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Our analysis suggests that, over 465 ha of mangrove area, almost 83% of aboveground tree biomass were harvested annually for commercial timber purposes, using a keyhole harvest pattern (Fig. S3b). Yet after 25 years of natural and human-induced regeneration, both field- and satellite-based assessments reveal that biomass carbon stocks and canopy cover had fully recovered. Our approach using space-for-time substitution indicates that manual selective logging did not significantly affect soil carbon stocks and rates of annual carbon burial. While the differences in soil carbon stock between sites may be due to the diverse hydro-geomorphic settings8,14 the mangrove root mass in the top 1-m were not disturbed by manual logging activities. Similar situation was found in Tampa Bay, Florida where peat formation from root mass has enhance carbon sequestration15. These findings reduce uncertainty around the effects of mangrove forest management on the long-term functional capacity of blue carbon storage and provide evidence that managed mangrove ecosystems may deliver nature-based climate solutions.Recovery of forest structure, canopy cover and species diversityAlong carbon stocks, forest structure and species diversity also demonstrated recovery (Fig. 2, Table S1). Seedling densities were significantly higher in 5 year-old mangrove plots than in plots at any other stage (F(5,13) = 28.321, p  More

Genetic diversity of E. chinense based on COIThe partial fragment of COI, 595 bp in length, was sequenced from 113 E. chinense individuals collected from the eleven sites in South Korea (Table 1). The resultant COI sequences were aligned together with 27 COI individual sequences12,13,14 retrieved from the NCBI GenBank (Table 1). The latter consists of 26 from six collection sites in South Korea and one from a Japanese site. Hence, 140 COI sequences of E. chinense were analyzed, representing 18 collection sites at the nine populations in South Korea and Japan (Table 1). Based on the alignment set (no indels) of these 140 COI sequences (Data S1), we obtained a total of 58 COI haplotypes, of which 43 were singleton, appeared in only a single site. The novel 41 out of 58 COI haplotypes obtained were registered under the GenBank accession nos. MW265437–MW265477 (Table S2). According to the sequence alignment of the 58 COI haplotypes (Fig. S2; Data S2), there were 71 polymorphic sites and 31 parsimoniously informative sites (Fig. 1C), among which four adenine/guanine hotspots at 207, 282, 354, and 420 were ascertained to articulately divide the haplotypes of E. chinense into four meaningful phylogenetic groups: (a) A(207)–A(282)–A(354)–A(420), (b) A–A–G–A, (c) G–A–G–A, and (d) G–G–G–G.Table 1 List of collection sites and the number of individuals of Ellobium chinense with genetic markers applied to each of the nine populations in South Korea and Japan.Full size tableBased on the COI haplotype sequence alignment (Fig. S2; Data S2), we reconstructed a ML tree using Ellobium aurisjudae as an outgroup. In the resultant tree topology (Fig. S3), it was confirmed that E. chinense appeared as a monophyletic group, but no distinction between the haplotypes from each geographical population was observed. To define detailed relationships among the COI haplotypes, the outgroup was removed and then an unrooted ML tree (Fig. 1D) was reconstructed. The resultant tree showed two distinctive phylogenetic groups, namely A–A–A–A and the other groups (including at least one G or more in the four positions), regardless of collection localities. The A–A–A–A group included 35 of the 58 COI haplotypes. The others could be divided into the A–A–G–A group (N = 12: ECH11, 12, 15, 16, 18, 23, 27, 28, 32, 33, 36, and 49), the G–A–G–A group (N = 1: ECH35), and the G–G–G–G group (N = 8: ECH01, 07, 19, 29, 41, 45, 48, and 54).As shown in Table S2 and Fig. S3, ECH01 was a dominant member of the G–G–G–G group with the most individuals (27), which appeared across all the South Korean populations examined here. As shown in Fig. 1D, the A–A–A–A group is likely to be an ancestral type because it was most frequently found in the other species within Ellobiidae (unpublished data) and its haplotype diversity was the highest among the four genetic groups. Given that the G–G–G–G group exhibited much lower haplotype diversity than the A–A–A–A group, and was not observed in any other ellobiid species (unpublished data), it is reasonable to suggest that the G–G–G–G group is a derived rather than an ancestral type. Thus, as shown in Fig. 1D, it is conceivable that unidirectional and stepwise A → G transition events from A–A–A–A to G–G–G–G may have been occurred in E. chinense. Within the A–A–A–A and A–A–G–A groups, parsimoniously informative A → G transition events were found at the sites 120 (ECH12, 15, 16, 18, 23, 38, 40, 44, and 49) and 183 (ECH3, 9, 10, 20, 43, 50, and 55), with a few exceptional cases of G → A at the sites 216 (ECH12, 15, 16, 18, and 23), 372 (ECH12, 15, and 23), and 429 (ECH37, 38, 46, and 47; ECH19 found in the G–G–G–G group).As indicated in Table 2, the nucleotide diversity (π) is relatively low among the nine populations of E. chinense, ranging from 0.00749 (population BG) to 0.01042 (SC) with an average of 0.00865, whereas the haplotype diversity was very high across these populations, ranging from 0.924 (YG) to 1.000 (SC and JB) with an average of 0.939). All values of Tajima’s D and Fu’s FS were congruently negative, with averages of − 1.87100 (P  More