'#
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.
1 978
16
pages typescript
c
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Circadian. Interdependence of different, organisms
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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. beedeen 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
mcm
teli4N
R:...\./T5.7E
: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
interdependence hardly ever seems to exist
ciradian
between two individual species only, but either between two groups of species
or between one individual'species on one side and a
other side of the complex.
grom of species on the
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/
•
f
However, it seems necessary to givenarrower definition of the subject.
B. RESEARCH DIFFICULTIES
Instead of one animal 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 insects looking for food.
Orchids
of the genus 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 animals 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.
composites.
Bumble-bees rest by hiding their heads in the flowers of
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 biotope 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
However, in humid forests and on rainy days it is active throughout
night.
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.
the
Like the opening of
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
1s4
-Phase
18
22
le-Phase
2
6
1.5
->
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 beetles. Even the hawkmoths (Sphingidae) with their long probosces 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 beetle Cyclocephala 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 longspined 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 (according to data given by CLOUDSLEY-THOMPSON
Activity of
the Vector
Filaria in the
Peripheral Vessels
Occurrence
Vector
the Philippines, .
the Fiji Islands,
Samoa, Africa
Aedes variegaduring the day
tus
Culex fatigans during the night
day and 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 predominantly, :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 LightActive Animals as Prey of Diurnal or Nocturnal Predators
(According to Data by UTTEND5RFER, 1952)
.Goshawk Hooby Peregrine Falcon
Barn Owl Tawny OwI Long-Eared Owl Eagle Owl Eurasian
Sparrowhawk
77,600
55,600
60,000
5,500
7,150
9,000
Glires
100
125
1
13
1
1
Chiroptera
113
128
14
3
1
1 Lacerta
41
(Lacerta,
Anguis,
Coronella,
Natrix)
Vertebrata
Reptilia
-
1 Lacerta
.
-
220
Common Buzzard
930
6,500
approx. 15,000
_
-
-
2
1 Lacerta
1
-
as a rule: Lacerta, Anguis,
Natrix, Coronella, Vipera
10
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.
it feeds its off-
out of the breeding season it largely lives on insects,
spring on migratory birds.
While
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)
Without fish (control group)
with fish
2.
LL
LD
97
86
81
3
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), bumblebees (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
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
223
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 culture 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
Sci Rep. Tohoku Univ.,
Relation to the Presence of the Dominant One.
Ser. IV (Biol.) 31, 207-215 (1965).
KULLENBERG, B.: Studies in Ophrys Pollination.
34, 1-340, 51 Fig. (1961).
Zool. Bidrag fran Uppsala
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 Insect, 104 p., Wittenberg:
PINNER, E.:
REMMERT, H.:
Deception Tactics of the Glowworms?
Neue Brehm-BüCherei, 1951
Naturw. Rundschau 17, 30, 1 966.
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
Thesis, Kiel, 74 p., 1965.
TISCHLER, W.:
1955,
Synecology of Terrestial Animals, 414 P., Stuttgart:
Fischer,
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).
226
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