Philippa Kaur

Changes of straw degradation characteristics at different culture stagesCorn straw degradation ratioCorn straw weight loss in M44 at F1 reached 35.90% at 15 ℃ for 21 days, which was greater than that at F5, F8, and F11 by 2.33%, 3.01%, and 3.35%, respectively. There were no significant differences between F8 and F11(Fig. 1).Figure 1Corn straw degradation ratio was measured at different culture stages. The same small letter means there was no significant difference, and different small letters indicate significant differences at p  More

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Each sedimentary sequence from the three excavated ponds (Tampolove [TAMP], Ankatoke [ANKA], and Andranobe [ANDR]) includes a layer of clay (defined as zone 2), which separates the surface soil formation (zone 1) from the underlying fossiliferous muddy sand and bedrock (zone 3, Figs. S4–S7 & S9). Details regarding the composition of this sediment and its microfossils are given in Appendix-Results-Excavation (Figs. S9–S12).Subfossils and chronologyCoastal survey recovered mostly zebu bones on exposed sandy surfaces, some pygmy hippo and giant tortoise bones on the margins of shallow ponds, and giant tortoise carapace under overhanging limestone outcrops (Appendix-Results-Survey, Fig. S3). A high proportion of surface bone failed 14C analysis (~ 55%, Table S1), yet the successfully analyzed specimens (n = 8) span up to 3390–3220 calibrated years before present (cal BP, PSUAMS 8681, 3150 ± 15 14C BP, a hippo molar). Pond deposits that are relatively deep include bones that cover a relatively long period of time (Figs. S14–S16, Dataset S6). This span ranges from ~ 6000 years at TAMP (~ 120 cm deep) to ~ 2500 years at ANDR (~ 100 cm deep), with the oldest bones present in the fossiliferous sedimentary zone 3 and scarce bones in the overlying clay (zone 2).Zone 3Most bones in this layer are relatively intact and include readily identifiable pygmy hippo long bones and cranial fragments (e.g., Fig. S13a,f), giant tortoise carapace and plastron fragments (Fig. S13d), ratite eggshell and long bones (Fig. S13c,m), and crocodile scutes, cranial fragments, and teeth (Fig. S13b). Scarce bones of a duck (genus Anas) were recovered at ANDR. Remains of subfossil lemurs were scarce or absent, but they may be represented by an unknown type of bone fragment identified through protein fingerprinting (ANDR-1-5-55, Dataset S3). The widespread success of collagen extraction from these bones attests to the excellent preservation of organics in this zone. ANKA also includes keratin (mostly in the form of crocodile claws, e.g., Fig. S13i), as well as two rounded agates found associated with ratite eggshell (Fig. S13m).Remains of a juvenile pygmy hippo were recovered from both TAMP and ANDR (a femur and tibia, respectively, Dataset S3). The epiphyses of some of the pygmy hippo long bones have gnaw marks (Fig. S13f), and none of the bones include chop marks. In association with these bones towards the top of this zone are some large ( > 1 cm diameter) charcoal fragments and scarce bones of bushpig (Fig. S13k) and zebu (Fig. S13e). Protein fingerprinting identified a screened fragment of a non-zebu bovid in ANKA zone 3 and confirmed that a tentatively identified bushpig canine fragment (ANKA 1-4-151) belonged to a hippo. This zone at TAMP and ANDR also includes occasional mangrove whelk (Terebralia palustris) shells (Fig. S13g). These whelks currently live at least ~ 500 m distant from these ponds, and whelk shells at ANDR each have an irregular hole above the operculum.The span of time represented by bones in zone 3 ranges up to ~ 4000 years (~ 6000–2000 cal BP at TAMP, Fig. S14). Confirmed introduced animal bones from zone 3 failed direct 14C analysis. There are multiple examples of directly 14C-dated bone in close stratigraphic association that nonetheless differ in age by  > 1000 years, and there are a couple of examples of bones from the same individual that are separated stratigraphically. For example, two giant tortoise carapace and plastron fragments from TAMP that have indistinguishable 14C ages are separated by 22 cm of sediment (PSUAMS 8670 comes from 112 cm depth, and PSUAMS 8668 comes from 90 cm depth).Although ANKA produced what is thus far the oldest directly 14C dated pygmy hippo bone from a coastal subfossil site (PSUAMS 9383, 4380 ± 25 BP, 5030–4840 cal BP), the mean calibrated age of hippos from the Tampolove excavations (n = 11, x̄ = 2858 cal BP, SD = 972 yr) is significantly less than that of the giant tortoises (n = 9, x̄ = 4582 cal BP, SD = 705 yr, t(18) = − 4.4, p  2000 years older than a closely associated charcoal sample (38 cm depth, PSUAMS 8849, 575 ± 30 14C BP, 630–510 cal BP), which makes this molar comparable in age to bone from zone 3. Consequently, the youngest directly 14C-dated ancient bone from the Tampolove excavations comes from the lowermost zone 3: a pygmy hippo’s vertebra recovered at 90 cm depth at TAMP (PSUAMS 8730, 1865 ± 15 14C BP, 1819–1705 cal BP). Though poorly constrained in time, the deposition of zone 2 sediment came sometime within the past two millennia, which witnessed marine regression and dry intervals recorded in both the δ18O record of a nearby speleothem27 and the salinization of a nearby pan36. Previously directly 14C-dated bone collected around Tampolove attests to the local persistence of at least pygmy hippos and giant tortoises until the start of the last millennium (n = 15), and an atlas from Lamboara/Lamboharana is in fact the most recent confidently dated pygmy hippo bone from the island (PSUAMS 5629, 1100 ± 15 14C BP, 980–930 cal BP).Figure 4Cutmarked pygmy hippo femur recovered from Tampolove during recent excavation at ~ 40 cm depth (TAMP-1-2-61, above), and previously-recovered and directly 14C-dated (~ 3500 and 1600 cal BP37) cutmarked pygmy hippo femora from the nearby site of Lamboara/Lamboharana that are currently housed in the National Museum of Natural History in Paris (MAD 1709 & MAD 1710, below). Four views highlight three locations of cutmarks on the broken shaft of TAMP-1-2-61, and the inset frames show 20 × magnification of these areas, with corresponding orientations given by red lines. Note that the false color insets of TAMP-1-2-61 are meant to highlight linear edges and crevices, and the overview photos of all three femur fragments are on the same scale.Full size imageZone 1A fragment of iron (from TAMP, 16 cm depth) and sparse ceramic fragments (from ANKA, 3 & 9 cm depth) are present only in zone 1, and three 14C dates from TAMP and ANKA suggest that these specimens span the past ~ 200 years (Figs. S14–S15).CharcoalThe directly 14C dated charcoal spans all three stratigraphic zones yet consistently dates to the past millennium (Figs. S14–16). Multiple charcoal samples from different excavated ponds have practically indistinguishable 14C ages (Table S2), and much of the charcoal from Tampolove formed during peaks in the deposition of macrocharcoal at nearby Namonte (17 km distant; Fig. 5A). The onset of directly 14C-dated charcoal deposition approximately coincides with a decrease in Asafora speleothem δ18O values and with multiple directly 14C-dated first and final local occurrences of large animals. While directly 14C dated charcoal is limited to the past millennium, microcharcoal particles were abundant in all TAMP sediment samples (x̄ ± SD = 2.0 × 106 ± 2.8 × 106 particles). Additionally, microcharcoal is relatively abundant near the bottom of TAMP and ANKA, which contains bones that span ~ 6000–2000 cal BP (Fig. 5B).Figure 5Records of fire, drought, and faunal turnover from the vicinity of Tampolove within the past 1200 years, with dashed horizontal lines for reference (5A), and macrocharcoal concentrations from the excavated ponds, with depth intervals containing directly 14C-dated charcoal that spans the past millennium marked in red (5B). The past 1200 years includes the entire summed calibrated distribution of the 10 directly dated prebomb charcoal fragments from the Tampolove excavations. The calibrated probability distributions associated with the latest dates from endemic megafauna bone (giant tortoises and pygmy hippos) and earliest dates from introduced animal bone (zebu cattle and bushpigs) are shown as black distributions, and 95% of each distribution is bracketed. Considering directly dated remains within the past 4 ka from hippos (n = 26), giant tortoises (n = 18), and zebu (n = 9) and the assumption that bones were deposited uniformly over time, the grey distributions and bracketed 95% credible intervals give estimates of extirpation and arrival times. As in Fig. 3, the red line on the Asafora record follows from BCPA.Full size image More

We used a hierarchical Bayesian modeling approach to evaluate the relationship between the spring phenology of tree and wildflower species and various climate drivers (see Methods). Following model selection, our final model structure included fixed effects of average spring (March–April) temperature and elevation, as well as species-level random effects. We show continental distributions of spring temperature values in Fig. 1b (means and standard deviations are listed in Table S2). We report estimates for spring temperature sensitivities from the final model structure in the main text. Parameter estimates for elevation sensitivities as well as the model performance of other potential drivers and combinations of drivers are reported in Tables S3 and S4. An extended discussion of model assumptions and limitations is included in the Supplementary Information.Sensitivity differences by strataTree leaf out phenology (LOD) was substantially more sensitive to average spring temperature in North America (mean = −3.62 days °C−1; 95% credible interval (CI) = [−3.76, −3.49]) than in Europe (mean = −2.79; CI = [−3.27, −2.30]) and Asia (mean = −2.62; CI = [−2.97, −2.26]; Fig. 2). These values are consistent with previously reported phenological sensitivities in North America7 (−5.5 to −3.3 days °C−1) and Europe8 (−4.1 to −3.0 days °C−1), as the credible intervals from our results overlap with the reported credible intervals of prior studies. However, the Asian LOD sensitivity was less sensitive than previously reported27 (−3.50 to −3.03 days °C−1), potentially owing to differences in species selection28 or model structure. Previously reported sensitivities were determined in separate studies using either observational data7,8 or long-term observation-based weather station data27. The general consistency between our findings suggests that phenology data from herbarium collections are good indicators of patterns in natural systems29,30,31, a point supported by a recent study of phenological sensitivity derived from herbaria and from observed citizen science data32. These herbarium-based results provide evidence that phenological sensitivity differs across the temperate forest biome (but see ref. 33 for evidence of differences in response to warming and chilling accumulation). To our knowledge, our study is the first to contrast overstory and understory phenology across multiple continents and, therefore, to find differences in phenological sensitivity between trees and forest wildflowers across continents. We recommend future studies explore these differences using alternative approaches and methodologies that focus on the physiological basis for and mechanisms that underlie these patterns.Fig. 2: Posterior estimated means and 95% credible intervals for spring temperature sensitivity.Shapes represent parameter estimates for wildflower First Flower Date (FFD, blue circles; n = 1418, 618, and 1060 for Asia, Europe, and North America, respectively) and canopy tree Leaf Out Date (LOD, yellow triangles; n = 899, 532, and 995, for Asia, Europe, and North America, respectively). Estimates are considered different from 0 if credible intervals do not overlap the dashed 0 line and are considered different from each other if credible intervals do not overlap.Full size imageIn contrast to trees, wildflower sensitivity to spring temperature was similar across all three continents and exhibited no strong differences (i.e., overlap in 95% Bayesian credible intervals) among continents (means and 95% credible intervals in brackets: North America = −3.14, [−3.28, −3.00]; Europe = −3.02, [−3.48, −2.56]; Asia = −3.12, [−3.36, −2.86]; Fig. 2). These values are also generally consistent with those reported elsewhere in the literature (i.e., 95% credible intervals overlap with those reported in other studies; −2.2, [−3.7, −0.76] days °C−1 in North America7 and −3.6, [−4.04, −3.18] days °C−1 in Europe9), although we are unaware of any studies that have estimated phenological sensitivity for Asian forest wildflowers in days °C−1. Ge et al.3 report herbaceous plant sensitivity of −5.71 days per decade in Asia (±7.90 standard deviation; based primarily on long-term observational data), which appears to be roughly consistent with our model results, but the difference in units makes this more speculative than the other comparisons. Discrepancies in mean responses between this study and others may be due in part to different types of data (herbarium specimens versus field observations) and to choice in focal taxa, as temperature sensitivity has been shown to vary widely across taxa28.Particularly noticeable in our results was that r2 coefficients of predicted versus observed phenology were much higher in North America (0.70 and 0.76 for wildflower and tree models, respectively) compared to Asian (0.40 and 0.44, respectively) and European models (0.41 and 0.25, respectively). This difference in model performance could be due to the higher interannual variability of spring temperatures in North America33, leading to greater selective pressure for strong sensitivity to spring temperatures in North American plants. This difference could explain why North American species exhibit higher correlation of phenology with average spring temperatures (Table S4). Alternatively, European and Asian species may have stronger phenological responses to alternative spring forcing windows, winter chilling temperatures, or photoperiod, relative to the March–April temperature period used in this study (see Methods). We think the latter explanation is unlikely, given the strong correlations of phenology with spring temperature across all continents (see Supplementary Information – Justification for March–April Temperature Window).Herbarium-based phenological models may be improved by accounting for spatial autocorrelation within the dataset. For example, Willems et al.9 found that including spatial autocorrelation significantly improved predictability of European herbaceous flowering phenology, even when accounting for multiple drivers of spring phenology. We followed a similar approach as their study and found similar improvements in model performance with the addition of spatial autocorrelation (Tables S3–S4) that had substantial positive effects on r2 values of Asian and European models. However, spatial distributions of specimens differed substantially among continents (see Figs. S2–S4), and these differences could lead to artifacts that make results unreliable to interpret (see Supplementary Information). Therefore, we focus here on results for models without spatial autocorrelation while acknowledging that spatial aggregation of herbarium specimens in Europe and Asia may be partially responsible for the relatively lower r2 values. We encourage other researchers to explore this question further both with our data set and other datasets.Climate change and spring light windowsThe relative difference between wildflower and tree sensitivity varied substantially among continents, with wildflowers being approximately equally as sensitive to spring temperature as trees in Asia and Europe but substantially less sensitive (i.e., 95% BCI do not overlap) than trees in North America (Fig. 2). Importantly, these differences were driven by changes in tree phenological sensitivities among continents and resulted in different expectations for spring light window duration (i.e., the difference in time between estimated wildflower flowering date and canopy tree leaf out date) on different continents under current climate conditions (Fig. 3), based on modeled leaf out and flowering under a climate scenario derived from average climate conditions from 2009–2018 (Fig. S5).Fig. 3: Current estimated phenological escape duration in northern temperate deciduous forests.Estimated mean difference between wildflower First Flower Date (FFD) and canopy tree Leaf Out Date (LOD) (in days) under current climate conditions (averaged from 2009–2018, see methods) in a Asia, b Europe, and c North America. Negative values indicate tree LOD is estimated to occur before wildflower FFD. Estimations were cropped by the estimated area of broadleaf and mixed-broadleaf forest (see methods). 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