 13 is rounded down; thus an individual caught on 12 Nov. which, according to the age criteria, is significantly younger than one year, has lived minimally from mid-July, and is therefore evaluated as 4 months old. From five age criteria used three can be ascertained when handling living individuals in the field. For the shortest time one can use the ossification of long bones which is finished at the age of about 4 months, like in other bats (Rybar 1971). The difference in the coloration of young individuals and the absence of dental abrasion can be observed approximately up to the end of the first hibernation, i. e., up to the age of about 8 months. From May onwards (material from April is missing), the coloration of young and old individuals is the same. Dental abrasion in the yearlings after the first hibernation can be smaller than in some old individuals, but this sign is not reliable. In Tab. 4 there are 6 females from May to July in which medium abrasion was found (degree 2). These females are of varying ages, from 10 months to almost 4 years. Also this conclusion corresponds to information concerning some other bats (cf., Barbour & Davis 1969). Thus, according to outer signs, the age of N. n. can be exactly determined at most till the spring of the year following the year of their births. Tab. 4. Age determination of females. Explanations as in Tab. 3. Date Weight CoSexual activ. OssifiEye Abrasion of teeth Dentin Age Date (g) lour Sexual activ. cation (mg) Abrasion of teeth layers years months 13 Jan. 21.0 2 T 3 1.11 1 (1) 0 6 9 March 19.0 2 + 3 1.16 1 (1) 0 8 12 May 30.0 3 + 3 1.58 2 4 3 10 10 12 May 26.5 3 + 3 1.58 2 3 2 10 10 12 May 25.0 3 + 3 1.43 2 2 1 10 10 12 May 23.0 3 + 3 1.10 2 1 0 10 10 12 May 27.0 3 + 3 1.42 2 3 2 1 10 0 10 July 22.5 3 — 3 1.36 2 1 2 1 10 0 10 July 28.5 3 + 3 1.52 3 3 3 0 0 6 Aug. 20.9 1 — 1 1.17 1 — 3 0 1 1 1 6 Aug. 22.4 1 — 1 1.20 1 0 0 1 1 1 9 Aug. 19.0 1 — 1 1.20 1 — 0 0 1 1 1 9 Aug. 29.0 1 — 1 1.01 1 — 0 0 1 2 2 10 Sept. 31.0 3 + 3 1.41 3 — 2 1 2 2 10 Sept. 28.0 3 _U 3 1.51 3 — 3 2 1 2 2 10 Sept. 30.0 3 + 3 1.45 3 — 3 2 2 2 10 Sept. 28.0 3 + 3 1.35 3 — 1 1 0 2 2 10 Sept. 33.0 3 + 3 1.31 3 — 1 1 0 2 2 2 10 Sept. 28.0 2 + 2 1.19 1 — 1 1 0 2 2 2 10 Sept. 25.3 2 + 2 1.20 1 0 0 0 2 2 2 13 Oct. 16.0 2 + 2 1.28 1 — 0 0 3 25 Oct. 26.0 2 + 2 1.35 1 — 0 4 12 Nov. 25.0 2 + 2 1.19 1 — 0 4 8 Dec. 24.0 2 + 3 1.21 1 0 5 The remaining two criteria can only be used in the laboratory at autopsy. As far as we are informed the mass of the lens has hitherto been used to determine the age groups only in Tadarida brasiliensis (Perry & Herreid 1969); this sign alone does not enable one to recognize the age in years. The number of the increment layers of dentin was first used by Christian (1956) in Eptesicus fuscus and later by Klevezal' & Klejnenberg (1967) in several species of the fauna of the U.S.S.R., including N.n. The latter 14 two authors published a number of microphotographs of transversal as well as longitudinal sections of the canines of N. n. of different ages whose minimum age had been known in advance according to the results of banding. According to further information (Rachmatulina, in litt.), the method was verified by Soviet researchers on ample material and it was established that the number of dentin layers gave the age in years. In our material (Tabs. 3, 4) there are 4 specimens in which no increment layer of dentin was found. According to all criteria and the date of catching, their maximum age is about 4 months. In further 5 specimens one forming layer of dentin was found. Their age — again according to all criteria — is 5 to 8 months. One distinct layer of dentin was found in 2 females; according to the date of catch, the first of them is 10, the other 12 months old. The remaining 7 specimens have 2 to 5 dentin layers. According to other signs their ages cannot be estimated, with one exception, viz., that they are older than 1 year. Besides the material used for the study of reproduction we obtained an individual whose minimum age was known thanks to banding: the male had been banded on 23 Jun., 1969 and found recently dead on 11 May, 1971. According to the date of banding its age must have been at least 11 months, but most probably 1 year, so that at the time of death its age was almost 3 years. In the section through its canine there are three distinct increment layers of dentin. All the above fact prove that the number of layers of dentin is, for the time being, the most reliable criterion of bat age. That is why we used it to verify the last criterion, the dry mass of the eye, as shown in Fig. 4. The graph includes also specimens in which tooth sections have not been made, but whose age is unquestionable — altogether young animals. The graph shows that the scatter of the values of dry eye mass is great, particularly with the youngest specimens, but the overall trend of mass increment is in correlation with age. This sign can, therefore, be used as an auxiliary one in specimens older than one year, where tooth sections could not be made. Since the combination of all criteria enables one to determine the age of the whole sample with great probability, we have reliable data available to follow Fig. 4 (left). Correlation between the mass of dry eye (ordinate) and age determined according to the date of collection and the number of dentin layers on section through Ct (abscissa). Fig. 5 (right). Sexual activity of males in the course of the year according to the mass of the testes, epididymides, and vesicular glands (= semin. vesicle). In comparison with Tab. 3 one old malq obtained later was added in January. 15 (Mislin & Vischer 1942, Butovskij 1974, and others), in rock cracks (Barbu & Sin 1968) as well as in hollow trees (Sluiter & al. 1973) is similar. The corresponding data concerning the microclimate of the summer shelters are not available so far. The choice of summer shelters is certainly influenced by the requirements for an environment offering suitable and sufficient food supply. That is why hollow trees remain the main shelters of summer colonies, particularly for nursery colonies which are the most conservative component of populations. On the whole, N. n. has well adapted itself to the civilization pressure and, in places, it gradually turns towards the hemisynanthropic way of life. Even if the changes in abundance of the species in time have not been reliably recorded, we can, with great probability, consider N. rt. to be one of the least endangered bats of Europe. Hunting activity Already Blasius, Koch, Altum, Kolenati, and other exact observers of the 19th century found that N.n. flies quickly with a comparatively si raight flight, mostly at the level of tree crowns or higher. The flight apparatuses fully adapted to that (Gaisler 1959). Besides general information in different monographs (Ryberg 1947, Kuz'akin 1950, N a t u s ch k e 1960, • j ? k™ore 1963) its ways of flying and the hunting activity are described in detad by Klawitter & Vierhaus (1975). According to those authors, the activity of N. n. starts earlier than in most of the other European bats, as a rule several minutes to 1 hr after sunset. In spring and in autumn, the flight activity starts earlier than in high summer, and at the end of September and in October it can begin even before sunset. The flight speed varies from 30 to 54 km per hr (16 measurements). In the wood environment N.n. hunts at the height of about 15 m; above open terrain such as fields or lakes, 20 to 70 m; on the average, 31 m (28 observations). Under certain circumstances, such as m hunting low-flying prey or in strong wind, these bats fly lower. This corresponds with the observations by Cranbrook & Barrett (1956) who mist-netted low flying N.n., taking house crickets as they flew from a municipal dump. A typical hunting ground of N.n. is in the wood, above all, in vvood clearings and on wood edges. Owing to their great action radius, these bats can be met as far as 6 km from the nearest wood over fields, water areas, or in outskirts of villages. Inside major towns, N. n. does not hunt even if its shelters are situated there. Our findings fully agree with the information obtained by the above-mentioned authors. Of all European bats, flying N.n. were comparatively most frequently observed during daytime. Due to the fact that sometimes it was more individuals and most cases like that were observed in late summer and in autumn some authors considered this to be correlated with migration. Although already Lohrl (19oo) was looking for a more probable explanation of this activity m hunting for food, reports on „daytime migrations" of N.n. have appeared even recently (Reichhc.If 1976). The problem was dealt with in detail by Krzanowski (1959) who listed a large number of observations but did not arrive at an unambiguous explanation. If the cause of the daytime activity were cold nights or otherwise unfavourable weather, there would be far mo e observations of this kind. On the ot'her hand, it is conspicuous that a Seat majority of the observations were made during clear sunny days. 28 In our opinion, it is necessary to differentiate according to the season of the year and the behaviour of the bats flying at daytime. The daytime flight activity in winter or in mid-summer can be due to the disturbance of individuals in their shelters, or possibly due to abnormal behaviour of aberrant (e. g., sick) individuals. On the other hand, flights on clear autumn or spring days, when individuals or whole groups are on the wing regularly and for a long time over a certain stretch, must be motivated by food. Such behaviour has been observed several times even with Pipistrellus pipistrellus and Eptesicus serotinus (Gaisler, unpublished). In no case do we see any reason why bats like N. n. should, in movements or migrations, fly at daytime when they are exposed to much greater danger from predators than at night. The timing of the night activity has not been exactly investigated; preliminarily it seems that under normal circumstances N. n. hunts mainly during the first half of the night. In the aspect of the whole year, the activity lasts approximately from April to the beginning of October. There are, of course, local differences. In the central zone of the European part of the U.S.S.R., populations of this species which are migratory stay from mid-April to late September (Kuz'akin 1950, Pan'utin 1963). In the southern regions of the U.S.S.R., individuals were observed on the wing from March to late October. In southwestern Germany, according to Roer (1977), N. n. search for winter shelters as late as from mid-November. Reproduction For learning about the population ecology of the species, the most important information concerns the time of sexual maturity, litter size, and the percentage of reproducing females. Already Bels (1952) noticed that "several one-year old females are found among the pregnant ones" and some of them are "very likely pregnant". He is, however, of the opinion that "it does not yet justify the conclusion that the females reach maturity after one year". Cranbrook & Barrett (1965) found, on the basis of nettings carried out for several years, that "five certainly and six probably out of fourteen females reached sexual maturity in the year of their birth". As for the males, they judge that they do not mature during the first year. Kleiman (1969), K lei man & Racey (1969), Racey (1970) and Racey & Kleiman (1970) published the results of their investigations on the reproduction of the species under laboratory conditions. Two out of five female N. n. born in captivity were observed to mate when they were about 3 months old, and both gave birth when one year old. In the males the first spermatogenesis takes place probably as late as at the age of 1 year, according to the above authors. From the papers by Pan'utin (1963, 1970), though they are in fact only abstracts of unpublished voluminous data, it is possible to judge that all yearling females take part in the reproduction process and bear young at the age of one year. The author does not give his opinion on the sexual maturation in males. These data prove that N. n. females can mature during the first year of their life., Basing on our own results, we believe that in fact all yearling females d o mature, but only part of them give birth to and bring up the young at the age of one year. To estimate the percentage of females giving birth to young at that time the available samples are too small. This fraction may constitute 40 to 100%, being smaller in England than in Central and 29 Eastern Europe. Still greater differences appear in judging the males, as, unlike British authors, we found conclusive signs of sexual activity in most yearling males studied in this respect. In spite of this fact, young males do not participate in the reproduction and remain in juvenile colonies. For the preservation of the population their participation is not necessary, since old males mate with several females each, as will be described when discussing the population. In one litter there are 1 to 2, rarely even 3 young (R y b erg 1947). According to Blackmore (1963) in England, litters with 1 young are the rule, whereas on the continent it is 2 young. In the material of Kleiman & Racey (1969) one parous female gives birth to an average of 1.3 young; one nulliparous female, 1.0 young. Sluiter & Heerdt (1966) published, among others, the results of their sampling nursery colonies in the latter half of July when the young already flew out of the cavities. In that sample (n = 155) there fall 1.2 young to 1 female. Even though the number can be reduced due to the fact that not all young flew out yet, this number obtained in Holland approaches the data from England. Pan'utin (1963, 1970) evaluated several samples: out of 15 females of unknown age, 60% had two and 40% one embryo; out of 9 yearling females, 22 % had two and 78 % one embryo; and 4 old females had 2 embryos each. In nurseries he found 1.8 to 1.9 young per 1 female. Our data, particularly the estimate according to the big sample from nurseries, approach most closely those of Pan'utin. As for the average number of reproducing females, the results obtained in England also differ from the observations in the Soviet Union and Czechoslovakia. Out of 58 females — mostly of unknown age — examined by Cranbrook & Barrett (1965), only 83 % participated in the reproduction in the particular year. In the material obtained by Racey & Kleiman (1969), the percentage of females bearing the young was even lower: 44% in parous and 25 % in nulliparous females, but the results can be distorted due to laboratory conditions. On the other hand, all females (n = 26) dissected by Pan' utin (1963, 1970). at the time of expected pregnancy were pregnant. Besides a small sample of females dissected in May (n = 5, all of them pregnant), we can use a big sample of living females examined in June, when the proof of pregnancy (according to their body mass and by the palpation method) or lactation (according to the state of their nipples) is doubtless. In this sample (n = 172), 90% of females were pregnant or lactant, 10% females without any signs of pregnancy or lactation. The females of the second group were nulliparous, and thus with greatest probability former yearlings. The first group must have also included a part of former yearling females, but their reliable distinguishing at that time is no longer possible. From the above analysis it is evident that the reproductive rate is lowest in England and probably also in the Netherlands and highest in the central belt of the European part of the U.S.S.R. (Pan'utin's material comes mostly from the Voronez State Reserve). The reproductive rate of the population of N. n. studied in Czechoslovakia lies between the two extremes, but it approaches more the situation in the U.S.S.R. On the whole, the reproductive rate is high as compared to many other bat species: quick sexual maturation, frequent births of twins, and a high percentage of reproducing females. This phenomenon is certainly in correlation with relatively high losses resulting from hibernation in little protected shelters and from great migration activity of the species. 30 Although no detailed work has been done on the reproduction cycle of N. n., there is a lot of information concerning the partial aspects of this process, particularly in the papers quoted above. If those data are supplemented by our own results, for the time being the most complete, we can reconstruct the whole reproduction cycle of the two sexes. The resulting picture is given in Fig. 9. • • • copulation ? epididymis vesicular gland testis copulation ? ovulation estrus subestrus subostrus '/Graaf. follicla '<'////// Graaf. follicle lactation / corpus luteum 1 j'f'm'a'm'j'j'a's'o'n'd1 Fig. 9. A schematic representation of the reproductive cycle of sexually mature males and females of N. noctula. The dashed parts represent the period of hibernation. Population Most^ data published on the composition of the population concern the sex ratio. The data on the sex ratio among grown-up individuals in the summer season are biased in favour of females. More realistic are data concerning the sex-ratio in the juveniles. Bels (1952) found, in a large sample (n = 384), 44 /0 of males and 56% of females. Sluiter & Heerdt (1966) found, in a July sample (n = 85), 43.5% of males and 56.5% of females; in an August sample (n —94), 49% males and 51% females; in September (71 = 23), 91% males and 9 0 females. Kleiman (1969) states, among young born in captivity (n — 19), 47% males and 53% females. The hitherto greatest sample of young was obtained by Pan'utin (1970), (n = 2,028), of which males constituted 51 /o and females 49%. These data, together with ours, show that the sex ratio among the young is more or less balanced. The increasing number of males in the juvenile colonies in the course of summer, found out concordantly by the Dutch authors and by ourselves, will be explained below. 31 From the summer material of grown-up individuals, particularly the sample of Cranbrook & Barrett (1965) is worth mentioning; it was not obtained by catching the bats from their shelters, but through mist-netting. As stated by the authors themselves, there is no reason why one sex should be mist-netted more frequently than the other. In the whole sample (n = 319), obtained in the course of three successive growing seasons, the males constituted 48 %, the females 52 %. This more or less balanced and expected sex ratio did not, however, concern the samples of the successive months: in June, and to a lesser extent also in July, females prevailed in the mist-netted sample; in August the sex ratio was balanced; and in September, but particularly in October the males prevailed. The authors explain this phenomenon by the fact that in the station most females appeared earlier in summer than males but, on the other hand, the males left the station later in autumn. For our consideration it is essential to state that the overall sample of the above authors very much approaches the ideal ratio, thus supporting the hypothesis of the even representation of males and females among grown-up individuals in the summer populations. Our small sample of mist-netted individuals (n = 19), in which the males predominate strongly, is distorted by the existence in the neighbourhood of a cavity used as a shelter by a male colony. Unlike the samples obtained from summer shelters the samples from the hibernacula should theoretically involve both sexes evenly, as the colonial roosting and formation of dense clusters is necessary for both sexes to survive in little protected places (S1 u i t e r & al. 1973). Lohrl (1936) caught 55 individuals from a shelter in a building, among which there were 53 % males and 47 % females. B e 1 s (1952) quotes a sample from a tree cavity (n = 34) with 71% males and 29% females. Kepka (1962) found, in two colonies in tree hollows (n = 120), 64 % males and 36 % females. It is interesting to note that one of the colonies contained admixed individuals of Pipistrellus pipistrellus, as in a colony mentioned by us (Sternberk castle) where, on the other hand, a smaller colony of N. n. was admixed in a big colony of P. pipistrellus. Gauckler & Kraus (1966) found, in a colony in a building (n = 53), 57 % males and 43% females, and in a colony situated in a hollow tree (n=31), 58 % males and 42 % females. Heerdt & Sluiter (1965) and S 1 u i t e r & Heerdt (1966) obtained material from four hibernating quarters in hollow trees (n = 209), in which the males constituted 59 % and the females 41 %. And, finally, Barbu & Sin (1968) in a sample from a rock crack (n = 228) found 36 % males and 64 % females. The survey shows that males predominate in all samples from Western and Central Europe, irrespective of whether they come from trees or buildings. This opposes the possibility conjectured by us in analysing our own material, viz., that the females should prefer to hibernate in trees, males in buildings. Another possiblity, viz., that part of the females — or more females than males — migrate for hibernation to the south, would be supported by the sex ratio found in Rumania (Barbu & Sin 1968). In this connection it is interesting to note that the Soviet authors (Strelkov 1969, Pan'utin 1970) believe that the regions at the northern limit of the range of the species, where hibernation is not possible, are populated mainly by females, whereas most of the males stay more to the south. This would be supported by the fact that long migration flights, 300 to 1,600 km, were evidenced chiefly in females, even if it is necessary to take into consideration the possible distortion due to the majority of females in the sample of banded individuals. Per analogiam with 32 ttTESrt rount of evidence °f Will be described in detail relations m the late summer samples, it frequently sits straight in the cavitv eniron sounds, the male most shelter. The whole behaviour is that of If Y he flieS around the sexually active females. y hlS ***** and lurin§ up to several weeks, whlreas the femat, 4 T*® f°r a long time' Besides the "singing"'malTaLe ^ * to 2 *** ing of 1 male and 1 to 20 females usuallv 4 to I f i o shelters consistsumnter, but in *53^ « fcjssta I are inseminated already at the beeinninJ„f c £®males °f the a conspicuous decrease of iuvenile fPmai September. This corresponds to ber, found concordantly by theDutehaut^ Tl aggregations in Septem^ that in Western anTcenSlEurnn^hI , u by US' We believe mating quarters, p^babT however hT * • th"t ^ m°TC <° th* females. ' some delay in comparison with old individuafs1 frbm t ™ * °f ®™p all authors, the explanations offered bv fhemWmS i-«S°n W3S establlshed by - - - they were born and disperse over a The females, however, return every year to the region where they were bom. The total number of grown-up males and females constituting the subpopulation studied is probably the same according to the Dutch authors. There is no conclusive evidence of whether or not the members of one sex are more bound to the place of their birth than those of the other sex; again, this question remains open to dispute. The only known fact is that the two sexes often change their summer shelters. According to Pan'u tin (1970), 25% of the young perish in the course of the first three weeks, and till autumn there survives approximately 1 young per 1 female. The yearly mortality rate of females is 46%. These numbers without any specification of the initial data do not have a significant character, especially if we know how the composition of the samples can be distorted when compared to the actual population structure in the respective area. Owing to an extraordinarily low percentage of recoveries of banded individuals of N. n., none of the authors has a representative sample to evaluate the age structure of the population. That is why the problem of mortality rate in the different age classes and the drawing up of life tables must be left for a later elaboration. Only then will it be possible to judge the population turnover and the production of the species. Yet there is one important production parameter available, viz., the population density. It is the more valuable, because the estimates of population densityhave so far been carried out in only a few species and areas concerning the European bats (cf., Gaisler 1975). In the case of N. n. there exist two more data besides that of this paper (Tab. 7). Pan'utin (1970) gives the population density of N. n. in the Voronez State Reserve on an experimental plot of 3.5 km2; in July there lived 75 to 80 individuals per km2, or about 0.77 ind. per ha. The author notes that the species is particularly abundant in that area, the average population density in the whole reserve being lower. Details of the calculation are not given. Pan'utin's estimate does not differ from ours as to the order of magnitude, and therefore one can assume that in the Voron£3: region the species is approximately as numerous as in southern Bohemia in summer. The values between 0.3 to 0.7 ind. per ha will probably hold for optimum habitats of the species in general. Still higher a value is given by Gaisler (FIBRC Abstracts, Nairobi 1975) who made an estimate on an area of 40 ha near the town of Sibiu in Rumania. On the area several colonies were found which were not disturbed and the estimate was made according to the number of flying individuals with concurrent netting. The observations were facilitated by the fact that the area is a camping site illuminated at night. The resulting value of 2 ind. per ha shows that in the locality there was an extraordinary concentration of the species in late summer (observed from 28 Aug. to 2 Sept.). Irrespective of a very suitable habitat — an old oak stand — the reason for this high concentration of N. n. can be seen in local movements and/or the abundance of food. In localities where N. n. reaches relatively high population densities it is often the most frequent bat. Bats of the genus Pipistrellus prevail in only some regions (Lichacev 1961, Pan'utin 1970). Due to the fact that N.n. is considerably bigger than other frequent species of forest bats, it contributes a great deal to the biomass of local bat communities. 34 Summary The ecology of N. n. was investigated in the area lying between 50°15' to 48°40' n. lat. and 13°10' to 18°15' e. long, on the basis of a sample of 1,383 individuals caught and further observations specified in the Methods. In describing the individual components of the populations the following terms are used: juveniles = young bats from the birth to the end of September; grown-ups = all individuals older than juveniles; yearlings = individuals younger than 1 year in the period from October to May; the old = all individuals older than yearlings. The main shelters of the species studied are in hollow trees and buildings. Colonies in hollow trees were found from March to October, individuals from June to October. The largest number of members of the summer colony was 53. Five types of colonies were found, out of which nursery colonies are bound exclusively to this type of shelter. Hibernating colonies were not found in trees in the period of investigation (1955 through 1977), but they are evidenced by earlier material. The preference of different tree species is discussed on the basis of the authors' own material and that of literary data; in the region under investigation, oak was the most frequently inhabited tree species. Entrances to cavities were 1 to 16 m high, most frequently up to 5 m. Colonies and individuals in buildings were found throughout the year with the exception of June and July. Winter colonies were always found in towns; most of the evidence is from prefabricated houses in Prague. Besides, two thanatocolonies were obtained numbering 200 and 166 mummies or skeletons respectively. One winter colony was found in a crack of a rock wall. In the discussion it is hypothesized that besides hollow trees, rock craks are the original shelters of the species. The population of buildings and further circumstances show that N. n. succeeded to adapt itself to the civilization changes of the environment. The hunting grounds of the species are lowland woods up to the elevation of 500 m surroundings of ponds, fields among woods, parks, lanes, and vicinity of human settlements. The flight activity starts 2 to 40 min. after sunset; in five cases hunting individuals were observed at daytime (April, September, October). N. n. flies swiftly, as a rule lu to 20 m high. The course of the reproduction process is expressed in Fig. 9. The comparison with SS?at""!1slY>ws that the reproductive rate of the species in Western Europe (England, Holland) is lower than in Central and Eastern Europe. In the region under investigation, f,f™a'eS, m?Sl m*l6S ™a*ure as earIy 38 at the a&e °f 3 months, thus passing from tw> • ?. ItT* } stage. Although all yearling females mate, only part of if sPvS.nl S brv!n£, UP,the y°Ung at the age of 1 yearYearling males - even 1 7 P biyT d° n.0t teke part in the mating. All females older than prown ,, -f f 1^% mid"June to early July. The average number of young per 1 I year'bear twins. t0 *** th3t * prevailinS number of females older than The composition of the population samples obtained by catching from shelters does not correspond to the actual population structure owing to different probability of ascertaining the various population components which, besides, change in the course of the year The available information enables us, however, to explain the apparent discrepancies and to JpW h i the Sltuatl°n as follows: The sex ratio at birth is 1 :1 and remains approximaba a"Ced also among yearlings and old individuals. In summer, most of the old males and Z fpUll0"1 aggregations of the females. Nursery colonies disintegrate in August and the females move to the mating quarters of old males which, at that time show territorial behaviour. After the disintegration of nurseries the young bats form juvenile moveietn mar t fS Predommate gradually, since even females born in that year imamW T , Quarters in late August and in September. The only fact that cannot be unambiguously explained is the predominance of males in all samples from hibernating quarters in Western and Central Europe; the possible causes are discussed. in fhn nnf ^ estimated basing on catches from summer shelters on two plots in the optimum habitat. The average value is 0.32 ind. per ha; details are given in Tab Os L'nTci npr hexpwct liat in.?ptimum habitats within the species' range there live view^of production Ecology!* "" m°S' "h""'8"' ^ *»» «» point"* 35 Pe3K)Me n) B TeqeHMe 20 noKasajio, to kojioHhh JeHMeSMMOBKaKoJoHuS HaxOAaTC51 B fly™ sepeBBeB, r*e iipohcxoahx pasMHO3tot BMJ1 OTMeneH KnvrnMM fn^to I13BeCTHa JIMLUb no crrapuM MaxepwajiaM. B SflaHHax MHOJKeHIIP N n OT '/-» MCK®OHeHJieM MKDHa M HIOJIS; He yCXaHOBJICHO pauCoScTBeHHbie naiiHMpHn "^CTax" °^Ha shmhjis kojiohmh Qbijia Haw^eHa B Tpemwue CKaji&T. pa3JIMHHbIX rnvnn R ' nrJf ^ J1HTepaJyPHble' n03B0JIHK>T Cflejiaxb BblBOfl 06 OTHOUieHMM OXO?™HX wa^K»TlSS?wK **5**°*" M 00 OCOSeHHOCTBX HX ITOBe^eHMB Ha flGCTHiaioT nojiopovi qnPn^ p^bHOM EBP°ne Bee caMKw m Sojisihhhctbo caMpoB Kf. n. S Tca^oTcrnnZ rnJ^ ** MeCHIiaM' TecPa3y n°cae okoHHaHHa pocra y 60^- Ha B^oe™ . f6oJI^ajtOCb no 2 «eTCHbima M opeflHee hhcjio flexeHbime* »e cooTHOiiieHHe nojmR m' C00TH0uieHIie.n0J10B cpe^H HOEopoxmeHHbix 1:1, Taicoe cs Ha cbct c nojiORwm,T ma' eMy MHeHW1°< coxpaHaexcx m no3«e. Mojioahsk noxBJiaeTflHHKO.M pacnanaioxm r HaHajia M10JIa M kojiohhm otpoawbiiimx caMOK c moji6BG3pacT0B a avnjra BpeMH rOHa nePBbIMM nepecejunoxcx caMKH cxapiuwx caMijbi b pa3MH0xceHnn Lm PPMT0PMajn>Hbix caiwpoB, a 33t©m cerojiexKH. Cerojiexicra ZJn?15™10/' HecMOTpx Ha nojiOBoapejiocxb, 11 b K>BexiOKHnaioT aeTHMf^A'fip^rMifro npeoSjiaaaiGT naA cajviKaMH. B okxh6Pc N. n. w nempajibHOH EBDonTcaviifu 2 ™ anpejI51" Bo Bcex CJiyiia»x shmoh b 3ana«HOfi ifreHOMena flHCKvccifnmTM rw npeo6jia«aioT Ha« caMKaMw, B03M0>KHbre npuHHHbk sxbr'o Omotohc cocxaBJiaex n 3? njI0TH°CTb nonyjiamm, MccjieflOBaHHaa b onxMMajibHOlv* npMcno%Cm,TnJf^ M W ra" XoTK Pbma* Bweprama jiecnoft Bm, oh cy^eji TponHbiM. 10 ^KBIIJI1I3ai^MM M B HCKOTopbix cjiynaax BBjiaexcx noaycMiiaHLiterature ABELENCEV, V. I., PIDOPLlCKO, I. T., .. POPOV, B. M., i956: Fauna Ukrainy, 1, Ssavci [Fauna of the Ukraine, 1, Mamirials]. Kijiv, 448 pp. (in Russian). BARBOUR, R. W. & DAVIS, W. H., 1969: Bats of America. Lexington, 286 pp. BARBU, P. & SIN, G., 1968: Observa[ii asupra hibernarii speciei Nyctalus noctula (Schreber, 1774) in faleza lacului Razelmcapul Dolo?man - Dobrogea. Stud. cere. Biol., Zool., 20: 291—297 (in Roumanian). BARRETT-HAMILTON, G. E. H., 1910: A history of British mammals, 1, Bats. * London, 450 pp. BELS, L., 1952: Fifteen years of bat banding in the Netherlands. Publ. Nat. Gen. Limburg, 5: 1—99. BIANCHI, C„ CAPORIACCO, L. di, MASSERA, M. G., VALLE A., 1949: Raccolte faunistiche della grotta della Spipola (Bologna). Com. Pont. Acad. Sc., 13: 493-527. BLACKMORE, M., 1963: Bats — noctule, Leisler's and serotine. London, 27 pp. BOJSEBAJEV, K., 1966: Ryzaja veCernica * (Nyctalus noctula L.) v orechovych lesach Juinoj Kirgizii [The noctule bat in the walnut forests of southern Kirghiz]. Zool. 2., 45: 1583 (in Russian). BRAAKSMA, S. & WIJNGAARDEN, A. van, 1969: Winterverblijfplaats van rosse 3* vleermuizen in een kasteel. Lev. Nat., 72 - 185—188 (in Dutch). BUTOVSKIJ, P. M., 1974: Temperaturnyje uslovija zimovki ryzich vecernic (Nyctalas noctula Schreb.) [Temperature conditions of hibernation in the noctule bats]. Mat. I. vses. sov. Chiroptera, Leningrad: 108-109 (in Russian). . CHRISTIAN, J. J., 1956: The natural history of a summer aggregation of the big brown bat, Eptesicus fuscus fuscus. Amer„ Midi. Nat., 55 : 66-95. CRANBROOK Earl of & BARRET, H. G., 1965: Observations on noctule bats (Nycta^ lus noctula) captured while feeding. Proc. Zool. Soc. London, 144: 1—24. DUMITRESCU, M„ ORGHIDAN, T., TANASACHI, J., 1955: Dou3 descoperiri intere— sante in pestera Cioclovina cu Ap3. Bull st. Acad. RPR, Biol., 7: 359-368 (in Roumanian). GAISLER, J., 1956: Faunistische Ubersicht der tschechoslowakischen Fledermause Ochr. Pfir., 11: 161-169. GAISLER, J., 1959: Beitrag zum vergleichenden Studium des Flugapparates der Fledermause (Microchiroptera). Zool. Listy, 8: 37-62. GAISLER, J., 1966: A tentative ecological classification of colonies of the European bats. Lynx, 6: 35-39. GAISLER, J., 1975: A quantitative study,of some populations .of bats in Czechoslovakia (Mammalia: Chiroptera). Acta Sc. Nat. Brno, 9 (5): 1-44. GAISLER, J. & HANAK, V., 1969: Ergebnisse der zwanzigjahrigen Beringung von Fledermausen (Chiroptera) in der Tschechoslowakei: 1948-1967. Acta Sc. Nat. Brno 3 (5): 1-33. • ; ' GAISLER, J. & KLlMA, M., 1968: Das Geschlechterverhaltnis bei Feten und Jungen einiger Fledermausarten. Z. Saug., 33: 352-357. GAUCKLER, A. & KAUS, M., ,1966: Winterbeobachtungen am Abendsegler (Nyctalus noctula Schreber, 1774). Saug. Mitt. 14: 22-27. ' GILES,,R. H. (editor), 1971: Wildlife mana- . gement techniques. Washington, 633 pp. HAENSEL, J., 1967: Notizen liber 1963—1966 insbesondere in Berlin aufgefundene Fledermause. Milu, 2 : 313-322. HANAK, V., 1967: Verzeichnis der Saugetiere der Tschechoslowakei. Saug. Mitt 15: 193-221. HANAK, V., GAISLER, J., FIGALA, J., 1962: Results of bat-banding in Czechoslo-' vakia, 1948—1960. Acta Univ. Carol., Biol., 1962 (1): 9-87. HEERDT, P. F. van & SLUITER, J. W., 1965: Notes on the distribution and behaviour of the noctule bat (Nyctalus noctula) in the Netherlands. Mammalia, 29: 463-477. HURKA, L., 1973: Ergebnisse der Fledermausberingung in Westbohmen in Jahren 1959-1972 mit Bemerkungen zum Vorkommen, Okologie und Ektoparasiten der Fledermause. Sbor. Zapces. muz. Plzeh, Pfir., 9: 1—84. KEPKA, O., 1962: Uber zwei Winterschlafgemeinschaften des GroBen Abendseglers Nyctalus noctula Schreb., in Graz. Mitt'. Natwiss. Ver. Steiermark, 92 : 42—43. KLAWITTER, J. & VIERHAUS, H.' 1975* Feldkennzeichen fliegender Abendsegler Nyctalus noctula (Schreber, 1774) und Breitflugelfledermause, Eptesicus serotinus (Schreber, 1774). Saug. Mitt., 23: 212-222. KLEIMAN, D., 1969: Maternal care, growth rate, and development in the noctule (Nyctalus noctula), pipistrelle (Pipistrellus pipistrellus), and serotine (Eptesicus serotinus) bats. J. zool. London, 157: 187—211 KLEIMAN, D. & RACEY, P. A., 1969: Observations of noctule bats (Nyctalus noctula) breeding in captivity. Lynx 10• 6577. KLEVEZAL', G. A. & KLEJNENBERG, s! E., 1967: Opredelenije vozrasta mlekopitajuSdich po sloistym strukturam zubov i kosti [Age determination of mammals by layered structure in teeth and bone]. Moskva, 144 pp. (in Russian). KRZANOWSKI, A., 1959: Daytime activity of Nyctalus noctula. Acta Ther., 2: .2'83284. KUZ'AKIN, A. P., 1950: Letucije my§i [Bats], Moskva, 443 pp. (in Russian). LICHACEV, G. N., 1961: Ispol'zovanije .letucimi mysami ptic'jich iskusstv^nnych gnezdovij [The use of bird nest boxes by bats]. Tr. Prioksk-terr. gos. zap., 3: 85^156 (in Russian). LOHRL, H., 1936: Der Winterschlaf von Nyctalus noctula Schreb. auf Grund von Beobachtungen am Winterschlafplatz. Z. Morph. Okol. Tiere, 32: 47-66. LOHRL, H., 1955: Mannchengesellschaften und Quartierwechsel bei Fledermausen. Saug. Mitt., 3: 103-104. LOHRL, H., 1955: Ziehende Fledermause. Saug. Mitt., 3: 128. MEISE, W., 1951: Der Abendsegler. Neue Brehm-Biich., 42: 1—42. MISLIN, H. & VISCHER, Lv 1942: Zur Biologie der Chiroptera II: Die Temperaturregulation der iiberwinternden Nyctalus noctula Schreb. Verh. Schw. Natf. Ges. 1942: 131-133. NATUSCHKE, G., 1960: Heimische Fledermause. Neue Brehm-Biich., 269: 1-146. PAN'UTIN, K. K., 1963: O razmnozenii ryzej vefernicy [On the reproduction in the noctule bat], U6. zap. Mosk. obi. ped. inst., Zool., 126: 63—66 (in Russian). PAN'UTIN, K. K., 1970: Ekologija letu£ich my§ej v lesnych land§aftach [Ecology of bats in forest regions]. Avtoref. kand. dis. Moskva: 1-24 (in Russian). PERRY, A. E. & HERREID, C. F., 1969: Comparison of the tooth-wear and lensweight methods of age determination in the guano bat, Tadarida brasiliensis mexicana. J. Mammal., 50: 357-360. RACEY, P. A., 1970: The breeding, care and management of vespertilionid bats in the laboratory. Lab. Anim., 4: 171-183 RACEY, P. A. & KLEIMAN, D. G., 1970: Maintenance and breeding in captivity of some vespertilionid bats, with special reference to the noctule. Int. Zoo Yearb. 1065—70. RACEY, P. A. & TAM, W. H., 1974: Reproduction in male Pipistrellus pipistrellus (Mammalia: Chiroptera). J. Zool. London 172: 101-422. REICHHOLF, J., 1976: Ein Zugstau (?) beim Abendsegler, Nyctalus noctula (Schreber 1774), im Spatherbst. 1974. Saug. Mitt., 24: SO. ROER, H., 1971: Weitere Ergebnisse und Aufgaben der Fledermausberingung in Europa. Decheniana-Beih., 18: 121-144. • ROER, H:, 1977:. Uber Herbstwanderungen • und Zeitpunkt des Aufsuchens der .Uber37 winterungsquartiere beim Abendsegler, Nyctalus noctula (Schreber, 1774), in Mitteleuropa. Saug. Mitt., 25: 225-228. RUMLER, Z., 1977: Predbezna informace o okolnostech nalezu samice netopyra pestreho (Vespertilio murinus Linnaeus, 1758) a o jejim porodu v zajeti [A preliminary report on the finding of Vespertilio murinus and its parturition in captivity]. Zpr. Cs. zool. Spol., 10-12: 16-17 (in Czech). RYBAR, P., 1971: On the problems of practical use of the ossification of bones as age criterion in the bats (Microchiroptera). Pr. stud. prir. Pardubice, 3: 97—121. RYBERG, O., 1947: Studies on bats and bat parasites. Stockholm, 330 pp. SAINT GIRONS, M.—C., 1973: Les mammif&res de France et du Benelux (faune marine exceptge). Paris, 481 pp. SKREB, N. & DULlC, B., 1955: Contribution 6 l'dtude des noctules (Nyctalus noctula Schreb.) en liberte et en captivity. Mammalia, 19: 335-343. SLUITER, J. W. & HEERDT, P. F. van, 1966: Seasonal habits of the noctule bat (Nyctalus noctula). Arch, neerl. zool., 16: 423-439. SLUITER, J. W., VOCTE, A. M„ HEERDT, P. F. van, 1973: Hibernation of Nyctalus noctula. Per. biol. Zagreb, 75: 181-188. STRATMANN, B., 1968: Unsere Methoden und Erfahrungen bei der Arbeit mi* Baumfledermausen am Ostufer der Miiritz. (1965-1967). Milu, 2 : 354-363. STRELKOV, P. P., 1969: Migratory and stationary bats (Chiroptera) of the European part of the Soviet Union. Acta zool. Crac., 14 : 393-439. TOSCHI, A. & LANZA, B., 1959: Fauna d'ltalia, 4, Mammalia [Generality, Insecti— vora, Chiroptera]. Bologna, 485 pp. WIMSATT, W. A., 1977: Biology of bats, Vol. III. N. York, S. Francisco, London, 651 pp. Autors' addresses: RNDr. Jifi G a i s 1 e r, CSc., Department of Animal and Human Biology, J. E. Purkyni University, Kotldfskd 2, 611 37 Brno, Czechoslovakia, RNDr. Vladimir Hanik, CSc., Department of Systematic Zoology, Charles University, ViniCnd 7, 128 44 Praha Czechoslovakia, Jan D u n g e I, prom, biol., Vondrdkova 32, 635 00 Brno, Czechoslovakia. 38 Acta Scientiarum Naturalium Academiae Scientiarum Bohemoslovacae Brno Dear Readers, ™,^ttLoS<;ioentiarwm Na,turalium appear in issues of approximately 50 pases each A vol..™* Volume VI (1972) No. 1 V. BARUS, E. KULLMANN, F. TENORA: Parasitische Nematoden aus Wirbeltieren Afghanistans. No. 3 V. BARUS, B. RYSAVY, J. GROSCHAFT, C. FOLK: The helminth fauna of Corvus frugilegus L. (Aves, Passeriformes) in Czechoslovakia and its ecological analysis. No. 4 J. KRISTEK: Die Blattwespen Euura leata und Euura mucronata in den Korbweidenanlagen Mahrens. No. 6 F. SMARDA: Pilzgesellschaften einiger Laubwalder Mahrens. No. 7 J. LIBOSVARSKY, S. LUSK, M. H. ELSEDFY: Fishery survey carried out at Lake Borullus, A. R. E., in the spring of 1971. No. 8 J. GAISLER, G. MADKOUR, J. PELIKAN: On the bats (Chiroptera) of Egypt. No. 9 R. OBRTEL: Soil surface Coleoptera in a reed swamp. No. 10 J. PROKOPIC: Biocenotical study on Cestodes of small mammals in various biotopes. No. 11 J. PELIKAN: Arvicola terrestris (L.), indexes of reproduction in Czechoslovakia. Volume VII (1973) No. 2 D. POVOLNY: Ergebnisse der zoologischen Forschungen von Dr. Z. Kaszab in der Mongolei (Nr. 267), (Tribus Gnorimoschemini, Lep., Gelechiidae). No. 7 I. 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JUR1K: Bionomics of fleas in birds' nests in the territory of Czechoslovakia. Volume IX (1975) N°- 1 LV. KNUTSON, R. ROZKOSNY, C. O. nf Ph^hil gy an£ immature stages of Pherbina and Psacadina (Diptera, Sciomyzidae). ^ ' No. 2 O. STERBA: Comparative anatomy of ideus ventralls and mrhomboNo. 3 J. PROKOPIC et K. KRlVANEC: Helminths of amphibians, their interaction and host-parasite relationships. No. 4 F MILLER et R. OBRTEL: Soil surface spiders in a lowland forest. No. 5 J. GAISLER: A quantitative study of some populations of bats in Czechoslovakia (Mammalia: Chiroptera). No. 6 K. HUDEC: Density and breeding of birds in the reed swamps of southern Moravian ponds. No. 7 M. SPURNY: Interaction of movement mechanisms in the growing hypocotyl of pine seedlings. No. 9 J. JAKRLOVA: Primary production and plant chemical composition in flood-plain meadows. No. 10 J. PEL1SEK et D. SEKANINOVA: Results of basic research of natural and anthropogenic soils in the North Bohemian Lignite District. Volume X (1976) No. 1 O. 5t£rba: Prenatal development of microtine rodents. No. 2 J. GLOSER: Photosynthesis and respiration of some alluvial meadow grasses: Responses to irradiance, temperature and COj concentration. No. 3 J KRATOCHVIL & al: Westareal der Verbreitung der Brandmaus [Apodemus agrarius (Pallas, 1778)], N°* 4 Pavlov^liA' DiS paia°utische StationNo. 5 F. TENORA: Tapeworms of the family Anoplocephalidae Cholodkowsky, 1902 Evolutionary implications. No. 6 Z. KRATOCHVIL: Das Postkranialskelett der Wildund Hauskatze (Felis silvestris und F. lybica f. catus). No. 7 F. DUSBABEK et B. ROSICK"?: Afgasid ticks (Argasidae, Ixodoidea) of Czechoslovakia. N°" 8 F-..l^ALAT: Fortpflanzungsokologie der hohlenbrutenden Vogel im sudmahrischen Kiefernwald. No. 9 J. HOLC1K et I. BASTL: Ecological effects of water level fluctuation upon the fish populations in the Danube river floodpiain in Czechoslovakia. No. 10 J. ZEJDA: The small mammal community of a lowland forest. N°' 11 «3^UKEC:T,Der Vogelbestand in der stadtischen Umwelt von Brno (CSSR) und seine Veranderungen. No. 12 R OBRTEL: Soil surface harvestmeii (Opiliomdea) in a lowland forest. Volume XI (1977) N°" 2 £i0HAVL^ * K• TJURLOV: Der Kleinvogelbestand des Baraba-Tieflands <SW Sibirien) in der Sommerzugpenode. No. 3 J. TRNKOVA: Die toxische Wirkung des Endnns auf aquatische Wirbellose. No. 4 J. GLOSER: Photosynthesis and respiration of some alluvial meadow grasses * Responses to soil water stress, diurnai and seasonal courses. No. 5 M. DUB: Die Wirkung des Endrins 20 auf Microtus arvalis. No. 6 E. WOHLGEMUTH: Toxicity of Endrin to some species of aquatic vertebrates. No. 7 M. PENAZ: Population analysis of the barb, Barbus barbus, from some Moravian rivers (Czechoslovakia). No. 8 V. HOLlSOVA et R. OBRTEL: Food resource allocation in four myomorpb. rodents coexisting in a reed swamp. No. 12 J. PIKULA et M. BEKLOVA: Ecological distribution of forest avifauna in the region of. constructed energetic system Dukovany-DaleSice. Volume XH (1978) N°' 1 : stand°rte des Hamsters (Cricetus cricetus L., Rodentia, Mamm .i in der Ostslowakei. No. 2 J. MALEK: Waldtypengruppen Sudwestmahrens in biogeozonologischer Auffassung. No. 3 J. HAVLIN: Die Vogelwelt landwirtschaftlicher Objekte. No. 4 J. KRATOCHVIL: Araignees caverni— coles des iles dalmates. No' 5 ™r,£FrNAZ:-' MPROKES, E. WOHLGEMUTH: Fish fry community of the Jihlava River near Mohelno. <•>•-.! No. 6 F. MORAVEC et A. AMIN: Some helminth parasites, excluding Monogenea. from fishes of Afghanistan. No. 7 J. LIBOSVARSKY et V. BARUS: Computed growth and survival of chub Leuciscus ceplialus, from the Rokytnh stream. No. 8 R. FIALA: . Underground organs of Typha angustifolia and Typha latifolid their growth, propagation and production. No. 9 J. PELISEK: Die Bodenverhaltnisse der Waldreservationen in der Tschechischen sozialistischen Republik (CSR). • . No. 10 J1. PIKULA et VI. KUBlK: Die Brutokologie der Turkentaube Streptopelia decaocto im Milieu der Stadt Brno. No. 12 J. ZIMA: Chromosome characteristics kiaVeSPertiii°nidae fr°m Czec^oslovaIn the case you are interested in some numbers, you may order them directly in the PuVL lishing-House AC AD EMI A, Vodifikova 40. 112 29 Praha 1-NovC MCsto, Czechoslovakia. A group of dwelling houses of the Prague quarter Zahradni MSsto. The arrow points to one of the openings of ventillation shafts which become frequent shelters of the species, namely in winter (above). A close-up showing the location of the entrance to a winter shelter of N. noctula (below). i PL. IIIRODOVEDNEJ V TAVU SKOSLOVENSKE ADEMIE VED BRN^ | ACTA SCIENTIARUM NATURALIUM SCIENTIARUM BOHEMOSLOVACAE BRNO ISSN 0032-8758 XIII • NOVA SERIES 1979 2 Alberto Coy Otero — Vlastimil Barus NEMATODES PARASITIZING CUBAN REPTILES JUL 3 „ ]ScfQ ACADEMIA • PRAHA