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Within-Species Relationship of Patchiness to Both Abundance And

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Within-Species Relationship of Patchiness to Both Abundance And Oecologia (2021) 196:1107–1117 https://doi.org/10.1007/s00442-021-04985-w COMMUNITY ECOLOGY – ORIGINAL RESEARCH Within‑species relationship of patchiness to both abundance and occupancy, as exemplifed by seagrass macrobenthos R. S. K. Barnes1,2,3 Received: 21 November 2020 / Accepted: 5 July 2021 / Published online: 9 July 2021 © The Author(s) 2021 Abstract For the frst time, intraspecifc relationships between the macroecological metrics patchiness (P) and both abundance (A) and occupancy (O) were investigated in a faunal assemblage. As a companion study to recent work on interspecifc P, A and O patterns at the same localities, intraspecifc patterns were documented within each of the more dominant invertebrates forming the seagrass macrobenthos of warm–temperate Knysna estuarine bay (South Africa) and of sub-tropical Moreton Bay (Australia). As displayed interspecifcally, individual species showed strong A–O patterns (mean scaling coefcient − 0.76 and mean R2 > 0.8). All P–O relations were negative and most (67%) were statistically signifcant, although weaker (mean R2 0.5) than A–O ones; most P–A ones were also negative but fewer (43%) achieved signifcance, and were even weaker (mean R2 0.4); 33% of species showed no signifcant interrelations of either O or A with P. No species showed only a signifcant P–A relationship. Compared with interspecifc P–A–O data from the same assemblages, power–law scaling exponents were equivalent, but R 2 values were larger. Larviparous species comprised 70% of the total studied, but 94% of those displaying signifcant patchiness interrelationships; 5 of the 9 showing no P–A or P–O relationships, however, were also larviparous. At Knysna, though not in Moreton Bay, larviparous species also showed higher levels of occupancy than non-larviparous ones, whilst non-larviparous species showed higher levels of patchiness. Dominant Moreton Bay species, but not those at Knysna, exhibited homogeneously sloped P–O relationships. Keywords Patchiness · Occupancy · Seagrass · Macrobenthos · Larvipary Introduction abundance per unit area are also well-known to be related to those of occupancy (i.e. frequency of occurrence), the Neither resources nor organisms are dispersed evenly across more widely an organism is distributed the more abundant space. In efectively all cases, some regions support concen- it is also likely to be (see Hansky 1982; Gaston et al. 2000, trations of any given item whilst other regions are areas of 2006). Indeed in a number of cases occupancy can be pre- scarcity. Areas of abundance occur across wide ranges of cisely estimated from knowledge of mean density and the spatial scales, and may vary in their size, in distance from associated variance (He and Gaston 2003; Gaston et al. other such concentrations, and in the supported intensity 2006). This relationship is an important link between eco- or density of the items concerned. Diferential levels of logical pattern and process (Freckleton et al. 2005), and as such is potentially of considerable importance in understand- ing ecological assembly processes, species distribution pat- Communicated by Bryan Brown. terns, and requirements for conservation of individual spe- * R. S. K. Barnes cies and whole assemblages, including in the prediction of [email protected] responses to such threats as global warming, habitat destruc- tion, etc. (dos Anjos et al. 2011; Manne and Veit 2020). 1 Department of Zoology and Entomology, Rhodes University, Areas of high relative abundance may constitute patches Makhanda, Eastern Cape, Republic of South Africa and several widely used measures of patchiness (i.e. the degree 2 School of Biological Sciences and Centre for Marine of inequality of spatial distribution of abundance) are also Science, University of Queensland, Brisbane, QLD, Australia based on expressions involving the same two metrics that may 3 Department of Zoology and Conservation Research Institute, permit the accurate estimation of occupancy, mean density University of Cambridge, Cambridge, UK Vol.:(0123456789)1 3 1108 Oecologia (2021) 196:1107–1117 and the associated variance (e.g. Morisita 1959, 1962; Lloyd Methods 1967). Recently the author sought to align such patchiness with the interspecifc (i.e. between-species) macroecological Fifty two datasets were available for analysis from two abundance–occupancy relationship (Barnes 2019, 2021). A localities: (A) 26 detailing the abundance of the 12 most preliminary study of the interspecifc relationship between numerous and widespread individual component species patchiness, abundance and occupancy was conducted using of the seagrass macrobenthos from apparently uniform the component macrobenthic species of the intertidal beds of (i.e. non-patchy) local areas of seagrass (each < 1 ha in seagrass, Zostera capensis, within the warm–temperate South size) along the Rainbow Channel coast of North Strad- African Knysna estuarine bay (Barnes 2019). This work was broke Island (Minjerribah) in the Moreton Bay (Quan- then broadened by investigating in comparable fashion the damooka) Marine Park, Queensland, Australia; and (B) equivalent macrobenthic assemblages associated with inter- the same number containing similar information for the tidal beds of six other seagrass species (Cymodocea serratula, 15 equivalent species in non-fragmented seagrass beds in Halodule uninervis, Halophila ovalis, Zostera muelleri and the Knysna estuarine bay, Garden Route National Park, Z. noltei) in sub-tropical Moreton Bay, Queensland, and on Western Cape, South Africa (see Electronic Supplemen- cool-temperate Scolt Head Island in the North Sea, together tary Information 1). All datasets from each locality were with subtidal beds of the Z. capensis at Knysna (Barnes 2021). located within a linear distance of 7 km, and were from Notwithstanding the great diferences in overall macrofaunal a single uninterrupted expanse of coastal seagrass across −2 abundance (> 60,000 to < 2000 ind m ), overall species rich- which dispersal throughout the whole locality was pos- ness (> 200 to < 30), species composition, and habitat features sible. The qualifcation for inclusion of a species was its of these assemblages, as well as their latitudinal and longi- presence in at least ten datasets in abundances of > 35 m−2 tudinal separation, all showed a consistent interspecifc rela- in low-density Moreton Bay or of > 80 m−2 in higher-den- tionship between patchiness, abundance and occupancy. The sity Knysna, and occurrences of > 1 ind. in any single core more abundant and widespread the species, the less was its sample in all qualifying datasets. Analysis was therefore patchiness, with a signifcant negative patchiness–occupancy confned to relatively common species, not only because relationship in the form of a power law with a mean scaling confdence in nature of dispersion assessed from relatively coefcient of − 0.76. Notwithstanding the close relationship few samples is low (Green 1966) but also because abun- of abundance and occupancy, that of patchiness to abun- dance–occupancy relationships may be diferent in rare dance, however, was less marked and not always statistically versus widespread species (Borregaard and Rahbek 2006; signifcant. Bijleveld et al. 2018), regression slopes being shallower Abundance–occupancy relationships within individual and coefcients of determination (R2) weaker in the rare species (i.e. intraspecifc patterns) are seemingly less com- category (Freckleton et al. 2006; Buckley and Freckleton mon and less uniform than interspecifc ones (Gaston 1999; 2010). Although the intertidal and immediately subtidal Gaston et al. 2001). Indeed Bijleveld et al. (2018) studying shores of the two localities were both dominated by con- an equivalent assemblage of Wadden Sea macrobenthic spe- tinuous swards of dwarf-eelgrasses (Zostera subgenus Zos- cies to those investigated by Barnes (2019, 2021) in Knysna terella = Nanozostera in the revision of Coyer et al. 2013), and Moreton Bay, found many very poor abundance–occu- there were diferences between them. At Knysna, shores pancy relationships within various individual polychaete, varied widely in form, from extensive, tide-washed fats gastropod and bivalve species. Such variation across species, of marine sands at the mouth to steeply sloping banks of however, has the potential to help understand causality of the soft estuarine mud at the head, whilst those in Moreton abundance–occupancy relationship (Buckley and Freckleton Bay were much more uniform, extensive mangrove-backed 2010; Gaston et al. 2001; Freckleton et al. 2005; Verberk et al. fats. 2010). As it may also illuminate relations with patchiness, this Sampling of these two seagrass macrobenthic assem- research revisits the databases underlying the earlier interspe- blages used the same methodology, involving series of cifc patchiness–abundance–occupancy analyses to present a core samples (0.0054 m2 area; 100 mm depth), with a corresponding investigation of the nature and magnitude of minimum 30 cores per dataset at Knysna and 45 in More- intraspecifc relationships within the individual component ton Bay. Intertidal samples were collected at low tide species of those same seagrass assemblages. before complete tidal ebb whilst the substratum was still covered by at least 15 cm of water, and the subtidal ones (some 1.5 m below low water spring level) by snorkel- ling. Subtidal and intertidal assemblages difered quan- titatively but not qualitatively (Barnes and Claassens 2020). Cores were gently sieved (’puddled’) through
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