Digestive mechanics and gluttonous feeding in the feather starOligometra serripinna (Echinodermata: Crinoidea)

The movement and digestion of food in the gut ofOligometra serripinna (Carpenter) were studied at Lizard Island (14°3842S; 145°2710E) in the austral winter of 1986. Feather stars in the laboratory were fed a brief, small meal of brine shrimp nauplii and killed at increasing time intervals thereafter. Histological reconstructions showed that the ingested nauplii progressed along the digestive tract surprisingly quickly. Some nauplii were found in the mid and hind intestine in only 30 min, and all of the nauplii had reached the hind intestine and rectum in 1 h. Digestion of the nauplii had started at 1 h, and only a few fragments of naupliar exoskeleton remained in the hind intestine and rectum 5 h after the start of feeding. Videotape analysis showed that no fecal pellets were released during this experiment. In the natural environment ofO. serripinna, ingested particles may similarly be transported quickly to the hind part of the gut and digested there — when feather stars were fixed in the field, most of the gut contents were found in the hind intestine and rectum.O. serripinna, which efficiently rejects inert particles before they are ingested, usually defecates infrequently (probably not more than once over a span of many hours) and differs from some other feather stars that ingest numerous inert particles and defecate much more frequently. When specimens ofO. serripinna were fed continuously on brine shrimp nauplii,Artemia sp. (San Francisco strain), in the laboratory, the feather stars fed gluttonously, packing their guts with several hundred nauplii in 1 to 2 h. Thereafter, superfluous feeding began (i.e., further ingestions appeared to force undigested nauplii, some of them still living, out of the anus). These observations suggest thatO. serripinna usually feeds at relatively modest rates in its natural habitat, but can feed gluttonously to take advantage of infrequent patches of highly concentrated, nutritious particles (e.g. copepod swarms, migrating demersal zooplankton, and invertebrate gametes from mass spawnings). It is likely that such patches of nutritious particles are usually small enough to drift out of reach of the feather stars before gluttonous feeding proceeds to superfluous feeding. Opportunities for superfluous feeding in nature are probably very infrequent (e.g. ingestion of coral gametes and embryos after a mass spawning), and the feather stars evidently have no behavior that stops further ingestions after the gut becomes filled to capacity.     

Frostschutz-Glykoproteine bei Mytilus edulis?

Fractions of glycoproteins preventing formation of ice crystals in water were isolated from the mussel Mytilus edulis, using ion-exchange procedure and gel filtration. The protein fractions depress the freezing point of water more than would be expected from their concentrations, taking into account their molecular weights (>10 000 Daltons). It is suggested that the occurrence of such antifreeze glycoproteins contributes essentially to the mechanism of freezing resistance in the mussel.

---

---

Mit finanzieller Unterstützung durch die Deutsche Forschungsgemeinschaft (Th 158/7).     

Neue Pogaina-Arten (Turbellaria,Dalyellioda) mit Zooxanthellen aus dem Mesopsammal der Nordsee- und Mittelmeerküste

Zusammenfassung Die Gattung Pogaina Marcus wird um 2 neue Arten aus dem marinen Mesopsammal erweitert. Pogaina kinnei nov. spec. stammt aus dem eulitoralen Sandwatt der Nordseeküste, Pogaina annulata aus sublitoralen Grobsanden des Golfes von Neapel.Wahrscheinlich liefert die Gattung Pogaina ein neues Beispiel für eine Endosymbiose zwischen Turbellarien und Diatomeen.

---

Two new species of the genus Pogaina Marcus are described. Pogaina kinnei nov. spec. lives in intertidal sand flats of the North Sea whereas Pogaina annulata nov. spec. was collected from sublittoral coarse sand of the Gulf of Naples. The genus Pogaina probably represents a new example of an endosymbiosis between turbellarians and diatoms.

---

---

Mit unterstützung der Deutschen Forschungsgemeinschaft.     

Hybridization and introgression between Indo-Pacific species of Diadema

We present the first case of hybridization between echinoid species evaluated through genetic markers and morphology. We examined the three tropical Indo-Pacific species of the genus Diadema: D. paucispinum A. Agassiz, D. savignyi (Audouin) Michelin and D. setosum (Leske). Specimens morphologically intermediate between two of these species, D. savignyi and D. setosumhave previously been noted. Fertile hybrids have also been produced in the laboratory. To determine extent of hybridization, we first assayed the allozyme products of 22 loci in individuals which, on the basis of morphology and collection locality, could be unambiguously assigned to one of the three species. We found four loci that were either diagnostic or semi-diagnostic between D. setosum and the other two species, and one locus semi-diagnostic between D. savignyi and D. paucispinum.We then assayed individuals of intermediate morphology to find out whether they had hybrid genotypes. In the Ryukyu Islands, where D. setosum and D. savignyi coexist, we found one specimen which on the basis of all four diagnostic loci was an F₁ hybrid, and several individuals that could be either F₂ (or later-generation) hybrids or progeny of backcrosses.We also found one individual that on both genetic and morphological grounds appeared to belong to D. paucispinum (even though this spcies has only been reported from Hawaii and Kiribati) and three other individuals that carried alleles characteristic of D. paucispinum. Thus, previous reports of hybridization between D. setosum and D. savignyi were correct; it is also possible that larvae of D. paucispinum occasionally arrive at localities outside Hawaii, reach sexual maturity, and hybridize with the other two species. Counts of pure and hybrid morphotypes in other populations across the western tropical Pacific revealed a low but widespread incidence of apparent F₁ hybrids and backcrosses of D.savignyi and D. setosum. However, the existence of diagnostic or semi-diagnostic loci, low interspecific gene-flow estimates based on F _st statistics, and the lack of Hardy-Weinberg or linkage disequilibria among individuals of pure morphology all suggest that gene introgression between the three species is limited.     

The highly specialised gut of Fritillariidae (Appendicularia: Tunicata)

The morphology and some functions of the gut of Fritillaria pellucida and Fritillaria formica (Fritillaridae) were investigated by light and electron microscopy, and also by means of histochemical and immunohistochemical techniques. Fritillarids, very important for their abundance and ecological impact in marine ecosystems, have a very simplified gut: a straight oesophagus connects the pharynx to the digestive nucleus, composed of globular stomach and rectum, connected dorsally through a very short proximal intestine. The latter is characterised by a few (two to four) extremely specialised cells, completely filled with mitochondria associated tightly with membrane infoldings showing strong ATP-ase activity, and probably involved in the osmoregulation of internal body fluids. The gut is formed of an extremely low number of cells, which, although poorly diversified, are very large in the stomach and rectum. Food transfer along the gut depends on and is regulated by well-developed cardiac and pyloric valves, and signs of general digestive and absorptive activity are recognisable all along the brush border of the main tracts. The macroscopic organisation and cytological characters of the gut in fritillarids are completely different from those of the oikopleurids. In particular, fritillarids lack specialised cells for endocytosis and intracellular digestion, like those described in the genus Oikopleura. The general simplification and specialisation observed in Fritillaria gut may account for their elevated growth rate and abundant diffusion in all oceans.     

Beziehungen zwischen Körperbau und Lebensweise bei Blenniidae (Pisces) aus dem Roten Meer. III. Morphologie des Auges

The morphology of the eyes of 3 salariin Blenniidae have been investigated and compared: Salarias fasciatus (inhabiting the sublittoral), Istiblennius edentulus (eulittoral), and Alticus kirkii (supralittoral). An effective protection against desiccation in A. kirkii is offered by the very thick cornea conjunctiva. Extension of the visual field in this species is achieved by protrusion of the eyes from the head, by recession of the dermal pigment, and by a relatively large lense which allows better perception of marginal rays. The retina of I. edentulus is adapted for amphibious vision by the existence of many prominent swellings and folds, and of a central depression into which the lens can be with-drawn. Thereby, the stronger rays of light retraction present on land are focussed on the swellings, since the retina of the depression is under-developed. The cornea conjunctiva and propria of A. kirkii are separated, and thereby constitute an additional eye chamber. Presumably this cavity possesses a lower refractive index than the cornea or water, and thereby enables sharp vision in air, since the light-concentrating effect of the curved cornea is thus diminished. Additionally, the shape of the cornea propria can be varied, since it is centrally connected with the lens; by this means the eye chamber can be extended, enabling displacement of the tocal point of the light rays to correspond with the momentary residence. The adaptations described here for I. edentulus and for A. kirkii have so far not been reported in other amphibious vertebrates.

---

---

Hit Unterstützung der Deutschen Forschungsgemeinschaft     

Das kommensalische verhältnis zwischen Periclimenes imperator (Decapoda: Palaemonidae) und Hexabranchus sanguineus (Nudibranchia: Doridacea)

The Red Sea nudibranch Hexabranchus sanguineus, rare in this region, was found in the Gulf of Aqaba together with a pair of commensal shrimp, Periclimenes imperator. The behaviour of the shrimp, especially as regards their adaptation to the host, was studied in the field and in an aquarium. The shrimp live on the sea slug, feeding on particles adhering to its mucous surface; they move on it even when it is swimming. The shrimp's colour is well adapted to that of the host. The observed specimens never left a healthy host willingly; when, in aquarium experiments, various red substrates or mucous sea cucumbers were offered beside H. sanguineus, the shrimp always chose the nudibranch. However, dead or injured and motionless slugs were abandoned. Two days after the death of a nudibranch, its commensal shrimp accepted the brown synaptid Euapta godeffroyi as host; on a previous occasion, a shrimp was found living on this latier species in the reef. The adaptation of P. imperator to living on the temporarily free-swimming H. sanguineus apparently represents a special evolutionary stage compared with that of other members of this family, which are mostly commensal with sessile hosts. Only palaemonid cleaner-shrimps feed briefly from free-swimming substrates.

---

---

Mit Unterstützung der Deutschen Forschungsgemeinschaft     

Ökologische bedeutung der schleimsekretion bei den planula-larven der hydroidengattung Eudendrium

Colonies of hydroid polyps of the genus Eudendrium Ehrenberg release planula larvae with a high yolk content. Immediately after hatching, they begin to excrete a slime rope. In the presence of water movements, larvae hatching simultaneously tend to join a trail of slime; in stagnant water, the larvae move down the rope separately. The planula larvae are unable to float. On the basis of aquarium observations, we suggest that slime ropes with larvae adhere to substrates in the vicinity of the mother colony; this prevents the larvae from being carried off too far into open waters. In view of the absence of medusae in Eudendrium, this response of the planula is interpreted as an adaptation to a purely sessile life cycle. Structure and distribution of the slime secretions are documented by means of light and electron-microscope photos.

---

Ökologische bedeutung der schleimsekretion bei den planula-larven der hydroidengattung Eudendrium

     

Accumulation and release of petroleum-derived aromatic hydrocarbons by four species of marine animals

Survival of Ctenodiscus crispatus during exposure to hypoxia (P _{O 2}<3 mm Hg) at 5°C is greater than that of any echinoderm reported in the literature, the LT₅₀ being 248 h; this is reduced to 236 h in the presence of hydrogen sulfide. Unlike Asterias vulgaris and A. forbesi, both of which lose the tube foot response to tactile stimulation long before death from hypoxia occurs, C. crispatus remains responsive until death. The extension of the highly protrusible epiproctal cone, which occurs in 75% of the mud stars simultaneously exposed to hypoxia and H₂S, serves to maintain burrow contact with the overlying water. The rate of oxygen consumption remains constant down to an ambient oxygen partial pressure of 10 to 25 mm Hg, becoming more oxygen-dependent after prior exposure of the asteroids to hypoxia. C. crispatus exhibits a clear oxygen-debt phenomenon as well as a compensatory reduction in the residual P _{O 2} (oxygen partial pressure at which oxygen consumption ceases) from 2.4 to 0.2 mm Hg after hypoxic exposure.     

Hypoxic life of intertidal acorn barnacles

Oxygen levels were monitored within the mantle cavities of three barnacle species ( Chthamalus stellatus, Semibalanus balanoides, Elminius modestus), using optode microsensors. Conditions were always hypoxic, even when barnacles were actively using the prosoma and cirri to pump aerated seawater into the mantle cavity. Mantle fluid oxygen concentrations were extremely variable; behaviour and oxygen concentrations were not closely coupled. Ventilation of the mantle cavity depended partially on external water flow, with higher and more stable mantle fluid oxygen concentrations being sustained when the water around barnacles was agitated. During emersion, barnacles initially pumped seawater between the mantle cavity and the cone above the opercular plates to achieve ventilation. As water was lost it was replaced by air bubbles, eventually resulting in an air-filled mantle cavity. In S. balanoides and E. modestus, once the mantle cavity was filled with air, the barnacle usually used up the oxygen within the bubble within 2–3 h and did not regain oxic conditions until the barnacle was reimmersed in seawater. In C. stellatus, the air bubble was repeatedly refreshed for many hours by pneumostome formation. In response to low environmental salinity, all three species closed the opercular plates firmly and rapidly used up oxygen within the mantle fluid.     

Natürliche Oberflächenfilme im Seegebiet vor Sylt

The physical and chemical properties of the sea surface can be altered considerably by the formation of slicks and the accumulation of particles (including plankton). Investigations of natural slicks near the island of Sylt consisted of analyses of the dissolved and particulate surface-active substances, as well as the identification of plankton and bacteria collected by a surface-film sampler. The analyzed surface material consisted primarily of phytoplankton (Prorocentrum micans in particular), bacteria, detritus and dissolved compounds of fatty acids. These fatty acid compounds alone would not account for the slick. The slick appeared to be caused by the calm weather, the active accumulation of P. micans at the surface, and the increase in bacteria associated with this accumulation. Here a decrease was observed in the dissolved surface-active substances compared with the concentration usually found in normal surface waters, of which the total fatty acid content in a sample was taken as indicator. In some cases a correlation was found between fatty acids and particulate matter in naturally occurring slicks. Particularly large variations were found in the fatty acid patterns in the filtrate and filter residue at a time when no slick was present and no particulate matter had accumulated at the surface.

---

---

Diese Arbeit wurde von der Deutschen Forschungsgemeinschaft im Rahmen des SFB 94, Hamburg, unterstützt.     

Möglicher einfluß von feinden auf das verhalten von Diadema-seeigeln

The diurnal rhythms of the sea urchin Diadema setosum differ in coral-reef and port populations at Eilat, Israel. In the port basin (built in 1967), predators of D. setosum are not present. The locomotory activity of reef and port sea-urchins was measured in the field, using SCUBA equipment. During the day, reef sea-urchins sit motionless in sheltered areas; at night, they crawl around and search for food. In contrast, port sea-urchins are active during 24 h. The activity pattern of D. setosum seems to depend on the presence or absence of predators. Apparently, the sea urchins which entered the newly-built harbour have adapted to their predatorless environment. It is not known how this fast adaptation came about; possible explanations are discussed.     

Coexistence and reproductive isolation of the sympatric echinoids<Emphasis Type="Italic"> Diadema savignyi</Emphasis> Michelin and<Emphasis Type="Italic"> Diadema setosum</Emphasis> (Leske) on Kenyan coral reefs

The sympatric echinoids Diadema savignyi and D. setosum coexist in shallow reef lagoons throughout East Africa. The reproductive strategies of these echinoids were studied to investigate reproductive isolation as a possible mechanism for maintaining the coexistence of these closely related species. The annual reproductive cycle and lunar periodicity were determined by gonad index measurements, histological examination of gametogenesis, and induction of spawning with injections of KCl. The peak reproductive period of D. savignyi coincided with the north-east monsoon period (when light and temperatures are high) as gonad indices were high (>8%) beginning in February and peaked at 9.7% in May. Gonad indices subsequently rapidly decreased (by 26%) in June at the beginning of the cooler south-east monsoon period. However, the presence of sperm and ova in most months of the year indicates continuous gametogenesis with reduced reproductive effort during the cooler months. The annual cycle for D. setosum showed less of a seasonal trend as gonad indices remained above 7% throughout much of the year but tended to be highest when temperatures were lower. This is the first confirmation of continuous reproduction in these two species at the equator. The reproductive patterns of both species remained consistent over 2 years of sampling. Both species exhibited a synchronized lunar spawning periodicity during the 3 months sampled, with D. setosum spawning on lunar days 8–10 and D. savignyi spawning after the full moon (lunar days 17–18). Whereas spawning in D. savignyi was very tightly synchronized, 20% of D. setosum individuals still spawned after the peak spawning period. The coexistence of these closely related species appears to be maintained by temporal reproductive isolation during the lunar spawning period reinforced by seasonal differences in reproductive effort.Communicated by O. Kinne, Oldendorf/Luhe     

Die verbreitung der uferwanzen (Heteroptera: Saldidae) im brackigen und marinen litoral der nordamerikanischen pazifikküste

Shore bugs (Heteroptera: Saldidae) were collected during summer 1973 along the Pacific Coast of North America between the arctic and the subtropical regions. The field studies were aimed at determining the species-specific upper and lower limits of distribution in the littoral, and the distribution range of these species in estuaries. The saldids inhabiting the littoral zones can be divided into two groups: those species which live mainly along the coast (coastal species), and those which live mainly inland (inland species). Species found in the supralittoral occur at inland localities as well, and thus can either tolerate limnetic-oligohaline conditions (Salda littoralis, S. provancheri, Saldula coxalis) or are confined to habitats along salt lakes (Pentacora signoreti). Inhabitants of the eulittoral can either occur in the supralittoral and inland localities as well and are holeuryhaline (Saldula palustris, S. pallipes), or have disconnected inland populations (S. nigrita), or they live exclusively in the intertidal zones. In the subarctic, coastal species are distributed from the intertidal zones to inland limnetic habitats. The increasing aridity of southern climatic zones may act as a limiting factor confining the distribution of coastal species to the coast. On the other hand, the distribution of inland species may be influenced by competition with the coastal forms.

---

---

Mit dankenswerter Unterstützung der Deutschen Forschungsgemeinschaft.     

237Pu experiments with the plaice Pleuronectes platessa

Some experiments have been made on the metabolism of ²³⁷Pu — a high specific activity, gamma-emitting isotope of plutonium — by the plaice Pleuronectes platessa L. Very little of the isotope was incorporated into fish after 2 months exposure to labelled water. The oral retention from a variety of labelled foods was also very poor, and apart from the gut itself no incorporation of the isotope could, in fact, be demonstrated. Injected ²³⁷Pu was eliminated very slowly, and some evidence was obtained for an inverse relationship between the rate of ²³⁷Pu loss and the rate of growth of the fish. Growing fish incorporated a relatively larger fraction of the ²³⁷Pu body burden into skeletal material, and this was attained at the expense of the isotopic content of the liver. Very little ²³⁷Pu was incorporated into muscle.     

Characteristic features of body composition and metabolism in some interzonal copepods

Protein, lipid, phosphorus, and organic carbon contents, as well as electron transport system (ETS) activity, lactatedehydrogenase activity, and gut evacuation rate, were measured in four interzonal species of Pacific copepods:Calanus australis, C. pacificus, Eucalanus inermis, andE. elongatus f.hyalinus, collected at the upwelling areas off Peru (8°S) and California (27°N), and in the middle of the North Pacific (30°N), from February to April 1987. The two Eucalanidae species —E. inermis andE. elongatus — have distinctive biochemical and elemental body composition and rates of main physiological processes. Relative protein, lipid, phosphorus, and organic carbon contents (µg mg^–1 wet weight) in these species were, respectively, ca. 1/7 to 1/10, 1/5 to 1/20, 1/5 to 1/10, and 1/5 those inCalanus spp. Likewise, oxygen uptake rate per unit of wet weight (based on ETS activity) inE. inermis andE. elongatus was 5 to 10% of that in calanids; a similar difference was found in phosphorus excretion rate. In addition, gut evacuation rates inE. inermis andE. elongatus were ca. one-fifth of those inCalanus spp. Based on these data, we considered the eucalanids as belonging to a distinctive physiological group, figuratively named jelly-body copepods. In contrast with calanids, active lactatedehydrogenase has been found in the bodies ofE. inermis andE. elongatus, apparently allowing them to survive for a long time in layers of extremely low oxygen content (<0.2 ml l^–1). The adaptive value of physiological features in these eucalanids and typical calanids is compared.     

Feeding and respiration rates of a planktonic copepod (<Emphasis Type="Italic">Calanus sinicus</Emphasis>) oversummering in Yellow Sea Cold Bottom Waters

The distribution, feeding and oxygen consumption of Calanus sinicus were studied in August 2001 on a transect across Yellow Sea Cold Bottom Waters (YSCBW) and two additional transects nearby. The distribution of C. sinicus adults and copepodites stage CV appeared to be well correlated with water temperature. They tended to concentrate in the YSCBW (>10,000 ind. m^–2) to avoid high surface temperature. Gut pigment contents varied from 0.44 to 2.53 ng chlorophyll a equivalents (chl a equiv.) ind.^–1 for adults, and from 0.24 to 2.24 ng chl a equiv. ind.^–1 for CV copepodites. We found no relationship between gut pigment contents and the ambient chl a concentrations. Although the gut evacuation rate constants are consistent with those measured for other copepods, their low gut pigment contents meant an estimated daily herbivorous ingestion of <3% of body carbon in the YSCBW and <10% outside the YSCBW. However, based on estimates of clearance rates, C. sinicus feeds actively whether in the YSCBW or not, so the low ingestion rates probably reflect shortage of food. Oxygen consumption rates of C. sinicus ranged from 0.21 to 0.84 l O₂ ind.^–1 h^–1, with high rates often associated with high temperature. From the oxygen consumption rates, daily loss of body carbon was estimated to be 4.0–13.7%, which exceeds our estimates of their carbon ingestion rates. C. sinicus was probably not in diapause, either within or outside the YSCBW, but this cold-water layer provides C. sinicus with a refuge to live through the hot, low-food summer.Communicated by T. Ikeda, Hakodate     

Natural food,foregut clearance-rate and activity of the crab Scylla serrata

The natural diet, rate of foregut clearance and diurnal activity of the crab Scylla serrata were determined. The gut volume is related to size of crab as gut volume (ml)=0.07e^0.033x , where x=carapace width in millimetres. Fifty per cent of crabs collected in Australia and South Africa contained molluscan remains and about 21% contained crustacean remains — chiefly grapsid crabs. Fish remains were rarely found, and it was concluded that S. serrata does not normally catch mobile forms such as fish and penaeid prawns. Gut clearance of organic tissue was rapid and almost complete after 12 h. Fish bone was retained for a mean time of 2 to 3 days, and shell for 5 to 6 days. Time-lapse photography, using infra-red light, was used to record activity. Visible light flashes reduced activity. S. serrata remained buried during the day, emerging at sunset to spend the night feeding, which occurred intermittently even when unlimited food was available. If no food was present the amount of time spent on the substrate surface was halved.     

Multifarious activities of gut epithelium in an appendicularian (Oikopleura dioica: Tunicata)

The functions of the various gut regions of Oikopleura dioica (oesophagus, left and right gastric lobes, vertical intestine, mid-intestine and rectum) were investigated by means of histochemical, histoenzymatic and immunohistochemical techniques at light and electron microscopes. Ciliary food progression is evidenced by the presence of ATPases on cilia, along the entire gut, with the exception of the cardiac valve, a passive device controlling food direction. Absorptive processes (alkaline phosphatase), active transport (ATPases) and nitrogen excretion (D-amino acid oxidase) occur along the entire gut, in both ciliated microvillar and globular cells. The latter, typical of the left gastric lobe and rectum, are also involved in endocytotic processes (exogenous peroxidase as tracer) and intracellular digestion (!-amylase, aminopeptidase M, acid phosphatase, 5'-nucleotidase, non-specific esterase). The giant cells of the gastric band participate in extracellular digestion; they contain secretory granules positive to various hydrolytic enzymes, the activity of which is also recognisable in faecal pellets inside the intestinal lumen. Lipid storage occurs mainly in the right gastric lobe and vertical intestine, whereas protein storage takes place in the rectal granular cells. Epithelial transport and possible osmoregulation occur along the entire gut, especially at the level of diffuse baso-lateral interdigitations, which increase the plasmalemma surface enormously, are often associated with mitochondria and possess numerous ATPase pumps. Data extend previous histological observations and hypotheses on the physiological role of the various gut regions. The remarkable and specific location of enzymatic activities and nutrient storage are in agreement with the high capacity of O. dioica to process a great quantity of food very rapidly and efficiently.     

Physiologische anpassungen eines marinen insekts. I. Die zeitliche steuerung der entwicklung

The Baltic populations of Clunio marinus (Chironomidae, Diptera) differ from their Atlantic counterparts in three ways: they neither pupate nor hatch in synchrony with a lunar rhythm; they live in a permanently submersed habitat; the structures of their gelatinous egg-masses are different. C. marinus produces two generations each year. The spring generation, which hibernates in the 4th larval stage, hatches in May; the summer generation hatches at the end of July or beginning of August; hatching periods last only a few days. In the laboratory, a permanent short-day period (8 h light, 16 h dark; LD 8:16) induces larval quiescence (oligopause), which can be terminated by long-day conditions (ca. LD 16:8). If larval populations are exposed to continuous light (LL) for 4 days every 30 days, particularly large numbers of imagos hatch about the time of the next LL-period. Increased illumination may influence the time of hatching. Raising the temperature for a few days does not affect hatching. The roles of photoperiod, illumination and temperature are discussed with regard to the temporal programming of development and the start of mass hatching in the natural habitat.

---

---

Herrn Professor Dr. D. Neumann danke ich für die Anregung und Betreuung dieser Arbeit, die Teil einer Dissertation ist.

---

Communicated by O. Kinne, Hamburg