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    Land and people

    Africa’s population is rapidly growing, with its share of the global population projected to increase from 17% in 2020 to 39% by 2100 (ref. 8). The continent is already grappling with low agricultural productivity and food security challenges. Tremendous efforts are needed to increase food production; however, arable land continues to undergo widespread degradation due to issues such as nutrient mining, erosion, overgrazing and pollution. Climate change and more frequent weather extremes, such as floods and droughts, further degrade land and reduce agricultural productivity.Some efforts to counteract low productivity, however, can increase greenhouse gas emissions and derail efforts to meet global climate targets. Poor water management, fertilizer application and residue burning in rice production are, for example, major sources of potent greenhouse gases such as methane and nitrous oxide9,10. To ensure that the United Nations sustainable development goals and the African Union’s Agenda 2063 for food and water security are realized at minimal environmental cost, science-based land management practices are needed to decouple agricultural productivity from greenhouse gas emissions.
    Credit: majimazuri21/PixabayThe Agriculture, Forestry and Other Land Uses (AFOLU) sector contributes the largest share of greenhouse gas emissions in Africa11. Thus, developing large-scale agronomic, livestock and forest management practices that increase productivity and reduce emissions is key to achieving enhanced production and environmental sustainability. However, it is impossible to effectively manage greenhouse gas emissions if there is limited capacity to quantify them in Africa.Improved data infrastructure and research are needed to quantify emissions associated with specific land management practices under different land uses. Similarly, land use mitigation strategies should be informed by existing and potential future land use changes and their impact on greenhouse gas emissions under different climate scenarios. However, past studies that examined land use changes at various temporal scales mainly used coarse resolution satellite imagery and suffered from limited availability or poor-quality of data, partly due to cost. Such challenges have resulted in limited knowledge of land management practices that reduce greenhouse gas emissions while increasing agricultural productivity.Improved greenhouse gas observation networks and in situ measurements12 will enable the development of country-specific emission factors (IPCC tier 2/3)13 and quantification and management of land use specific greenhouse emissions. It will reduce uncertainties in emissions inventory data on Agriculture, Forestry and Other Land Uses14, which are currently estimated using emission factors extracted from default value databases (tier 1 methodologies).Free earth observation data, such as those from the European Space Agency and United States Geological Surveys, are becoming increasingly available. Together with improvements in cloud-based computing infrastructure, this presents an opportunity to advance research into current and future land use and vegetation dynamics. Coupled with accurately quantified greenhouse gas emissions, this can support current and future land management practices that contribute to mitigation and adaptation objectives of countries. More

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    Liolophura species discrimination with geographical distribution patterns and their divergence and expansion history on the northwestern Pacific coast

    Sample collection of Liolophura japonica and the genetic diversity of COI barcoding regionTo examine genetic lineage divergence within L. japonica on the northwestern Pacific coast, we newly collected a total of 342 L. japonica samples from 12 sampling localities in the intertidal coasts of the Korean Peninsula and Japanese Archipelago (Fig. 1; Table S1). From the collected L. japonica samples, we amplified the COI barcoding region using PCR, and then sequenced the 635-bp PCR products. As a result, a total of 75 COI haplotypes based on COI sequences obtained from 342 individuals of L. japonica were detected via the present study (Table S2). In addition to this, we extracted 31 COI haplotypes based on COI sequences from 127 individuals of L. japonica (also known as Acanthopleura japonica) previously reported in the NCBI GenBank database, consisting of two Japanese and 29 Chinese COI haplotypes. Finally, we gathered 106 COI haplotypes from 469 L. japonica individuals collected in 15 localities of South Korea, Japan, and southern China (Tables S1, S2). The average haplotype (h) and nucleotide diversities (π) were 0.808 and 0.04936, respectively; the highest haplotype diversity was observed in Tsushima (TS; h = 0.963), and the highest nucleotide diversity was found in Wando (WD; π = 0.04581), located in the South Sea of the Korean Peninsula. As shown in Table S3, the population distribution pattern of COI haplotypes revealed that all collection sites had site-specific haplotype(s) except for Busan (BS), Wando (WD), Sinan (SA), and Jeju Island (JJ). The most abundant haplotype was A1, which was found in 170 (39.4%) out of the COI sequences obtained from 469 L. japonica individuals.Figure 1A map showing sampling localities and photos of a habitat landscape and wild samples of Liolophura japonica inhabiting coastal areas of the Korean Peninsula (N = 249), the Japanese Archipelago (N = 57), and southern China (N = 125) in the northwestern Pacific Ocean. (a) A map showing twelve direct sampling localities for L. japonica in coastal areas of the northwestern Pacific Ocean. The sampling localities of one southern Chinese (ZJ) and two Japanese (EH and MY) previously catalogued haplotype sequencing studies retrieved from NCBI are also depicted. Table S1 and S2 contain more accurate information on the populations and individuals. The basic map is from a free map providing site (https://d-maps.com), which is modified with Adobe Illustrator v.25.2. (https://www.adobe.com). (b) Photos of a habitat landscape and wild samples of L. japonica, taken from Seogwipo-si, Jeju Island, South Korea, photographed by Mi Young Yeo, Bia Park, and Cho Rong Shin. The photos were edited using Adobe Photoshop v.22.2 (https://www.adobe.com).Full size imagePhylogenetic and population genetic analyses based on COI
    We constructed a nucleotide sequence alignment set with 106 COI haplotypes of L. japonica (Data S1), and identified 95 polymorphic sites (15.0%, Table S4) and 68 parsimoniously informative sites (10.7%). To elucidate phylogenetic relationships among the populations of L. japonica, we performed molecular phylogenetic analyses, including maximum likelihood (ML), Bayesian inference (BI), and neighbor-joining (NJ) analyses, based on these 106 COI haplotypes with the outgroup Acanthopleura spinosa (Fig. 2a, Figs. S1, S2). The resultant phylogenetic trees clearly revealed the existence of three distinct genetic lineages within the monophyletic group of L. japonica (100 BP in ML, 1.00 BPP in BI, and 100 BP in NJ): Lineage N (91 BP, 1.00 BPP, and 100 BP), Lineage S1 (79 BP, 0.82 BPP, and 98 BP), and Lineage S2 (98 BP, 1.00 BPP, and 100 BP). Among these three genetic lineages, Lineages S1 and S2 were grouped with high node confidence values (94 BP, 1.00 BPP, and 95 BP). We additionally conducted a phylogenetic network analysis using a neighbor net algorithm without an outgroup (Fig. 2b), which confirmed that these sequences were distinctly divided into three genetic lineages, in agreement with the topology of the rooted phylogenetic trees (Fig. 2a, Fig. S1).Figure 2Phylogenetic, TCS network, and PCoA analyses based on 106 COI haplotypes from 469 individuals of Liolophura japonica inhabiting coastal areas of the northwestern Pacific Ocean, suggesting the existence of the three different genetic lineages: Lineage N, Lineage S1, and Lineage S2. (a) Maximum likelihood tree showing the three different genetic lineages for L. japonica: Lineage N members are most likely from the populations inhabiting a wide range of South Korea and Japan, Lineage S1 members from the populations inhabiting southern coastal areas of South Korea and Japan only, and Lineage S2 members from the southern Chinese population. As shown in Fig. S1, Acanthopleura spinosa was used as an outgroup. Numbers on branches indicate node confidence values: BP in ML, BPP in BI, and BP in NJ in order. (b) A phylogenetic network reconstructed using the neighbor net algorithm without an outgroup, showing three different genetic lineages for L. japonica inhabiting the northwestern Pacific coast: Lineages N, S1, and S2. The COI sequence alignment set used is shown in Data S1. Detailed information of the 106 COI haplotypes used in this phylogenetic analysis is summarized in Table S1 and S2. (c) An unrooted TCS network showing three distinct genetic clusters, corresponding to Lineages N, S1, and S2. Three different genetic groups correspond to the three genetic lineages shown in the phylogenetic tree (a), respectively. The haplotype frequency is displayed by the circle size. (d) A two-dimensional PCoA plot showing the three distinct genetic groups corresponding to Lineages N, S1, and S2 shown in the phylogenetic tree (a). The score on the first two axes (Axis 1 = 79.05% and Axis 2 = 15.32%) from the matrix of genetic distances estimated with the 106 COI haplotypes are indicated.Full size imageConsistently, the TCS network analysis (Fig. 2c) and principal coordinate analysis (PCoA) (Fig. 2d) showed the existence of three distinguished genetic groups among L. japonica, in accordance with the phylogenetic analyses (Fig. 2a − b). The TCS network (Fig. 2c) revealed that Lineages S1 and S2 were separated by 18 mutation steps, which is far shorter than the distance between Lineages N and S1 (37 mutation steps) or between Lineages N and S2 (60 mutation steps), indicating that Lineages S1 and S2 have a close affinity and had only recently diverged from each other. The overwhelming dominancy of the A1 haplotype implies a recent and rapid population expansion of Lineage N. In addition to A1, it was found that haplotypes B2 for Lineage S1 and C21 for Lineage S2 were dominant. In the PCoA plot (Fig. 2d), the three genetic groups of L. japonica were also observed, as in the phylogenetic (Fig. 2a,b) and TCS network (Fig. 2c) analyses. Lineage N was distantly located from Lineages S1 and S2, while Lineages S1 and S2 were spatially much closer.Sample collection of L. japonica and the genetic diversity of 16S rRNA
    The 342 individuals of L. japonica from 12 localities in the intertidal coasts on the Korean Peninsula and Japanese Archipelago (Fig. 1) were subjected to PCR amplification of a partial region of 16S rRNA (506 bp) (Tables S5, S6). Of these, only 299 samples were successfully amplified and sequenced. Based on 299 individual 16S rRNA sequences, a total of 23 16S rRNA haplotypes of L. japonica were detected (Tables S5, S6). Combined with 11 haplotypes extracted from 16S rRNA sequences of 125 L. japonica individuals known previously in southern China, we totaled 34 16S rRNA haplotypes from 425 L. japonica individuals in 13 collection localities. The average haplotype (h) and nucleotide (π) diversities were 0.702 and 0.02093, respectively; the highest haplotype diversity was found in Geojedo (GJ; h = 0.833), and the highest nucleotide diversity in Wando (WD; π = 0.02244), located in the South Sea of the Korean Peninsula. Overall, the average haplotype and nucleotide diversities of 16S rRNA were lower than those of COI (Table S1). As shown in Table S7, the population distribution pattern of 16S rRNA haplotypes revealed that most of the collection sites had site-specific haplotype(s), except for BS, GJ, WD, SA, JJ, and TT. The most abundant haplotype was RA1, which was found in 186 (48.1%) out of the 16S rRNA sequences obtained from 425 L. japonica individuals.Phylogenetic and population genetic analyses based on 16S rRNA
    We constructed a nucleotide sequence alignment set with 34 16S rRNA haplotypes of L. japonica (Data S2), and identified 35 polymorphic sites (6.9%; Table S8) and 24 parsimoniously informative sites (4.7%). Phylogenetic analyses, including ML, BI, and NJ analyses, were conducted with the outgroup Acanthopleura echinata (Table S6). The resultant phylogenetic trees (Fig. S2) and unrooted phylogenetic network (Fig. 3a) consistently supported the three distinct genetic lineages of L. japonica, with the phylogenetic relationship between Lineages S1 and S2 being much closer than those inferred from the results of COI (Fig. 2a,b; Fig. S1). The TCS network (Fig. 3b) revealed that Lineages S1 and S2 were closely connected with only with 4–5 mutation steps between them, while Lineages N and S1 or Lineages N and S2 were remotely distanced by 18 mutation steps. Also, the overwhelming dominance of the RA1 haplotype implied a recent and rapid population expansion of Lineage N. In addition to RA1, haplotypes RB1 for Lineage S1 and RC1 and RC2 for Lineage S2 were dominant (Fig. 3b; Table S7). Consistent with this, in the PCoA plot (Fig. 3c), the three genetic groups of L. japonica were spatially separated. Lineage N was distantly located apart from Lineages S1 and S2, while Lineages S1 and S2 were spatially much closer.Figure 3The results of phylogenetic and population genetic analyses based on 34 16S rRNA haplotypes from 425 individuals of Liolophura japonica inhabiting coastal areas of the northwestern Pacific Ocean. (a) Phylogenetic network reconstructed using the neighbor net algorithm, showing three different genetic lineages for L. japonica: Lineage N, Lineage S1, and Lineage S2. The 16S rRNA sequence alignment set used is shown in Data S2. Detailed information of 34 16S rRNA haplotypes used in these analyses is summarized in Table S5 and S6. (b) An unrooted TCS network. There are distinctly observed three different genetic groups, corresponding to the three genetic lineages shown in the phylogenetic network (a). The haplotype frequency is displayed by the circle size. (c) A two-dimensional PCoA plot showing the three distinct genetic groups, corresponding to Lineage N, Lineage S1, and Lineage S2. The score on the first two axes (Axis 1 = 87.77% and Axis 2 = 4.4%) from the matrix of genetic distances estimated with the 34 16S rRNA haplotypes are indicated.Full size imageExamination of species discrimination of L. japonica based on COI and 16S rRNA
    Using the Automatic Barcode Gap Discovery (ABGD), we performed distribution of pairwise genetic divergences, ranked pairwise difference, and automatic partition analyses based on COI and 16S rRNA of L. japonica, respectively (Fig. 4a–c), which confirmed that there were distinct barcoding gaps between intraspecific and interspecific variations, strongly supporting the possibility of species discrimination of L. japonica. the COI-based analysis yielded two different barcoding gaps, while the 16S rRNA-based analysis revealed only a single barcoding gap (Fig. 4a–c). The results of automatic partition at each value of the prior intraspecific divergence (P) divided L. japonica into three groups by COI and two groups by 16S rRNA, respectively (Fig. 4a–c). We also implemented two DNA taxonomy approaches to evaluate the possibility of species discrimination based on COI: the general mixed Yule coalescent (GMYC) approach (Fig. S3) and a Bayesian implementation of a Poisson Tree Processes model (bPTP) (Fig. S4). The results consistently and robustly supported the possibility that L. japonica can be divided into three different species, as shown in the results of ABDG (Fig. 4a–c).Figure 4Distribution of pairwise genetic divergences, ranked pairwise difference, and automatic partition based on COI and 16S rRNA haplotypes of Liolophura japonica and a COI-based NJ tree showing the phylogenetic relationship with a congeneric species L. tenuispinosa. (a) Distribution patterns of pairwise genetic divergences observed in COI and 16S rRNA for L. japonica. The horizontal axis represents intervals of pairwise Kimura-2-parameter (K2P) genetic distance in percentage, and the vertical axis represents the number of individuals associated with each distance interval. (b) The results of ranked pairwise differences based on COI and 16S rRNA, ranked by ordered value, which is similar to the distribution of pairwise genetic divergence in (a). The horizontal axis indicates a ranked ordered value based on K2P genetic distance, and the vertical axis represents the K2P genetic distance in percentage. (c) The results of automatic partition analyses based on COI and 16S rRNA. The horizontal axis represents the prior maximum intraspecific divergence (P), and the vertical axis represents the number of groups inside the partitions (primary and recursive). (d) A COI-based NJ tree with L. tenuispinosa. Refer to Fig. S3 and Data S3.Full size imageThe molecular variance analyses using analysis of molecular variance (AMOVA), based on COI and 16S rRNA, were conducted to evaluate the degree of genetic differentiation among Lineages N, S1, and S2 (Tables S9, S10). According to the results, supposing that there are three genetic lineages (N, S1, and S2) or two genetic lineages (N and S1/S2), almost all variation in both cases is attributed to variation among groups (= among lineages), whereas variations within populations (within lineages) exhibit negative values in common. We confirmed that there was a high degree of genetic differentiation among Lineages N, S1, and S2, which supports the results of the COI barcoding gap analysis shown in Fig. 4a–c, although this was not statistically significant (P  > 0.05; Table S9). When we assumed only two genetic groups, Lineages N and S1/S2, the genetic differentiation between the two groups was statistically significant (P  0.05) (Tables S9 and S10). The discrepancy between the number of barcoding gaps inferred from COI and 16S rRNA may have been affected by different gene evolutionary rates of the molecular markers11; nucleotide substitution rate of 16S rRNA is known to be generally slower than that of COI (which is especially fast in the third codon positions: 105 out of 127 polymorphic sites). When an ML tree was constructed based on 22 polymorphic sites, which are found only in the first and second codon positions of COI that are much more conserved than the third codon position, the three genetic lineages were retained in the resultant tree (Fig. S5), but Lineage S2 was nested within Lineage S1, as in the trees inferred from 16S rRNA (Fig. S2). Reflecting the powerful resolution of the COI barcoding marker well known from animals12 and the high degree of variation among the three genetic lineages (Fig. 4, Figs. S3, S4), we suggested that L. japonica could be categorized into three different species: L. koreana, Yeo and Hwang, sp. nov. for Lineage N, L. japonica for Lineage S1, and L. sinensis Choi, Park, and Hwang, sp. nov. for Lineage S2. To examine whether it is reasonable to give these a species-level taxonomic status, as shown in Fig. 4d, we reconstructed a COI-based NJ tree with one congeneric species L. tenuispinosa13, which was originally described as a subspecies-level taxon of L. japonica14,15 and was then revised as an independent species closely related to L. japonica by Saito & Yoshioka16 in 1993. The resultant tree (Fig. S6 and Data S3) showed that L. tenuispinosa forms a sister group with L. japonica (Lineage S1). This likely indicates that L. koreana and L. sinensis have taxonomic status as independent species.Morphological comparison and geographical distribution of the three Liolophura speciesWe compared morphological characteristics among Liolophura koreana, sp. nov. (Lineage N), L. japonica (Lineage S1), and L. sinensis, sp. nov. (Lineage S2). Their morphological appearances are shown in Fig. 5a–c, which indicated that black spots on the tegmentum (Fig. 5d–e) and shapes of spicules on the perinotum (Figs. 5f–k, 6e–f) represent key morphological characteristics to distinguish them from each other. Although black dots in pleural areas, which are between the middle and lateral areas of the tegmentum on valves II–VII (or VIII), are commonly shared in all three lineages (Fig. 5a–c), other black spots on the valves exhibit a high degree of variation in morphology (Fig. 5a–c, Fig. S7). Herein, we described a new species of genus Liolophura, that is, L. koreana Yeo and Hwang from South Korea and Japan, with detailed descriptions of morphological characteristics observed by light microscopy (M205, Leica Camera AG, Germany) and FE-SEM (SU8220, Hitachi, Japan). In addition, we suggested the divergence of a new species, L. sinensis Choi, Park, and Hwang from southern China, with simple remarks based on distinct genetic difference (mainly COI barcoding gaps), with possible unique morphological characteristics as follows.Figure 5Morphological comparison of Liolophura koreana, sp. nov., L. japonica, and L. sinensis, sp. nov. (a–c) Photos of dorsal views of the individuals belonging to L. koreana (Lineage N), L. japonica (Lineage S1), and L. sinensis (Lineage S2) in order. (d,e) Morphological comparison of pleural and lateral black spots on valves III and IV of the tegmentum of L. koreana (d; holotype) and L. japonica (e). (f,g) Morphological comparison of spicules on the perinotum of L. koreana (f; holotype) and L. japonica (g). (h–k) Morphological comparison of the spicule of L. koreana (h,i; paratype) and L. japonica (j,k) in lateral and dorsal views. The scale bar marks 2.0 mm (d,e), 1.0 mm (f,g), and 0.5 mm (h–k). The photos were edited using Adobe Photoshop v.22.2 (https://www.adobe.com).Full size imageFigure 6Microstructural comparison of Liolophura koreana, sp. nov. and L. japonica using field emission scanning electron microscopy (FE-SEM). (a,b) Middle and lateral areas on the tegmentum of the holotype of L. koreana. (c,d) Middle and lateral areas on the tegmentum of L. japonica. Arrows indicate that morphological difference of the posterior valve margin of the valve II between two species. The scale bar marks 1.0 mm. (e,f) The occurrence frequency, and shape and structure differences of the spicules on the perinotum between the holotype of L. koreana (e) and L. japonica (f). The scale bar marks 1.0 mm and 0.2 mm, respectively. The photos were edited using Adobe Photoshop v.22.2 (https://www.adobe.com).Full size image
    Liolophura koreana Yeo and Hwang, sp. nov. (Figs. 5, 6; Figs. S7, S8)(urn:lsid:zoobank.org:act:4418355E-F55C-44FA-B4CE-585589FDCD23).Type specimens examined[Holotype] SOUTH KOREA: 1 specimen, Jeju-do, Seogwipo-si, Seongsan-eup, Seopjikoji, 3.XI.2020, UW Hwang, B Park & CR Shin (LEGOM040501); [Paratypes] SOUTH KOREA: 1 specimen, Gyeongsangbuk-do, Pohang-si, Guryongpo-eup, Janggil-ri, 27.VII.2008, UW Hwang (LEGOM040502); 3 specimens, Gyeongsangbuk-do, Ulleung-gun, Seo-myeon, Namyang-ri, Ulleungdo Island, Namtong tunnel, 12.VI.2007, UW Hwang (LEGOM040503–0505); 1 specimen, Gyeongsangbuk-do, Ulleung-gun, Namyang-ri, Ulleungdo Island, Namyang tunnel, 5.X.2007, UW Hwang (LEGOM040506); 2 specimens, Gyeongsangnam-do, Geoje-si, Nambu-myeon, Dapo-ri, 28.IV.2009, MY Yeo (LEGOM040507,0508); 4 specimens, same data as the holotype (LEGOM040509–0512); 2 specimens, Jeollanam-do, Yeosu-si, Hwajeong-myeon, Sado-ri, Sado Island, 8.IV.2008, MY Yeo (LEGOM040513,0514); 4 specimens, same data as the holotype (LEGOM040515–0518); JAPAN: 6 specimens, Tottori Prefecture, Hakuto, 24.V.2009, UW Hwang (LEGOM040519–0524); 1 specimen, Tottori Prefecture, Iwato, 25.V.2009, UW Hwang (LEGOM040525).DescriptionBody small-sized and broad oval- to oval-shaped (Fig. 5a; Fig. S7); length 3.9 (1.9–12.3) mm and width 2.4 (1.2–7.1) mm. Tegmentum entirely brown (dark brown or black entirely, or each valve with black line anteriorly or white line laterally), with black dots on the pleural areas of valves II–VII (or VIII) (Fig. 5a,d, Fig. S7); articulamentum entirely black (dark brown); whitish and blackish spicules on the perinotum scattered irregularly, sometimes forming a band besides each valve (Fig. 5a, Fig. S7). Surface of the tegmentum in middle and lateral areas as in Fig. 6a,b and Fig. S8; posterior margin of the head valve nearly straight; dorsal shape of intermediate valves round-backed and side slopes slightly convex; the posterior valve margin with a distinct central apex, its shape subtriangular to triangular (rounded or linear), particularly valve II, mainly with a strong projection (Fig. 6a). Perinotum covered with large, solid, slightly curved, and obtusely pointed spicules (rarely with smooth and radial ribbed spicules apically), its density relatively lower than that of L. japonica (Figs. 5f,h,i, 6e).DistributionSouth Korea, Japan; avobe 33°24′ N (Seogwipo, JJ) in South Korea and TT and MY in Japan (Fig. 7).Figure 7Geographical distribution of Liolophura koreana, sp. nov., L. japonica, and L. sinensis, sp. nov. inhabiting coastal areas of the northwestern Pacific Ocean. A COI-based map showing geographical distribution of L. koreana, L. japonica, and L. sinensis on the northwestern Pacific coast. L. koreana are found in a wide range of South Korea and Japan above ca. 33°24′ N (JJ), L. japonica in mainly southern coastal areas of South Korea and Japan below ca. 35°53′ N (TT), and L. sinensis in ZJ of southern China around ca. 27°02′ N–28°00′ N. The sympatric distribution of L. koreana and L. japonica is shown between 33°24′ and 35°53′ N. Table S1–S3 contain the full names of localities and detailed haplotype information. The question mark indicates that collection of Liolophura samples from such coastal areas in Japan is required to clarify distribution patterns of L. koreana and L. japonica in the East Sea (= Sea of Japan). The basic map was obtained from a free map-providing site (https://d-maps.com), which was modified using Adobe Illustrator v.25.2. (https://www.adobe.com).Full size imageHabitatThis new species appears to be attached to rocks in coastal areas with strong waves, or a calm inner shore in the northwestern Pacific Ocean (Fig. S9).EtymologyThe species is named per the locality of the new species.RemarksWe found that Liolophura koreana, sp. nov. has no black spots on lateral areas of the tegmentum (Fig. 5d), and large, slightly curved, and obtusely pointed spicules on the perinotum compared to those of L. japonica (Figs. 5f,h,i, 6e). On the other hand, L. japonica from southern South Korea and southern Japan has two black spots on the lateral areas of valves II–VII (or VIII) (Fig. 5e), and small, almost straight, and cylindrical spicules compared to those of L. koreana (Figs. 5g,j,k, 6f).As shown in Fig. 7, L. koreana (Lineage N) was observed in all the South Korean and Japanese populations examined here, except for the EH population in Japan (refer to Tables S1, S2), which were found from JJ at 33°24′ N to MY at 38°32′ N. On the other hand, L. japonica (Lineage S1) was found from the southern coastal areas of South Korea and Japan, which were found only between JJ at 33°24′ N and TT at 35°53′ N. Interestingly, we found only L. koreana north from the latitude of 35°10′ N (BS) such as UL/DD (37°24′ N) and PH (36°02′ N) in South Korea. In Japan, there was found only L. koreana at MY (38°32′ N) too, but it remains to be explored to clarify its distribution range in Japan with much more sample collections covering northern Japanese coastal areas through further study. It was also confirmed that L. koreana and L. japonica show a sympatric distribution pattern between JJ at 33°24′ N and TT at 35°53′ N on the southern coastal area of South Korea and Japan.
    Liolophura sinensis Choi, Park, and Hwang, sp. nov. (Fig. 5c).(urn:lsid:zoobank.org:act:72DF7E75-1853-4F23-AC12-3AB8CD054187).Type specimens examined[Holotype] CHINA: 1 specimen, Zhejiang Province, Dongtou Island, 27°49′57.44″ N, 121°10′19.13″ E, 2017; [Paratypes] CHINA: 1 specimen, Beiji Island, 27°37′08.82″ N, 121°11′47.82″ E, 2017; 1 specimen, Beilongshan Island, 27°40′08.56″ N, 121°58′51.56″ E, 2017; 1 specimen, Chaishi Island, 27°25′40.36″ N, 121°04′54.45″ E, 2017; 1 specimen, Daleishan Island, 27°29′39.48″ N, 121°05′24.50″ E, 2017; 1 specimen, Daqu Island, 27°47′29.92″ N, 121°05′23.97″ E, 2017; 1 specimen, Dazhushi Island, 27°49′12.87″ N, 121°12′48.74″ E, 2017; 1 specimen, Dongce Island, 27°45′32.04″ N, 121°09′01.38″ E, 2017; 1 specimen, Dongxingzai Island, 27°02′40.36″ N, 121°02′47.98″ E, 2017; 1 specimen, Houjishan Island, 27°28′26.96″ N, 121°07′40.71″ E, 2017; 1 specimen, Luxi Island, 27°59′33.43″ N, 121°12′50.70″ E, 2017; 1 specimen, Nanji Island, 27°27′30.57″ N, 121°03′06.28″ E, 2017; 1 specimen, Nanpanshan Island, 28°00′15.29″ N, 121°15′33.62″ E, 2017.DistributionSouthern China; Zhejiang Province around ca. 27°02′–28°00′ N (Fig. 7).HabitatThis new species appears to be attached to rocks in coastal areas of the northwestern Pacific Ocean.EtymologyThe species is named after its locality.RemarksL. sinensis, sp. nov. was examined and established mainly by molecular data such as the COI barcoding gap (Fig. 4a–c) presented in this study and photos provided from Prof. Yong-Pu Zhang (Wenzhou University, Zhejiang Province, China) without any direct real sample observation. The morphology of this new species is very similar to that of a previously known species, L. japonica. L. sinensis from southern China has black spots on the lateral areas of valves II–VII similar to L. japonica, but black bowtie-shaped spots anterior to valves II–VII (Fig. 5c) are a unique characteristic for L. sinensis. L. sinensis (Lineage S2) was found in Zhejiang Province (ZJ) in southern China, around ca. 27°02′–28°00′ N (Fig. 7; Tables S1, S5).Demographic history and divergence time estimation analysesMismatch distribution analyses (MDA) based on COI were performed for L. koreana, L. japonica, and L. sinensis, respectively. The MDA results (Fig. 8a) showed a unimodal curve for each of the three lineages. In addition, when neutrality tests were performed with COI and 16S rRNA (Table S11), all three showed statistically significant negative values in both Tajima’s D and Fu’s Fs, except for the Tajima’s D values in COI (L. sinensis) and 16S rRNA (L. japonensis and L. sinensis), implying that these had experienced population expansions. Bayesian skyline plot (BSP) analyses with COI (Fig. 8b) were performed to examine the fluctuation patterns in effective population sizes for L. koreana, L. japonica, and L. sinensis, respectively. The effective population sizes of L. japonica and L. sinensis had gradually grown between ca. 100 Ka and ca. 50 Ka, while those of L. japonica had grown between ca. 80–50 Ka, L. sinensis had grown between 100–60 Ka, and that of L. koreana had begun to rapidly expand ca. 85 Ka and ceased ca. 75 Ka. This indicated that population expansion had occurred more dramatic in L. koreana than in L. japonica and L. sinensis, following the last interglacial age, called the Eemian (129–116 Ka). As shown in Fig. 8c and Fig. S10, according to the molecular clock analysis by the BEAST program, it was estimated that L. japonica and L. koreana shared their most recent common ancestor about 3.37 Ma, around the mid-Pliocene warm period (3.30–3.00 Ma), before the extensive glaciation in the late Pliocene (ca. 3.00 mya). L. japonica and L. sinensis likely diverged around 1.84 Ma, around the beginning stage of the Early Pleistocene Transition (EPT; 1.85–1.66 mya). The augmentation of haplotype diversity in L. japonica, L. sinensis, and L. koreana might have intensified in the interglacial stages during the late-middle (0.35–0.126 Ma) and late Pleistocene (0.126–0.012 Ma), before the last glacial maximum (LGM: 0.026–0.019 Ma).Figure 8The results of mismatch distribution analyses (MDA), Bayesian skyline plots (BSPs), and molecular clock analysis performed with COI haplotypes for Liolophura koreana, sp. nov., L. japonica, and L. sinensis, sp. nov. (a) MDA plots resulting in a unimodal curve for L. koreana, L. japonica, and L. sinensis. Dotted lines indicate the observed distribution of mismatches, and solid lines represent the expected distribution under a demographic expansion model. (b) BSP results showing the demographic history of population expansions of L. koreana, L. japonica, and L. sinensis. The graph in gray depicts sea level changes during the last 330 Ka. (c) Time-calibrated Bayesian tree reconstructed using BEAST with the inference of ancestral areas under the Bayesian binary MCMC (BBM) model implemented in RASP ver 3.2. Ancestral areas were hypothesized based on the distribution range of the fossil records of Mopalia and the contemporary distribution of L. koreana, L. japonica, and L. sinensis. LGM indicates the last glacial maximum (0.026–0.019 Ma; blue vertical bar) and three interglacial periods are indicated by light green boxes during the late-middle and late Pleistocene. The pictures were edited using Adobe Illustrator v.25.2. (https://www.adobe.com).Full size image More

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    Eco-environmental assessment model of the mining area in Gongyi, China

    Technical criterion for ecosystem status evaluationOn March 3, 2015, the Ministry of Ecology and Environment of the People’s Republic of China approved the “Technical Criterion for Ecosystem Status Evaluation” as the national environmental protection standard. This standard is based on the former standards released in 2006, and 48 relevant documents from 2006 to 2012 were searched to propose new standards and factor weights based on actual utilization effects and expert guidance. The eco-environment assessment uses a comprehensive index (eco-environmental status index, EI) to reflect the overall state of the regional eco-environment. The indicator system includes the biological abundance index, vegetation coverage index, river density index, land stress index, and pollution loading index. These indexes reflect the abundance of organisms in the evaluated area, the level of vegetation coverage, the abundance of water, the intensity of land stress, and the extent of the pollution load. Each indicator was calculated according to its weight to obtain an eco-environment assessment map (Table 1). All parameters involved in the calculation are derived from this standard.Table 1 Weights of the evaluation indicators.Full size tableThe calculation of the eco-environment status is as follows:$$begin{aligned} EI & = 0.35*biological ; abundance ; index + 0.25*vegetation ; coverage ; index hfill \ & quad + 0.15*river ; density ; index + 0.15*(1 – land ; stress ; index) hfill \ & quad + 0.1*(1 – pollution ; loading ; index) hfill \ end{aligned}$$
    (9)
    Biological abundance indexThe biological abundance index refers to the number of certain organisms in this area. The calculation method is as follows:$$Biological , abundance , index , = , left( {BI , + , HQ} right)/2$$
    (10)
    In this formula, BI is the biodiversity index and HQ is the habitat quality index. When the biodiversity index does not have dynamic data updates, the change in the biological abundance index is equal to the change in the HQ.Biodiversity is a general term for the complexity of species and their genetic variation and ecosystems in space over time. Biodiversity plays an important role in maintaining soil fertility, ensuring water quality, regulating the climate, stabilizing the environment, and maintaining ecological balance.The BI method is as follows:$$BI = NPP_{mean} *F_{pre} *F_{tem} *(1 – F_{alt} )$$
    (11)
    NPPmean is the net primary productivity. Fpre is the annual average precipitation. Ftem is the temperature parameter. Falt is the altitude parameter.NPP refers to the amount of organic matter accumulated per unit area and unit time of green plants. NPP is the remainder of the total amount of organic matter produced by photosynthesis after deducting autotrophic respiration and is usually expressed as dry weight. In this study, the estimation of NPP was based on the absorbed photosynthetically active radiation (APAR) and actual light-use efficiency (LUE) (ε) of the CASA ecosystem model40. The CASA model is a process-based remote sensing model that couples ecosystem productivity and soil carbon and nitrogen fluxes, driven by gridded global climate, radiation, soil, and remote sensing vegetation index datasets41. The model can be expressed generally as follows:$$NPP(x,t) = APAR(x,t)*varepsilon (x,t)$$
    (12)
    The entire study area is divided into 11,303 pixels on a 30 * 30 m grid. x indicates the location of each pixel, and t indicates time; the data were collected once a month. APAR(x,t) represents the photosynthetically active radiation absorbed by pixel x in that month (gC * m−2* month−1). Ɛ(x, t) is LUE (gC * MJ−1) of the vegetation42.Estimation of the fraction of APAR using RS data is based on the reflection characteristics of the vegetation on the infrared and near-infrared bands. The value of APAR is determined by the effective radiation of the sun and the absorption ratio of the vegetation to the effective photosynthetic radiation. The formula is as follows:$$APAR(x,t) = SOL(x,t)*FPAR(x,t)*0.5$$
    (13)

    where SOL(x,t) represents the total amount of solar radiation at pixel x in month t, FPAR(x,t) represents the absorption ratio of the vegetation layer to the incident photosynthetically active radiation, and a constant of 0.5 indicates the ratio of the effective solar radiation that can be utilized by the vegetation to the total solar radiation.Since there is a linear relationship between FPAR and NDVI within a certain range, this relationship can be determined according to the maximum and minimum values of a certain vegetation type NDVI and the corresponding FPAR maximum and minimum values.$$FPAR(x,t) = frac{{(NDVI(x,t) – NDVI_{i,min } )}}{{(NDVI_{i,max } – NDVI_{i,min } )}}*(FPAR_{max } – FPAR_{min } ) + FPAR_{min }$$
    (14)

    where NDVImax and NDVImin correspond to the NDVI maximum and minimum values of the ith planting type, respectively. There is also a good linear relationship between FPAR and the simple ratio index (SR) of vegetation, which is represented by the following formula:$$FPAR(x,t) = frac{{(SR(x,t) – SR_{i,min } )}}{{(SR_{i,max } – SR_{i,min } )}}*(FPAR_{max } – FPAR_{min } ) + FPAR_{min }$$
    (15)

    where the values of FPARmin and FPARmax are independent of vegetation type and are 0.001 and 0.95, respectively; SRi,max and SRi,min correspond to the 95% and 5% percentiles, respectively, of the ith NDVI. SR(x,t) is represented by the following formula:$$SR(x,t) = frac{1 + NDVI(x,t)}{{1 – NDVI(x,t)}}$$
    (16)
    A comparison of the estimated results of FPAR-NDVI and FPAR-SR shows that the FPAR estimated by NDVI is higher than the measured value, while the FPAR estimated by SR is lower than the measured value, but the error is less than that estimated directly by NDVI. As a result, these two values can be combined, and their weighted average value is taken as an estimate of the estimated FPAR, while ɑ means weight:$$FPAR(x,t) = alpha FPAR_{NDVI} + (1 – alpha )FPAR_{SR}$$
    (17)
    Light use efficiency (LUE) refers to the ratio of chemical energy contained in organic dry matter produced per unit area over a certain period of time to the photosynthetically active radiation absorbed by plants projected onto the same area at the same time. Different vegetation types and the same types of vegetation have different light energy utilization rates in different living environments43. The differences are mainly due to the characteristics of the vegetation itself, temperature, moisture, and soil44. Vegetation has the highest utilization rate of light energy under ideal conditions, but the maximum light energy utilization rate in the real environment is mainly affected by temperature and moisture, which can be expressed as follows:$$varepsilon left( {x,t} right) = T_{varepsilon 1} left( {x,t} right) cdot T_{varepsilon 2} left( {x,t} right) cdot W_{varepsilon } left( {x,t} right) cdot varepsilon_{max }$$
    (18)
    where Tε1(x,t) and Tε2(x,t) represent the stress effects of low temperature and high temperature on light energy utilization, respectively, Wε(x,t) is the effect of water stress on the maximum light energy utilization under ideal conditions, and εmax is the maximum light energy utilization under ideal conditions (gC * MJ−1). The maximum solar energy utilization rate εmax varies depending on the vegetation type. In this study, the maximum light energy utilization rate of different land use types simulated by an improved Carnegie-Ames-Stanford Approach (CASA) model is used as the input parameter of light energy utilization in the CASA model (Table 2). The monthly maximum light energy utilization rate is determined in three steps: first, calculate the APAR, temperature, and water stress factors of all pixels; then, select the NPP measured data of the same time period in the study area; finally, simulate the εmax of vegetation according to the principle of minimum error45. Figure 5 shows the calculation process of NPP. The weight of each habitat type in the HQ is shown in Table 3. The weight value is derived from the official document30. To facilitate the calculation, this paper normalizes the calculation results from 0 to 1 (Fig. 6a).Table 2 Maximum LUE rates of different land use types.Full size tableFigure 5NPP calculation process.Full size imageTable 3 Weight of each habitat type in the HQ.Full size tableVegetation coverage indexThe vegetation coverage index was obtained from the NDVI, which is a simple, effective, and empirical measure of surface vegetation status. The vegetation index mainly describes the difference between the reflection of vegetation in the visible and near-infrared bands and the soil background. This index also reduces the solar elevation angle and noise caused by the atmosphere and is thus the most widely used and effective calculation method. Each vegetation index can be used to quantitatively describe the growth of vegetation under certain conditions. The expression is as follows:$$NDVI = frac{NIR – R}{{NIR + R}}$$
    (19)

    where NIR and R are reflectance values in the near-infrared and red bands, respectively.NDVI values are obtained by processing the RS images of the Landsat 8 satellite. This satellite is equipped with an operational land imager (OLI) that includes nine bands with a spatial resolution of 30 m, including a 15-m panchromatic band. To facilitate the calculation, this paper normalizes the calculation results from 0 to 1 (Fig. 6b).Figure 6Eco-environment assessment indexes and evaluation rating map (the first quarter was used as an example). (a) Biological abundance index; (b) vegetation coverage index; (c) river density index; (d) land stress index; (e) pollution loading index; and (f) environmental status classification. The Figure is created using ArcGIS ver.10.3 (https://www.esri.com/).Full size imageRiver density indexThe river density index refers to the total length of rivers, lakes, and water resources in the assessed area as a percentage of the assessed area, which is used to reflect the abundance of water in the assessed area and is calculated as follows:$$begin{gathered} River ; density ; index = (A_{riv} *river ; length / area + A_{lak} *water ; area/area hfill \ + A_{res} {*}amount ; of ; resources/area , )/3 hfill \ end{gathered}$$
    (20)

    where Ariv is the normalization coefficient of river length, with a reference value of 84.3704, Alak is the normalization coefficient of the lake area, with a reference value of 591.7909, and Ares is the normalization coefficient of water resources, with a reference value of 86.387. Finally, the calculation results were normalized from 0 to 1 (Fig. 6c).Land stress indexThe land stress index is the degree to which the land quality in the assessment area is under stress. The weight of the land stress index evaluation is shown in Table 4.Table 4 Weight of the land stress index evaluation.Full size tableThe calculation method is as follows:$$begin{gathered} Land ; stress ; index = A_{ero} *(0.4*severe ; erosion ; area + 0.2*{text{mod}}erate ; erosion ; area hfill \ + 0.2*construction ; land ; area + 0.2*other ; land ; stress ; area)/area hfill \ end{gathered}$$
    (21)

    where Aero is the normalization coefficient of the land stress index, with a reference value of 236.0436. According to the “Classification criteria for soil erosion”46, the influencing factors of soil erosion, vegetation, soil texture, landform, and precipitation are ranked according to importance. In the calculation of the land stress index, all the land is divided into three categories, in which the weight of severe erosion is 0.4, the weight of non-erosion is 0, and the other erosion types such as moderate erosion and construction land are 0.2. The areas with severe erosion include vegetation coverage less than 30% and areas of soil erosion greater than 3.7 mm/a due to human activities. These areas are generally developed on highly erosive-sensitive soils. Cinnamon soil and loess soil in the study area are highly erosive-sensitive soils. Therefore, the industrial and mining areas of cinnamon and loess soil types are regarded as severe erosion areas. Areas with vegetation coverage greater than 50% are non-eroded areas, so water bodies and woodlands are divided into non-erodible areas. All areas except these two types have a weight of 0.2. Finally, the calculation results were normalized from 0 to 1 (Fig. 6d).Pollution loading indexThe pollution loading index refers to the load of pollutants in a certain area or an environmental element. In this study, the AQI was used to calculate the pollution loading index, and the results were normalized from 0 to 1 (Fig. 6e).The eco-environmental evaluation score was calculated based on the national environmental protection standard according to the weight of each indicator (Fig. 6f).Improved evaluation system and intelligent evaluation modelImproved evaluation systemConsidering that the evaluation factors in the national environmental protection standards are applicable to ordinary areas, areas affected by mines should have more evaluation factors than those in the standards. Thus, an improved evaluation system was proposed. The improved evaluation system has added factors that affect the environment of the mine based on the factors of the original system. The impact of the mine on the environment is reflected in the pollution of the atmosphere, such as dust from open pits and industrial waste from concentrators; the occupation of land by solid waste, such as ore piles and coal piles; soil pollution, such as the diffusion of heavy metals from coal piles, coal mine concentrator plants, and mines; and the increased likelihood of geological disasters, such as collapse caused by underground mining, spontaneous coal combustion and landslides caused by open-pit mining surfaces. Therefore, the improved evaluation system adds an air pollution range, a solid waste area, a geologic hazard range, and a metallic and non-metallic mine soil pollution buffer to the national environmental protection standards.The area of air pollution in mining areas is generally near open pits and concentrator plants. Therefore, the air pollution range was selected within 50 m around the open pits and the concentrator plants. Due to the dilution and dispersion of the air itself, an estimate of the pollution is the reciprocal of the wind speed (Fig. 7a).Figure 7New factors in the improved evaluation system. (a) Air pollution range; (b) solid waste area; (c) geological hazard range; (d1) non-metallic mine soil pollution buffer; and (d2) metal mine soil pollution buffer. The Figure is created using ArcGIS ver.10.3 (https://www.esri.com/).Full size imageMine solid waste pollution includes a large amount of waste rock from open-pit mining and pit mining, coal gangue produced by coal mining, tailings from beneficiation and slag from smelting. These solid wastes are generally piled up near the mining area. They not only occupy large areas of land and induce geological disasters such as landslides and mudslides but also cause chemical pollution, spontaneous combustion, and radiation from radioactive materials due to long-term stacking. This may affect the health and safety of humans and other biological organisms. The scope of the solid waste area is determined by the ore piles, coal piles, and dumping sites (Fig. 7b).Mine geological disasters are caused by a large number of mining wells and rock and soil deformation, as well as serious changes in the geological, hydrogeological, and natural environments of the mining area, endangering human life and property and destroying mining engineering equipment and mining resources. In this study, the geologic hazard range consists of areas with underground mining stopes and coal piles (Fig. 7c).After the pollutants generated by the mining operation enter the soil, physical and mechanical absorption, retention, colloidal physicochemical adsorption, chemical precipitation, bioabsorption, etc. of the suspended pollutants through the soil continue to accumulate in the upper soil. When pollutants reach a certain maximum, they cause deterioration of the soil composition, cycle, properties, and functions and begin to accumulate in plants, which affects the normal growth and development of plants, decreases crop yield and quality, and ultimately affects human health. Metallic and non-metallic minerals have different effects on soil pollution. The pollution of soil by non-metallic minerals mainly occurs in coal mines and coal piles, and the buffer zone is centred on the coal mines and coal piles. Coal production activities can cause heavy metals in coal piles to enter the soil and cause pollution. Due to different types of heavy metals, the range of soil contamination is different47. Combining the non-metallic mineral industrial squares and coal mine-based non-metallic minerals around the heavy metal soil pollution range, the buffers are graded at 30, 200, and 1000 m (Fig. 7d1)43. The metal mines in Gongyi are mainly aluminium ore and iron ore. Referencing the spread range of heavy metal pollution in the soil of aluminium ore and iron ore mines48,49,50,51, the buffers are graded at 50, 100, 300, and 500 m (Fig. 7d2).The four new elements in the improved evaluation system are normalized from 0 to 1 during the calculation.Intelligent evaluation modelArtificial neural networks, decision trees, and SVMs were calculated using IBM SPSS Modeler software to find an intelligent model suitable for environmental assessment of the mine in the study area. Then, several models with high evaluation accuracy were selected. The SVM, CART, and C5.0 models were chosen for further comparison. The sampling points were selected randomly; 700 sampling points were selected from the area away from the mining area; 100 sampling points were selected from the mining area after random sampling by mine type, and these points were used as training samples. Non-mining evaluation scores were based on the national environmental protection standard, while the mining area scores were based on field investigation. In the field investigation, preliminary scoring of the sampling points was conducted according to mining type, mining intensity, air quality, and surrounding environment. Then, a photo of the field was taken at every sampling point in the mine, and experts were invited to further score the area according to the photo. This score is the relative score obtained by referencing the national environmental protection standard.The index layers of the training samples were used as input, and the scores were used as the output to train the machine learning models. The trained models were applied to the entire study area, and all points except the training sample points were used for verification. After further comparison with SVM, CART, and C5.0, the evaluation accuracy rates of the three methods in the mining area and non-mining area were obtained. In the non-mining area, the model evaluation results of various land use types were compared with the national environmental protection standards. The accuracy in various land use types is shown in Table 5. In the mining area, the model evaluation score is compared with the score from the experts, and the obtained accuracy table is shown in Table 6.Table 5 Accuracy of each algorithm in various land use types in non-mining areas.Full size tableTable 6 Accuracy of each algorithm in the mining area.Full size tableIn non-mining areas, the accuracy of the SVM model is significantly better than that of the other two methods. However, in the mining area, the accuracy of the CART model is higher. Therefore, the SVM model was used to evaluate the area away from the mine, and the CART model was used to evaluate the mining area. The evaluation results of these two models were combined to obtain the evaluation map of the entire study area. More

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    These authors contributed equally: Celso H. L. Silva Junior, Nathália S. Carvalho, Ana C. M. Pessôa.Tropical Ecosystems and Environmental Sciences Laboratory (TREES), São José dos Campos, São Paulo, BrazilCelso H. L. Silva Junior, Nathália S. Carvalho, Ana C. M. Pessôa, João B. C. Reis, Aline Pontes-Lopes, Juan Doblas, Wesley Campanharo, Henrique Cassol, Yosio E. Shimabukuro, Liana O. Anderson & Luiz E. O. C. AragãoInstituto Nacional de Pesquisas Espaciais (INPE), São José dos Campos, São Paulo, BrazilCelso H. L. Silva Junior, Nathália S. Carvalho, Ana C. M. Pessôa, Aline Pontes-Lopes, Juan Doblas, Wesley Campanharo, Henrique Cassol, Luciana Gatti, Ana P. Aguiar, Yosio E. Shimabukuro & Luiz E. O. C. AragãoUniversidade Estadual do Maranhão (UEMA), São Luís, Maranhão, BrazilCelso H. L. Silva JuniorCentro Nacional de Monitoramento e Alertas de Desastres Naturais (CEMADEN), São José dos Campos, São Paulo, BrazilJoão B. C. Reis & Liana O. AndersonUniversity of Bristol, Bristol, UKViola Heinrich & Joanna HouseInstituto de Pesquisa Ambiental da Amazônia (IPAM), Brasília, Distrito Federal, BrazilAne Alencar, Camila Silva & Paulo BrandoUniversidade Estadual de Campinas (UNICAMP), Campinas, São Paulo, BrazilDavid M. LapolaEcología del Paisaje y Modelación de Ecosistemas (ECOLMOD), Universidad Nacional de Colombia (UNAL), Bogota, ColombiaDolors ArmenterasUniversidade de Brasília, Brasília, Distrito Federal, BrazilEraldo A. T. MatricardiUniversity of Oxford, Oxford, UKErika BerenguerLancaster University, Lancaster, UKCamila Silva, Erika Berenguer & Jos BarlowSouth Dakota State University, Brookings, SD, USAIzaya NumataEmpresa Brasileira de Pesquisa Agropecuária (EMBRAPA) Amazônia Oriental, Belém, Pará, BrazilJoice FerreiraUniversity of California, Irvine, CA, USAPaulo BrandoWoodwell Climate Research Center, Falmouth, MA, USAPaulo BrandoInstituto Nacional de Pesquisas da Amazônia (INPA), Manaus, Amazonas, BrazilPhilip M. FearnsideJet Propulsion Laboratory (JPL), Pasadena, CA, USASassan SaatchiUniversity of California, Los Angeles, CA, USASassan SaatchiUniversidade Federal do Acre (UFAC), Cruzeiro do Sul, Acre, BrazilSonaira SilvaUniversity of Exeter, Exeter, UKStephen Sitch & Luiz E. O. C. AragãoStockholm Resilience Centre, Stockholm University, Stockholm, SwedenAna P. AguiarSchool of Forest, Fisheries, and Geomatics Sciences, University of Florida, Gainesville, FL, USACarlos A. SilvaEuropean Commission, Joint Research Centre (JRC), Ispra, VA, ItalyChristelle Vancutsem, Frédéric Achard & René BeuchleCenter for International Forestry Research (CIFOR), Bogor, IndonesiaChristelle Vancutsem More

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    Selecting when to bloom

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