Philippa Kaur

Sample procurement and data analysisTo establish a diachronic dataset of Galápagos tortoise dietary stable isotope ecology, we selected samples from five sources (see Supplemental Text): the American Museum of Natural History, New York, New York, (2) the California Academy of Sciences, San Francisco, California, (3) the Natural History Museum, London, England, (4) the National Museum of Natural History, Smithsonian Institution, Washington, D.C., and (5) the Thompson’s Cove (CA-SFR-186H) archaeological site in San Francisco, California. We provide details regarding sample provenience information and date-of-death as supplemental information. From these collections, we obtained single or multiple isotope samples from a total of 57 individual tortoises representing the following subspecies (n = 10) and islands: five C. n. abingdonii (Pinta Island), one C. n. becki (Volcán Wolf, Isabela Island), five C. n. chathamensis (San Cristóbal Island), four C. n. darwini (Santiago Island), thirteen C. n. duncanensis (Pinzón Island), four C. n. guentheri (Sierra Nega, Isabela Island), six C. n. hoodensis (Española Island), one C. n. microphyes (Volcán Darwin, Isabela Island), four C. n. niger (Floreana Island), nine C. n. porteri (Western Santa Cruz Island), one C. n. vicina (Cerro Azul, Isabela Island), one unknown Isabela Island tortoise, two C. n. vicina tortoises which were transported, lived and collected on Rabida Island, and one unknown tortoise (Chelonoidis niger ssp.; unknown Island—the San Francisco Gold Rush sample). The two earliest collected tortoises in our sample date to1833 and the latest tortoise is from 1967, representing a period of 134 years.To understand tissue-specific isotopic variation and fractionation for the purposes of reconstructing long-term dietary ecology, we sampled tortoise bone collagen (n = 57), bone apatite (n = 23), scute keratin (n = 8) and skin (n = 2) for carbon (δ13Ccollagen and δ13Capatite), nitrogen (δ15N), hydrogen (δD) and oxygen (δ18Oapatite) stable isotopes. All samples were drilled or cut using a Dremel rotary tool with either a blade or diamond spherical bit attachment and were transported to the University of New Mexico, Center for Stable Isotopes (UNM-CSI), Albuquerque, NM, for preparation and analysis. All statistical and metric data analysis and visualization occurred in R (4.0.4) and RStudio (2022.02.4). We provide reproducible source code supplemental to the text35.Bone collagen δ13C, δ15N and δDAnalysis of bone collagen, skin and scute keratin for carbon, nitrogen and hydrogen stable isotopes followed standardized protocols (e.g., see36). For bone collagen, we cut and demineralized a small portion of bulk bone in 0.5 N hydrochloric acid (HCl) at 5 °C for 24 h prior to rinsing all samples to neutrality using deionized water. For lipid extraction, we immersed the samples in a solution of 2:1 chloroform:methanol (C2H5Cl3) for 24 h (repeated three times) while also sonicating samples for 15 min to ensure complete chemical saturation. Preparation of skin and scute keratin samples was only included this during the later lipid extraction step (i.e., no demineralization required). After 72 h we rinsed all samples to neutrality and lyophilized the tortoise samples for another 24 h. We then measured approximately 0.5–0.6 mg of bone collagen/skin/scute tissue into tin capsules for carbon (δ13Ccollagen) and nitrogen (δ15N) stable isotope analysis. We also measured approximately 0.2–0.3 mg of bone collagen/skin/scute tissue into silver capsules for hydrogen (δD) isotope analysis. We report isotope values in delta (δ) notation, calculated as: ((Rsample/Rstandard) − 1) × 1000, where Rsample and Rstandard are the ratios (e.g., 13C/12C, 15N/14N) of the unknown and standard material, respectively. Delta values are reported as parts per thousand (‰).Carbon and nitrogen samples were measured on a Costech 4010 elemental analyzer (Valencia, California, USA) coupled to a Scientific Delta V Plus isotope ratio mass spectrometer by a Conflo IV, and hydrogen samples were measured on a Finnigan high-temperature conversion elemental analyzer (TC/EA) coupled to a Thermo Scientific Delta V Plus mass spectrometer by a Conflo IV at UNM-CSI (see37 for details on the high temperature conversion method for hydrogen analysis). All nitrogen and carbon isotope data are reported relative to atmospheric N2 and V-PDB, respectively. The data were corrected using lab standards with values of δ15 N = 6.4‰ and δ13C =  − 26.5‰ (casein protein), and of δ15N = 13.3‰ and δ13C =  − 16.7‰ (tuna muscle) that have been calibrated relative to the universally accepted standards: IAEA-N1, USGS 24, IAEA 600, USGS 63, and USGS 40.To ensure equilibrium between the exchangeable hydrogen in tissue samples and local atmosphere38, we weighed hydrogen standards and samples into silver capsules and allowed both to sit in the laboratory for at least 2 weeks before analysis. Hydrogen data were corrected using three UNM-CSI laboratory keratin standards (δDnon-ex =  − 174‰, − 93‰, and − 54‰) of which the δDnon-ex values were previously determined through a series of atmospheric exchange experiments. These standards were also calibrated to USGS standards CBS and KHS values of − 178.8‰ and − 47.5‰, respectively (see39,40 for details and updated values). To quantitate any error imparted to our collagen data through correction with keratin standards, a UNM-CSI cow (Bos taurus) bone collagen standard was analyzed in every run over a 6-month period (July 2017–January 2018) and gave an inter-run standard deviation of 3.9‰, suggesting the difference in percent exchangeable hydrogen between collagen and keratin tissues did not significantly impact our results. All hydrogen isotope data are reported relative to Vienna-Standard Mean Ocean Water (V-SMOW). The H3 factor was between 8 and 8.5 for all runs.Collagen precision (standard deviation; SD) for within-run analyses is  More

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MAG binning and general featuresFrom the four hot springs, we assembled four associated metagenomes and then binned a total of 42 MAGs. We recovered 12 MAGs from RB10 hot spring, 13 from RB13, 14 from RB32 and 3 from RB108. Out of these 42 MAGs, 7 were of high-quality, 25 of nearly-high quality, 9 of medium quality and 1 of low quality (Table 1) based on metagenomic standards26. The GC% was quite variable, ranging from 25.76 to 70.35% among all MAGs and between 32.15 and 69.21% only among the high- and near high-quality MAGs. With the exception of RB108 from which we only recovered bacterial MAGs, we retrieved both bacterial and archaeal MAGs in the other hot springs. Two thirds of the MAGs (26/42) were assigned to the domain Bacteria and the rest to the domain Archaea (16/42) (Table 2).Table 1 General characteristics of the 42 MAGs obtained from RB10, RB13, RB32 and RB108 samples.Full size tableTable 2 Classification of the MAGs based on the taxonomic classification of GTDB-Tk software (v2.1.0) and the Genome Taxonomy Database (07-RS207 release).Full size tableTaxonomic and phylogenomic analyses of MAGsThe taxonomic affiliation of the MAGs was investigated in detail through the workflow classify of GTDB-Tk (v 2.1.0; GTDB reference tree 07-RS207) (Table 2) and through de novo phylogenomic analyses (Fig. S1a–i). We also tried to classify MAGs on the basis of overall genome relatedness indices (OGRI), which is detailed in supplementary material (Text S1, Table S2, Fig. S2).De novo phylogenomic analyses globally confirmed the positioning of MAGs provided by GTDB-Tk, with high branching support. For Bacteria, GTDB-Tk analyses allowed us to place the MAGs in the following clades: six in the phylum Aquificota from the four different springs, comprising four MAGs belonging to the genus Hydrogenivirga (family Aquificaceae) (RB10-MAG07, RB13-MAG10, RB32-MAG07, RB108-MAG02), and two belonging to the family ‘Hydrogenobaculaceae’ (RB10-MAG12, RB32-MAG11) (Table 2, Fig. S1a). Their closest cultured relatives originated either from hot springs or from deep-sea hydrothermal vents27. Three MAGs from three geothermal springs belonged to the phylum Armatimonadota (RB10-MAG03, RB13-MAG04, RB32-MAG03) and had no close cultured relatives. Seven MAGs have been classified into the phylum Chloroflexota: three MAGs belonging to the genus Thermoflexus from three different springs (RB10-MAG04, RB13-MAG05, RB32-MAG02), one affiliating with the genus Thermomicrobium (RB32-MAG08), one falling into the family Ktedonobacteraceae (RB108-MAG03), one belonging to the class Dehalococcoidia (RB32-MAG04) and another one whose phylogenetic position is more difficult to assert because it is a MAG of medium quality (RB32-MAG14). Six MAGs from four various hot springs belonged to the phylum Deinococcota, and to the genera Thermus (RB10-MAG08, RB10-MAG11, RB13-MAG09, RB32-MAG10, RB108-MAG01) and Meiothermus (RB13-MAG13). One MAG belonged to the family ‘Sulfurifustaceae’ (RB13-MAG01), in the phylum Proteobacteria (Gamma-class). The MAG referenced as RB32-MAG13 was classified into the phylum ‘Patescibacteria’, in the class ‘Paceibacteria’, and was distantly related to MAGs originating from groundwater and from hot springs. Finally, two MAGs from two different springs belonged to the phylum WOR-3, in the Candidatus genus ‘Caldipriscus’ (RB32-MAG12, RB10-MAG09).For Archaea, almost all the MAGs reconstructed in this study, e.g. 15 of the 16 archaeal MAGs, belonged to the phylum Thermoproteota. Among them, four belonged to the genus Ignisphaera (RB10-MAG05, RB13-MAG08, RB13-MAG11, RB32-MAG05), three to the genus Infirmifilum (RB10-MAG06, RB13-MAG03, RB32-MAG09), two to the genus Zestosphaera (RB10-MAG02, RB13-MAG06), three to the family Acidilobaceae (RB10-MAG01, RB13-MAG02, RB32-MAG01) and two to the order Geoarchaeales (RB10-MAG10, RB32-MAG06). Additionally, one belonged to the family Thermocladiaceae (RB13-MAG07). Lastly, the MAG belonging to another phylum (RB13-MAG12) was affiliated with the ‘Aenigmatarchaeota’, class ‘Aenigmatarchaeia’, and was distantly related to MAGs from hot springs and from deep-sea hydrothermal vent sediments28,29.Out of these 42 MAGs, at least 19 MAGs corresponded to different taxa at the taxonomic rank of species or higher according to GTDB (Table 2). Eighteen of them belonged to lineages with several cultivated representatives including the species Thermus thermophilus. 13 new genomic species within the GTDB genera Hydrogenivirga, HRBIN17, Thermoflexus, SpSt-223, CADDYT01, Zestosphaera, Ignisphaera, Infirmifilum, Thermus, Thermus_A, Meiothermus_B, JAHLMO01 and Caldipriscus, and 6 putative new genomic genera belonging to the GTDB families Hydrogenobaculaceae, Acidilobaceae, WAQG01, Thermocladiaceae, Sulfurifustaceae and HR35 could be identified (Table 2). In addition, 9 MAGs belonged to lineages that are predominantly or exclusively known through environmental DNA sequences. Thus, these 42 MAGs comprised a broad phylogenetic range of Bacteria and Archaea at different levels of taxonomic organization, of which a large majority were not reported before.The approaches implemented here were not intended to describe the microbial diversity present in these sources in an exhaustive way or to compare them in a fine way, and cannot allow it because of a 2-year storage at 4 °C. This long storage has probably led to changes in the microbial communities and to the selective loss or enrichment of some taxa. As a result, no analysis of abundance or absence of taxa can be conducted from these metagenomes and the results are discussed taking this bias into account. However, they do provide an overview of the microbial diversity effectively present. If we compare the phylogenetic diversity of the MAGs found in the four hot springs, we can observe that 3 shared phyla (Deinococcota, Aquificota and Chloroflexota: phyla names according to GTDB), 2 shared families (Thermaceae and Aquificaceae), and one shared genus (Hydrogenivirga) were found among the four sources (Fig. 2). In addition, hot springs RB10, RB13 and RB32, that are geographically close ( More

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