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And Macrozooplankton Time-Series 1974 to 2004: Lessons from 30 Years of Single Spot, High Frequency Sampling at the Only Off-Shore Island of the North Sea

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And Macrozooplankton Time-Series 1974 to 2004: Lessons from 30 Years of Single Spot, High Frequency Sampling at the Only Off-Shore Island of the North Sea Helgol Mar Res (2004) 58: 274–288 DOI 10.1007/s10152-004-0191-5 ORIGINAL ARTICLE Wulf Greve Æ Frank Reiners Æ Jutta Nast Sven Hoffmann Helgoland Roads meso- and macrozooplankton time-series 1974 to 2004: lessons from 30 years of single spot, high frequency sampling at the only off-shore island of the North Sea Received: 26 September 2003 / Revised: 29 June 2004 / Accepted: 10 July 2004 / Published online: 11 November 2004 Ó Springer-Verlag and AWI 2004 Abstract The Helgoland Roads meso- and macrozoo- subjects in ecosystem research. The degree of the plankton time-series 1974 to 2004 is a high frequency understanding of such processes is expressed in analyt- (every Monday, Wednesday and Friday), fixed position ical and operative models predicting changes in abun- monitoring and research programme. The distance to dance distributions. The rules of change concern the coastline reduces terrestrial and anthropological physicochemical forcing, physiological response mecha- disturbances and permits the use of Helgoland Roads nisms and trophodynamic processes. Significant time- data as indicators of the surrounding German Bight scales can cover several orders of magnitude. Therefore, plankton populations. The sampling, determination and causal time-series analysis has to rely on data sets from IT methodologies are given, as well as examples of an- various sources. While experimental time-series may nual succession, and inter-annual population dynamics operate informatively on less complex systems (e.g. of resident and immigrant populations. Special attention Greve 1995a), in situ time-series are real and their doc- is given to the phenology and seasonality of zooplank- umentation and analysis are most important (Perry et al. ton populations. The influence of winter sea surface 2004). The Helgoland Roads station enables the con- temperature on the seasonality of spawning of the tinuous monitoring of marine processes in an offshore common sole Solea solea is given as an example for location with little terrestrial and anthropogenic distur- merozooplankton populations. bance and, due to the Biologische Anstalt Helgoland, affordable sampling capacity is available. Methods Introduction The Helgoland Roads time-series on meso- and macro- A single observation of nature permits the definition of zooplankton was started in 1974 as a spin-off data col- the momentary state. Repeated observations in space or lection stemming from the daily live-food catches from time permit the recognition of distributional rules and the small cutters of the Biologische Anstalt Helgoland variances. Local repetitions of observations produce (BAH) marine station. These zooplankton samples were time-series as a documentation of the system dynamics. provided to the laboratory for cultivation experiments Time-series enable the development of analytical models and for eventual observation. Phytoplankton and defining state variables and rates of processes, and nutrients had been measured every work day since 1962 testing hypothetical rules of change. The definition of (Hickel et al. 1993); the expected availability of this the rules, and variances determining abundance distri- information, the then dominant bottom-up theory and butions of populations and their changes, are the main the measurement of meso- and macrozooplankton led to the expectation that even in a hydrographically complex Communicated by K. Wiltshire locality the trophic controls of zooplankton population abundance and distribution might become evident. The & W. Greve ( ) Æ S. Hoffmann aim was the production of time-series which would German Centre for Marine Biodiversity Research (Senckenberg Research Institute), Germany permit the testing of simulation models on the tropho- E-mail: [email protected] dynamics of the pelagic ecosystem then believed to be F. Reiners Æ J. Nast Æ W. Greve the decisive mechanism for population control. The Federal Marine and Hydrographic basis of the intended modelling in those days was a Agency of Germany, Germany conceptual model of the IBP type (Greve 1972). 275 Sampling, preservation, sorting and counting Table 1 Alphabetic lists of taxonomic and ontogenetic entities (76 mesozooplankton and 367 macrozooplankton) determined and enumerated since 1 April 1994. Further variables have been sepa- The plankton nets commonly used for the sampling of rately documented. Meso- and macrozooplankton have been zooplankton on the small cutter ‘‘Ellenbogen’’ until sampled every Monday, Wednesday and Friday since April 1 1974 April 1974 were not equipped with a counter to and the macrozooplankton counted at least every Wednesday until measure the volume filtered, nor with a conical design 2001 that avoided escape responses of zooplankton Mesozooplankton: approaching the net. These nets were replaced by the Acartia spp. HYDROBIOS quantitative collection device 438040 Actinotrocha (150 lm hand-net, aperture 17 cm, net length 100 cm, Alaurina composita FlowMeter, ballast weight) and the HYDROBIOS ring Amphiura filiformis pluteus Appendicularia trawl net 438700 (Calcofi Net; mesh size 500 lm, Asterias rubens bipinnaria 100 cm aperture, length 400 cm, depressor and Flow- Asterias rubens brachiolaria Meter). Both nets were used for oblique hauls under Autolytus spp. the prevailing conditions of Helgoland Roads, gener- Beroe spp. juvenile Branchiostoma spp. ally not permitting vertical hauls due to a depth of Calanoida only 6 to 8 m. Calanus spp. Sampling was generally carried out at the position Carcinus maenas megalopa 54°11¢18¢¢N7°54¢ E) during morning hours, thereby Carcinus maenas zoea Centropages shifting the sampling by 45 min daily through the tidal Centropages hamatus female cycle. Samples were live-counted until 1981 on a daily Centropages spp. basis. In 1982, sampling was reduced to Mondays, Centropages typicus female Wednesdays and Fridays. Samples are preserved in 4% Cirripedia nauplius Cladocera formaldehyde. As some species (e.g. Bolinopsis infun- Copepoda dibulum) cannot be preserved, occasional live counts Copepoda nauplius were continued for limited time-periods on the basis of Corycaeus spp. plane transports from Helgoland to Hamburg. Crangon crangon zoea The taxonomic and numeric biodiversity was inves- Ctenophora Cyclopina spp. tigated and documented to the earliest possible date, Cyclopoida handling meso- and macrozooplankton separately. Cyphonautes While mesozooplankton was determined by F. Reiners Decapoda and W. Greve only, macrozooplankton was determined Echinodermata Euterpina acutifrons + Tisbe spp. by W. Greve, J. Nast, S. Hoffmann and various other Evadne spp. assistants and students. The list of species and stages Fish egg counted varied with taxonomic changes and increasing Fish larva professional expertise. Fish larvae which were at first Fritillaria borealis Galathea spp. zoea intentionally excluded from taxonomic analysis due to Gastropoda larva the institutional organisation, were determined and Harpacticoida counted for 7 years from 1990. Holoplankton Metazoa continuously change their characteristics Hydrozoa Lamellibranchia larva with growth. Fixation does not preserve the pigments of Lanice spp. fish larvae; they become bleached. Therefore, the taxo- Magelona spp. nomic treatment of the complete size class, especially of Meroplankton macrozooplankton, requires expertise that can only be Microsetella spp. obtained after a long training period. Though this Mitraria Monstrilla spp. training could be supported by the experiences gained, Noctiluca scintillans individual inconsistencies cannot be excluded. Obelia spp. The training was supported by selected literature (e.g. Oikopleura dioica Sars 1903; Zimmer 1909; Grimpe 1911–1942; Pesta 1928; Oithona spp. Ophiura spp. pluteus Russell 1953, 1970, 1976; Williamson 1957; Kermack Pagurus bernhardus zoea 1966ff; Jones 1976; Pierrot-Bults and Chidgey 1988; Paracalanus parvus + Pseudocalanus elongatus Ingle 1992), photographs and drawings, samples from Pectinaria spp. the sorting exercises and a collection of reference sam- Penilia avirostris Pilidium ples. Pleurobrachia pileus juvenile The plankton was sub-sampled according to the Podon spp. number of organisms present. An aliquot of 60 indi- Polychaeta viduals of each key species was regarded as an accept- Polychaeta larva undeterminedo able threshold level. Determination and counting were Protochordata Psamechinus miliaris pluteus carried out in counting chambers using microscopes. 276 Table 1 (Contd.) Table 1 (Contd.) Pseudocalanus elongatus female Branchiostoma spp. Rathkea octopunctata + Lizzia blondina Calanoid copepods sum Rathkea octopunctata + Lizzia blondina not differentiated Calanus spp. copepodit Rotatoria Calanus spp. female Sagitta sum Calanus spp. male Sagitta elegans Calanus spp. not differentiated Sagitta setosa Calanus sum Sagitta spp. Caligidae Spatangoidea pluteus Caligus spp. Spioniden metatrochophora Callianassa subterranea zoea not differentiated Steenstrupia nutans Callianassa subterranea zoea sum Temora longicornis Callianassa subterranea megalopa Temora longicornis nauplius Callianassa zoea sum Macrozooplankton: Callianassa zoea and megalopa sum Agastra mira Cancer pagurus zoea not differentiated Aglantha digitale sum Cancer pagurus zoea sum Agonus cataphractus larva Cancer pagurus megalopa Amphinema dinema Cancer pagurus zoea and megalopa sum Amphinema rugosum Candacia armata not differentiated Amphinema spp. Candacia armata sum Amphinema sum Candacia spp. sum Amphipoda undetermined Candacia sum Amphipoda sum Caprella spp. Anchialis agilis Carcinus
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