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Genetic variationThe best fitting substitution models for COI codons 1–3 were identified as TN + F + G4, F81 + F + I, and TN + F, respectively. The maximum likelihood (ML) and Bayesian inference (BI) trees showed similar topologies of the main nodes, although the BI tree displayed greater resolution of the ingroup branches (Fig. 1). Furthermore, the BI tree revealed three monophyletic clades, while two of those clades were merged in the ML tree. Two of the three molecular species delimitation methods (ASAP and TCS) recovered three groups in the BI tree as distinct taxa (Fig. 1), corresponding to the three previously described ESUs27,28. The third method (bPTP) recovered between 8 and 43 groups (mean = 28.03) suggesting that there is evidence of additional genetic differentiation within the three groups identified by ASAP and TCS. The outputs of the three methods are provided in the Supplementary information. The molecular diagnosis uncovered several fixed nucleotide differences COI characters for each taxon (Table 1: “W. carteri” I = 10; “W. carteri” II = 3; “W. carteri” III = 5). There were also 13 fixed nucleotide differences in W. carteri for the 16S gene. The remaining two taxa had no fixed nucleotide differences for the 16S gene.Figure 1Phylogenetic trees obtained by maximum likelihood (left) and Bayesian inference (right) analysis of “Westralunio carteri” mtDNA COI sequences, including support values for the major genetic clades [ultrafast bootstrap values (left) and Bayesian posterior probabilities (right)]. Colour coded bars show support for the three major clades by the species delimitation methods (ASAP = dark shade; TCS = lighter shade). Green = WcI = “W. carteri” I; blue = WcIII = “W. carteri” III; red = WcII = “W. carteri” II. Results of bPTP analysis not shown (see supplementary data). Haplotype names correspond to Benson et al.28. Outgroup taxa are Velesunio ambiguus (Philippi, 1847) (Hyriidae: Velesunioninae) and Cucumerunio novaehollandiae (Gray, 1834) (Hyriidae: Hyriinae: Hyridellini).Full size imageTable 1 Molecular diagnoses of “Westralunio carteri” Evolutionarily Significant Units (ESUs) from southwestern Australia (after Bolotov et al.122 with reanalysis of data from Klunzinger et al.27 and Benson et al.28).Full size tableVariation in shell morphologyBased on results from analyses of variances (ANOVAs), shells of “W. carteri” I were significantly larger (for size metrics total length (TL), maximum height (MH), beak height (BH) and beak length (BL)) and more elongated (i.e., had a lower maximum height index (MHI)) than shells of “W. carteri” II and “W. carteri” II + III combined (Table 2). However, there was no difference in size or shape metrics between “W. carteri” I and “W. carteri” III (Table 2). The lack of significant differences in beak height index (BHI) and beak length index (BLI) among any of the taxa (Table 2) indicates that wing and anterior shell development was not discernibly different between any of the ESUs.Table 2 Shell size metrics [mm], shape indices [%] and scores for the first two principal components (PC) obtained by Principal Component Analysis of shape indices and 18 Fourier coefficients generated by Fourier Shape Analysis for each “Westralunio carteri” species and subspecies-level Evolutionarily Significant Units (ESUs): n, number of specimens measured; minimum (min) to maximum (max) and mean (± standard error (SE)).Full size tableThis pattern was partly confirmed in the principal component analysis (PCA) of these three shell shape indices, where PC1, largely explained by variation in BLI (Fig. 2A), did not differ between the two species (i.e., “W. carteri” I vs. “W. carteri” II + III) or among the three taxa (Table 2). The PC2, largely explained by variation in MHI and BHI (Fig. 2A), differed significantly between “W. carteri” I and “W. carteri” II (Table 2). Accordingly, 70% (70% jack-knifed) of specimens were assigned to the correct species in the corresponding discriminant analysis (DA), whilst this was true for only 55% (54%) at the MOTU-level.Figure 2Scatterplots of the first two PC axes obtained by PCA on (A) calculated shape indices based on shell measurements, and (B) 18 Fourier coefficients for “Westralunio carteri” I, “W. carteri” II and “W. carteri” III. 95% Confidence Intervals are displayed at the species level, i.e., for “W. carteri” I (full line) and “W. carteri” II + III (dashed line). Extreme shell outlines in (B) are depicted to visualise trends in sagittal shell shape, along PC axes.Full size imageThe difference in shell elongation between “W. carteri” I and “W. carteri” II was confirmed by Fourier shape analysis. As visualised by synthetic outlines in Fig. 2B, shell elongation is expressed along the PC1 (explaining 15% of total variation in Fourier coefficients). The PC1 as well as PC2 scores differed significantly between the two species (i.e., “W. carteri” I vs. “W. carteri” II + III) as well as between “W. carteri” I and “W. carteri” II, respectively (Table 2). Combined with synthetic outlines, this indicated a tendency towards a more elongated, somewhat wedge-shaped shell in “W. carteri” I, whilst “W. carteri” II shells tended to be relatively high with a stout anterior margin (Fig. 2B). An analysis of similarities (ANOSIM) analysis on all Fourier coefficients revealed no significant difference between the two species (i.e., “W. carteri” I vs. “W. carteri” II + III; ANOSIM: R = − 0.018, p = 0.097), but did indicate a significant difference between the three ESUs (ANOSIM: R = 0.0625, p = 0.0051). Specifically, “W. carteri” I differed significantly from “W. carteri” II (Bonferroni-corrected p = 0.0009). Only 66% and 65% (62% and 62% jack-knifed) of specimens were assigned to the correct species and taxon in DAs on that dataset, respectively.Taxonomic accountsClass: Bivalvia Linnaeus, 175831.Subclass: Autobranchia Grobben, 189432.Infraclass: Heteroconchia Gray, 185433.Cohort: Palaeoheterodonta Newell, 196534.Order: Unionida Gray, 185433 in Bouchet & Rocroi, 201035.Superfamily: Unionoidea Rafinesque, 182036.Family: Hyriidae Parodiz & Bonetto 196337.Genus: Westralunio Iredale, 19349.Type species: Westralunio ambiguus carteri Iredale, 19349 (by original designation).Redescription: Westralunio carteri (Iredale, 1934)SynonymyUnio australis Lamarck38: Menke39, p. 38, specimen 219. (Non Unio australis Lamarck, 181938).Unio moretonicus Reeve40: Smith41, p. 3, pl. iv, Fig. 2. (misidentified reference to Unio moretonicus Reeve, 186540).Hyridella australis (Lam.38): Cotton & Gabriel42 (in part), p. 156. (misidentified reference to Unio australis Lamarck, 181938).Hyridella ambigua (Philippi26): Cotton & Gabriel42 (in part), p. 157. (misidentified reference to Unio ambiguus Philippi, 184726).Westralunio ambiguus carteri: Iredale, 19349, p. 62.Westralunio ambiguus (Philippi26): Iredale9, p. 62, pl. iii, Fig. 8, pl. iv, Fig. 8. (Non Unio ambiguus Phil. 184726), Iredale43, p. 190.Centralhyria angasi subjecta Iredale, 19349, p. 67 (in part), Iredale43, p. 190.Westralunio carteri Iredale9: McMichael & Hiscock10pl. viii, Figs. 1, 2, 3, 4, 5, 6 and 7, pl. xvii, Figs. 4, 5.Type materialLectotype: AMS C.61724 (Fig. 3A) Westralunio ambiguus carteri Iredale, 19349.Figure 3(A) Westralunio ambiguus carteri Iredale, 1934, Lectotype: Victoria Reservoir, Darling Range, 12 mi E of Perth, AMS C.061724. Detail of fusion in anterior muscle scars from either valve represented by dashed lines and black polygons. Bottom image showing detail of hinge teeth. Photos provided with permission by Dr Mandy Reid, AMS. (B) Valves and detail of sculptured umbo of a juvenile W. carteri from Yule Brook, Western Australia, UMZC 2013.2.9. Photo by Dr Michael W. Klunzinger. (C) Glochidia of W. carteri from Canning River, Western Australia. Photo by Dr Michael W. Klunzinger.Full size imageParalectotypes: AMS C.170635 Westralunio ambiguus carteri Iredale, 19349 (n = 4).Type locality: Victoria Reservoir, Darling Range, 12 miles east of Perth, Western Australia (Fig. 4A).Figure 4(A) Victoria Reservoir, Canning River, near Perth, Western Australia, type locality for W. carteri. Photo by Corey Whisson. (B) Goodga River, Western Australia, type locality for W. inbisi inbisi, at vertical slot fishway where holotype of W. inbisi inbisi was collected from. Photo provided with permission by Dr Stephen J. Beatty. (C) Margaret River, Western Australia, type locality for W. inbisi meridiemus, at Canebreak Pool. Photo by Dr Michael W. Klunzinger.Full size imageLectotype: BMNH 1840–10-21–29 Centralhyria angasi subjecta Iredale (selected by McMichael & Hiscock10).Type locality: Avon River, Western Australia.Material examined for redescription: For W. carteri (= “W. carteri” I), molecular data examined included 52 and 61 individual COI mtDNA and 16S rDNA sequences, respectively, for species delimitation. Additionally, Fourier shell shape outline analysis and traditional shell morphometric measurements were examined from 238 and 290 individuals, respectively. Complete details on all specimens examined are provided in Supplementary Table S1.ZooBank registration: urn:lsid:zoobank.org:act:6B740F4D-40C3-4D6A-8938-B0FD7FD1F6D7.Etymology: The species name carteri is most likely named after the surname of the collector who provided original type specimens to the Australian Museum, although Iredale9 did not specify this as the case. We have applied ICZN Articles 46.1 and 47.144, designating W. carteri as the nominotypical species.Revised diagnosis: Specimens of W. carteri are distinguished from other Australian Westralunio taxa by having shell series that are significantly larger and more elongated than Westralunio inbisi inbisi subsp. nov., but not different from Westralunio inbisi meridiemus subsp. nov. The species has 10 diagnostic nucleotides at COI (57 G, 117 T, 210 G, 249 T, 255 C, 345 G, 423 T, 447 T, 465 A, 499 T) and 13 at 16S (137 T, 155 C, 228 C, 229 T, 260 G, 290 A, 305 G, 307 T, 310 A, 311 C, 321 T, 330 A, 460 A), which differentiate it from its sister taxa, W. inbisi inbisi and W. inbisi meridiemus (each described below) using ASAP and TCS species delimitation models.RedescriptionThis species is of the ESU “W. carteri” I27,28.Shell morphology: Shells of relatively small to medium size, generally less than 70 mm in length, but to a maximum length of approximately 100 mm10,45, MHI 46–89%; anterior portion of shell with moderate development, BLI 22–49%; larger shells with abraded umbos scarcely winged; wing development variable, generally decreasing with size, BHI 76–104% (Table 2). Shell outline oblong-ovate to rounded; posterior end obliquely to squarely truncate, anterior end round; ventral edge slightly curved, nearly straight in larger specimens; hinge line curved, hinge strong. Umbos usually abraded in specimens  > 20 mm in length; unabraded umbos with distinctive v- or w-shaped plicated sculpturing (Fig. 3B and Zieritz et al.46). Shell substance typically thick; shells of medium width with pronounced posterior ridge; periostracum smooth, dark brown to reddish, with fine growth lines. Pallial line less developed in smaller specimens and prominent only in large specimens (e.g.,  > 60 mm TL). Lateral teeth longer and blade-like, slightly serrated to smooth and singular in left valve, fitting into deep groove in right valve; pseudocardinal tooth in right valve coarsely serrated, thick, and erect, fitting into deeply grooved socket in left valve. Anterior muscle scars well impressed and anchored deeply in larger specimens; anterior retractor pedis and protractor pedis scars both small and fused with adductor muscle scar; posterior muscle scars lightly impressed; dorsal muscle scars usually with two or three deep pits anchored to internal umbo region.Anatomy: Supra-anal opening absent, siphons of moderate size, not prominent but protrude beyond shell margin in actively filtering live specimens, pigmented dark brown with mottled lighter brown to orange splotches; inhalant siphon aperture about 1.5 times size of exhalant and bearing 2–4 rows of internal papillae (Fig. 5A); ctenidial diaphragm relatively long and perforated. Outer lamellae of outer ctenidia completely fused to mantle, inner lamellae of inner ctenidia fused to visceral mass then united to form diaphragm; palps relatively small, usually semilunar in shape; marsupium well developed as a distinctive swollen interlamellar space in the middle third of the inner ctenidium of females. Outer ctenidia in both sexes thin, with numerous, short intrafilamentary junctions and few, irregular interlamellar junctions; in females similar, but marsupium has numerous, tightly packed, well-developed interlamellar junctions. Thus, brooding in females is endobranchous.Figure 5Live specimens of actively filtering freshwater mussels in the burrowed position. (A) Westralunio carteri (Iredale, 1934), Canning River at Kelmscott, Western Australia, inhalant siphon with 2–4 rows of papillae oriented toward substrate. Photo by Dr Michael W. Klunzinger. (B) Westralunio inbisi meridiemus subsp. nov., Canebreak Pool, Margaret River, Western Australia; inhalant siphon edges lined with protruding papillae facing towards water surface, away from substrate. Photo by Dr Michael W. Klunzinger.Full size imageLife history: Sexes are separate in W. carteri, and hermaphroditism appears to be rare47,48,49. Males and females both produce gametes year-round but brooding of glochidia appears to be seasonal and ‘tachyticitc’ (i.e., as defined by Bauer & Wächtler19, fertilisation and embryonic development occurring in late winter/early spring and glochidia release in early summer)50. Glochidia are released within vitelline membranes, embedded in mucus which extrude from exhalant siphons of females (i.e., ‘amorphous mucus conglutinates’) during spring/summer. Glochidia attach to host fishes and live parasitically on fins, gills or body surfaces for 3–4 weeks while undergoing metamorphosis to the juvenile stage. Host fishes which have been shown to support glochidia metamorphosis to the juvenile stage in the laboratory include Afurcagobius suppositus (Sauvage, 188051), Gambusia holbrooki (Girard, 185952), Nannoperca vitttata (Castelnau, 187353), Pseudogobius olorum (Sauvage, 188051) and Tandanus bostocki Whitley, 194454 but not Carassisus auratus Linnaeus, 175831 or Geophagus brasiliensis (Quoy & Gaimard, 1824 More

Effects of plant genetic diversity on multiple trophic groupsWe found that plant genetic diversity (i.e. diversification of cropping or plant cultivation systems; see Methods and Supplementary Table 15) decreased the overall performance of plant antagonists (effect size = −0.539, t = −2.070, P = 0.039) and several of its components (i.e., herbivores (effect size = −0.606, t = −4.127, P  More

General description of sequencesAfter the quality filtering step, removal of chimeric fragments, and read merging, a total of 3,378,323 reads with 3007 different features was obtained, with an average of 27,244 sequences per individual sample. After quality filtering, none of the samples was excluded from the analysis of microbial communities.Amoxicillin and thiamphenicol treatments influence microbial diversity and the abundance of specific taxaUsing 16S rRNA NGS, the gut microbial community composition of the chicks in each group was characterized at different time points. At phylum level, microbiota composition varied with age rather than with treatment (Supplementary Fig. S1). Proteobacteria were the most abundant phyla at 1 day of age (d.o.a.), Firmicutes became dominant at later stages, while Bacteroidota were highly abundant in caecum samples collected at 46 d.o.a. Similar dynamics were observed also at family level, since Enterobacteriaceae and Clostridiaceae were significantly more abundant at 1 d.o.a. in all groups, Lactobacillaceae, Lachnospiraceae, and Ruminococcaceae seemed to bloom at 8 d.o.a., and Rikenellaceae were the dominant family in the caecum samples collected at 46 d.o.a. (Fig. 1; Supplementary Fig. S2).Figure 1Heatmap representing the microbial community composition at family level. The heatmap was generated in R (version 4.2.1) (https://www.r-project.org/) using package pheatmap (version 1.0.12).Full size imageEarly-age administrationIn both α-diversity indices (Fig. 2A,B), there was a trend towards increasing diversity from early to late time points in all groups; however, the only significant differences were between the group treated with amoxicillin (AMX1) and the other groups on day 21 post treatment (p.t.), and within AMX1 group between day 21 p.t. and the other time points. PERMANOVA showed that the microbial community was significantly different between the group treated with thiamphenicol (THP1) and the other two groups (i.e. AMX1 and control) on day 1 p.t. (p  More

Total catch control regulation does not lead to the recovery of fisheries and the maintenance of community functionTo contain the decline of wild capture fisheries by overfishing, a series of management regulations have been in place in China to mitigate the fishing impacts as much as possible and maintain sustainable stocks. The “zero-growth” policy is one of the most outstanding representatives. The results showed certain achievements after the implementation of the policy. Simulating the status without the “zero-growth” policy, B/Bmsy fell below 0.5 by 2010 and close to zero by 2019, indicating the impossibility for recovery. However, the policy is not enough for fishery recovery and community health, failing to stop the degradation of fishery resources. Under the implementation of the “zero-growth” policy, B/Bmsy was in a healthy state in 1998, fell below 1 for the first time in 2003, and dropped to 0.52 in 2019, accompanying by F/Fmsy as 1.60. If fishing pressure were maintained at the level of 2019 (F = 1.56 Fmsy), the resource would decline to the depletion state by 2030 (B/Bmsy close to zero, F/Fmsy = 3.64, catch = 35 T). Therefore, a great degree of negative production growth as well as the strict implementation is extremely important. A rapid reduction in the catch control under 0.5 Fmsy scenario would expect to achieve a quick recovery with B/Bmsy over 1 in 2025. Nevertheless, a significant reduction in production would lead to the decline of fishery economics, livelihood difficulties for fishermen and a series of derivative social problems28. An alternative of 1.0 Fmsy would be feasible, under which B/Bmsy could rise to 1 by 2030 with a production of 11.64 MT, close to MSY.The “zero growth” policy faces some inherent challenges, at least from the point of view of ensuring the sustainable use of individual species stocks. Attention should be also paid at the catch quota control of individual species. Because the variation of the intrinsic growth rate of different species, the B is dynamic, and the F changes with the change of B. In a constant production, r-strategic species could remain a higher B/Bmsy than 1 even at a large proportion in catch, but K-strategic species did not show the same fortune. The control of total catch volume rather than individual species could not prevent the community structure from becoming fragile, with the exhaustion of high-trophic species and the decrease of mean trophic level.Individual species have different responses to overfishing that highly associated with their biological characteristicsHigh trophic level species can be sensitive to overfishing, and difficult to rebuild stocks after collapseHairtails Trichiurus spp. are the largest contribution group to China marine capture fisheries, at 0.90 MT about 8.3% of the total production in 20202. They are carnivorous and aggressive with a mean trophic level of 4.4, mainly feeding on fishes in the adult stage, and Mysidacea and Euphausiacea in the juvenile stage29,30. The spawning seasons of Trichiurus spp. are mainly from April to June, and from September to November in Chinese waters31.China coastal areas are excellent foraging and spawning grounds for Trichiurus spp, sustaining a large stock size. If the “zero-growth” policy was not implemented since 1999, the resources of Trichiurus spp. would be exhausted by 2027, having no possibility to recovery at 1.0 Fmsy. Although the total fisheries production has been controlled, and the fishing moratorium period partly covered the spawning seasons of Trichiurus spp., their resource continuous declined into a “destroying” state in 2007, due to the time-lag effect of fishing on high trophic level predators characterized by long population doubling time-consuming32. Under intensive fishing pressure, Trichiurus spp. have showed astonishing fisheries-induced adaption33 by reducing the age and size of maturity, which effectively alleviates the decline rate of B value, resulting the maintenance of Trichiurus spp. capture production. Under the rebuilding scenario of fishing pressure as 1.0 Fmsy, Trichiurus spp. B/Bmsy rose to 0.87 by 2030, lower than the recovery rate of national total capture fisheries, suggesting the recovery rate of high trophic level species could be slow34. Furthermore, in this study fisheries rebuilding only considers the responses of species to fishing pressure, irrespective of a series of factors sensitive to high trophic level species such as pollution and climate change, which indicated a longer period is needed for resource recovery.Middle trophic level species seems non sensitive to total catch control policyAs a representative of middle trophic level species, L. polyactis performed different from Trichiurus spp. Under high fishing pressure. It forms spawning and over-wintering aggregations between nearshore and offshore waters, as well as vertical migration, rising at dusk and falling at dawn35. The spawning season is from mid-February to early May, prior to the national fishing moratorium, indicating young juveniles are in effective protection rather than spawning stock. In the 1950s, L. polyactis was one of the few important species in domestic marine capture fisheries in Chinese waters, producing more than 100,000 T annually5. The catch volumes then showed a downward trend and fell significantly to less than 50,000 T in the 1960–1980s. After 3 decades low catch volumes, the annual capture production rebounded significantly to more than 200,000 T and maintained at such high levels for 2 decades5, showing high resilience to overfishing.Despite many concerns on the risk of resource exhaustion of L. polyactis stocks5,36, official statistics showed that the annual catch remains high. The L. polyactis production broke through 150,000 tons in 1995, and was above 300,000 tons after 2005. There is likely to have a large offshore stock of L. polyactis, which gradually joined the catch under increasing fishing efforts offshore. Furthermore, the L. polyactis stocks can be resilience to high pressure for several reasons: (1) its miscellaneous diet makes them be able to receive sufficient food sources; (2) size and age at sexual maturity reduced37,38; and (3) the over consumption of top predators relieves the prey pressure on middle trophic level species, such as L. polyactis, snappers, and flatfishes. A good job is the difficulties of artificial propagation and seedling breeding of small yellow croaker were broken for the first time in 201539 and the whole artificial cultivation was successfully realized in 2020 (https://www.chinanews.com.cn/cj/2020/07-02/9227715.shtml), which would effectively alleviate the market demand and wild stock sustain of small yellow croaker.Pelagic small fish stocks may not recovery quickly as early cognitionSmall pelagic fishes enjoy assembling in large schools of tens of thousands of individuals, and are more vulnerable to predators. Species S. sagax mainly filter plankton with a low trophic level about 2.8. It spawns in May–June, with high fecundity (an absolute fecundity of 30,000–100,000 pelagic eggs) and fast growth, and has short generation time of 1.4 years40. S. sagax shows strong phototropy, and can be caught using light purse seine, gill net, and fixed net fishing at night41,42.In 1989, the biomass of S. sagax was about twice of Bmsy. With the decreasing capture production of traditional economic fishes, S. sagax became a target species using specific fishing methods43, resulted in catch increase accompanied with B/Bmsy decline into a state of extremely unhealthy in 2019. Recovery of small pelagic species stocks would be delayed by the total catch control policy, mainly because the removal of large numbers of predator species left more opportunities for their feeding objects44. Resource rebuilding of S. sagax was not as quick as expected, as small pelagic species had to endure increasing predation pressure from the recovery of high-trophic species under the total catch control. At 1.0 Fmsy scenario, B/Bmsy would be only 0.88 by 2030, in need of a longer period to healthy state.Well-planned restocking can enhance resource recoverySwimming crab P. trituberculatus has high reproductive capacity, with a female can release two to three batches of eggs during a breeding season, and a batch contains about 1–6 million eggs45. Under the complementary of existing management measures and restocking programmes, the production of P. trituberculatus was kept in a certain amount close to a healthy state, and there is not an urgent need for its stock rebuilding. Since the 1990s, restocking of hatchery-produced larvae of P. trituberculatus has been promoted in coastal waters of China. Large-scaled restocking programmes were documented: 33 million larvae were released into the Yellow Sea by Shandong Province in June 2013 (http://hyj.shandong.gov.cn/xwzx/sjdt/201311/t20131120_507389.html); 50.3 million larvae with carapace width over 6 mm were released in the northern Yellow Sea by Liaoning Province in June 2020 (http://nync.ln.gov.cn/fwzx/zxdt/202007/t20200707_3902016.html); 16.1 million larvae were released into the East China Sea by Daishan County of Zhejiang Province in June 2021 (http://www.daishan.gov.cn/art/2021/6/8/art_1383064_59012675.html). What should be of concern is when, where, and how many seedlings are released46,47,48, to maximumly utilize the environmental resources without encroaching on the benefits of other species.Short-living species can be resilience to overfishingThe main cephalopod species in Chinese fisheries are Sepiella maindroni, mainly distributes in the East China Sea35 and Sepia esculenta, mainly distributes in the Bohai Sea, the Yellow Sea and the East China Sea49. As a 1-year lifespan species with fast growth rate, S. maindroni forms spawning migration from deep water to shallow nearshore bays in spring, partly within the fishing moratorium period. Due to the positive phototaxis, the cuttlefishes can be captured by light seining. Sepiella esculenta was the most important cephalopod economically in the northern coastal seas and one of the four major fisheries in the Bohai Sea and the Yellow Sea until the 1970s50. The abundance of this species has been greatly reduced with continuous fishing pressures and dwindling spawning grounds51.Total catch control and fishing moratorium showed significant output on the short lifespan cuttlefishes. Without the implementation of the “zero-growth” policy, the cuttlefishes resources would have been exhausted by 2015 and impossible to rebuild. According to the current state of resources, by 2030 the cuttlefish stocks can be recovered under the 1.0 Fmsy scenario. Moreover, the extent of cuttlefishes stock recovery relies on food supply.Ways to sustain fisheriesThe conflict between rising demand for fishery products and declining resources under multiple pressures including overfishing, climate change, and marine pollution has put heavy pressures at a global scale52. Chinese government has undertaken serious reforms to effectively replan the fishery industry.The effective recovery and rational utilization of resources depend on the support by sufficient reliable data. China started fishery statistics right after the foundation of the People’s Republic of China, completed by MOA (1949–2017) and MARA (since 2018). However, the statistical dataset has been questioned internationally53. According to the explanation by FAO54, before 2000s, especially from 1979 to the late 1990s, as the central government raced to meet the increasing demand for seafood and to grow the domestic production, the local governments had frequently overreported their local catch. In addition, fishermen may falsely claim to increase their production for surplus compensation, after the government introduced fishing subsidies. On the contrary, the production might have been underreported since the early 2000s55,56, which could be attributed to the existence of a large number of “black ships” (fishing vessels without relevant legal permits). Moreover, the lack of professionals in the early period and inaccurate knowledge of species identification by fishermen also lead to data uncertainty. Reasonable fisheries data should be consistent with the species functional traits and life history characteristics. However, in the actual fishing activities, the intentional and high-intensity selective fishing of species may greatly deviate the catch data from the data predicted by models. The Chinese government has been trying to improve the statistical system, including data coefficient adjustment, training of fishermen and professional, and supervision of statistical authorities5. In this study, selected objects are inshore species: the species are familiar to fishermen; the fishing vessel supervision is in place; the data collection is relatively rational and complete; all these are conducive to the reliability of the results.The zero-growth policy, which has been implemented since 1999, is an important measure in the history of marine fishery development and management in China. That is, the total catch of marine fisheries in the current year cannot be higher than that of the previous year. However, the “12th Five-year Plan” for national fishery development (2011–2015) issued by the Ministry of Agriculture canceled the mandatory targets of controlling the production but to encourage more catches of marine fisheries (http://www.moa.gov.cn/gk/ghjh_1/201110/t20111017_2357716.htm). In 2013, the State Council published the first state-level marine fishery development document as “Several Advices on Promoting Marine Sustainable and Healthy Development”, incorporating marine fishery development into the strategy of building a maritime power (http://www.gov.cn/zwgk/2013-06/25/content_2433577.htm). This policy shift was clearly reflected in the significant increase in the national annual catch from 12 to 14 MT. Until the “13th Five-year Plan” for national fishery development (2016–2020) issued in 2016, the zero-even negative-growth policy was revalidated, and the volume of annual output control was clearly proposed as 8–10 MT57, which was determined by multiplying the fishing coefficient by the total stock size derived from the assessment of surveys on the zoning of fisheries and the supplementary survey of marine biological resources in the exclusive economic zone and the continental shelf7. To achieve the target of keeping fishing capacity at a high level of sustainability, significant reductions in fishing pressures over a period of time are required, as well as rational updates of control policies.Many policies were introduced together or around the same time as the “zero-growth” policy, such as summer fishing moratorium, fishing license system, and fishing fuel subsides. However, the achievements are far from satisfactory. The fishing fuel subsidy policy together with the license system induced the direct fishing vessel construction boom which resulted in fewer but bigger and more powerful fishing vessels. Fishing moratorium is the most promising policy, by leaving enough time and space for fish to successfully reproduce. However, the truth is that, right after the fishing closure season, almost all fishing vessels immediately rush into the sea and fishermen try their best to fish as much as possible within the gears and engine power permission of their fishing licenses, attempting to earn a year’s income in a short period of 2–4 months. As a result of such high fishing effort, the achievements of seasonal fishing bans were largely offset and resource densities fell to low levels after autumn. The number of legally binding standards for mesh size is not enough, only 6 at present of at least 40 fishing target species and over 10 fishing gears, leaving many fishing gears and fish species outside the regulation of existing standards6,58. Ideally, standards of mesh size should be updated corresponding to the changes of species traits, however, it is a challenge because the main fishing mode is multiple species fishery by bottom trawling. Moreover, species in China seas are diverse, and the spawning period of different species may not fall into the fishing closure season5. The lack of specificity to sufficiently cover all the species may result an unbalance of community composition. Another system “Double Control” aims to limit both the numbers of fishing vessels and the total power. Unfortunately, the inspections of fishing vessels and their power are not very strict, due to the need of developing local economy and guaranteeing the fishermen’s income, e.g., under a nominal power mask the low-power engines have been replaced by high-power engines, some fishing vessels do not have the fishing licenses28. The limitation of the license number and engine power also stimulate the technological improvement for more catch7.The structure adjustment of fisheries composition is the main management measure at present. The high degree of self-sufficiency in fishery products in China has been achieved through overfishing of domestic fishery resources, resulting in the rapid depletion of fisheries in China’s coastal waters59. Aquaculture, accounting for more than 70% of China’s total fisheries production2, is identified as a successful way. Accompanying by aquaculture development, a series of problems also arise, particularly, the demand of low-value/trash fish and fish meal that significantly drives further expansion of capture fisheries60. Cooperation with other countries to promote regional aquaculture may be an alternative way to meeting global growing demand for seafood and combating overfishing61,62. Seeking resources from the high seas and EEZs of other countries is also a choice, of course, on the premise of taking full account of ecology, maritime, and food security of other countries63,64,65.In addition, this study pointed out a new focus for fisheries management, in which differences in species biological traits, including species vulnerability, population multiplication, and resilience to environmental pressures, should be given full consideration. On this basis, more detailed and targeted management schemes are supposed to propose to achieve the dual purpose of recoverable fisheries resource and balanced species composition, so as to become a truly sustainable fishery. In short, the effective implementation of various management measures is an indispensable guarantee. More

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Landscape use and anthropogenic influenceThe site could have had a specific and maybe extraordinary position in the microregion or in the trade networks41,42. The idea for creating trenches may have spread along trade routes—either as a habit of migrating people or as an ideology in the area of South and West Bohemia, Southern Germany, and the Austrian Land Salzburg55,56,57.Creeks along the settlement were major landscape elements. The settlement itself is entirely situated in the landscape periphery2. Steep slopes above Židova strouha creek and Blatenský potok brooks fundamentally limit agricultural use of the hinterland on the Březnice site, based on a model of reconstruction of the landscape potential (Fig. 6). The slopes may have been covered with sparse forest or shrubs. They were also forested in the nineteenth century, at the time of maximum agricultural load on the landscape as historical maps prove (Fig. 7).Figure 7Březnice and Hvožďany: the map of the second military mapping. Site catchements81 are according to the walk distance83 are shown hatched.Full size imageFieldsIn terms of human nutrition, the fields were crucial. The arable field area consisted of the actually cultivated fields and fallows. Analysis of plant macroremains provides us with knowledge of the grown species and the weed spectrum. The potential area and location of fields are reconstructed by a model that combines the agricultural potential of the landscape and previously published knowledge of the economic needs of the economic unit2,5,60,61,62,63.There is a possibility to assume, according to the SCA, the location of fields in relatively drier parts of the settlement area. Areas suitable for fields were probably located eastward and northward of the site, about 10–15 min walking distance (Fig. 6). The burial site was located beyond the northern border of the area where our analysis predicted the existence of fields93.Areas located eastward and northward of the settlement are even drier nowadays. The wetter fields may have been located in the north and northeast of the settlement, in its immediate vicinity. Moist soil is still present in these places today. The seeds and fruits of weed plants appear to have been transferred into the settlement together with the harvest. After being cleaned they were deposited as waste or used for further purposes, e.g. as an organic ingredient in ceramics or in daub4. The drier fields could correspond to finds of the following plant species: Arenaria serpyllifolia, Clinopodium acinos, Galeopsis augustifolia, Geranium cf. columbinum, Medicago lupulina, Rumex acetosella, Scleranthus annuus. Conversely, the following plants may have grown in the wetter fields, as documented in features on the settlement: Echinochloa crus-galli, Fumaria officinalis, Persicaria lapatifolia, Rumex cf. acetosa, Stachys arvensis.Synanthropic vegetation and ruderal habitatsArchaeobotanical analysis recorded many plant species characteristic for ruderal vegetation (most frequented Chenopodium album, Atriplex sp., Galium spurium, Polygonum aviculare, Chenopodium ficifolium, Fallopia convolvulus, Galium aparine). One could expect the presence of ruderals in the settlement area and its nearest surroundings in places that have been intensively used by humans and animals. The plants on the site could have reached the buildings by direct sedimentation and accidental charring, use of the ruderal plants, or as a result of waste burning.Deforested grazing areasGrazing took place in the enclosures and in the forests, which were made more open. The grazing of domestic animals had to be regulated in order to avoid crop damage and free movement around the settlement area. Winter fodder for animals had to be obtained within the reach of the settlement area, which contributed to the further lowering density of the forest. The archaeobotanical data reflect the grazing habitats in forest and deforested areas. Detrended correspondence analysis shows two clusters of plant species compatible with such environment (Fig. 4). The question is the process by which the plants reached the settlements. Species which appear in the ordinary space between the grassland and woodland—shrub positions could have grown on grasslands and light forests (e.g. Lychnis flos-cuculi, Dianthus cf. armeria, Galium palustre, Festuca ovina, Juncus sp., Campanula cf. glomerata) species in the ordinary space between “ruderal” and “grassland” could have grown at both habitats, e.g. at the transition of the settlement to the open countryside (e.g. Achillea millefolium, Alopecurus pratense, Asperula cynanchica, Briza media, Festuca cf. pratensis, Galium cf. verum, Ranunculus cf. bulbosus, Silene vulgaris, Stellaria graminea, Trifolium pratense). Taxa displayed between the “field” and “grassland” could have grown for example on fallow lands or abandoned fields that have successively overgrown (e.g. Clinopodinum acinus, Plantago lanceolata, Trifolium repens, Polycnemum arvense, Trifolium arvense). Taxa typical for “field” and “woodland-shrub” significantly differ in Březnice (Fig. 8).Figure 8Březnice: detrended correspondence analysis (DCA) Displayed samples and botanical taxa: the first axis explains 44.57% variability, the first and the second axis together 50.47%.Full size imageThe archaeobotanical analysis captured multiple grassland types. Both drier and wetter environments can be reconstructed. Wetter areas were represented by e.g. Alopecurus pratense, Alopecurus geniculatus, Carex cf. hirta, Carex cf. vulpina, cf. Euphorbia palustris, Galium cf. palustre, Juncus sp., Lychnis flos-cuculi, Myosotis sp., Persicaria lapatifolia, Plantago lanceolata, Stachys cf. palustris, Stellaria graminea, Urtica dioica. Drier areas were represented by e.g. Asperula cynanchica, Briza media, Campanula cf. glomerata, Carex cf. contigua, Clinopodium acinos, Dianthus cf. armeria, Phleum sp., Festuca cf. ovina, Galeopsis augustifolia, Galium cf. verum, Medicago lupulina, Polycnemum arvense, Ranunculus cf. bulbosus, Scleranthus annuus, Silene vulgaris, Solanum nigrum, Spergula arvensis, Trifolium arvense, Vicia tetrasperma, Vicia cf. villosa (Fig. 8).The existence of grasslands is associated with long-term human activities94. The Bechyně region has been apparently continuously settled since the end of the Early Bronze Age34. The landscape around the settlements has always been influenced by human activity and a large part of it has been deforested or covered with a sparse pastoral forest. However, not all the settlement areas were occupied permanently3, and those which were unoccupied became overgrown.Meadows and pastures are much more suitable for the grazing of herbivores than a forest with a dense canopy. Forest-steppe or significantly open forest is a convenient combination ensuring sufficient grazing for animals and wood production. Grazing increased soil fertility, reduced weeds on ruderal sites, and prevented forest growth95. Our study recorded a wide spectrum of charred macroremains of plants, which grew in the grasslands. They could have reached the site in several ways. In the excrements of the animals coming from a grazing area96, as raw materials collected by humans for further use in the settlement economy (e.g. food, medicinal plants, dyeing plants, bedding, admixture of screed and ceramic earth and daub, etc.). Studies1,3,32 assume, that the area in the immediate vicinity of the site was probably forestless. Forests at least half an hour’s walking distance from the site was significantly influenced by human activity. With an increasing distance from the centre of the site, the forest was probably less affected by human activities. The character of woodland usually clearly corresponded with the environmental conditions of the location31. The current forest area is extremely unsuitable for usage (slopes, wetlands). We assume that the occurrence of woodlands and shrubs in the Late Bronze Age was much more widespread, even in less extreme habitats.Shrubs and forestSpecies of herbs from different forest and shrub environments were also frequently recorded in the archaeobotanical assemblage. In the environment of wet forests could have grown e.g. Alliaria petiolata, Galium cf. palustre, Galium odoratum, Galium sylvaticum, Lychnis flos-cuculi, Persicaria lapatifolia, Solanum dulcamara, Stachys cf. palustris. In the coastal shrubs and edges of wet forests could have occured e.g. Cuscuta cf. europea, cf. Euphorbia palustris, Chelidinium majus, Impatiens nolitangere, Juncus sp., Myosoton aquaticum, Urtica dioica, Veronica hederifolia. Suitable locations could have been along the streams that flowed around the settlement and were within a quarter-hour walk. On the edges of the forests and their glades could have grown e.g. Atropa bella-donna, Festuca cf. ovina, Galium aparine, Prunella vulgaris, Rumex acetosella, Silene dioica, Thymus sp. Light forests and slopes were suitable for e.g. for Campanula cf. glomerata, Carex cf. contigua, Dianthus cf. armeria, Geranium cf. columbinum (Fig. 8).The areas for hunting and harvesting of wild crops were also economically important. The fruits that could have been collected included Corylus avellana, Crataegus sp., Atropa bella-donna, Prunus spinosa, Quercus sp., Rubus ideaus, Rubus fruticosus, Sambucus nigra, Solanum nigrum, Solanum dulcamara; their remains were found in the infills of features. The source of the collected fruits was located mostly in the sparse forest, forest edges and shrubs.The forest was also a source of building material and firewood3. From this acreage, the firewood for one farm could have been collected from 10 hectares. The rest would be used for collecting fodder and forest grazing7. The map of the potential natural vegetation92 predicts acidophilous oak forests (Quercetea robori-petraeae, Fig. 7) for the majority of both settlement areas. These species-poor woodlands are characteristic of Quercus dominance and in places mixed with Betula, Pinus, Sorbus, and Tilia on both dry and wet acidic soils, and Fagus, Abies, or Picea at higher altitudes. The results of our anthracological analysis clearly documented the predominance of this vegetation type in the vicinity of both archaeological sites.In the valleys of the streams and rivers were reconstructed alluvial forests with Alnus and mesophilous oak-hornbeam woods. The archeobotanical analysis of charcoals and fragments of fruits detected presence of Quercus, Tilia, Corylus, Crataegus, and Carpinus. These macroremains indicate existence of mesophilous forests. The hornbeam is rare in southern Bohemia97, it is the first of the archaeobotanical finds from prehistory. Due to the structure of taxa, which was captured by archaeobotanical analysis in Březnice, meadows and alder tree woods may be assumed there. Results of archaeobotanical analysis also documented the presence of Salix/Populus, Alnus.The most dominant tree species discovered in the trench-like features was oak which was mainly used as a construction material (Fig. 5). Firs were used as construction wood, which is predominantly present in stake pits in Březnice. In Hvožďany, trench 1 contained a cultural layer with apparent remains of a destructed building with charcoals of fir, spruce, and pine which in this case also served as construction wood34. The material commonly available in the forests surrounding the settlement area served as firewood (Figs. 4, 5, 8, 9).Figure 9Hvožďany: detrended correspondence analysis (DCA) Displayed samples and botanical taxa: the first axis explains 64.08% variability, the first and the second axis together 72.12%.Full size imageTime of housing: landscape potential vs. human needsThe homestead management (construction, abandonment, destruction, reconstruction etc.) during the settlement´s lifespan is a long-term studied question98,99. The existence of a hierarchized Late Bronze Age settlement network was evident in the lowland settlement areas of the Czech Republic with the continuity of occupational activity. Two main types of settlement are usually recognized there: (1) long-term large settlements and (2) short-term small settlements100,101. Agricultural productivity, exploitation of natural resources in settlements areas, and trade networks differed in cases of small or large settlements102. From the archaeological evidence perspective, the South Bohemia region was sparsely populated and the presence of long-term large settlements areas was very rare34.Previous research (excavations and magnetometry survey) has led to the conclusion that the 70 trenches are depositions of 70 houses and each trench is a deposition of one original house4,5,58. Based on such data, there could be many settlement forms differing in the space and time. The possible size of the settlement could be derived from the comparison of demands for fields, pastures, and forests with carrying capacity.SCA model and prediction model when compared to the possible demand7 of the community show that forest and pastures were not limiting factor for the settlement sustainability. In case of fields, there could be four variants of the possible extent of the settlement connected with different intensity of landuse. (1) The optimal acreage of fields (69 ha) with optimal land-use (7.5 ha/household); (2) the maximal extent of the fields 104 ha with optimal land-use or optimal extent of the field systems with intensive land-use (5 ha); (3) the maximal extent of the fields 104 ha and intensive land-use (5 ha); (4) sub-optimal land-use and fields located outside of the reach and optimal soils (Table 2). This model is an ideal prediction. For better yield the farmer could travel longer time than is expected however poor soils on a sloped terrain in the close vicinity were probably used rather as pastures.Table 2 Březnice: possible duration of the settlement based on four land use strategies: light green-optimal extent of the fields (69 hectares), with 7.5 hectares of fields per homestead; dark green-maximal extent of the fields (104 ha) or more intensive use of the fields (5 ha/homestead); maximal use and maximal extent; red—not sustainable agriculture or location of fields on places outside predicted optimal areas.Full size tableDrawing upon the typological and radiocarbon dating, it is often impossible to find out what was the lifespan of the settlement on the actual site. In this case, the uncertainty of 14C dates gives us a maximum possible span 73–264 years (95% probability), probably for 107–192 years (68% probability) (Supplementary Table 1, Fig. 2). Typological dating indicates 100–150 years (1150–1000 BC).The model described above indicates that the hinterland of Březnice could have sustained up to 20 houses at the same time in case of the maximal extent of the fields and intensive land-use. In this case, the settlement would have lasted only 90 years. If the land was used extensively it could have bore maximum of 14 houses at the time. That would correspond to a duration of roughly 126 years. Optimal areas of field systems in combination with sufficiently large fallows could have been used by a maximum of nine houses present at the time (192 years). The crucial part of the model is ritual burning and rebuilding houses after one generation58.Models of potential spatial and temporal characteristics of the settlement derived from prediction modeling cannot be tested. Therefore we need to compare our predictions with the radiocarbon model. The shortest duration of the settlement based on prediction is 90 years which corresponds with the 72 years modelled from 14C data. Since the model does not reflect the maximal duration of dwelling, this limit has to be based only on 14C model (262 years at 95% probability. At the maximum possible landuse levels, the settlement could have lasted from 72/90 to 262 years. The optimal duration of the settlement based on prediction could be 192–262 years. Extensive but more demanding land-use could support the duration of the settlement from 126 to 262 years (Table 2).Březnice and Hvožďany: the interpretation of both settlement areas from an archaeobotanical perspectiveThe two similarly dated settlement areas in one microregion with high quality archaeobotanical data allow (based on archaeobotanical material) a detailed study of the behaviour of communities in the Late Bronze Age. Archaeobotanical assemblages bring the reconstruction of the environment where the communities of the settlements drew plant resources from. Although the number of plant remains from both sites is significantly different, the interpretation of the environment does not differ in broad terms. For both sites, a similar share of fields and ruderals was documented. The spectrum of cultivated species was also identical41. Both settlements were self-sufficient in plant production—both waste and production parts of cultivated plants were found in the assemblages21,34,41. Animal bones were not preserved due to the acidic soil. However, for the Late Bronze Age sites the types of the domestic103 and the hunted104 animals are known.According to the environmental model, a greater proportion of species in Březnice came from grassland rather than from woodland and shrubs (Fig. 4). According to the analysis of plant macroremains more deforestation was recorded (i.e. more fields and pastures) in Březnice than in Hvožďany (Figs. 4, 5, 8, 9). Predicted areas for fields were in case of Hvožďany from 27 to 130 ha. Hvožďany site could possibly have larger field systems, but further away than in case of Březnice settlement. In Hvožďany there have been documented many taxa typical also for ruderal sites and fields. Several taxa could have grown either on ruderal sites or grasslands. Three reconstructed environments (ruderals, fields, grasslands) in Hvožďany significantly differ from woodland—shrub (Figs. 8, 9). The large volume of analysed samples from Březnice brought a number of botanical taxa which was mostly found in only a few specimens but ultimately brought the opportunity to reconstruct the surroundings of the site in more detail. In Hvožďany, a common spectrum of plants was found (Fig. 9), which usually occurs at similar South Bohemian sites, e.g. Černýšovice, Rataje, Zhoř, Oldřichov, Písek—Bakaláře105,106. Nevertheless, it brings the possibility to reconstruct the surroundings at least in rough features.The archaeological field data does not allow us to reconstruct how many houses were on the Hvožďany site at the same time. Total inhabited area of ​​the settlement in Březnice is approximately 13 ha, at Hvožďany site it is altogether 5 ha. It suggests two explanations: either more people lived in Březnice than in Hvožďany or the settlement had a longer span (or both possibilities). However, both options mean greater deforestation in Březnice. The carrying capacity and landscape potential of the settlement in Hvožďany could not have been exhausted (Fig. 6). The area of high quality soil in a quarter/half hour’s walk from the site is sufficient for 3.6–25 houses (27–130 ha). Two community areas could have been separated by the Lužnice river (walking distance within one hour). The agricultural systems of the settlements were probably very similar. According to our models, both settlement sites would have only needed to exploit natural resources in their immediate hinterland, within an hour walking radius. The limiting factor is the availability of suitable land for fields.According to the archaeobotanical results, the landscape in Březnice was more affected by human activity than the one in Hvožďany. A greater number of species were found, evidenced by light woodland and shrubs and different types of grassland. In the vicinity of the settlement from which people drew resources, a light landscape can be assumed. So far there is no pollen profile available. Approximately 2 m of accumulated clay and sand without organics were sampled in the floodplain of the Židova strouha. About 20 km away from Březnice, the analysis was performed in Sepekov, which base could have corresponded to the Bronze Age (2920 ± 410 BP). The character of the vegetation based on the profile could be interpreted as wet and relatively nutritious fir woodland or fir alder woodland situated on a relatively small spring area at the edge of the water meadow of the Smutná river. The palaeobotanical record in this phase does not record any effect of the settlement on the vegetation present34. The profile containing the pollen record from the Borkovická blata is located about 10 km away from Březnice. As well as the profile from Sepekov, it reflects local peat bog vegetation of the subboreal character without significant indicators of human activity107.The conditions and availability of resources in the hinterland of both settlements were probably overall so good that the details did not matter much. In the vicinity of both settlements, there were a sufficient number of areas for fields, pastures, and cultural forests. The settlement areas of the Late Bronze Age in South Bohemia were probably in separate deforested niches. More