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    Evaluating sea cucumbers as extractive species for benthic bioremediation in mussel farms

    Avdelas, L. et al. The decline of mussel aquaculture in the European Union: Causes, economic impacts and opportunities. Rev. Aquac. 13, 91–118. https://doi.org/10.1111/raq.12465 (2021).Article 

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    Anthropogenic interventions on land neutrality in a critically vulnerable estuarine island ecosystem: a case of Munro Island (India)

    Land vulnerability of an area is directly related to the natural as well as anthropogenic activities involved in the geomorphological unit. Being one of the most vulnerable ecosystems, the estuaries and estuarine islands are delicately affected by both ecological processes of the sea and land and have pressures from multiple anthropogenic stressors and global climate change42,43,44. Ecological vulnerability and ecological sensitivity are similar and both originated from the concept of ecotone10,45. The geomorphologic concept of landscape sensitivity was first proposed by Brunsden and Thornes, who argued that the sensitivity indicated the propensity to change and the capacity to absorb the effects of disturbances10,46,47. Landscape sensitivity is studied by many researchers such as Allison and Thomas, Miles et al., Harvey, Knox, Usher, Haara et al., Thomas, Jennings and Yuan Chi8,47,48,49,50,51,52,53,54, through different case studies. Based on their findings Yuan Chi summarized the important characteristics of the landscape sensitivity are: a, the change of the landscape ecosystem; it involves the change likelihood, ratio, and component, as well as the resistance and susceptibility to the change, b, the temporal and spatial scales; which determine the occurrence, degree, and distribution of the change, c, the external disturbances that cause the change; the disturbances included natural and anthropogenic origins with different categories and intensities, and d, the threshold of the landscape sensitivity; it refers to the point of transition for the landscape ecosystem8. The environmental vulnerability of the Munroe Island has been studied based on the characterization of the geomorphological and sociocultural dynamics of the region based on the above characteristics.Bathymetric surveys in Ashtamudi lake and the Kallada riverThe present study shows that the geomorphic processes occurring on the Munroe Island are affected by anthropogenic disturbances in the morpho-dynamics of the Kallada river, Ashtamudi backwaters and associated fluvio-tidal interactions. A detailed bathymetric survey of both water bodies up to the tidal-influenced upper limit of the Kallada river27 was conducted with 200 m spaced grid references (Fig. 5). Bathymetry shows that the deepest point of the Ashtamudi backwater system is in Vellimon lake (13.45 m), the SE extension of Ashtamudi lake. The eastern side of Ashtamudi lake is deeper than the western side of this backwater system. The depth of the backwater decreases towards the estuary, and most parts of the lakebed are exposed here at the mouth of the inlet during the low tide. Compared to Ashtamudi lake, the Kallada river is deeper, and the riverbed area is recorded as the average depth is greater than 13 m. The deepest part of 14.9 m is recorded near Kunnathoor bridge, which is 12 km upstream from Munroe Island. Except for a few spots of hard (resistant) rocks, the river fairly and consistently follows a higher depth throughout its course.Figure 5Bathymetric profile of Ashtamudi lake and adjoining Kallada river (Figure was generated by Arc GIS 10.6).Full size imageOnce the Kallada river supplied very fertile alluvium during its flooding seasons (monsoon/rainy season), and most of this alluvium is deposited in the floodplains of the Munroe Island and the Ashtamudi lake. With a vast river catchment area from elevated lands of Western Ghats and a shorter course of 121 km33,55 and a higher elevation gradient of 12.6 m/km56, the Kallada river has a higher transporting capacity. The eroded surface and mined river/lakebeds at lower courses were replaced by the sediment load supplied by the Kallada river during each flood season until dam construction. During the focus group discussions with residents of the Island, they had described that they were crossing the Kallada river on foot in the 1990s or even earlier during the dry seasons. The construction of the Thenmala reservoir dam in 1980s across the river drastically choked the sediment supply of the Kallada river. In addition, excessive commercial sand mining without any regulation from the riverbeds of Kallada and Ashtamudi waterbodies accelerated the deepening of waterbodies. It increased the erosion of surface and subsurface soils through fluvial and hydraulic action. This, in turn, drastically reduced the deposition of fertile alluvium over the low-lying Munroe Island. The current bathymetry shows that the river channel has deepened its course to 14 m compared to 5–6 m of 1980s. When comparing the bathymetric data of 200127, it is interesting to note that no considerable changes occurred in the bathymetry of Ashtamudi lake over the last two decades.Dams indeed alter aquatic ecology and river hydrology, upstream and downstream, affecting water quality, quantity, breeding grounds and habitation22. The other significant impact of the damming of the Kallada river is the saline water intrusion towards upstream of Ashtamudi lake and the Kallada river. The freshwater discharge is regulated after the construction of the Thenmala reservoir, and the water is being diverted to the reservoir and associated canals. There is a decline in sedimentation over the floodplains and catchment area as a result of the increased tidal effects and associated running water dynamics, which may accelerate the erosion trend of the nearby places.Lithological characterization of the Munroe IslandThe Munroe Island is a riverine delta formation by the Kallada river at the conjunction of river and backwater systems. To understand the micro-geomorphological processes of the study area, the near-surface geology of the Munroe Island had been studied in detail with the help of resistivity meter surveys and borehole datalogs from different locations. As per the current resistivity survey, it is evident that the Munroe Island is formed by recent unconsolidated loose sediments more than 120 m thick succession below ground level (Figs. 6 and 7). The electrical resistivity tomography of identified locations within the deltaic region shows a meagre resistance value to its maximum penetration (Fig. 6), which proves that the sedimentary column with intercalations of sand and carbonaceous clays of varying thickness extends to a depth of 120 m, in turn indicating the process of enormous sedimentation happened during the recent geological period. Loose wet soils of saline nature records a lower resistance value for an electric circuit. The layers formed in the diagram (Fig. 6) represent the seasonal deposition of unconsolidated soils as thin sequence. The Mulachanthara station of the resistivity meter tomography, which is situated at a more stable location of the Island, has a higher resistivity value than the West Pattamthuruth location, which is located at the exact alluvial flood plain.Figure 6Electrical resistivity profiles of Munroe Island.Full size imageFigure 7Geomorphological map showing litho-log of north (Kannamkadu); middle (Konnayil Kadavu); and south (Perumon bridge) locations of Munroe Island (borehole data source: PWD, Govt of Kerala) (Software used: Arc GIS 10.6).Full size imageThe Public Works Department (PWD), Kerala State carried out soil profile studies through Soil Penetrating Test (SPT) borehole drilling method as part of constructing bridges at three different locations up to a depth of 62 m, i.e., one across the Kallada river (north side)57, one across Ashtamudi lake in southern Munroe Island58 and one at the central part of Munroe Island (across a canal)59 (Fig. 7). The hard rock is found only on the southern side of the lake at a depth of 45 m. The litho-log shows that unconsolidated loose sediments of significantly higher thickness occur in the entire Munroe Island (Fig. 7). Anidas Khan et al.60 studied the shear strength and compressibility characteristics of Munroe Island’s soil for two different locations with disturbed and undisturbed samples. They classified the soil of Mundrothuruth into medium compressibility clay (CI) and high compressibility clay (CH) with natural moisture contents of 44.5% and 74%, respectively. The unconfined compressive strengths of the undisturbed and remolded samples for the first location are 34.5 kN/m2 and 22.1 kN/m2, respectively, while they are 13 kN/m2 and 9 kN/m2 respectively for the second location60. Such compressive strength indicates that the soils of Munroe Island are soft or very soft in nature.Land degradation: a morphological analysisTo decrease the impact of the monsoon floods and to distribute the alluvium to the southern part of the island, Canol Munroe, the then Diwan of the Thiruvithamkoor Dynasty, made an artificial man-made canal during the 1820s connecting the Kallada river with the eastern extension of Ashtamudi lake, and this river is known as “Puthanar” (meaning a new river). During the last few decades, (after 1980s) the estuarine island ecosystem of Munroe Island has faced several structural deformities. The natural sedimentation and flooding happening in the Islands were very limited and hence, the normal events happened over the past several decades disturbed and significantly affected the land neutrality. These islands, once known as the region’s rice bowl, now devoid of any paddy cultivation mainly because of the increased soil salinity. According to the Cadastral map prepared by the revenue department (1960s) there were many paddy fields, locally named as Mathirampalli Vayal (Vayal is the local name for paddy field), Thekke Kothapppalam Vayal, Mattil Vayal, Kottuvayal, pallaykattu Vayal, Konnayil Vayal, Vadakke Kundara Vayal, Thachan Vayal, Thekke Kundara Vayal, Kizhakke Oveli Vayal, Thekke Oveli Vayal, Odiyil Vettukattu Vayal, Nedumala Vayal, Madathil Vayal, Karichal Vayal, Moonumukkil Vayal, Arupara Vayal, Kaniyampalli Vayal, Manakkadavu Vayal, Panampu Vayal, Pattamthuruth Vayal etc. The recent satellite images shows that no paddy cultivation exist now, which is further confirmed by the field observations conducted through our study. The annual report published by Gramapanchayat39 indicate that the paddy field of region was reduced from 227 to 8 acres (from 1950 to 1995) and now about in 2 acres only (2018). Most of the paddy fields of northern and northwestern regions are severely affected by land degradation due to erosion, saline water intrusion and flooding and are entirely or partially buried under the backwater system. Figure 8 depicts the morphological degradation of the severely affected areas of Munroe Island from 1989 to 2021 through different satellite images. Some paddy fields are converted into filtration ponds to take the benefit of frequent tidal flooding. The coconut plantations were later introduced in place of paddy fields, and they eventually replaced the paddy fields. However, during the last decades, it has been observed that these coconut plantations are also under threat mainly because of degradation of the soil fertility, which directly bears the quality and quantity of production (Fig. 9).Figure 8Morphological changes in the study area from the satellite images (a) 1989 (aerial photograph); (b) 2000 (Landsat); (c) 2011 (World View—II); (d) 2021 (Sentinel) (the modified maps of (a) is obtained from National remote Sensing Centre (NRSC), Hyderabad, (b) is downloaded from https://earthexplorer.usgs.gov/ (c) is obtained from Digital Globe through NRSC and (d) is downloaded from https://scihub.copernicus.eu/. Figures were generated using Arc GIS 10.6).Full size imageFigure 9Threatened coconut plantations indicating the low productive regime. Photographs taken by Rafeeque MK.Full size imageOver the study area the most affected alluvial plain of the Peringalam and Cheriyakadavu island are taken separately to study the morphological changes over the decades. This area is named Puthan Yekkalpuram (which means new alluvium land), and the north side of the Kallada river (the northward extension in the Mundrothuruth GP) is demarcated as old alluvium land (Pazhaya Yekkalpuram) as per the revenue department’s cadastral map. The study shows that total 38.73 acres of land has lost from the Peringalam and Cheriyakadavu Islands during the last 32 years, which is equivalent to 11.78% and 46.95% of the total geographical area of the Peringalam and Cheriyakadavu Islands, respectively. The land degradation details over the last three decades are given in the Table 2. Many other locations, such as Nenmeni and West Pattamthuruth, are also severely affected by land degradation. However, these areas are landlocked and less affected by running water or floods. Hence, the land degradation experienced is the settling of the topsoil and subsidence of structures such as houses and bridges. The sinking of basements of many houses and even the subsidence of railway platforms are well observed during field visits, indicating the alarming land degradation issues (Figs. 1 and 10) to be addressed its deserving importance. There are also clear indications of the gradual formation of new waterlogged areas in the islands, which may further deteriorate and forms the part of the backwater system which eventually affects total land area of the Munroe Island.Table 2 Land degradation of Peringalam and Cheriyakadavu region for the past 32 years.Full size tableFigure 10Various environmental degradations in Munroe Island. Photographs taken by Rafeeque MK.Full size imageThe island population also shows a negative growth over the years. According to the census report of 201138, the total population of Gramapanchayat has decreased to 9440 person/km2 in 2011 from 10,013 person/km2 of 2001 and 10,010 person/km2 of 1991 census reports. Frequent flooding (especially tidal flooding), the lack of drinking water, and migration in search of a better livelihood are the main reasons for the observed population reduction as revealed through the survey. The high intrusion of saline water into the cultivated land through tidal flooding and the lack of flushing of surface saline soils by monsoon floods (freshwater) decreased agricultural productivity of the area, and hence, now people are more dependent on fishing and backwater activities for their livelihood. Lack of proper transportation to the nearby markets limits their fishing activities to a daily subsistence level. Due to the flooding caused by subsidence/tidal surges and land degradation during the last few decades, more than 500 households have vacated their houses38,39.Tidal Flooding and Estuarine ProcessesIn Mundrothuruth, the major environmental degradation problems where occurring due to tidal flooding and saline water intrusion into the freshwater ecosystem. Mathew et al. studied the tidal and current mechanisms of the Ashtamudi backwater in 200161. They reported that the Kallada river plays a vital role in determining the eastern lake’s circulation pattern. In addition, the increased discharge from the north Chavara canal and the south Kollam canal also influences the local circulation of the Ashtamudi backwater. The current velocity reaches up to 100 cm/s at the estuary entrance, but it rapidly diminishes in the eastern parts, where the speed is generally less than 30 cm/s. One of the critical observations made during the field study, which corroborates with the acquaintance of local people as well, is that the flooding on Munroe Island is not related to the spring tide of the open ocean. The disappearance of the semidiurnal tide in the central lakes occurs due to frictional resistance and the time lags for the tide to travel across the estuary61. At the shorter semidiurnal period of approximately 12 h, the tide is more dissipated than the more extended constituents of 24-h duration. The survey conducted with the island inhabitants also reiterates these views.As per the experience of local inhabitants, tidal flooding in Munroe Island was not frequent in earlier times. The comparison of the bathymetry data collected during 200058 and 2017 (Fig. 5) in and around the regions of Munro Islands shows that there is not much change in bathymetry during the period. Hence, changes in basin geometry are not having a significant role in tidal dynamics in imparting the variations as observed. In addition to the bathymetric survey, the data on tide measurements at four locations corresponding to three seasons were also collected. The tide data measured during the pre-monsoon period is shown in Fig. 11a. The figure shows that the tidal range in the inland area is almost the same even during the spring and neap tides. As discussed earlier, the tidal flooding in Munro Island is not related to spring tide in the ocean, and there may be the influence of specific complicated dynamics in the basin for this flooding that needs to be studied more profoundly. Further the data pertaining to tidal dynamics were inadequate; we established three tide gauges in selected locations in and around Munro Island. From the analysis of tide gauge data, it is found that the signature of anomalous variability in water column height, which is not at all linked to the tidal dynamics.Figure 11(a) Salinity variation of bottom water at selected locations in Kallada river during monsoon and post monsoon. (b) Observed tide during pre-monsoon months.Full size imageThe water quality analysis for three time periods, during the year of the cyclonic storm, Okhi (2017), was conducted to understand river run-up impact on salinity in and around Munroe Island (Fig. 11). The riverbed is lowered below the baseline of erosion, and dense saline water is trapped in the deeps during high tide. This has been confirmed during the bathymetric survey of the Kallada river and Ashtamudi backwaters, which showed a significant increase in water depth, particularly within the river channel. The high-density saline water is trapped in the basins and trenches created in the river channel due to uncontrolled sand mining, which leads to the degradation of the quality of sediments and groundwater in the region. Nevertheless, the samples collected immediately after Okhi (when the dam’s shutter was opened due to heavy rainfall in the catchment area) show that the high runoff replaced the trapped saline water with fresh water. After ten days of the first sampling, the water became saline nature after the closure of the dam’s shutter. This proves that because of dam construction, the river runoff in the Kallada river was reduced significantly, and extensive human interactions especially sand mining activities increased the riverbed deepening and formation of pools beyond the base level of running water.Conservations and management strategiesConsidering the facts discussed above, the Munroe Island may continue to be badly affected unless suitable sustainable management strategies are not evolved. Construction and associated activities, such as the damming of reservoirs, sand mining and landfilling, are indispensable for any nation’s economic and social development. United Nations’s member states have formulated 17-point Sustainable Developmental Goals (SDGs) to better the world sustainably. Local and national governments pertaining to the Munroe Island need to develop a sustainable management plan to protect this Ramsar-listed wetland. The environmental issues of Mundrothuruth can be controlled, and land degradation may be monitored through a well-drafted working plan. All aspects of earth and social sciences may be integrated to draft such a management plan of reverse landscaping. The reverse landscaping (i.e., recalling the degrading landscape to its geomorphic isostatic state) method is a must-considered sustainable solution for land degradation and other environmental issues.The deep courses of Kallada river must be upwarped through a well-planned artificial sedimentation to eradicate the saline banks of deep basins. The sediments deposited in the Thenmala reservoir and the sediments removed through the digging of boat channels may be utilized in a periodic monitoring method. Sand mining from Ashtamudi lake and the Kallada river may be strictly controlled, and the minimum freshwater flow should be ensured. The construction methods practiced in Mundrothuruth are outdated and technically nonexistent. Well-studied engineering methods suitable for an environmentally fragile area must be implemented with a proper understanding of the soil characteristics, such as shear strength and compressibility rate, and hydrodynamics, such as tidal and fluvial actions. Soil fertility must be increased by supplying additional fertile soil and freshwater, at least for a minimum period. The inhabitants’ socioeconomic well-being is strengthened by advancing technology and providing easy access to the market and other social amenities. More

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    Isolation and infection cycle of a polinton-like virus virophage in an abundant marine alga

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    Using size-weight relationships to estimate biomass of heavily targeted aquarium corals by Australia’s coral harvest fisheries

    Establishing size-weight relationships for heavily targeted coral species is an important first step towards informing sustainable harvest limits19. Placing coral harvests into an ecological context is a core requirement for implementing a defensible stock assessment strategy, and this need is particularly critical given escalating disturbances and widespread reports of coral loss7,17,25. Using these relationships, managers can now easily sample and calculate biomass per unit area. It is important to point out that all sites sampled in our study represent fished locations, and there is no information available to test whether standing biomass has declined due to sustained coral harvesting at these locations. While these data may now provide a critical baseline for assessing the future effects of ongoing fishing, it is also important to sample at comparable locations where fishing is not permitted or has not occurred (where possible), to test for potential effects of recent and historical harvesting.Biomass per unit area data presented herein highlights the highly patchy abundance and biomass of targeted coral species14, which is evident based on the often vastly different mean and median values (Table 2). Examining biomass per unit area estimates for C. jardinei for example, which returned some of the highest biomass estimates, the 33.75 kg·m−2 maximum estimate from a transect stands as an extreme outlier, with 12 of the 16 other transects being below 0.2 kg·m−2. This indicates the challenges of managing species that occur in patchily distributed concentrations, particularly in a management area the size of the QCF. It is also important to note, these estimates are generated only on transects where the target species occurred, and therefore, should technically not be considered as an overall estimate of standing biomass. While the estimation of size-weight relationships is a step towards a standing biomass estimate, many challenges remain in terms of sampling or reliably predicting the occurrence of these patchily distributed species. Bruckner et al.14 attempted to overcome this management challenge in a major coral fishery region of Indonesia by categorising and sampling corals (in terms of coral numbers) in defined habitat types, and then extrapolating to estimated habitat area based on visual surveys and available data. This approach, utilising size-weight relationship derived biomass per unit area estimates (instead of coral numbers), may be a viable method for the QCF, however much more information is needed to understand the habitat associations (e.g., nearshore to offshore), and environmental gradients that influence the size and abundance of individual corals. Fundamentally, it is also clear that much more data is required to effectively assess the standing biomass of aquarium corals in the very large area of operation available to Australian coral fisheries.These corals are found in a range of environments, and it is important to consider available information on life history if attempting to use coral size-weight relationships to inform management strategies via standing biomass estimation. All corals in this study can be found as free living corals (at least post-settlement) in soft-sediment, inter-reefal habitats, from which they are typically harvested by commercial collectors19. However, only four of the 6 species are colonial (C. jardinei, D. axifuga, E. glabrescens, M. lordhowensis) while the remaining two species (H. cf. australis and T. geoffroyi) are more typically monostomatous or solitary. As indicated in previous work24, if larger colonial corals were to be fragmented during harvesting instead of removed entirely, fishery impacts would likely be lessened24. Given the power relationship between coral maximum diameter and weight, larger corals contribute disproportionately to the total available biomass of each species in a given area. The potential environmental benefit of leaving larger colonies (at least partially) intact is not limited to impacts on standing biomass, as this practice would likely be demographically beneficial given the greater reproductive potential (i.e., fecundity) of larger colonies, which also do not need to overcome barriers to replenishment of populations associated with new recruits (i.e., high mortality during and post-settlement26). This conclusion was drawn largely from data on branching taxa (e.g., Acropora), which are relatively resilient to fragmentation and commonly undergo fragmentation as a result of natural processes27,28,29. D. axifuga can be considered to exhibit a relatively similar branching growth form, however, the growth form of E. glabrescens and C. jardinei changes with size, moving from small discrete polyps to large phaceloid and flabello-meandroid colonies, respectively19. While larger colonies of E. glabrescens and C. jardinei may be relatively resilient to harvesting via fragmentation, the same may not be true for smaller colonies, or species with massive growth forms such as M. lordhowensis. Typically, for each species, the average reported weight was quite low, coinciding with the lower end of the sampled maximum diameter range. For colonial species, the harvested smaller maximum diameters (if fragments) are ideal from an ecological perspective as this will have the least impact possible on standing biomass, and may also leave a potentially mature breeding colony intact. Ultimately, in light of these considerations, the development of uniform and standardised industry-wide harvest guidelines to balance economic and ecological outcomes may be necessary. The development of these guidelines would require consultation with commercial harvesters, as well as considerable additional work in measuring ecological impacts and better understanding the cost of these impacts from an economic perspective. Conversely, if whole colonies are collected, which is necessarily the case for solitary species such as H. cf. australis and T. geoffroyi (and potentially smaller colonies of other species such as E. glabrescens and C. jardinei); smaller colonies may be collected before they reach sexual maturity, hindering their ability to contribute to population replenishment. Therefore, collection of small fragments should be encouraged for colonial species; while for monostomatous species where this is not possible, introduction of a minimum harvest size based on sexual maturity should be considered.Additionally, the need for further consideration of the selectivity of ornamental coral harvest fisheries3,4,30 when assessing standing biomass is evident. Due to various desirable traits, the majority of available biomass may not be targeted by collectors. As emphasised in this study, the focus on smaller corals is indicative of the trend towards collection of most of these species at the lower portion of their size range, at least compared to some of the maximum sizes recorded on transects (e.g., see Tables 1 and 2, section b). However, it is also important to consider that transects were conducted in areas subject to commercial collection and are likely to skew results and prevent clear conclusions relating to size selectivity. Sampling of unfished populations (i.e., any residing outside of permitted fishing zones) and/or spatial and temporal matching of catch data and transect data across a larger sample of operators will be required to properly address industry size selectivity trends. For instance, only 17.5% of C. jardinei corals measured on transects fell within the diameter range represented by data obtained from collectors, with 81.9% of corals measured on transects exceeding this range. If it is viable to collect fragments from larger colonies (which does appear to be the case for some corals such as C. jardinei), then a larger proportion of standing biomass outside of this size range could be targeted by fishers. As an additional consideration, only desirable colour morphs of these corals will be harvested, and due to lack of appropriate data, the prevalence of these morphs remains unclear. H. cf. australis and M. lordhowensis for example often occur in brown colour morphs, which are far less popular in markets where certain aesthetic qualities (e.g., specific, eye-catching colours or combinations of colours) are desired, such as the ornamental aquarium industry. Even without delving into further considerations such as heritability of phenotypic traits, management conclusions drawn from standing biomass estimates may be ineffective in the absence of efforts to account for selectivity in this fishery.The relationship between size and weight was found to differ between all corals, with the exception of C. jardinei and E. glabrescens. There can be some moderate similarity in skeletal structure between these two species, particularly between small colonies, reflecting the similar maximum diameter range of sampling in the current study. Subsequently, inherent physiological constraints may be imposed on corals that prevent the maintenance of growth rates between corals of smaller and larger sizes, for example, as the surface area to volume ratio declines with growth31. In the current study, all corals, with the exception of C. jardinei, showed evidence of allometric growth, as exhibited by an estimated exponent value different to 3. Sample size for C. jardinei was greatly limited, as this species typically forms extensive beds, and are rarely brought to facilities as whole colonies. Therefore, the lack of evidence for allometric growth may reflect higher error for the species coefficient parameter due to the comparatively small sample size for this species. This suggests that mass would not increase consistently with changes in colony size in 3 dimensions31, which seems likely considering the change in exhibited form described for E. glabrescens and C. jardinei previously. In the current context, this indicates that the estimated ‘a’ and ‘b’ constants are likely to vary as the sample range increases, reflecting the changes in the size-weight relationship between smaller and larger samples of these species. Therefore, ideally, these models should incorporate data that reflect the maximum diameter range of the species in the region of application to allow increased accuracy of biomass estimation. To achieve this will require additional fishery-independent sampling, as large colonies are rarely collected whole, though may be collected as fragments depending on the species. Sampling may be challenging for some species given the difficulty of physically collecting and replacing large whole colonies, particularly for inter-reefal species such as M. lordhowensis, which can occur in deep, soft sediment habitat, subject to strong currents. Importantly, obtaining ex situ or in situ growth rate data should be considered a priority for the management of heavily targeted species. This data is likely to be another necessary component (in conjunction with size-weight relationships) of any stock assessment model developed for LPS corals, and may also eliminate the need to collect large sample colonies to improve estimated size-weight relationships.The disproportionate focus on smaller corals (i.e., corals in the current study averaged between 4.28 and 11.48 cm in maximum diameter) is likely to lead to an underestimation of weight in corals at greater diameters when used as inputs for size-weight models. This may explain the apparent minor underestimation observed in some species (e.g., M. micromussa, T. geoffroyi). In the current context, this represents an added level of conservatism with estimates obtained from these equations. While the relationship between size and weight was particularly strong for some species, (mainly D. axifuga and T. geoffroyi), for other species, such as M. lordhowensis, growth curves tended towards underestimation at larger diameter values. As the mass of a coral is reflective of the amount of carbonate skeleton that has been deposited32, the coral skeleton may increase disproportionately to coral diameter if or when corals start growing vertically. For example, in massive corals such as M. lordhowensis, vertical growth (i.e., skeletal thickening) is often very negligible among smaller colonies, with thickening of the coral skeleton only becoming apparent once the coral has reached a threshold size in terms of horizontal planar area. Additional fisheries-independent sampling outside of the relatively narrow size range of harvested colonies will be required to address this source of error in future applications. Ecological context in the form of fishery independent data on stock size and structure is essential for effective management, especially in ensuring that exploitation levels are sustainable and appropriate limits are in place. Coral harvest fisheries offer managers an ecologically and biologically unique challenge, as the implementation of standard fisheries management techniques and frameworks is hampered by their coloniality and unique biology, as well as a general lack of relevant data for assessing standing biomass and population turnover, not to mention the evolving taxonomy of scleractinian corals33. Similarly, fishery-related management challenges such as the extreme selectivity in terms of targeted size-ranges and colour-morphs, plus the potentially vast difference in the impact of various collection strategies (i.e., whole colony collection vs fragmentation during collection) also complicates the application of typical fisheries stock assessment frameworks. The relationships and equations established in the current work offer an important first step for coral fisheries globally by laying the groundwork for a defensible, ecologically sound management strategy through estimation of standing biomass, thus bridging the gap between weight-based quotas and potential environmental impacts of ongoing harvesting. It is important to note that the species selected for the current work do not represent the extent of heavily targeted LPS corals. For example, Fimbriaphyllia ancora (Veron & Pichon, 1980), Fimbriaphyllia paraancora (Veron, 1990), Cycloseris cyclolites (Lamark, 1815), and Acanthophyllia deshayesiana (Michelin, 1850) are examples of other heavily targeted corals of potential environmental concern19, and management would also benefit from the estimation of size-weight relationships for these species. Moving forward, the next challenge for the coral harvest fisheries will be to comprehensively document and track the standing biomass of heavily targeted and highly vulnerable coral stocks, explicitly accounting for fisheries effects and also non-fisheries threats, especially global climate change. More

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