New species and records of Laboulbeniales from the subantarctic islands of New Zealand

Until now Rhachomyces kenodactyli Balazuc & W. Rossi has been the only species of Laboulbeniales known to occur on Coleoptera in the Bounty, Antipodes, Auckland, Campbell and Snares Islands, which lie 48° to 35° S. Four new species (Diphymyces depressus, Diphymyces leschenii, Laboulbenia subantarctica and Laboulbenia loxomeri) and five new records for the subantarctic (Cucujomyces phycophilus, Diphymyces penicillifer, Laboulbenia sp. 1, Rhachomyces sp. 1 and Teratomyces sp. 1) are reported, increasing the known number of taxa tenfold. An expanded geographic range for Rhachomyces kenodactyli is reported. A relatively high percentage (12%) of known beetle species in the subantarctic serve as hosts for Laboulbeniales. This host utilization rate is higher than that in tropical and north temperate regions. The high proportion of intertidal coleopteran taxa in the subantarctic fauna probably accounts for the greater number of host species utilized. Fungi on intertidal beetles (Omaliinae [Staphylinidae], Oopterus [Carabidae] and Kenodactylus audouini [Carabidae]) are known from many host individuals and collections, while those on terrestrial species are known from few, and in some cases, a single collection or host. The sporadic occurrence of some species encountered increases the likelihood that a few species of Laboulbeniales on Coleoptera probably remain undiscovered in the region.     

A marine protected area network does not confer community structure resilience to a marine heatwave across coastal ecosystems

Marine protected areas (MPAs) have gained attention as a conservation tool for enhancing ecosystem resilience to climate change. However, empirical evidence explicitly linking MPAs to enhanced ecological resilience is limited and mixed. To better understand whether MPAs can buffer climate impacts, we tested the resistance and recovery of marine communities to the 2014–2016 Northeast Pacific heatwave in the largest scientifically designed MPA network in the world off the coast of California, United States. The network consists of 124 MPAs (48 no-take state marine reserves, and 76 partial-take or special regulation conservation areas) implemented at different times, with full implementation completed in 2012. We compared fish, benthic invertebrate, and macroalgal community structure inside and outside of 13 no-take MPAs across rocky intertidal, kelp forest, shallow reef, and deep reef nearshore habitats in California's Central Coast region from 2007 to 2020. We also explored whether MPA features, including age, size, depth, proportion rock, historic fishing pressure, habitat diversity and richness, connectivity, and fish biomass response ratios (proxy for ecological performance), conferred climate resilience for kelp forest and rocky intertidal habitats spanning 28 MPAs across the full network. Ecological communities dramatically shifted due to the marine heatwave across all four nearshore habitats, and MPAs did not facilitate habitat-wide resistance or recovery. Only in protected rocky intertidal habitats did community structure significantly resist marine heatwave impacts. Community shifts were associated with a pronounced decline in the relative proportion of cold water species and an increase in warm water species. MPA features did not explain resistance or recovery to the marine heatwave. Collectively, our findings suggest that MPAs have limited ability to mitigate the impacts of marine heatwaves on community structure. Given that mechanisms of resilience to climate perturbations are complex, there is a clear need to expand assessments of ecosystem-wide consequences resulting from acute climate-driven perturbations, and the potential role of regulatory protection in mitigating community structure changes.     

Distribution and abundance of fish in a rocky reef environment at the subantarctic Auckland Islands,New Zealand

Summary The distribution and abundance of fish in a rocky reef environment were investigated at the subantarctic Auckland Islands of New Zealand, in June 1986. Fish were counted in transects and specimens were taken. The diversity and abundance of species of large reef fish was low; a total of eight species were observed. Most fish were benthic carnivores. These findings are similar to studies of reef fish in subantarctic waters of Chile. Large differences in species composition were found among locations at the Auckland Islands. The number of species and their abundance was highest near exposed headlands. Few fish were found on reefs at more sheltered inlets. At most locations the nototheniids Paranotothenia angustata and P. microlepidota ranked first in abundance at deep and shallow sites. There were some species that showed differences in abundance with depth. Highest abundance of Bovichthys variegatus, Latridopsis cilaris, Latris lineata and Pseudolabrus cinctus were in deep water on reefs. Large numbers of juvenile P. microlepidota were found only in shallow water. No depth related patterns were found for P. angustata and Mendosoma lineatum. Specimens, other than those counted in transects, were also collected. There were small fish, 4 species of tripterygiids, 1 gobiescocid and 2 syngnathids. The Auckland Islands had closer zoogeographic affinities with islands of temperate and subantarctic New Zealand than with other landmasses of the westwind drift. Although most fish found at the Auckland Islands are found also in temperature New Zealand, the converse of this pattern was not found. The proportional representation of species and trophic groups differed markedly between these regions.     

The influence of coastal upwelling on the functional structure of rocky intertidal communities

Summary Relationships between organisms at all trophic levels are influenced by the primary productivity of the ecosystem, and factors which enhance rates of primary production may modify trophic relationships and community structure. Nutrient enrichment of intertidal and nearshore waters leads to enhanced production by intertidal algae, and it was hypothesized that where rocky shores are washed by nutrient-rich upwelled waters, the intertidal communities should show a characteristic functional structure, based on the effects of enhanced primary production. Study sites were chosen on rocky shores in southern Africa, central Chile and the Canary Islands, in areas with and without coastal upwelling, and mid-shore community structure at these sites was analysed in terms of the abundance of certain functional guilds of organisms.It was found that algal cover and the biomass of herbivorous limpets supported per unit area on rocky shores were significantly greater in regions of coastal upwelling than in regions where upwelling did not occur. Ground cover by sessile filter-feeding organisms was significantly greater on shores in non-upwelled areas. However, correspondence analysis showed no functional aspect of intertidal community structure that was characteristic of coasts washed by upwelled waters. Primary reasons for this are probably the large variations in the nature of nutrient enrichment that accompanies upwelling, and in the nutrient status of non-upwelled areas. Other factors are man's exploitation of intertidal organisms and differences in the genetic origins of the intertidal species involved.     

Wetlands of the Pacific Island region

The wetlands of 21 countries and territories of the Pacific Islands region are reviewed: American Samoa, Cook Islands, Federated States of Micronesia, Fiji, French Polynesia, Guam, Kiribati, Marshall Islands, Nauru, New Caledonia, Niue, Northern Mariana Islands, Palau, Papua New Guinea, Samoa, Solomon Islands, Tokelau, Tonga, Tuvalu, Vanuatu, and Wallis and Futuna. The regions’ wetlands are classified into seven systems: coral reefs, seagrass beds, mangrove swamps, riverine, lacustrine, freshwater swamp forests and marshes. The diversity of species in each of these groups is at near global maxima at the west of the region, with decline towards the east with increasing isolation, and decreasing island size and age. The community structure is unique in each country, and many have endemic species with the habitat isolation that epitomises this island region. There remain, however, some serious gaps in basic inventory, particularly in freshwater biodiversity. Threats to wetlands include introduced freshwater species, loss of wetlands adjacent to urban growth, downstream effects of mining and land clearance, and over-use of mangrove, seagrass and coral reef resources by predominant subsistence economies that remain in this region. Only five countries are signatories to the Ramsar convention on wetlands, and this only recently with seven sites. Wetland managers have identified the need for community education, baseline surveys and monitoring, better legislation and policy for wetland management, and improved capacity of local communities to allow the wise use of their wetlands.     

SPATIAL AND TEMPORAL COMPARISON OF ALGAL BIODIVERSITY AND BENTHIC COVER AT GARDNER PINNACLES,NORTHWESTERN HAWAI`IAN ISLANDS1

Papahānaumokuākea Marine National Monument in the Northwestern Hawai`ian Islands is the second largest marine protected area in the world, providing an opportunity for scientists to understand natural ecosystem fluctuations in subtropical marine communities with little anthropogenic impact. Gardner Pinnacles is composed of two emergent basaltic rocks and has the smallest land area of any island in the Northwestern Hawai`ian Island chain but has among the largest submerged reef area. Gardner Pinnacles is also among the least anthropogenically impacted island in the Hawai`ian Archipelago, although a thriving lobster and bottomfish industry existed in the area for many years. This study assesses nearshore algal biodiversity and percent cover at Gardner Pinnacles to examine interannual differences in community dynamics and places them in an ecosystem context by also examining associated invertebrate and fish communities. Biodiversity surveys increased the number of known eukaryotic algal species occurring in marine environments immediately adjacent to the emergent portion of Gardner Pinnacles from 31 to 77. Algal percent cover, specifically populations of the green alga Microdictyon setchellianum M. Howe, varied dramatically between sampling years, possibly in response to seasonal differences. Towed‐diver surveys revealed that macroalgae covered >90% of the substrate during the 2003 sampling period but returned to 2000 levels (1%–35% cover) by 2004 without any detectable damage to other reef organisms. Fish communities remained statistically similar between sampling years, and percent cover of live coral around the island did not exceed 7%.     

The structure of Mediterranean rocky reef ecosystems across environmental and human gradients, and conservation implications

Historical exploitation of the Mediterranean Sea and the absence of rigorous baselines makes it difficult to evaluate the current health of the marine ecosystems and the efficacy of conservation actions at the ecosystem level. Here we establish the first current baseline and gradient of ecosystem structure of nearshore rocky reefs at the Mediterranean scale. We conducted underwater surveys in 14 marine protected areas and 18 open access sites across the Mediterranean, and across a 31-fold range of fish biomass (from 3.8 to 118 g m(-2)). Our data showed remarkable variation in the structure of rocky reef ecosystems. Multivariate analysis showed three alternative community states: (1) large fish biomass and reefs dominated by non-canopy algae, (2) lower fish biomass but abundant native algal canopies and suspension feeders, and (3) low fish biomass and extensive barrens, with areas covered by turf algae. Our results suggest that the healthiest shallow rocky reef ecosystems in the Mediterranean have both large fish and algal biomass. Protection level and primary production were the only variables significantly correlated to community biomass structure. Fish biomass was significantly larger in well-enforced no-take marine reserves, but there were no significant differences between multi-use marine protected areas (which allow some fishing) and open access areas at the regional scale. The gradients reported here represent a trajectory of degradation that can be used to assess the health of any similar habitat in the Mediterranean, and to evaluate the efficacy of marine protected areas.     

Systematic and conservation implications of geographic variation in pipits (Anthus: Motacillidae) in New Zealand and some offshore islands

The New Zealand Pipit Anthus novaeseelandiae occurs as a single species in Europe (as Richard's Pipit), Asia, Africa, New Guinea, Australia and New Zealand. In the New Zealand region, subspecific status has been accorded to allopatric populations on the mainland (A. n. novaeseelandiae) , on the Auckland and Campbell Islands (A. n. aucklandicus) , on the Antipodes Islands (A. n. steindachneri) and on the Chatham Islands (A. n. chathamensis) . Analyses of 23 allozyme loci and morphometric variation of populations on mainland New Zealand and on the Auckland, Campbell, Antipodes and Chatham Islands showed an appreciable and significant divergence between mainland and island populations. The magnitude of the allozyme difference is sufficient to suggest full species designation for the island birds, a conclusion supported by morphometric analyses and in sharp contrast to current taxonomy. However, formal taxonomic change will not be undertaken until the two New Zealand groups have been compared with more distant populations of this wideranging genus. The cryptic variation found in this species has conservation management implications for the small offshore populations.     

Structure of reef fish communities in Catalinas Islands and Ocotal beach, North Pacific of Costa Rica

The reefs are heterogeneous systems that maintain a high diversity of organisms. Fish community structure varies within and among reefs, so it would be expected that reef structure and heterogeneity should affect fish communities inhabiting reefs. Four reef patches at Catalinas Islands (Sur, La Pared, Roca Sucia and Sombrero) and one in Ocotal beach (10 degrees 28'45" N; 85 degrees 52'35" W) were studied with visual censuses (July-December 2003). The structure and composition of fishes between Catalinas islands and Ocotal beach were different, and habitat structure and composition explain most of the variance founded. The presence of the fleshy algae Caulerpa sertularioides in Ocotal, and the corals Tubastrea coccinea and Pocillopora spp. at Catalinas Island explained the variability among sites and how it affected fish community structure and composition. The butterfly fish Johnrandallia nigrirostris, damselfish Microspathodon dorsalis, and surgeon fish Prionurus punctatus were directly correlated with the ahermatipic coral Tubastrea coccinea in Roca Sucia reef, while the angel fish Holacanthus passer was associated to reefs with a major percentage of rocky substrate. Other species such as the damselfish Abudefduf troschelli and Halichoeres dispilus were more abundant at Ocotal, where the algae C sertularioides dominated. The number and abundance of reef fishes was directly correlated with the rugosity index at the reefs of Roca Sucia and Ocotal, but not at reefs of La Pared and Sombrero.     

Early Devonia benthic communities of the Alexander Terrane, southeastern Alaska

Emsian (upper Lower Devonian) limestones exposed on Kasaan, Round, and Wadleigh Islands in southeastern Alaska are part of an allochthonous suitc of island are deposits preserved within the accrctionary Alexander Terrane. Ten benthic marine communities, including several new brachiopod associations, are defined on the basis of field data correlated with a cluster analysis. Biologic and sedimentologic evidence is integrated with community group assignments in order to substantiate the paleoecology and evolutionary history of each community. These faunas were adapted to a spectrum of quiet and rough water habitats, including restricted, shallow subtidal areas nearshore, offshore biostromal banks, and open and restricted portions of a lagoon or shelf. Comparisons with coeval assemblages from the Cordillera and elsewhere reveal that these communities are unique to southeastern Alaska. Their origin within an early Paleozoic island are appears to have led to biogeographic isolation from other parts of the Cordillera and the development of a distinct biota at a site offshorc from the ancient continental margin. ▭ Emsian, paleocommunities, community groups, cluster analysis, Alexander Terrane.     

Implications of reef ecosystem change for the stability and maintenance of coral reef islands

Coral reef islands are among the most vulnerable environments on Earth to climate change because they are low lying and largely constructed from unconsolidated sediments that can be readily reworked by waves and currents. These sediments derive entirely from surrounding coral reef and reef flat environments and are thus highly sensitive to ecological transitions that may modify reef community composition and productivity. How such modifications – driven by anthropogenic disturbances and on‐going and projected climatic and environmental change – will impact reef island sediment supply and geomorphic stability remains a critical but poorly resolved question. Here, we review the unique ecological–geomorphological linkages that underpin this question and, using different scenarios of environmental change for which reef sediment production responses can be projected, explore the likely resilience of different island types. In general, sand‐dominated islands are likely to be less resilient than those dominated by rubble grade material. However, because different islands typically have different dominant sediment constituents (usually either coral, benthic foraminifera or Halimeda) and because these respond differently to individual ecological disturbances, island resilience is likely to be highly variable. Islands composed of coral sands are likely to undergo major morphological change under most near‐future ecological change scenarios, while those dominated by Halimeda may be more resilient. Islands composed predominantly of benthic foraminifera (a common state through the Pacific region) are likely to exhibit varying degrees of resilience depending upon the precise combination of ecological disturbances faced. The study demonstrates the critical need for further research bridging the ecological–geomorphological divide to understand: (1) sediment production responses to different ecological and environmental change scenarios; and (2) dependant landform vulnerability.     

Marine biological community baselines in unimpacted tropical ecosystems: spatial and temporal analysis of reefs at Howland and Baker Islands

Howland and Baker Islands are two small, isolated reef and sand islets located near the equator in the central Pacific Ocean that are situated approximately 60 km apart. In 2004 and 2006, species-level monitoring at multiple sites, coupled with towed-diver surveys in 2002, 2004, and 2006 on both of these federally protected islands, revealed diverse fish, coral, macroinvertebrate, and algal assemblages. This study examines inter- and intra-island spatial and temporal differences in community composition among sites and presents baseline biological community parameters for two of the least impacted reef systems in the world. Despite similarities in species composition, permutational multivariate analysis of variance (PERMANOVA) and multidimensional scaling ordinations (nMDS) suggest biological communities at the two islands are distinct with Baker Island containing a greater percent cover of branched Acroporid corals and turf algae and Howland Island containing a greater percent cover of crustose coralline red algae and small, compact genera of coral. Both islands also contained considerable cover of non-invasive macroalgae. PERMANOVA further revealed benthic and fish species composition to differ between forereef and reef shelf sites from different sides of each island. When islands were considered as a whole, temporal changes were not noted between 2004 and 2006; however, temporal changes at select sites did occur, with coral cover decreasing significantly along the west side of Baker Island from 2004 to 2006.     

The ‘island mass’ effect on the phytoplankton and primary production of the Hawaiian Islands

A 170 station oceanographic reconnaissance in the nearshore waters of the Hawaiian Islands has emphasized the ‘island mass’ effect in increasing primary production and chlorophyll a standing crop as well as in altering community assimilation ratios. The distribution of oceanographic parameters and micronutrients indicate that one of the enriching mechanisms is turbulence and vertical mixing in island channels.The species composition of the net phytoplankton communities characteristic of inshore, offshore, and oceanic stations is given and the relative importance of the nannoplankton component discussed. The nannoplankton dominated in all regions, with a tendency for the net/nannoplankton ratio to increase toward the base of the euphotic zone. The net phytoplankton consisted chiefly of open ocean diatom species with neritic species restricted to a narrow inshore environment.     

The vegetation of the Farasān Islands,Red Sea,Saudi Arabia

Abstract. Classification of the vegetation of the Farasān Islands using TWINSPAN technique resulted in the recognition of seven community types associated with seven different habitat types: silty runnels, palm orchards, rocky plains, rocky plateau crevices, coastal sand dunes, sand plains, and mangroves. These communities were dominated or co-dominated by 13 perennial species; 87 associate species were recorded in the study area with chamaephytes dominating the life-form spectrum. Canonical Correspondence Analysis (CCA) indicated that organic carbon, soil moisture, silt, electrical conductivity and calcium carbonate were the major edaphic gradients controlling the distribution of the plant communities on the Farasān Islands. Higher species richness was recorded in the plant communities inhabiting the palm orchards, the crevices of the rocky plateau, silty runnels, rocky plains and rocky plateau, while those of the mangrove and sand formations (dunes and plains) showed a lower species richness. Soil texture and organic matter are the main factors promoting species diversity in the more diversified habitats, while high salinity and calcium carbonate are the main factors associated with lower species diversity in the less diversified habitats of the study area.     

An all-taxon microbial inventory of the Moorea coral reef ecosystem

The Moorea Coral Reef Long Term Ecological Research (LTER) Site (17.50°S, 149.83°W) comprises the fringe of coral reefs and lagoons surrounding the volcanic island of Moorea in the Society Islands of French Polynesia. As part of our Microbial Inventory Research Across Diverse Aquatic LTERS biodiversity inventory project, we characterized microbial community composition across all three domains of life using amplicon pyrosequencing of the V6 (bacterial and archaeal) and V9 (eukaryotic) hypervariable regions of small-subunit ribosomal RNA genes. Our survey spanned eight locations along a 130-km transect from the reef lagoon to the open ocean to examine changes in communities along inshore to offshore gradients. Our results illustrate consistent community differentiation between inshore and offshore ecosystems across all three domains, with greater richness in all domains in the reef-associated habitats. Bacterial communities were more homogenous among open ocean sites spanning >100 km than among inshore sites separated by <1 km, whereas eukaryotic communities varied more offshore than inshore, and archaea showed more equal levels of dissimilarity among subhabitats. We identified signature communities representative of specific geographic and geochemical milieu, and characterized co-occurrence patterns of specific microbial taxa within the inshore ecosystem including several bacterial groups that persist in geographical niches across time. Bacterial and archaeal communities were dominated by few abundant taxa but spatial patterning was consistent through time and space in both rare and abundant communities. This is the first in-depth inventory analysis of biogeographic variation of all three microbial domains within a coral reef ecosystem.     

Arthropoda of the subantarctic islands of New Zealand

Ischyroplectron isolatum (Hutton) is redescribed from the Bounty Islands, south‐east of New Zealand. The relationships of the subantarctic Orthoptera are discussed. A key is given to the four genera of Rhaphidophoridae.     

Depth-associated patterns in the development of benthic assemblages on artificial substrata deployed on shallow, subtropical reefs

Changes in the structure of many benthic habitats occur across a number of physical gradients and result in corresponding changes in the structure of associated epifaunal assemblages; however, investigations of faunal assemblages are often confounded by variation in the morphology of habitats. In this experiment, identical nests of nylon pan scourers were employed to examine changes in the structure of epifaunal assemblages across a depth gradient at two island sites within the Solitary Islands Marine Park (SIMP), NSW, Australia. Artificial substratum units (ASUs) were anchored to rocky reef at 8, 16 and 24 m for a period of five months over summer and winter. Data were subjected to univariate and multivariate statistical analyses to determine the similarity of assemblages across Depths, Islands and Times. A number of species displayed a distinct fidelity with depth across both islands and times. Although significant interactions between factors were apparent for most variables, very few significant differences across the main effects were identified for univariate analyses of summary community variables (S, N, H′), major taxonomic groups (bivalves, amphipods, polychaetes) or individual species analysed. In contrast, multivariate analyses revealed significant differences in assemblage structure for all comparisons of depth during each sampling period. Although the experiment was conducted both over summer and again over winter, depth-associated patterns were maintained at each island during each sampling period. The results highlight the importance of depth as a structuring factor for epifaunal assemblages of subtropical rocky reefs.     

Controls on coral-ground development along the northern Mesoamerican Reef tract

Coral-grounds are reef communities that colonize rocky substratum but do not form framework or three-dimensional reef structures. To investigate why, we used video transects and underwater photography to determine the composition, structure and status of a coral-ground community located on the edge of a rocky terrace in front of a tourist park, Xcaret, in the northern Mesoamerican Reef tract, Mexico. The community has a relatively low coral, gorgonian and sponge cover (<10%) and high algal cover (>40%). We recorded 23 species of Scleractinia, 14 species of Gorgonacea and 30 species of Porifera. The coral community is diverse but lacks large coral colonies, being dominated instead by small, sediment-tolerant, and brooding species. In these small colonies, the abundance of potentially lethal interactions and partial mortality is high but decreases when colonies are larger than 40 cm. Such characteristics are consistent with an environment control whereby storm waves periodically remove larger colonies and elevate sediment flux. The community only survives these storm conditions due to its slope-break location, which ensures lack of burial and continued local recruitment. A comparison with similar coral-ground communities in adjacent areas suggests that the narrow width of the rock terrace hinders sediment stabilization, thereby ensuring that communities cannot escape bottom effects and develop into three-dimensional reef structures on geological time scales.     

Effects of physical and biological disturbances on algal turfs in Kaneohe Bay, Hawaii

Disturbance in coral reef environments commonly results in an algal community dominated by highly productive, small filamentous forms and cyanobacteria, collectively known as algal turf. Research on the types of disturbance responsible for this community structure has concentrated mainly on biological disturbance in the form of grazing, although physical and other forms of biological disturbances may be important in many coral reef areas. On the reef flat in Kaneohe Bay, Oahu, Hawaii, algal turfs grow primarily upon coral rubble that tumbles with passing swells. We manipulated the frequency of rubble tumbling in field experiments to mimic the effects of physical disturbance by abrasion and light reduction on algal biomass, canopy height, and community structure. Treatments approximated a gradient of disturbance intensities and durations that occur on the reef flat. Although sea urchins and herbivorous fishes are not widespread and abundant on the reef flat, biological disturbances to algal turf communities in the form of herbivory by small crabs and abrasion by tough macroalgae contributed significantly to the variation in algal turf biomass. Within all experiments increasing disturbance significantly reduced algal biomass and canopy heights and the community structure shifted to more disturbance-tolerant algal forms. This study shows that the chronic physical disturbances from water motion and biological disturbances other than grazing from large herbivores can control algal communities in coral reef environments.     

Responses of algal communities to gradients in herbivore biomass and water quality in Marovo Lagoon, Solomon Islands

Settlement tiles were used to characterise and quantify coral reef associated algal communities along water quality and herbivory gradients from terrestrial influenced near shore sites to oceanic passage sites in Marovo Lagoon, the Solomon Islands. After 6 months, settlement tile communities from inshore reefs were dominated by high biomass algal turfs (filamentous algae and cyanobacteria) whereas tiles located on offshore reefs were characterised by a mixed low biomass community of calcareous crustose algae, fleshy crustose algae and bare tile. The exclusion of macrograzers, via caging of tiles, on the outer reef sites resulted in the development of an algal turf community similar to that observed on inshore reefs. Caging on the inshore reef tiles had a limited impact on community composition or biomass. Water quality and herbivorous fish biomass were quantified at each site to elucidate factors that might influence algal community structure across the lagoon. Herbivore biomass was the dominant driver of algal community structure. Algal biomass on the other hand was controlled by both herbivory and water quality (particularly dissolved nutrients). This study demonstrates that algal communities on settlement tiles are an indicator capable of integrating the impacts of water quality and herbivory over a small spatial scale (kilometres) and short temporal scale (months), where other environmental drivers (current, light, regional variability) are constant.