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Altogether nine polished pieces of the lower Cenomanian Kachin amber from northern Myanmar (Figs. 1A–D, 2A–E) were examined in this study (depository: Russian Museum of Biodiversity Hotspots, N. Laverov Federal Center for Integrated Arctic Research of the Ural Branch of the Russian Academy of Sciences, Arkhangelsk, Russia). A brief description of each amber piece is given below.Figure 1Lower Cenomanian Kachin amber samples with specimens and borings of †Palaeolignopholas kachinensis gen. & sp. nov. from northern Myanmar used in this study. (A) RMBH biv1115 (frontal view with the holotype). (B) RMBH biv1101 (lateral view with two paratypes and a shell fragment). (C) RMBH biv1116 (frontal view with the fossilized paratype). (D) RMBH biv1100 (frontal view with borings). The red frames indicate position of the type specimens (holotype and some paratypes). The red arrows indicate bivalve borings. Scale bars = 5 mm. (Photos: Ilya V. Vikhrev).Full size imageFigure 2Lower Cenomanian Kachin amber samples with borings of †Palaeolignopholas kachinensis gen. & sp. nov. from northern Myanmar used in this study. (A) RMBH biv1102 (frontal view). (B) RMBH biv1103 (frontal view). (C) RMBH biv1114 (frontal view). (D) RMBH biv1118 (frontal view). (E) RMBH biv1117 (frontal view). The red arrows indicate bivalve borings. Scale bars = 5 mm. (Photos: Ilya V. Vikhrev).Full size imageRMBH biv1115: Size 8.5 × 5.8 × 8.1 mm (Fig. 1A). Inclusions: articulated shell of †Palaeolignopholas kachinensis gen. & sp. nov., “floating” in the resin (the holotype).RMBH biv1101: Size 15.6 × 6.4 × 11.5 mm (Fig. 1B). Inclusions: two complete articulated shells (paratypes) and a shell fragment of †Palaeolignopholas kachinensis gen. & sp. nov., “floating” in the resin.RMBH biv1116: Size 22.5 × 8.3 × 16.5 mm (Fig. 1C). Inclusions: fossilized shell of †Palaeolignopholas kachinensis gen. & sp. nov. (paratype), borings of this species (filled with fine gray sand), unidentified fly specimens (Insecta: Diptera), and unidentified organic fragments (probably, plant debris).RMBH biv1100: Size 17.5 × 4.9 × 12.0 mm (Fig. 1D). Inclusions: borings of †Palaeolignopholas kachinensis gen. & sp. nov. (filled with fine gray sand), and an unidentified caddisfly specimen (Insecta: Trichoptera).RMBH biv1102: Size 15.6 × 5.1 × 12.7 mm (Fig. 2A). Inclusions: borings of †Palaeolignopholas kachinensis gen. & sp. nov. (filled with fine gray sand), and unidentified organic fragments (probably, plant debris).RMBH biv1103: Size 19.6 × 4.7 × 14.3 mm (Fig. 2B). Inclusions: borings of †Palaeolignopholas kachinensis gen. & sp. nov. (filled with fine gray sand), an unidentified beetle specimen (Insecta: Coleoptera), and unidentified organic fragments (probably, plant debris).RMBH biv1114: Size 33.1 × 7.8 × 21.7 mm (Fig. 2C). Inclusions: multiple borings of †Palaeolignopholas kachinensis gen. & sp. nov. (filled with fine gray sand), and unidentified plant remains.RMBH biv1118: Size 25.1 × 8.4 × 14.3 mm (Fig. 2D). Inclusions: separate borings of †Palaeolignopholas kachinensis gen. & sp. nov. (filled with fine gray sand), a plant fragment with a cluster of borings around, and an unidentified insect specimen.RMBH biv1117: Size 15.5 × 3.9 × 10.7 mm (Fig. 2E). Inclusions: borings of †Palaeolignopholas kachinensis gen. & sp. nov. (filled with fine gray sand), and an unidentified insect specimen.Additionally, six amber samples containing adult and sub-adult specimens of †Palaeolignopholas kachinensis gen. & sp. nov. were examined using photographs in published works as follows: BMNH 20205 (Department of Palaeontology, Natural History Museum, London, UK)15, NIGP 169623 and NIGP 169624 (Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, China)20, RS.P1450 (Ru D. A. Smith collection, Kuala Lumpur, Malaysia)19, and AMNH (Division of Invertebrates, American Museum of Natural History, New York, NY, United States of America)16.Based on morphological analyses of the fossil piddock shells, it was found to be a genus and species new to science, which is described here.Systematic paleontologyPhylum Mollusca Linnaeus, 1758Class Bivalvia Linnaeus, 1758Family Pholadidae Lamarck, 1809Subfamily Martesiinae Grant & Gale, 1931†Palaeolignopholas gen. novLSID: http://zoobank.org/urn:lsid:zoobank.org:act:1D686DCE-A5E9-41DA-9504-2EC58C93D988Type species: †Palaeolignopholas kachinensis gen. & sp. nov.Etymology. This name is derived from the prefix ‘Palaeo-’ (ancient), and ‘-lignopholas’, the name of a recent genus of estuarine and freshwater piddocks boring into wood, mudstone rocks, brickwork, laterites, etc.11,13. Masculine in gender.Diagnosis. The new monotypic genus is conchologically similar to several other piddock genera such as Lignopholas, Martesia, and Diplothyra Tryon 1862 but can be distinguished from these taxa by the following combination of characters: mesoplax relatively small, triangular, divided longitudinally, posterior slope without concentric sculpture, sculptured valve with concave parallel ridges (Martesia-like “rasping teeth”) curved anteriorly, periostracal lamellae dense, fine, hair-like. The fossil genus †Opertochasma Stephenson, 1952 shares a divided mesoplax but it clearly differs from both †Palaeolignopholas gen. nov. and Lignopholas by having two radial grooves on the shell surface21.Distribution. Kachin State, northern Myanmar; Upper Cretaceous (lower Cenomanian)15,19,22.Comments. Both †Palaeolignopholas gen. nov. and Lignopholas appear to be closely related to each other because they share a longitudinally divided mesoplax and periostracal lamellae, which are considered diagnostic features distinguishing this clade from Martesia + Diplothyra. Based on available conchological characters, we assume that †Palaeolignopholas gen. nov. might be placed on the ancestral stem lineage of the Lignopholas clade, although a possibility of homeomorphy could not entirely be excluded.†Palaeolignopholas kachinensis gen. & sp. nov = Plant Antheridia or Fungal Sporangia indet. sensu Grimaldi et al. (2002): 9, fig. 2a,b (bivalve specimens), fig. 3 (borings), fig. 5 (shell reconstruction of an immature specimen), figs. 6 and 7 (SEMs of borings surface showing rasped ornament at different magnifications)16. = Palaeoclavaria burmitis Poinar & Brown (2003): 765, figs. 1–4 (borings) [this fungal taxon was introduced using a trace fossil (boring) as the holotype]17; Poinar (2016): 2, figs. 10, 15, 16 (borings)18. = Martesiinae indet. sensu Smith & Ross (2018): 4, figs. 1a–c, 2a,b, 3a–d (borings), 4a,b, 5a–e (bivalve specimens)19. = Pholadidae indet. sensu Mao et al. (2018): 99, figs. 8a–f (borings), 8g,h (bivalve specimens)20. = Martesia sp. 2 sensu Mayoral et al. (2020): 10, figs. 4a (borings), 7b, 8a–l (bivalve specimens)15. = Pholadidae indet. sensu Balashov (2020): 623.Figures 1, 2, 3, 4, 5, 6 and 7.Figure 3Holotype and a paratype of †Palaeolignopholas kachinensis gen. & sp. nov. from lower Cenomanian Kachin amber, northern Myanmar. (A) Holotype: ventro-lateral view of articulated shell. (B) Paratype: anterio-lateral view of fossilized shell. VN ventral margin; DR dorsal margin; AN anterior margin; PS posterior margin; d disc; rs rasping surface of the valve; uvs umbonal ventral sulcus; pg pedal gape; pl periostracal lamellae. Scale bars = 500 µm. (Photos: Ilya V. Vikhrev).Full size imageFigure 4Paratypes of †Palaeolignopholas kachinensis gen. & sp. nov. from lower Cenomanian Kachin amber, northern Myanmar. (A) Paratype: dorsal view of articulated shell. Scale bar = 500 µm. (B) Paratype: dorsal view of articulated shell. The detached and deflected umbonal paired fragment of the valves is framed by red square. The blue contour indicates the lifetime position of this fragment. The blue arrows show the shell breakages. Scale bar = 200 µm. (C) Umbonal paired fragment of the holotype valves (inner view). The blue arrows show the shell breakage. Scale bar = 200 µm.  VN ventral margin; DR dorsal margin; AN anterior margin; PS posterior margin; ms longitudinally divided mesoplax (inner view); pr prora; d disc; rs rasping surface of the valve; uvs umbonal ventral sulcus; pg pedal gape; pl periostracal lamellae; sb shell breakage. (Photos: Ilya V. Vikhrev).Full size imageFigure 5Rasping surface of †Palaeolignopholas kachinensis gen. & sp. nov. shell. (A) Holotype shell. The red frame marks position of the enlarged area. (B) Undulated micro-sculpture of the rasping surface. Scale bar = 100 µm. (Photos: Ilya V. Vikhrev).Full size imageFigure 6Schematic reconstruction of †Palaeolignopholas kachinensis gen. & sp. nov. from lower Cenomanian Kachin amber, northern Myanmar based on the type series and other fossil material15,16,19,20. (A) Lateral view of adult specimen. (B) Dorsal view of adult specimen. (C) Ventral view of adult specimen (based on a paratype BMNH 2020515). (D) Anterio-ventral view of immature specimen. (E) Dorsal view of immature specimen. (F) Mesoplax of adult specimen. (G) Mesoplax of immature specimen. d disc; mt metaplax; ms mesoplax; hp hypoplax; ca callum; uvs umbonal ventral sulcus; pg pedal gape; pl periostracal lamellae. Scale bars = 1 mm (A–C). (Line graphics: Yulia E. Chapurina).Full size imageFigure 7Clavate borings of †Palaeolignopholas kachinensis gen. & sp. nov. from lower Cenomanian Kachin amber, northern Myanmar. (A) Cluster of borings. It marks drilling of immature piddocks into soft resin from the unidentified plant (wood?) fragment. (B–D) Clavate borings of adult piddocks. Scale bars = 1 mm. Abbreviation: bg a characteristic bioglyph indicating the shell rotation inside hardening resin. (Photos: Ilya V. Vikhrev).Full size imageLSID: http://zoobank.org/urn:lsid:zoobank.org:act:F6659EBF-B0A4-4B21-A99B-2C56BDB7EC9B.Common name. Kachin Amber Piddock.Holotype. RMBH biv1115, the adult shell with length 3.07 mm and width 1.13 mm “floating” in the resin (Figs. 1A, 3A, 5A,B), local collector leg., Russian Museum of Biodiversity Hotspots, N. Laverov Federal Center for Integrated Arctic Research of the Ural Branch of the Russian Academy of Sciences, Arkhangelsk, Russia.Paratypes. RMBH biv1116, the fossilized adult shell with length 4.05 mm and width 1.83 mm (Figs. 1C, 3B); RMBH biv1101, the immature specimen with articulated shell (width 1.86 mm) sharing a detached and deflected umbonal paired fragment of the valves due to the shell breakage (Figs. 1B, 4B,C); RMBH biv1101, the other immature specimen with shell length 2.68 mm and shell width 2.52 mm in this amber piece (Figs. 1B, 4A); BMNH 20205, adult specimen [illustrated in Mayoral et al. (2020): fig. 7B15], Department of Palaeontology, Natural History Museum, London, UK; NIGP 169623, adult specimen [illustrated in Mao et al. (2018): 100, fig. 8G20], and NIGP 169624, two adult specimens [illustrated in Mao et al. (2018): 100, fig. 8H20], Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, China; RS.P1450, two sub-adult specimens [illustrated in Smith & Ross (2018): 5, fig. 4A,B19], Ru D. A. Smith collection, Kuala Lumpur, Malaysia.Type locality and strata. The Noije Bum Hill mines, Hukawng Valley, near Tanai (26.3593°N, 96.7200°E), Kachin State, northern Myanmar; Upper Cretaceous (lower Cenomanian; absolute age of youngest zircons in enclosing marine sediment: 98.79 ± 0.62 Ma)19,22.Etymology. The name of this species reflects its type locality, which is situated in the Kachin State of Myanmar.Diagnosis. As for the genus.Description. Shell small (up to 9.3 mm in length15,19,20), conical, with a rounded anterior margin, tapering posteriorly (Figs. 3A,B, 4A–C, 6A–E); its shape is similar to those in the recent Lignopholas, Martesia, and Diplothyra. Valve sculptured, with concave parallel ridges (Martesia-like “rasping teeth”) curved anteriorly (Fig. 5A,B). The ridges share a characteristic wave-like micro-sculpture (Fig. 5B). Sulcus deep (Figs. 3A, 4C, 6A–C). Mesoplax longitudinally divided, relatively small, triangular, tapering or lobed anteriorly (Fig. 3A, 6B,F), in immature specimens sometimes with lateral lobes (Figs. 4C, 6E,G). Metaplax and hypoplax long, narrow, not longitudinally divided but sometimes slightly bifurcated posteriorly (Fig. 6A–C). Periostracum densely covered by fine, hair-like lamellae (Figs. 4B,C and 6D). Umbonal reflection with large flattened ridge. Pedal gape presents in immature (Figs. 4A,B, 6D) and some adult specimens (Fig. 3A) but it is covered by callum in older specimens (Figs. 3B, 6C). Morphological details of the new species were also presented in a series of micro-CT images published Mayoral et al. (see Fig. 8 in that paper15) and in the reconstruction of Grimaldy et al. (see Fig. 5 in that work16).Figure 8Recent freshwater piddock Lignopholas fluminalis (Blanford, 1867) in the middle reaches of the Kaladan River, Rakhine State, Myanmar13. (A) Habitat of the freshwater piddock: river pool with siltstone rocks at the bottom, a possible modern analogue of the Mesozoic riverine ecosystem with †Palaeolignopholas. (B) Siltstone rock fragment with living freshwater piddocks inside their clavate borings. (C) Ethanol-preserved piddock (dorsal view). (D) Living piddock with fully developed callum (ventral view). (E) Living piddock with pedal gape (ventral view). Abbreviations: d disc; mt metaplax; ms mesoplax; ca callum; uvs umbonal ventral sulcus; pg pedal gape; pl periostracal lamellae. Scale bar = 2 mm. (Photos: Olga V. Aksenova).Full size imageBorings and corresponding ichnotaxon. The borings produced by †Palaeolignopholas kachinensis gen. & sp. nov. represent club-shaped (clavate) structures (Figs. 1C,D, 2A–E, 7A–D), sometimes with a characteristic bioglyph revealing the shell rotation in hardening resin (Fig. 7C). These borings were illustrated in detail15,16,17,19,20, and were considered belonging to Teredolites clavatus Leymerie, 184215. Initially, the trace fossils produced by the Kachin amber piddock were described as sporocarps of Palaeoclavaria burmitis Poinar & Brown, 2003, a non-gilled hymenomycete taxon17. The holotype of this taxon represents a club-shaped piddock crypt labelled as follows: “Amber from the Hukawng Valley in Burma; specimen (in piece B with accession number B-P-1) deposited in the Poinar amber collection maintained at Oregon State University (holotype)17”. Hence, Palaeoclavaria Poinar & Brown, 2003 and P. burmitis Poinar & Brown, 2003 must be considered ichnogenus and ichnospecies, respectively. New ichnotaxonomic synonymies are formally proposed here as follows: Teredolites Leymerie, 1842 (= Palaeoclavaria Poinar & Brown, 2003 syn. nov.), and Teredolites clavatus Leymerie, 1842 (= Palaeoclavaria burmitis Poinar & Brown, 2003 syn. nov.). More

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Experimental designThis experiment was conducted at the Science and Technology experimental site (125° 27′ 5″ N, 49° 33′ 35″ E) of the North Corporation of Sinograin, Nenjiang, Heilongjiang, China. The soil in the tested area was classified as Black soil according to Chinese Soil Taxonomy (as Mollisol according to USDA classification system) with a thick humus layer and clay texture. The area has a mid-temperate continental monsoon climate with an average annual temperature of − 1.4 to 0.8 °C, precipitation of 450 mm, a frost-free period of 115 days, and an effective accumulated temperature of 2150 °C. The basic physicochemical properties at 0–20 cm plow layer of soil were shown in Table 2. The experiment included two modes: maize straw mulching (FG) and straw returning combined with rotary tillage. In the test plot, there were six 10-m ridges per treatment, and each treatment area was 39 m2. Each treatment has three replicate. The experiment started in 2012 under the continuous planting of maize, and the straw was mechanically crushed and returned to the field in the fall. In accordance with the adjustment in the C/N of the straw, the amount of applied fertilizer was N 150 kg hm−2, P2O5 75 kg hm−2, and K2O 75 kg hm−2. Five treatments were included: (1) stubble remaining in the field, which served as the control (CK); (2) full straw mulching (FG); (3) full straw returning combined with rotary tillage (1 XG) ; (4) 1/3 of full straw returning combined with rotary tillage (1/3 XG); and (5) half of full straw returning combined with rotary tillage (1/2 XG).Table 2 Basic physicochemical properties of surface soil.Full size tableSample collectionSoil samples were collected after the maize (Demeiya 2) was harvested in November 2019. Five soil cores (diameter 5 cm) were randomly taken from 0 to 20 cm depth in each plot, mixed thoroughly, and packed into cloth bags. After the crop roots and other debris were removed, the samples were air-dried for analyzing the content and chemical structure of SOC.Determination of total soil organic carbonThe content of total SOC was measured using a TOC (total organic carbon) analyzer (NC2100, Jena, Germany,) after air-dried soil samples were passed through a 100-mesh sieve.Purification of soil organic carbonFive grams of air-dried soil was added to a 100 mL plastic centrifuge tube, followed by the addition of 50 mL of hydrogen fluoride (HF) solution (10% v/v). After the tube was capped, the solution was shaken for 1 h and centrifuged for 10 min (3000 r min−1), and the supernatant was removed. Subsequently, the residue in the tube was treated with HF solution and then followed the above shaking and centrifuging steps. A total of eight repeats (according to the conditions of the actual samples) were performed with different duration of shaking (4 × 1 h, 3 × 12 h, and 1 × 24 h). Lastly, the residue in the tube was washed with double-distilled water four times, mainly to remove the residual HF in the soil sample. The detailed steps were as follows: 50 mL of double-distilled water was added into tube, shaken for 10 min and centrifuged (3000 r min−1) for 10 min, and then the supernatant was removed. The purified samples with free-HF were dried in an oven at 40 °C, ground through a 60-mesh sieve, and stored in a Zip-lock bag for NMR measurement.Determination of the chemical structure of soil organic carbonThe 13C solid-state NMR spectrum was collected on a Bruker AV400 NMR spectrometer (Switzerland). The cross-polarization magic-angle spinning (CPMAS) technique was used, the 13C NMR frequency was 400.18 MHz, the magic angle spinning frequency was 8 kHz, the contact time was 2 ms, the delay time was 3 s, and the number of data points was 3000. The chemical shift was calibrated based on the external standard sodium 2, 2-dimethyl-2-silapentane-5-sulfonate (DSS), the integrated area was automatically given by the instrument, and the relative content of organic C in each functional group of SOC was expressed as the percentage of the integrated area of a chemical shift interval to the total integrated area. The C structures corresponding to the chemical shift of the main 13C signal of SOC (Table 3) were as follows: alkyl C region (0–45 ppm), alkoxy C region (45–110 ppm), aromatic C region (110–160 ppm), and carbonyl C region (160–220 ppm)4,19.Table 3 13C solid-state NMR determination of organic carbon functional groups and corresponding high-molecular-weight compounds.Full size tableData analysisNMR spectra (CPMAS 13C-NMR) were analyzed using MestReNova professional software. After analyzing and extracting the source data, Microsoft Office Excel 2010 and Origin 8.0 software were used for data processing and plotting. The data in the “Available Data” were plotted using Origin by overlapping the fitted curve, and SPSS 17.0 (SPSS Inc., Chicago, USA) statistical analysis software was used to test for significant differences (Duncan’s method) and for correlation analysis. More