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    Fractal features of soil grain-size distribution in a typical Tamarix cones in the Taklimakan Desert, China

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    Evolutionary implications of new Postopsyllidiidae from mid-Cretaceous amber from Myanmar and sternorrhynchan nymphal conservatism

    Systematic palaeontologyOrder Hemiptera Linnaeus, 1758Suborder Sternorrhyncha Amyot et Audinet-Serville, 1843Superfamily Protopsyllidioidea Carpenter, 1931Family Postopsyllidiidae Hakim, Azar et Huang, 2019Genus Megalophthallidion Drohojowska et Szwedo, gen. nov.LSID urn:lsid:zoobank.org:act:A6F71390-9B8E-4A19-8F30-C2A024B6EFB1Type speciesMegalophthallidion burmapateron Drohojowska et Szwedo, sp. nov.; by present designation and monotypy.EtymologyGeneric name is derived from Classic Greek megas (μέγας)—large, ophthalmos (ὀφθαλμός)—an eye and Greek form of generic name Psyllidium. Gender: masculine.Type localityNorthern Myanmar: state of Kachin, Noije bum 2001 Summit Site amber mine in the Hukawng Valley, SW of Maingkhwan.Type stratumLowermost Cenomanian, Upper Cretaceous (‘mid-Cretaceous’).DiagnosisHead capsule with 12 stiff setae on tubercles (18 setae in Postopsyllidium); fore wing without pterostigma (tiny pterostigma, widening of ScP + RA present in Postopsyllidium); vein CuP not thickened distally (distinctly thickened distally in Postopsyllidium); profemur with a row of ventral (ventrolateral) setae (two rows in Postopsyllidium).Megalophthallidion burmapateron Drohojowska et Szwedo, sp. nov.LSID urn:lsid:zoobank.org:act:F3F971F4-AE04-4F41-98B0-9A0A04470625.(Figs. 1A–F, 2A–I).Figure 1Megalophthallidion burmapteron gen. et sp. nov., holotype (MAIG 6687), imago. (A) Photo of body, ventral side; (B) photo of right antennae and (C) drawing of antenna; (D) drawing of body, dorsal side; (E) drawing of thorax structure with sclerites marked: red—pronotum; orange—mesopraescutum; yellow—mesoscutum; light green—mesoscutellum, dark green—mesopostnotum; light blue—metascutum; dark blue—metascutellum; violet—metapostnotum; (F) photo of thorax dorsal side. Scale bars: 0.5 mm (A), 0.2 mm (B–D), 0.1 mm (F).Full size imageFigure 2Megalophthallidion burmapteron gen. et sp. nov., holotype (MAIG 6687), imago. (A) Photo of right fore wing; (B) photo of right wings; (C) photo of antenna and proleg; (D) photo of proleg and mesoleg, and (E) photo of femur of proleg, and (F) photo of right metatarsus and left mesotarsus in the background, and (G) photo of right mesotarsus of mesoleg, and (H) Photo of tarsi; (I) photo of male genital block. Scale bars: 0.5 mm (A–D), 0.2 mm (B,E,F,H), 0.1 mm (G,I).Full size imageMaterialHolotype, number MAIG 6687 (BUB 96), deposited in Museum of Amber Inclusions (MAIG), University of Gdańsk, Poland. Imago, a complete and well-preserved male. Piece of amber 8 × 6 × 3 mm, cut from larger lump, polished flat on both sides.Type localityNorthern Myanmar: state of Kachin, Noije bum 2001 Summit Site amber mine in the Hukawng Valley, SW of Maingkhwan.Type stratumLowermost Cenomanian, Upper Cretaceous (‘mid-Cretaceous’).DiagnosisAs for the genus with the following additions: three ocelli distinct, antennomere IX the longest, about as long as pedicel, antennomeres III–VII and XI of similar length, antennomere XII the shortest, subconically tapered in apical portion. Paramere lobate, ventral margin with acute, small process, apical and dorsal margins rounded. Aedeagus geniculately bent at base, directed dorsally, tapered apicad.DescriptionMale (Figs. 1A–F, 2A–I). Head with compound eyes distinctly wider than pronotum (Fig. 1D–F). Compound eyes subglobular, protruding laterally. Vertex short in midline, about 2.5 times as wide as posterior margin and as long in middle; trapezoidal, anterior margin slightly arched, lateral margins diverging posteriad, posterior margin shallowly arched, disc of vertex with distinct setae on large tubercles: four setae at posterior margin, two at anterior angles of compound eyes, two medial, over the median ocellus. Three ocelli present, median ocellus distinct, visible from above, lateral ocelli near anterior angles of compound eyes. Frons about as wide as long in midline, two rows of setae on tubercles, upper row at level of median ocellus, lower one, below half of compound eye height. Clypeus, elongate, triangular, in lower portion roof-like; two setae on tubercles near upper margin. Genae very narrow. Rostrum reaching slightly beyond mesocoxae, apical segment slightly shorter than subapical one, darker. Antennae bases placed at lower margin of compound eyes; antennal fovea elevated; scapus shorter than pedicel, cylindrical; pedicel cylindrical; antennomeres IIIrd–VIIth and XIth of similar length, VIIIth slightly longer than VIIth, as long as Xth antennomere, IXth the longest, XIIth the shortest, tapered apically; rhinaria absent.Thorax (Fig. 1D–F): pronotum quadrangular, about as long as mesothorax; pronotum with anterior and posterior margins parallel, merely arcuate, disc with transverse groove in the median portion, lateral margins slightly arcuate, two distinct setae on tubercles in anterolateral angle, two setae on tubercles anterior margin at distance1/3 to median line, three distinct setae on tubercles in posterolateral angles. Mesopraescutum subtriangular, with apex widely rounded, about 0.4 times as wide as pronotum, about 0.4 times as long as wide, delicately separated from mesoscutum. Mesoscutum as wide as pronotum at widest point, distinctly narrowed medially, anterior angles rounded, anterolateral margin sigmoid, lateral angle acute, posterior angles wide, posterior margin V-shape incised, posterolateral areas of mesoscutum disc declivent posteriorly; disc with two setae on tubercles, at 1/3 of mesoscutum width. Mesoscutellum about as long as wide, diamond-shape, anterior and lateral angles acute, posterior angle rounded. Mesopostnotum in form of transverse band, slightly widened in median portion. Metascutum narrower than mesoscutum, anterior angles widely rounded, lateral angles acute, anterolateral margin concave, posterior margin arcuate, with deep median arcuate incision. The suture between metascutum and metascutellum weakly visible, metascutellum subtriangular, longer than wide at base.Parapteron with three distinct setae.Fore wing (Fig. 2A,B) membranous, narrow, elongate, about 3.5 times as long as wide, widest at 2/3 of length. Anterior margin merely arcuate, slightly bent at very base, anteroapical angle widely arcuate, apex rounded, posteroapical angle widely arcuate, tornus arcuate, claval margin straight, with incision between terminals of Pcu (claval apex) and A1. Stem ScP + R + MP + CuA slightly arcuate, very short stalk ScP + R + MP + CuA leaving basal cell, stem ScP + R oblique, straight, forked in basal half of fore wing length, branch ScP + RA, oblique, reaching anterior margin slightly distally of half of fore wing length, slightly distally of ending of CuA2 branch; branch RP slightly arcuate, a little more curved in basal section, reaching margin at anteroapical angle; stalk MP + CuA slightly shorter than basal cell; stem MP almost straight, forked in apical half of fore wing, at about 2/3 of fore wing length, with three terminals reaching margin between apex and posteroapical angle; stem CuA shorter than branches CuA1 and CuA2, about half as long as branch CuA1; claval vein CuP weak at base, not thickened distally; claval vein Pcu straight, claval vein A1 straight. Basal cell present, subtriangular, about twice as long as wide, basal veinlet cua-cup oblique, no other veinlets present; cell r (radial) very long, longer than half of fore wing length; cell m (medial) the shortest, shorter than cell cu (areola postica). Margins of fore wing with fringe of long setae, starting on costal margin near base of fore wing, ending at level of middle of cell cu; longitudinal veins with distinct, scarcely but evenly dispersed, movable setae; terminal section of CuP with two setae; costal margin with row of short, densely distributed setae, apical margin, tornus and claval margin with rows of scaly setae.Hind wing (Fig. 2B) membranous, shorter than fore wing, 3.23 times as long as wide. Costal margin bent at base, then almost straight up to the level of ScP + RA end and wing coupling lobe, then straight to anteroapical angle, anteroapical angle widely arcuate, apex arcuate, posteroapical angle arcuate, tornus straight, claval margin merely arcuate, posteroclaval angle angulate; stem ScP + R + MP bent at base, then straight, stem ScP + R short, branch ScP + RA short, about as long as stem ScP + R, branch RP arcuate basally than straight, reaching apex; stem MP arcuate, forked slightly distad CuA1 terminus level, branch MP1+2 slightly arcuate, reaching margin at posteroapical angle, branch MP3+4 straight, reaching tornus; stem CuA slightly bent at base, then straight, forked slightly distad ScP + R forking, branch CuA1 arcuate, branch CuA2 short, straight, slightly oblique, reaching tornus; claval vein CuP weak, visible only at base, claval vein Pcu slightly arcuate; wing coupling apparatus (fold) with a few short setae.Legs slender, relatively long, profemora armed (Fig. 2C–H). Procoxa as long as profemur, narrow, flattened. Protrochanter scaphoid, elongate, with long apical and subapical setae. Profemur flattened laterally, about as long as protibia, ventrally armed with four large setae on elevated plinths; dorsal margin with row of short, decumbent setae. Protibia narrow, rounded in cross section, covered with short setae, a few longer setae in distal portion. Protarsus—single, long tarsomere, plantar surface with row of semi-erect setae; tarsal claws long, straight, directed ventrally, no arolium nor empodium.Mesocoxa elongate, narrow, slightly flattened. Mesotrochanter scaphoid. Mesofemur slender, flattened laterally, dorsal margin with short setae. Mesotibia subequal to mesofemur, slender, covered with setae, two apical setae slightly thicker and longer. Mesotarsus with three tarsomeres, basimesotarsomere the longest, shorter than cumulative length of mid- and apical mesotarsomere, plantar margins with setae, two apical setae slightly longer and thicker; midmesotarsomere the shortest, 1/3 of basimesotarsomere length, a few setae on plantar surface; apical tarsomere shorter than basimesotarsomere, twice as long as midmesotarsomere, plantar surface with a few, scarcely dispersed setae, tarsal claws long, narrow, directed ventrally, no arolium nor empodium.Metacoxa conical, narrow. Metatrochanter scaphoid, elongate. Metafemur slender, laterally flattened, longer than mesofemur, dorsal margin with row of short setae. Metatibia, long, slender, 1.6 times as long as metafemur, with suberect setae of different size, two larger and longer and two shorter setae subapical setae. Metatarsus slightly less than half of metatibia length, with three tarsomeres, basimetatarsomere the longest, more than twice as long as apical metatarsomere, 1.5 times as long as combined length of mid- and apical metatarsomere, plantar surface with scarce decumbent setae; mid metatarsomere the shortest, 1/4 of basimetatarsomere length, plantar surface with a few setae, two apical ones slightly thicker; apical metatarsomere about 0.4 of basimetatarsomere length, with scarcely dispersed setae on along plantar surface; tarsal claws, long, slender, other pretarsal structures absent.Abdomen (Fig. 1F) narrowly attached to thorax, tergite segment shorter, 2nd tergite distinctly longer, 3rd to 8th tergites of similar length; pygofer narrowing apicad, ventral margin strongly elongated posteriorly; anal tube short, directed posterodorsad, anal style shorter than anal tube. Paramere lobate, ventral margin with acute, small process, apical and dorsal margins rounded. Aedeagus (Fig. 2I) geniculately bent at base, directed dorsad, tapered apicad.Female. Unknown.Megalophthallidion sp. (5th instar nymph)(Figs. 3A–D, 4A–F)Figure 3Megalophthallidion sp. (MAIG 6688), nymph. (A) Photo of body, dorsal side and (B) drawing of body dorsal side; (C) photo of body dorsal side and (D) drawing of body ventral side. Scale bars: 0.5 mm (A–D).Full size imageFigure 4Megalophthallidion sp. (MAIG 6688), nymph. Photo of clypeus and (B) drawing of clypeus; (C) photo of proleg, and (D) photo of mesoleg, and (E) photo of metaleg; (F) photo of posterior part of abdomen ventral side. Scale bars: 0.1 mm (A–F).Full size imageMaterialNymph, 5th instar, MAIG 6688 (BUB 1799), deposited in Museum of Amber Inclusions (MAIG), University of Gdańsk, Poland. Piece of amber 13 × 6 × 2 mm, cut from larger lump, polished flat on one side, more convex on the other.Diagnostic charactersThe nymph of Megalophthallidion gen. nov. is similar in general body shape to the only known fossil protopsyllidioidean nymph described from Lower Cretaceous Lebanese amber—Talaya batraba Drohojowska et Szwedo, 2013. The nymph of Talaya batraba is 2nd or 3rd instar, therefore some features are difficult to compare with this last instar nymph of Megalophthallidion gen. nov. The morphological states observed in those two specimens are: head covered with strongly expanded disc and expanded disc of pronotum, however shapes and ratios of these structures differ; compound eyes on ventral side of head, shifted laterad (ommatidia on cones in T. batraba, while ventroposterior expansions are present in Megalophthallidion gen. nov.); compound eyes visible from above as short, stout cones in fissure between posterior margin of disc (hypertrophied vertex) and anterior margin of pronotum (compound eyes (?) are visible on dorsal side of Permian Aleuronympha bibulla Riek, 1974); in Megalophthallidion gen. nov. rostrum reached mesocoxa, while in Talaya batraba distinctly exceeds length of the body; abdomen with 9 segments; tergites of abdominal segments 5th–9th expanded posterolaterad in form of fan-like expansion; 9th abdominal segment short, placed ventral; anal tube short, cylindrical, epiproct (?) globular.DescriptionNymph, 5th instar (Figs. 3A–D, 4A–F). Body oval shaped, dorso-ventrally flattened, 1.5 times longer than wide with segmentation visible; on the ventral side slightly concave. Length of body c. 1.56 mm long, outline, in dorsal view, maximum width of body 0.94 mm; length of head and pronotum (cephaloprothorax) c. 0.46 mm in midline, width c. 0.83 mm; cumulative length of mesonotum + metanotum c. 0.25 mm; abdomen c. 0.8 mm long. Dorsal side (Fig. 3A,B) with distinct median line (ecdysial line), not reaching anterior or posterior margin of the body, the line distinctly roof-like in abdominal portion. Anterior margin of head (cephaloprothorax) disc arcuate, lateral angles rounded; anterior margin of pronotum arcuate, lateral margins arcuately diverging posteriad, posterior margin distinctly arcuate, anterior angles widely rounded, posterior angles acutely rounded, disc elevated, convex, lateral portions declivitous; the fissure between posterior margin of head disc and anterior margin of pronotum narrow, widened medially, with stalked compound eyes popping out.Head partly separated from prothorax, wide in ventral view. Bases of antennae protruding anterolaterally, wide, anterior margin arcuate, with a small lump extending anteriorly connecting margin with vertex expansion. Suture separating anteclypeus and postclypeus visible in ventral aspect (Fig. 4A,B). Postclypeus about three times as long as wide, oval, slightly swollen, without any setae; weak traces of salivary pump muscle attachments visible. Anteclypeus about as long as postclypeus, widened in upper section below clypeal suture, convex, carinately elevated in lower section, with sides distinctly declivitous, clypellus long, carinately elevated. Lora (mandibulary plates) distinct, separated from anteclypeus by shallow suture, with upper angles at half of postclypeus length, lower angles at half of anteclypeus length, about as wide as half of postclypeus width. Maxillary plates narrow. Genal portion of head enlarged, medial portion arcuately convex; lateral sections narrowing laterally, terminally encircling bases of compound eyes. Antennae short (Fig. 3C,D), placed in front of genal portion. Antennal flagellum indistinctly subdivided into four segments. Rostrum (Fig. 4A,B) three-segmented, 0.2 mm long, with apex reaching apex of mesocoxae; apical segment about 2.5 times as long as subapical one.No lateral sclerites on meso- and metathorax, only one plus one large medial sclerite on both meso- and metathorax. Mesothoracic and metathoracic wing pads distinct, wide, subtriangular, with posterior apices directed posteriorly; lateral portions of mesothoracic wing pads arcuate. Fore wing pad 0.6 mm long, with small, straight humeral lobe, forming a right angle, not protruding anteriorly. Mesothoracic tergites slightly larger than metathoracic segments (respectively c. 0.14 mm and c. 0.12 mm long in midline, 0.26 mm and 0.27 mm in lateral lines); mesothoracic tergum with distinct median elevation (low double crest with ecdysial line in between), slightly wider than long in midline, anterior margin arcuate, lateral margins straight, subparallel, posterior margin concave. Metathoracic wing pad apex slightly exceeding mesothoracic wing pad. Metathoracic tergum wider than long, slightly shorter than mesothoracic tergum, with distinct elevation in the middle.Legs relatively long (Figs. 3C,D, 4C–E). Coxae of legs placed near the median axis of the body. Prolegs: procoxal pit with margins elevated, procoxa conical (c. 0.1 mm long), protrochanter scaphoid, about as long as procoxa, profemur c. 0.13 mm long, slightly flattened laterally, merely thickened, protibia longer than profemur, c. 0.23 mm long; tarsus shorter than protibia, basiprotarsomere about as long as apical protarsomere, the latter with distinct tarsal claws, and wide arolium. Mesoleg similar to proleg, mesocoxa conical (c. 0.1 mm long), mesotrochanter scaphoid, mesofemur (c. 0.13 mm) slightly flattened laterally, mesotibia slightly longer than mesofemur (c. 0.18 mm), mesotarsus slightly shorter than mesotibia, three-segmented, basimesotarsomere the longest (c. 0.07 mm), about as long as combined length of mid- and apical mesotarsomeres (c. 0.04 mm respectively), arolium wide, tarsal claws distinct. Metaleg: metacoxa conical (c. 0.1 mm), metatrochanter scaphoid, about as long as metacoxa (c. 0.12 mm). Metafemur (c. 0.17 mm) slightly more thickened than pro- and mesofemur, metatibia slightly longer (0.19 mm) than pro- and mesotibiae. Metatarsus three-segmented: basimetatarsomere about as long (0.08 mm) as combined length of mid- and apical metatarsomeres (0.04 mm respectively), arolium lobate, wide, tarsal claws distinct, widely spread.Abdomen (Fig. 3A–D) 9-segmented, narrow at base, widening fan-shape posteriorly, 1st segment visible from above, segmentation visible, abdominal terga 5th–9th expanded posterolaterally. Tergites carinately elevated in the middle, separated by ecdysial line. 1st sternite visible in ventral view, sternites 2nd–4th fused medially, sternites 5th–9th separated; 9th abdominal segment short (Fig. 4F), placed ventrally, under tergal expansion; anal tube short, cylindrical, epiproct (?) globular. More

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    Influence of green technology, green energy consumption, energy efficiency, trade, economic development and FDI on climate change in South Asia

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    Spatial distribution characteristics and evaluation of soil pollution in coal mine areas in Loess Plateau of northern Shaanxi

    Analysis of contents of heavy metals in wasteland soilThe test results show (Table 5) that the contents of Hg, Cd, As, Pb, Cr, Zn, Ni and Cu in the surface soil within Shigetai Coal Mine vary from 0.043 to 0.255, 0.44 to 2.23, 2.66 to 18.40, 11.80 to 42.80, 40.50 to 118.60, 18.90 to 70.10, 4.31 to 28.10, 4.96 to 46.25 mg/kg, respectively; the average contents of Hg, Cd, As, Pb, Cr, Zn, Ni and Cu are 0.128, 1.03, 4.73, 23.08, 76.22, 46.94, 16.11 and 12.10 mg/kg, respectively. The average contents of Hg, Cd, Pb and Cr in soil within the research area are 2.03, 1.36, 1.11 and 1.23 times of the soil background values in Shaanxi Province, respectively. The average contents of As, Zn and Cu are lower than the soil background value in Shaanxi Province, but the maximum contents of these three elements are 1.65, 1.01 and 2.16 times of the soil background values in Shaanxi Province, respectively. It is reported that the average concentration of lead in agricultural soil affected by coal mines is relatively high (433 mg kg−1)38. Lead is usually related to minerals in coal and occurs mainly in the form of sulfide such as PbS and PbSe39. In addition, aluminosilicate and carbonate also contain lead40. Chromium is a non-volatile element, which is related to aluminosilicate minerals41. In the mining process, chromium may be accumulated in coal, gangue or other tailings, and then enter the soil or water body through rain leaching42.Table 5 Statistics of contents of heavy metals in wasteland soil (n = 79).Full size tableThe coefficient of variation (CV) of Hg and Cd contents in soil within the research area is 0.050 and 0.37, respectively, with moderate variation, indicating that the content of these two heavy metals is less affected by the external factors; the coefficient of variation (CV) of As, Pb, Cr, Zn, Ni and Cu contents is 2.81, 7.46, 18.00, 13.51, 5.44 and 5.64, respectively, with strong variation (CV  > 0.50)43, indicating that the content of these eight heavy metals may be affected by some local pollution sources. The skewness coefficient (SK) ranges from − 3 to 3, and the larger its absolute value, the greater its skewness. When SK  > 0, it is positive skewness; when SK  More

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    Behaviour dominates impacts

    The impacts of climate change on host–parasite dynamics are particularly complex to predict, as they involve an interplay of both physiological and behavioural factors, from both host and parasite. For example, while warming may increase parasite developmental rates and thus increase transmission, excessive heat may instead exceed thermal limits, leading to higher parasite mortality. Transmission also relates to both the distribution and abundance of host species, which may also shift under changing climates. More

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    Phytoplankton responses to changing temperature and nutrient availability are consistent across the tropical and subtropical Atlantic

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    Ecosystem productivity affected the spatiotemporal disappearance of Neanderthals in Iberia

    Fauna, culture and chronology datasetsA geo-referenced dataset of chronometric dates covering the late MIS 3 (55–30 kyr cal bp) was compiled from the literature (dataset 1). The dataset included 363 radiocarbon, thermoluminescence, optically stimulated luminescence and uranium series dates obtained from 62 archaeological sites and seven palaeontological sites. These chronological determinations were obtained from ten palaeontological levels and 138 archaeological levels. The archaeological levels were culturally attributed to the Mousterian (n = 75), Châtelperronian (n = 6) and Aurignacian (n = 57) technocomplexes. A number of issues can potentially hamper the chronological assessment of Palaeolithic technocomplexes from radiocarbon dates, such as pretreatment protocols that do not remove sufficient contaminants or the quality of the bone collagen extracted. Moreover, discrepancies in cultural attributions or stratigraphic inconsistencies are commonly detected in Palaeolithic archaeology. Information regarding the quality of date determinations and cultural attribution or stratigraphic issues is provided in the Supplementary Information.Our dataset also included the presence of herbivore species recovered from each archaeo-palaeontological site (hereafter referred to as local faunal assemblages (LFAs)), their body masses and their chronology. The mean body mass of both sexes, for each species, was obtained from the PHYLACINE database53 and used in the macroecological modelling approach described below (see ‘Carrying capacity of herbivores’). For visual representation purposes, the herbivore species were grouped into four weight categories: small (500 kg). The chronology of the occurrence of each herbivore species was assumed to be the same as the dated archaeo-palaeontological layer where the species remains were recovered. Thus, to estimate the chronological range of each species in each region, all radiocarbon determinations were calibrated with the IntCal20 calibration curve54 and OxCAL4.2 software55. The BAMs were run to compute the upper and lower chronological boundaries at a CI of 95.4% of each LFA (see ‘Chronological assessment’ for more details). One of the purposes of the current study was to estimate the potential fluctuations in herbivore biomass during the stadial and interstadial periods of the late MIS 3. Accordingly, the time spans of the LFAs were classified into the discrete GS and GI phases provided by Rasmussen et al.51.Geographic settingsThe Iberian Peninsula locates at the southwestern edge of Europe (Fig. 1). It constitutes a large geographic area that exhibits a remarkable diversity of ecosystems, climates and landscapes. Both now and in the past, altitudinal, latitudinal and oceanic gradients affected the conformation of two biogeographical macroregions with different flora and fauna species pools: the Eurosiberian and Mediterranean regions13,46. In the north, along the Pyrenees and Cantabrian strip, the Eurosiberian region is characterized by oceanic influence and mild temperatures in the present day, whereas the Mediterranean region features drier summers and milder winters (Fig. 1). Between the Eurosiberian and Mediterranean regions, there is a transitional area termed Submediterranean or Supramediterranean. Lastly, the Mediterranean region is divided into two distinctive bioclimatic belts: (1) the Thermomediterranean region, located at lower latitudes, with high evapotranspiration rates and affected by its proximity to the coast; and (2) the Mesomediterranean region, with lower temperatures and wetter conditions (Fig. 1).Previous studies have shown that zoocoenosis and phytocenosis differed between these macroregions in the Pleistocene13,46. However, flora and fauna distributions changed during the stadial–interstadial cycles in the Iberian Peninsula, which suggests potential alterations in the boundaries of these biogeographical regions. The modelling approach used in this study to estimate the biomass of primary consumers is dependent on the reconstructed NPP and the herbivore guild structure in each biogeographical region. To test the suitability of the present-day biogeographical demarcations of the Iberian Peninsula during MIS 3, we assessed whether the temporal trends of NPP and the composition of each herbivore palaeocommunity differed between these biogeographical regions during the MUPT.Chouakria and Nagabhusan56 proposed a dissimilarity index to compare time series data by taking into consideration the proximity of values and the temporal correlation of the time series:$${rm{CORT}}(S_1,S_2) = frac{{mathop {sum}nolimits_{i = 1}^{p – 1} {left( {u_{left( {i + 1} right)} – u_i} right)} (v_{(i + 1)} – v_i)}}{{sqrt {mathop {sum}nolimits_{i = 1}^{p – 1} {(u_{(i + 1)} – u_i)^2} } sqrt {mathop {sum}nolimits_{i = 1}^{p – 1} {(v_{(i + 1)} – v)^2} } }}$$
    (1)
    where S1 and S2 are the time series of data, u and v represent the values of S1 and S2, respectively, and p is the length of values of each time series. CORT(S1, S2) belongs to the interval (−1,1). The value CORT(S1, S2) = 1 indicates that in any observed period (ti, ti+1), the values of the sequence S1 and those of S2 increase or decrease at the same rate, whereas CORT = −1 indicates that when S1 increases, S2 decreases or vice versa. Lastly, CORT(S1, S2) = 0 indicates that the observed trends in S1 are independent of those observed in S2. To complement this approach by considering not only the temporal correlation between each pair of time series but also the proximity between the raw values, these authors proposed an adaptive tuning function defined as follows:$$d{rm{CORT}}left( {S_1,S_2} right) = fleft({{rm{CORT}}left( {S_1,S_2} right)} right)times dleft( {S_1,S_2} right)$$
    (2)
    where$$fleft( x right) = frac{2}{{1 + exp left( {k,x} right)}},k ge 0$$
    (3)
    In this study, k was 2, meaning that the behaviour contribution was 76% and the contribution of the proximity between values was 24%57. Hence, f(x) modulates a conventional pairwise raw data distance (d(S1,S2)) according to the observed temporal correlation56. Consequently, dCORT adjusts the degree of similarity between each pair of observations according to the temporal correlation and the proximity between values. This function was used to compare the reconstructed NPP between biogeographical regions during MIS 3 in the Iberian Peninsula. However, two different biogeographical regions could have experienced similar evolutionary trends in their NPP, even though their biota composition was different. Therefore, this analysis was complemented with a JSI to assess whether the reconstructed herbivore species composition in each palaeocommunity differed among biogeographical regions during the late MIS 3. The JSI was based on presence–absence data and was calculated as follows:$${rm{JSI}} = frac{c}{{(a + b + c)}}$$
    (4)
    where c is the number of shared species in both regions and a and b are the numbers of species that were only present in one of the biogeographical regions. Therefore, the higher the value the more similar the palaeocommunities of both regions were.Chronological assessmentPivotal to any hypothesis of Neanderthal replacement patterns by AMHs is the chronology of that population turnover. To this end, we used three different approaches to provide greater confidence in the results: BAMs, the OLE model and SPD of archaeological assemblages. As detailed below, each of these approaches provides complementary information about the MUPT.First, we built a set of BAMs for the Mousterian, Châtelperronian and Aurignacian technocomplexes in each region during the MIS 3. As stated above, we compiled the available radiocarbon dates for Iberia between 55 and 30 kyr cal bp. However, not all dates or levels were included in the Bayesian chronology models. Radiocarbon determinations obtained from shell remains were incorporated in the dataset (dataset 1); however, the local variation of the reservoir age was unknown from 55 to 30 kyr bp. Because of uncertainties related to marine reservoir offsets, all BAMs that incorporated dates from marine shells were run twice: including and excluding these dates. All of the archaeological levels with cultural attribution issues or stratigraphic inconsistencies were excluded. The Supplementary Note provides a detailed description of the sites, levels and dates excluded and their justification. All BAMs were built for each technocomplex using the OxCAL4.2 software55 and IntCal20 calibration curve54.Bayesian chronology models were built for each archaeological and palaeontological level. Then, the dates associated with each technocomplex were grouped within a single phase to determine each culture’s regional appearance or disappearance. Our interest was not focused on the chronological duration of the Mousterian, Châtelperronian and Aurignacian cultures, but on the probability distribution function of the temporal boundaries of these cultures in each region. Thus, this chronological assessment aims to provide an updated chronological frame for Neanderthal replacement by AMHs in Iberia. For this reason, we did not differentiate between proto- and early Aurignacian cultures, since both are attributed to AMHs.In each BAM, we inserted into the same sequence the radiocarbon dates associated with a given technocomplex within a start and end boundary to bracket each culture, which allowed us to determine the probability distribution function for the beginning and end moment of each cultural phase6. The resolution of all models was set at 20 years. We used a t-type outlier model with an initial 5% probability for each determination, but when more than one radiocarbon date was obtained from the same bone remain, we used an s-type outlier model and the combine function. The thermoluminescence dating likelihoods were included in the models, together with their associated 1σ uncertainty ranges. When dates with low agreement ( More

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