The Helgoland Roads time series is one of the richest temporal marine data sets available. Running since 1962, it documents changes for phytoplankton, salinity, Secchi disc depths and macronutrients. Uniquely, the data have been carefully quality controlled and linked to relevant meta-data, and the pelagic time series is further augmented by zooplankton, intertidal macroalgae, macrozoobenthos and bacterioplankton data. Data analyses have shown changes in hydrography and biota around Helgoland. In the late 1970s, water inflows from the south-west to the German Bight increased with a corresponding increase in flushing rates. Salinity and annual mean temperature have also increased since 1962 and the latter by an average of 1.67°C. This has influenced seasonal phytoplankton growth causing significant shifts in diatom densities and the numbers of large diatoms (e. g. Coscinodiscus wailesii). Changes in zooplankton diversity have included the appearance of the ctenophore Mnemiopsis leidyi. The macroalgal community also showed an increase in green algal and a decrease in brown algal species after 1959. Over 30 benthic macrofaunal species have been newly recorded at Helgoland over the last 20 years, with a distinct shift towards southern species. These detailed data provide the basis for long-term analyses of changes on many trophic levels at Helgoland Roads.
The paper briefly summarizes what is known about long-term changes (facts, causes, consequences) in the macrozoobenthos of intertidal and subtidal hardbottom communities around the island of Helgoland (German Bight, North Sea). There is increasing observational evidence that these communities (spectrum and abundances of species) are changing on a long-term temporal scale. The reasons are diverse and mainly anthropogenic. A shift in North Sea climate towards more oceanic conditions may be among the most important factors driving the recent changes in species spectrum. Many of the species which have been recorded as new to the Helgoland area during the past decade are southern (oceanic) species which may be considered as indicators of a warming trend.
Factors controlling the timing of reproduction in laboratory cultures of the polychaete Typosyllls prolifera. Typosyllis prolifera (Krohn) from Pore~ (Yugoslavia) has been cultured for 12 successive generations. The life cycle of the species in the laboratory is described briefly. During their life individuals reproduce several times (up to 15) by stolonization which, under constant laboratory conditions (LD 16:8, 20 °C), is cyclic and takes place about every 30 days. Based on the investigations of Durchon (1959) and Wissocq (1966), experiments on extirpation and transplantation of the proventriculus have been carried out. The results suggest that an endocrine system anatomically connected to the proventriculus is important in the control of reproduction. Most likely, the endogenous reproductive cycle of an individual is controlled by periodical changes of the activity of this system. During the period following stolonization, the endocrine system of the proventriculus, which at this time shows its maximal activity, inhibits sexual development and enables regeneration of the segments lost as stolon. A subsequent decrease of the hormonal activity induces sexual maturation and epitokous metamorphosis, thus leading to further stolonization. Exogenous factors influencing the timing of reproduction probably affect the endocrine function of the proventriculus. Short-day photoperiods (LD 10:14) and low temperatures (12 °C) given simultaneously (i.e. winter conditions) totally suppress reproduction. Under normally favourable conditions (LD 16:8, 20 °C),reproductive processes can be prevented by starving or amputation of caudal segments. In all these cases, however, stolonization can be induced by removing the proventriculus. Exogenous factors also play a decisive role in synchronizing reproductive events within the species population. Under field conditions reproduction shows a lunar periodicity. The endogenous reproductive cycles of cultured specimens can be synchronized by a periodical light regime simulating the change of the moon.
EINLEITUNG
A laboratory-based study was performed to assess the impact of climate warming on the recruitment of the endangered population of the European lobster (Homarus gammarus) at Helgoland (North Sea, German Bight). Egg-bearing females collected in situ just after spawning in late summer were subjected to various seasonal temperature regimes. Regimes with elevated temperatures (mild winters) resulted in a strong seasonal forward shift of larval hatching. Hatching took place at significantly lower temperatures than under regimes with normal winters. Experiments on larval development across a range of constant temperatures showed that no successful larval development occurred at temperatures below 14°C. Larval survival increased from 9% at 14°C to 80% at 22°C, while duration of larval development decreased correspondingly from 26 to 13 days. We hypothesize that an ongoing warming of the North Sea will strongly affect the recruitment success of the Helgoland lobster, mainly resulting from a decoupling of the seasonal peak appearance of larvae from optimal external conditions (temperature, food availability) for larval development.
Habitat segregation among competing species is widespread yet very little is know how this is achieved in practice. In a case study, we examined short-term effects of conspecific and congeneric density on habitat selection in two competing marine isopod species, Idotea emarginata and Idotea baltica. Under semi-natural conditions in large outdoor cylindrical tanks (4 m high; volume 5.5 m 3 ), animal groups of different size and composition had the choice between a set of relevant habitat samples (surface-floating seaweed, the water column, seaweed on the bottom). Habitat selection in both I. baltica and I. emarginata proved to be largely independent of conspecific density (level of intraspecific competition). In single-species treatments, both species showed a similar and stable pattern of distribution, with a clear preference for seaweed on the bottom. In mixed-species treatments (MST), however, the species were largely separated by habitat. While the distribution of I. emarginata was completely unaffected by the mere presence of interspecific competitors, habitat selection of I. baltica changed notably when I. emarginata was present. The habitat use patterns observed in MST conformed to those realized in geographical areas where the two species overlap in distribution: I. emarginata is dominant among decaying seaweed on the sea floor, and I. baltica is the dominant species among surface-floating seaweed. Our findings suggest that habitat segregation between the two species is essentially interactive, resulting from rapid decision-making of I. baltica with respect to habitat selection. The underlying mechanism is discussed. I. emarginata is highly superior to I. baltica in interference competition and rapidly eliminates the latter from one-habitat systems which do not allow I. baltica to escape from this interaction. In more natural, heterogeneous environments, however, I. baltica seems to be able to coexist with the superior competitor due to its broader habitat niche, flexibility in habitat selection, and a behavioural disposition to avoid normally preferred habitats when these are occupied by I. emarginata.
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