Historical baselines and a century of change in the demersal fish assemblages on South Africa's Agulhas Bank

Doctoral Thesis

2017

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University of Cape Town

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Accurate interpretation of recent variability observed in fish populations, species compositions, distribution ranges or ecological indicators, depends on knowledge of their past dynamics and historical states. The onset of systematic fisheries data collection typically lagged decades or even centuries behind modern fishing exploits. As a consequence, pre-disturbed reference points and descriptions of subsequent change are rarely available. A remarkably detailed set of historical trawl survey data from South Africa provided such a rare opportunity. Government-funded exploration of the Agulhas Bank fishery potential resulted in meticulously-documented trawl survey data from 1897-1904, when prior human impacts on those resources were negligible. Although they used less effective technology, the information recorded and methods used were similar to modern surveys. This thesis investigated change in demersal fish fauna of the Agulhas Bank and documents comparisons between historical trawl surveys and modern re-enactments at the same locations. In comparing trawl survey catches over multiple decades or among different periods, unquantified changes in fishing power pose a key challenge. The shape, size, materials, mesh sizes and speed at which trawl nets are dragged, interact with the behaviour, size and shape of fish, influencing fishing performance. To accurately compare current catch rates with those of historical trawl surveys, the same trawl gear and methods were carefully replicated in repeat surveys at three sites. An investigation of literature and photographs of the original vessel and equipment were conducted to support the construction of a replica 'Granton' otter trawl net. The net was composed of Manila hemp with a headline length of 27 m (90 ft) and was attached to flat wooden trawl boards. The historical towing speed was estimated as 1.34 m s⁻¹ (2.6 knots). Three parts of the shallow Agulhas Bank that were surveyed 111 years prior, were re-surveyed in 2015. Species composition was contrasted between the historical and re-survey periods by way of unconstrained ordination, permutational multivariate analysis of variance and tests of the homogeneity of multivariate group dispersions. Taxa discerning between periods were identified with similarity percentage analyses. Changes of standardised catch between periods were tested for 27 taxa, using a non-parametric bootstrap approach. Proportions among size-classes, recorded for three taxa, were tested using Fisher's exact test. Results revealed a substantially transformed demersal catch assemblage, where the period effect explained almost half of the measured variance among samples. These changes included the disappearance or heavy depletion of kob (Argyrosomus spp., absent in re-surveys), panga (Pterogymnus laniarius; 2.4% of historical catch abundance) and east coast sole (Austroglossus pectoralis; 4.6% of historical catch abundance), which had jointly contributed 70-84% of historical catch composition. Average re-survey catches were largely made up of gurnards (Chelidonichthys spp.; 3 792% of historical abundance), horse mackerel (Trachurus trachurus; 4 738% of historical abundance), spiny dogfish (Squalus spp.; 3 121% of historical abundance), hake (Merluccius capensis; 558% of historical abundance) and white sea catfish (Galeichthys feliceps; 13 863% of historical abundance). Analysis of available length information confirmed the expectation that fish sizes (specifically M. capensis and A. pectoralis) had declined. This implies that comparisons by weight would be more severe for declined abundances and less severe for those that increased, relative to contrasts of numerical abundance. Habitat preferences as well as geographic and depth distribution appeared to separate the taxa that increased from those that declined. These factors, together with reproductive and growth characteristics, as well as indirect trophic impacts, likely shaped the responses of demersal fauna to fishing and other human impacts during the 111 years between trawl surveys. An assessment of distribution changes of 44 common demersal taxa was undertaken. These analyses were restricted to the last 30 years of trawl survey data as the units and spatio-temporal resolution of prior data were incompatible. Standardised catches were used from annual spring and autumn south coast trawl surveys conducted by the government fisheries department. Geostatistical delta-generalised linear mixed models were used to predict species distribution functions, which were used to calculate annual estimates of latitude/longitude centres of gravity and effective areas occupied by each population. Average trends over the study period (1986-2016) were assessed using a Bayesian state-space model. Of nine species found to have a trend in average location, six moved westward or south-westward, while three moved eastward or north-eastward. Two species showed a trend of contracting spatial extent and one showed an expansion. Across the entire assemblage combined, there was a significant contraction in extent and a westward shift in average location. These assemblage-wide average trends are interpreted to be driven by climate forcing. Fishing impacts are expected to have contributed to the eastward movement in centre of gravity for kob, lesser sandshark (Rhinobatos annulatus) and white stumpnose (Rhabdosargus globiceps). Interpretation of these distribution shifts is hampered by a lack of knowledge on subsurface hydrographic trends on the Agulhas Bank, which is identified as a research priority. My research revealed substantial change in demersal fish communities on South Africa's Agulhas Bank and adds novel insight to the history underlying current states of demersal ecosystems. Valuable additions include a) documentation of the extent to which demersal assemblages have transformed during the last century at three representative inshore sites; b) estimates of current abundances relative to pre-disturbed baselines at those sites, which c) highlights drastic local depletion for certain taxa and d) reveals substantial abundance increases of certain species during the post-industrial fishing period; e) novel evidence of distribution trends in south coast demersal species; and f) identification of trends in the average distribution of the demersal fish assemblage, suspected to be climate-forced. Long-term comparisons, using minimally-disturbed baselines, revealed drastic transformation of the fish assemblage during a century of industrialisation, which points to trawling-induced alteration of benthic habitats and substantial changes in ecosystem structure. Besides the provision of novel historical context for current and future studies and decision-making, this work counters the erosive nature of shifting baselines in South Africa's marine environment.
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