'#

FISHERIES AND MARINE SERVICE

Translation Series No. 4377

et.

Circadian interdependence of different organisms

by H. Remmert

Original tile Tageszeitliche Verzahnung der Aktivitat verschiedener Organismen

From: Oecologia (Berlin) 3: 214-226, 1969

Translated by the Translation Bureau (wH) Multilingual Services Division Department of the.Secretary of State of Canada

Department of the Environment Fisheries and Marine Service Pacific Biological Station Nanaimo, B.C.

1978

16 pages typescript c

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Circadian. Interdependence of different, organisms

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Tageszeitliche Verzahnung der Aktivitat verschiedener Organismen

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CIRCADIAN INTERDEPENDENCE OF DIFFERENT ORGANISMS 214

by

, HERMANN REMMERT

Il. Zoological Institute of Erlangen-Nuremberg University

Summary. Synchrony of diurnal activity patterns seem to-have evolved entirely

between groups of species. No well established case of synchrony is known

which involves only two species. The interdependence of activity patterns

based on diurnal rhythms is a phenomenon well known in autecology, e:g. be-

edeen flowers and their pollinators, parasites and their hosts, predators

and their prey.

At different daytimes there are completely different food chains in

one and the same biotope.

The few existing quantitative investigations reveal that

1. strong selection pressure can limit the diurnal activity of a species;

2. the productivity in a biotope may reach a maximum when the daily feeding

time of its predators is restricted. This seems to hold, e.g., for the

marine plancton.

A. INTRODUCTION

A unilateral or mutai dependence of two organisms in the course of a

F". 11) 1:•or in i\ ION R:...\./T5.7E mcm teli4N :s-.., ul•Dt-n•-,:-nt • 2 day has been described time and again. However, there is only scant evidence for most of the examples. Hardly ever has the participation of a biological clock been proven; nor do we know very much about the timers. Besides, a ciradian interdependence hardly ever seems to exist between two individual species only, but either between two groups of species or between one individual'species on one side and a grom of species on the other side of the complex. Finally, the definition of the subject is difficult:

Bats developed ultrasonic direction-finding in adaptation to their activity in the dark. As a response to the special hunting method of the bats, the

Noctuidae developed tympanic organs. A mite (Myrmonyssus phaelenodectes) parasitizing the tympanic organs, developed a special mode of'attack which always leaves one tympanic organ functioning and hence ensures that the butterflies are largely safe from their predators (TREAT, 1958).

It would be possible to produce a large number of similar examples.

Ns a/ • However, it seems necessary to givenarrowerf definition of the subject.

B. RESEARCH DIFFICULTIES

Instead of one two organisms have to be studied. Each individual form of activity has to be registered on its own, the field conditions have to be borne in mind.

At different times of the day an animal may be looking for food, reproducing itself or making dispersal flights. Each of these forms of activity may be persued by different species of organisms at different times of the day. As a rule, flowers depend on looking for food. Orchids of the Ophrys, on the other hand, required Gorytes males willing to copulate (KULLENBERG, 1961). Thus the selection of activity registration is difficult. If one lures noctuid moths (Noctuidae), one gets an activity 3 maximum immediately after sunset, which fades away slowly but rather evenly throughout the night. If the same species is lured by using bright light, they appear considerably later; furthermore, a second maximum is obtained after midnight. Hence, a predator hunting flying Noctuidae, may have - totally different activity patterns (MOHRENBERG, 1964; WOHLRAB, 1969).

Black flies (Simuliidae) only attack their prey for blood-sucking purposes during the day; however, they are regularly caught in light traps

(KURECK, 1969). A specific enemy may be permanently active; a predator hunting on the hosts of the Simuliidae, however, may only be light-active.

An interdependence, as it is covered by our subject, may also exist between a resting and an active species: bugs (Cimex lectularius) depend on humans resting in the dark for blood-sucking.

Animals caught by the same methods at different times of the day, do not show a picture of mutual diurnal dependence. It is true, that when netting is done in one and the same area, different species of are caught at different times of the day (MARCHAND, quoted by TISCHLER, 1955). 216

But this method does not give any indication regarding the mutual dependencies.

Animals resting during the day will, for the most part, conceal themselves in hiding places from their optically oriented enemies. Day.'.active insects, on the other hand, often rest during the dark in clearly visible places, in which, however, vibrations can easily be perceived. Thus many Hymenoptera get a firm hold of thin swaying pistils. Syrphids sleep on the tips of blades Of grass. Bumble-bees rest by hiding their heads in the flowers of composites. Hence, when using nets at night, bumble-bees will be caught as well as some parasitic Hymenoptera and syrphids (i.e. light-active forms) in greater numbers than during the day (LEWIS and TAYLOR). Therefore, as far as we are concerned, only those catches may be taken into consideration, where use was 4

made of an animal's active performance. EVen specific properties of .a bio- tope may make the situation more difficult. The window fly (Phryne fenestralis) is light-active and requires a high amount of humidity. Hence, it is found feeding in dry areas only during very short periods in the morning and - at night. However, in humid forests and on rainy days it is active throughout the day. A predator which specifically goes for window flies in dry areas only needs to have an activity maximum early in the morning and another one late in the evening. An interdependence can hardly be proven.

C. Results

1. Autoecological Aspects

a) Insects and Flowers

A circadian coordination between different organl-Sms definitely exists in flower ecology. A sufficient number of examples is known (cf. e.g. OLBERG,

1951). Thus I only wish to present one. Arum (Arum maculatum) regularly opens its pitfall flower about 2 p.m. At this time the moth flies (Psychodidae) which do the pollination, are active. They enter the pitfall flower. They are held back by the trap hairs and remain trapped. The following day about

10 am. the pollen is ejected and poured over the moth flies. Subsequently, the trap hairs dry up and the Psychodidae can leave their prison. At the same time other arums will have opened. The pollen-laden moth flies enter their flowers, pollinate them and leave the day after. The trap hairs will dry up, regardless of whether pollination comes about or not. Like the opening of the flower it is part of the daily rhythm . As to the timing, the opening of the flower is exactly adjusted to the moth flies' time of activity

(STRUCK, 1965). 5

6 là ï4 lb 22 2 6 lb 14s 18 22 2 6 1.5

-Phase le-Phase ->

Fig. 1. Pollination of arum maculatum by Psychodidae. The arrows designate the opening of the flower and the drying up of the trap hairs. The solid line indicates the number of moth flies found in the trap. Abscissa: Time. According to data by STRUCK

In the absence of moth flies, the arum can be pollinated by other mosquitos and . Even the hawkmoths (Sphingidae) with their long pro- bosces can be replaced at their specific flowers - e.g. by horseflies

(Tabanida), which in some cases have an enormous proboscis looking for flowers.

With regard to Germany Pangonius micans could be mentioned as an example.

Phanerogams of non-European origin bloom in European gardens and produce seeds.

They are pollinated in our country. This even holds true if apparently a rather exactly functioning relationship exists in nature, such as was described by GESSNER (1960) for the castanea and the water lily

Victoria regia. In our hothouses Victoria regia develops normal flowers and fruit. It has to be pollinated in a different way as compared to the Amazon region. Extremely close ties exist between a pollinating species and a flowering plant, such as those between different species of Ophrys (orchids) and specific Gorytes males (Hymenoptera). In this case, however, there is no 6

diurnal coordination. Ophrys permanentlyemanates the pheromonoid substance which attracts the Gorytes males. For 2 weeks the flower may be in full bloom. Typical hawkmothflowers such as Lilium regale or Lonicera periclymenum only have a very low daily rhythm (production of odorous substances). - In the case of extremely close ties, obviously, a double check is introduced in that the plant can make use of a flying pollinator even at an unusual time of the day.

b) Partnerships and Symbioses

A number of animal species can populate certain biotopes on the basis of partnerships only, which, in addition, require diurnal synchronization.

Thus in the daytime a good number of Red Sea fish depend on the presence of long-spined sea urchins, among whose spines they find protection against their enemies. During the night these fish do not seem to have any enemies. They sleep in the open water. During the day they try to catch passing prey from within the protecting spines of the long-spined 'sea urchins. These long- spined sea urchins also live together with crabs with which, during the day, they jointly look for hiding places in cracks, on stones between moles, and the like. Here the crabs do not find any food. At night, when the long- spined sea urchins wander about in search of food, they are accompanied by the crabs, and these seek their food protected by the spines. They can only populate the areas concerned because they are protected day and night by the long-spined sea urchin, and because the long-spined sea urchin migrates reguiarly, thus carryingthe crab to biotopes where it can find food (MAGNUS,

1964).

c) Parasites and their Hosts

The example of filariae and their vectors (Table 1) is generally known.

Another example occasionally quoted is that of the cercariae of the lung 7

fluke (Paragonismus westermanni) which, during the night, at the main time

of activity of their intermediate hosts (Decapoda), leave the snail and then

look for the decapods. Experience made by my working group reminds us to

be cautious (DITTMANN, 1966; EISFELD, 1967; REMMERT in litt.). The sporocysts

of Leuchloridium macrostomum, which parasitize the amber snail (Succinea),

form very long tubes which during the day extend into the feelers of the

snail and here make characteristic pumping movements. The tubes of the

sporocysts have coloured rings and increase the feelers to five times its

normal size. Birds like to peck at such feelers. They are under high pressure 219

and empty their contents, consisting of infectious cercariae, into the bill

of a pecking bird. During the night, when there are no birds around, the

tubes withdraw to the inner part of the snail's body (HECKER and THOMAS, 1965).

. Table 1 Wichereria bancrofti. Reactions between Vectors and Filaria (accord-

ing to data given by CLOUDSLEY-THOMPSON

Occurrence Vector Activity of Filaria in the the Vector Peripheral Vessels

the Philippines, . Aedes variega- the Fiji Islands, tus during the day day and night Samoa, Africa Culex fatigans during the night during the night

Loa loa, Africa Chrysops during the day during the day

d) Competitors Pollinating, heavy and robust flies such as Eristalis can largely

keep other species, which are not as strong, away from very productive flowers.

Only . if Eristalis is not present,' the inferior species can be found on these

flowers (KIRUCHI, 1965). However,.these inferior species have not yet been

influenced in their proper daily rhythm, they merely turn to less favourable -

flowers. This effect will be even stronger with species which (such as Vespa) 8

occasionally assault and eat a small fly. Due to a synchronous activity .

phase small species are kept away from particularly productive food sources.

e) Prey and Predator

Most predators merely find their prey if it is moving, i.e. if - it is

active. Only few sensory functions make it possible for a predator to detect

inactive prey. Especially, if the prey has retired to a hiding place, e.g.

to a hole, it is practically safe from any attack. Accordingly, one will

find extremely nocturnal species among mammals (dormice, bats) occur predominant-

ly, :in the food of owls hunting at night. They are hardly even captured by

birds hunting during the day. On the other hand, 'reptiles (such as lizards,

snakes) which depend on sun in the extreme, are found primarily as prey of

diurnal birds of prey. Common buzzards regularly feed on such reptiles. The

existence of a specialist (the serpent eagle) for these animals is another

indication. Of the owls only the tawny owl catches reptiles on a regular

basis. Of the species presented (barn owl, long-eared owl and eagle owl),

it is the owl which hunts least in the dark (Table 2). Amphibians and fish

are equally available both day and night; as a result suitable specialists have

developed among diurnal predators (osprey, tern) and nocturnal predators

(fishing owls and fishing bats).

This does not hold true everywhere. In areas around the Mediterranean

bats often are active long before sunset; they are regularly found among

the prey of diurnal birds of prey. An observation on a Eurasian sparrow-

hawk in Egypt showed, that diurnalbirds of prey can adapt to a very high degree

• to such conditions (UTTENDÔRFER, 1952).

It need not be discussed here (cf. e.g. ROEDER, 1968, for nocturnal

predators) that the senses of day-active predators function quite differently .

than the senses of night-active predators and that a similar situation exists 9

also in the prey species.

What is essential, is the question of whether the prey can evade a predator by shifting its time of activity. Cases of this kind are well established with regard to such highly developed forms like our major mammals which in those areas where they are hunted by man have switched over to night activity. There are similar reports on the robber crab (Birgus latro), on

Ocypode and on the land hermit crab (Coenobite). They too are said to be generally night-active in those areas where they are hunted by man, in others, however, they are day-active. The results established with regard to Coenobita do not confirm this hypothesis, though so far they do not give a distribution of its activity in its natural biotope (NIGGEMANN, 1968). The same holds true for OcypOde (cf. NIGGEMANN, 1968, and LINSEMAYR, 1968). Nor can RÜPPELL's hypothesis, according to which the night activity of terrestial Amphipoda is relatable'to the number.of hunting birds, be readily considereclas shaving been proven if TONGIORGI's Mediterranean findings are taken into consideration.

KIKUCHI (cf. p. 219) makes reference to the displacement of some animals from favourable feeding grounds by predators with a synchronous activity.

In the Red Sea the fish Istiblennius rivulatus, living in the surface layers of water goes ashore for nocturnal rest and hides in cracks and holes.

Individuals that ha.ve remained too close to the water are seized by octopuses patrolling the shores at night with a keen eye on prey, drawn into the water and eaten (MAGNUS, 1965).

Given lack of food, female glowworms use the signal of a different species of glowworms to lure their males and eat them. Females ready to copulate emit the signal of their own species and thus lure males of their kind (LLOYD, quoted from PINNER, 1966).

Two examples permit quantification. Every autumn the Mediterranean

Table 2: Extremely Dark-Active and Extremely Light- 220 Active Animals as Prey of Diurnal or Nocturnal Predators (According to Data by UTTEND5RFER, 1952)

Barn Owl Tawny OwI Long-Eared Owl Eagle Owl Eurasian .Goshawk Hooby Peregrine Falcon Common Buzzard Sparrowhawk

Vertebrata 77,600 55,600 60,000 5,500 7,150 9,000 930 6,500 approx. 15,000

Glires 100 125 1 13 1 1 _ - -

Chiroptera 113 128 14 3 1 - 2 1 -

Reptilia 1 Lacerta 41 - 1 Lacerta - 1 La- - as a rule: La- (Lacerta, certa certa, Anguis, Anguis, . Natrix, Coro- Coronella, nella, Vipera Natrix) 1 0

is crossed by approx. 600 million migratory birds on their way from Europe to Africa. In general these migratory birds migrate during the night. On small rocky Mediterranean islands the Eleonora's falcon (Falco eleonora) is the only European bird to breed in spite of photoperiods getting shorter and shorter, i.e. precisely at the time of the autumnal bird migration. While out of the breeding season it largely lives on insects, it feeds its off- spring on migratory birds. The falconshover in a phalanx in the air in front of the islands; hardly any bird, which crosses the Mediterranean during the day, escapes them (WALTER, 1968). STRESEMANN (1968) computated that about 750,000 migratory birds are captured by the Eleonora's falcons every autumn. They can only hunt during the day. Hence they practically capture every migratory bird migrating during the day. Thus, there is a strong selection pressure towards nocturnal migration.

Aquatic insects generally emerge from the pupa after sunset (REMMERT,

1962). Almost all pupae which pass through a body of water to its surface are eaten by fish during the day. The pupae, on the other hand, which rise at night, reach the surface of the water almost without exception, and can metamorphose into imagines (FISCHER and ROSIN, 1968, Table 3). It is true that in warm regions they may still be caught by skimmers (Rynchops), which are active at dusk; however, the losses are minimal in every case. Animal species whose young are eaten whenever they meet adults, are segregated into young and adult animals as if they were different species. Shoals of young fish are to be found in totally different places of the bank than shoals of adult fish; their diurnal migrations differ from those of the adult fish.

Thus meetings of these shoals are virtually excluded. For example, this is the case in the black cod (Gadus virens) (HEMPEL, 1957). Table 3: Emergence of Chironomus in the Presence of Fish given Light-Dark

and Light Light Conditions (According to FISCHER and ROSIN, 1968)

LL LD

Without fish (control group) 97 86

with fish 3 81

2. Synecological Aspects

Totally different food Chains exist in the same biotope during the day and during the night. If one follows the food cycle in the ecosystem, different aspects have to be taken into consideration for day and night, respectively. They may differ to a higher degree in the same biotope than in adjacent biotopes at the same time of the day. --

In spring, the catkins (Salix) are frequented by bees (Apis), bumble- bees (Bombus), and above all by flies of the Egle genus, which here find food in the form of pollen and effect pollination. These insects are hunted by the wood warbler, which is the first to return from the south in spring, the chiffchaff (Phylloscopus collybita), arriving precisely at the time when the willow bushes are in flower. And about this time the sparrow hawk

(Accipiter nisus) comes back to its nesting places. With regard to the chiffchaff the sparrow hawk is the predator. During the night one finds a 223 profusion of other insects on the same flowers. Above all butterflies of the noctuid family, which have survived the winter as imagines (they emerged from the pupa at the beginning of winter) and now eat their first food, sit on the catkins. Here too one will regularly see the first bats of the year hunting. Visa-vis the noctuids they are predators: in many cases the tawny owl (Strix aluco) plays the role of a specialist vis-a-visthe bats. Thus 12

totally different sets of relations exist during the day and during the night.

However, we have no quantifiable data. Finally one example, which clarifies the enormous importance of biological production to a considerable degree, shall be briefly outlined. Marine zooplankter stay in deeper layers of water during the day and move to the surface during the night. Phytoplankter stay at the surface during the day, for only here they find the light which they recuire for their photosynthesis (LORENZEN, 1963). They are only eaten by the predatory zooplankters during the night - i.e. at a time when they only repire part of the substances produced by photosynthesis. A thorough analysis on a mathematical basis done by MACALLISTER (1968) showed that the primary and secondary production of a body of water reaches a maximum, if the producers (in this case the phytoplankter) are eaten during the night only.

But it decreases considerably, if the same amount is eaten continuously.

If.

* . y " • H 441i (

Fig. 2. Food chains relating to catkins (Salix) which differ from day to night. During the day: fly (ER1e), (Phylloscopus collybita), bird (Accipiter nisus); bird the night: butterfly (Taeniocampa/Monima), bat, during bird (Strix aluco) 13

Thus production is increased considerably by zooplankter occurring periodically. 224

Nothing can be said so far about similar phenomena ashore. The few individual examples show too heterogeneous a picture.

J J J 'A' S' J

Fig. 3. The total secondary production in the North Pacific calculated on the basis of field data for the various months of the year. A respiration rate of 6% of the body-weight of the zooplankter per day was assumed. Ordinate: amount of production expressed in terms of C-incorporation per m 2 . The different curve apply, assuming different feeding times but the same amount of food, a) feeding time during the first hours of the night, b) feeding time equal to the first half of the night, c) feeding time equal to the whole night, d) feeding time equally distributed over day and night. According to MACALLISTER, schematized. 14

10.0 8.0 6.0 4,0

2.0

1.0 0.8 06 0.4

7 10 11 12 13 14 15 16 17 18 Toge

Fig. 4. Effect of constant feeding and of nightly feeding only (given the saine amount of food) upon the development of a phytoplankter culture. Ordinate: plant cells per unit of volume; abscissa: successive days. Nights are marked by black dashes. The cul- ture in which the predators feed during the night only increases considerably (upper curve), the other one remains nearly constant. According to MACALLISTER, schematized.

BIBLIOGRAPHY

CLOUDSLEY-THOMPSON, C.L.: Rhythmic Activity in Animal Physiology and Behaviour, p. 236. London and New York: Academic Press, 1961. DITTMANN, F.: Activity Rhythms in the Shore Crab Carcinus maenas, Diploma Thesis, Kiel, 33 4- VII, 1966. EISFELD, F.: The diurnal Rhythm of the American Freshwater Crayfish (Cambarus affinis), Diploma Thesis, Kiel, 39 p., 1967. FISCHER, J. and S. ROSIN: Influence of Light and Temperature upon the Activity of Emergence of Chionomus nuditarsis. Str. Rev. Suisse Zool. 75, 538-549 (1968) GESSNER, F.: The Opening of the Flower of Victoria regia in its Relation to Light, Planta (Berl.) 54, 453-465 (1960). HECKER, U., and E. THOMAS: On Sporocyst Tubes of Leucochloridium macrostonum. Rud. Verh. Dtsch. Zool. Ges. 1964 Kiel, 444-456 (1965) HEMPEL, G.: Ecological Studies on the Diurnal Behaviour of Saltwater Fish. Verh. Dtsch. Zool. Ces. 1956 Hamburg, 415-421 (1957). • - Diurnal Variations in Catch, Feeding and Swimming Activity of Plaice (Pleuronectes platessa). Cons. PeLmanent Intern. Explor. de la Mer; Rapports et Proc. Verbaux 155 (1964). KIRUCHI, T.: Studies on the Coaction among Insects Visiting Flowers VII: Diurnal Rhythm of the Appearance of the Subordinate Syrphid Fly in 15

Relation to the Presence of the Dominant One. Sci Rep. Tohoku Univ., Ser. IV (Biol.) 31, 207-215 (1965). KULLENBERG, B.: Studies in Ophrys Pollination. Zool. Bidrag fran Uppsala 34, 1-340, 51 Fig. (1961). KURECK, A.: Diurnal Rhythms of Simuliidae (Diptera) in Lappland. Oecologia (Berl.) 2, 385-410 (1969) LEWIS, T., and E.R. TAYLOR: Introduction to Experimental Ecology, 401 p., London and New York: Academic Press, 1 967. LINSENMAYR, K.E.: Construction and Signal Function of the Beach Pyramid of Ocypode saratan. Forsk (Decapoda). Z. Tierpsychol. 24, 403-456 (1967) LORENZEN, C.J.: Diurnal Variation in the Photosynthetic Activity of Natural Plancton Populations. Limnol. and Oceanogr., 56-62 (1963) MAGNUS, D.B...: On the Problem of Partnerships in Long-Spined Sea Urchins. Verh. Dtsch. Zool. Ces., München 1963, 404-417 (1964). - Types of Movement of Lophalticus kurkii magnusi KLAUSEWITZ (Pisces, Slariidae) in its Biotope. Verh. Dtsch. Zool. Ges., Jena 1965, 542-545 (1966) McALLISTER, C.D.: Zooplancton Rations, Phytoplancton Mortality and the Estimation of Marine Production. Symposium on Marine Food Chains. Arhus, July, 23-26, 1968, MS. MORRENBERG, J.: The Catching of Noctuid Moths on Artificial Lights - its Problems and its Scientific Importance. Diploma ThesU, Kiel, 79 p., 1964. NIGCEMANN, R.: On the Biology and Ecology of the Land Hermit Crab Coenobite scaevola Forskal at the Red Sea. Oecologia 1, 236-264 (1968). OLBERG, G.: Flower and , 104 p., Wittenberg: Neue Brehm-BüCherei, 1951 PINNER, E.: Deception Tactics of the Glowworms? Naturw. Rundschau 17, 30, 1 966. REMMERT, H.: The Emergence Rhythm of Insects, 73 p., Wiesbaden: Steiner, 19 62. ROEDER, K.R.: Neural Foundations of Behaviour, 283 p., Bern and Stuttgart: Huber, 1968. RUPPEL, G.: Diurnal and Long-Term Faunal Shifts.in the Marine Supralittoral. Z. Morph. Ôkol. Tiere, 60, 338-375 (1967). SCHMIDT, U.: Contributions to the Biology of the Coal Fish (Gadus virens L.) in the Icelandic Waters. Ber. dtsch. Komm. Meeresforsch., 14, 46-82 (1955). STRESEMANN, E.: The Interference of the Eleonora's falcon with the Autumnal Bird Migration. J. Orn. (Berl.), 109, 472-474 (1968). STRUCK, S.: The Insects in the Pitfall Flowers of Arum Maculatum. Diploma 226 Thesis, Kiel, 74 p., 1965. TISCHLER, W.: Synecology of Terrestial Animals, 414 P., Stuttgart: Fischer, 1955, TONG1ORGI, P.: Richerche ecologishe sugli artropodi di una spiaggia sabbiosa del litorale tirrenico. I. Redia (Firenze) 48, 165-177 (1963). TREAT, A.E.: A Case of Peculiar Parasitism. Nat. Hist. (N.Y.) 67, 366-373 (1968). UTTEND6RFER, O.: New Results concerning the Food of Accipitres and Owls, 228 p., Stuttgart: Eugen Ulmer, 1952. WALTER, H.: On the Dependence of the Eleonora's Falcon (Falco eleonorae) on the Mediterranean Bird Migration. J. Ore. (Berl.) 109, 323-365 (1968) WOHLRAB, R.: Investigations of Insects on Fermenting Fruit. Diploma Thesis, Kiel, 58 p., 1969.

Prof. Dr. H. REMMERT II. Zoolog. Inst. der Universitâte 8520 Erlangen, Bismarckstr. 10