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FISHERIES and MARINE SERVICE Circadian Interdependence Of '# 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 DEPARTMENT OF THE SECRETARY OF STATE SECRÉTARIAT D'ÉTAT TRANSLATION BUREAU BUREAU DES TRADUCTIONS MULTILINGUAL SERVICES DIVISION DES SERVICES CANADA DIVISION MULTILINGUES /./ 3 7 7 TRANSLATED FROM — TRADUCTION DE INTO — EN German English AUTHOR — AUTEUR Hermann Remmert TITLE IN ENGLISH — TITRE ANGLAIS Circadian. Interdependence of different, organisms TITLE IN FOREIGN LANGUAGE (TRANSLITERATE FOREIGN CHARACTERS) TITRE EN LANGUE ÉTRANGÉRE (TRANSCRIRE EN CARACTÈRES ROMAINS) Tageszeitliche Verzahnung der Aktivitat verschiedener Organismen REFERENCE IN FOREIGN LANGUAGE (NAME OF 8 001( OR PUBLICATION) IN FULL. TRANSLITERATE FOREIGN CHARACTERS. RÉFÉRENCE EN LANGUE ÉTRANSE'RE (NOM DU LIVRE OU PUBLICATION), AU COMPLET, TRANSCRIRE EN CARACTÉRES ROMAINS. Oecologia REFERENCE IN ENGLISH — RÉFÉRENCE EN ANGLAIS PUBLISHER— EDITEUR PAGE NUMBERS IN ORIGINAL DATE OF PUBLICATION NUMÉROS DES PAGES DANS DATE DE PUBLICATION not shown LORI SI NAL 214-226 YEAR ISSUE NO. VOLUME PLACE OF PUBLICATION ANNÉE NUMÉRO NUMBER OF TYPED PAGES LIEU DE PUBLICATION NOMBRE DE PAGES DACTYLOGRAPHIÉES Berlin 19 69 3 1-16 REQUESTING DEPARTMENT DFE TRANSLATION BUREAU NO. 1486936 MINISTÈRE-CLIENT NOTRE DOSSIER Ne) BRANCH OR DIVISION Fisheries TRANSLATOR (INITIALS)..... WQ. DIRECTION OU DIVISION TRADUCTEUR (INITIALES) • NI; 2 51978 PERSON REQUESTING Allan_ T. Reid DEMANDÈ PAR YOUR NUMBER UNEDITED TRANSLATION VOTRE DOSSIER NCI Foe inicnInation only DATE OF REQUEST TRADUCTION NON REvisa DATE DE LA DEMANDE 17 July 1978 Inforn-la;i ,Dn seulement SOS•200-10-6 (REV. 2/68) 7 630-21-0284533 3 ascreey Spnrearlat I Q : 3 b ct.'W TRANSLATION 13UREAU BUREAU DES TRADUCTIONS MULTILINGUAL SERVICES DIVISION DES SERVICES DIVISION MULTILINGUES CLIENTS NO. DEPARTMENT DIVISION/BRANCH CITY VILLE N 0 OU CLIENT MINISTiRE . DIVISION/DIRECTION .. DFE Fisheries Ottawa BUREAU HO. LANGUAGE TRANSLATOR (INITIALS) N 0 DU SUREAU LANGUE TRADUCTEUR (INITIALES) AUG 2 3 I97a 1486936 English W.H. 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 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. 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 beetles. 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 beetle Cyclocephala castanea and the water lily Victoria regia.
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