Philippa Kaur

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These authors contributed equally: Laura H. Antão, Benjamin Weigel.These authors jointly supervised this work: Tomas Roslin, Anna-Liisa Laine.Research Centre for Ecological Change, Organismal and Evolutionary Biology Research Programme, Faculty of Biological and Environmental Sciences, University of Helsinki, Helsinki, FinlandLaura H. Antão, Benjamin Weigel, Giovanni Strona, Maria Hällfors, Elina Kaarlejärvi, Otso Ovaskainen, Marjo Saastamoinen, Jarno Vanhatalo, Tomas Roslin & Anna-Liisa LaineDepartment of Biological Sciences, University of South Carolina, Columbia, SC, USATad DallasDepartment of Biology, Lund University, Lund, SwedenØystein H. OpedalFinnish Environment Institute (SYKE), Helsinki, FinlandJanne Heliölä, Mikko Kuussaari, Juha Pöyry & Kristiina VuorioNatural Resources Institute Finland (Luke), Helsinki, FinlandHeikki Henttonen, Otso Huitu, Andreas Lindén, Päivi Merilä, Maija Salemaa & Tiina TonteriSection of Ecology, Department of Biology, University of Turku, Turku, FinlandErkki KorpimäkiFinnish Museum of Natural History, University of Helsinki, Helsinki, FinlandAleksi LehikoinenKainuu Centre for Economic Development, Transport and the Environment, Kajaani, FinlandReima LeinonenUniversity of Helsinki, Helsinki, FinlandHannu PietiäinenDepartment of Biological and Environmental Science, University of Jyväskylä, Jyväskylä, FinlandOtso OvaskainenCentre for Biodiversity Dynamics, Department of Biology, Norwegian University of Science and Technology, Trondheim, NorwayOtso OvaskainenHelsinki Institute of Life Science, University of Helsinki, Helsinki, FinlandMarjo SaastamoinenDepartment of Mathematics and Statistics, Faculty of Science, University of Helsinki, Helsinki, FinlandJarno VanhataloSpatial Foodweb Ecology Group, Department of Agricultural Sciences, University of Helsinki, Helsinki, FinlandTomas RoslinSpatial Foodweb Ecology Group, Department of Ecology, Swedish University of Agricultural Sciences, Uppsala, SwedenTomas RoslinDepartment of Evolutionary Biology and Environmental Studies, University of Zürich, Zürich, SwitzerlandAnna-Liisa Laine More

PTM concentrationsIn Table 1, the descriptive statistics of PTMs in 50 dust samples from Qom city are described. The background values were used based on the concentrations of metals in the Upper Continental Crust. The mean concentration of As, Cd, Cu, Mo, Pb, Sb, and, Zn exceeded the background value. Also, Cd, Cu, Mo, Pb, Sb, and, Zn had a coefficient of variation (C.V.) greater than 50%, indicating a severe variability in PTMs concentrations in the atmospheric dust of the studied area2. Metals with C.V.  Pb  > Zn  > As  > Cd  > Cu  > Mo  > Cr  > Mn  > Ni = Co. Antimony (38.55) and Pb (35.13) had the highest average EF values, which means they were enriched very high in the windowsill dust. Also, they had a wide range of EF values in the 50 stations: from 4.0 to 227.0 for Sb, and from 8.3 to 140.8 for Pb which might reflect the existence of discrete multiple sources in the studied area. The degree of enrichment for Pb and Sb in the industrial sector was extreme and in the commercial sector was very high; also, the other sectors were significantly enriched. Zinc and As had a more homogenous enrichment in the area. In all the functional sectors, 95% and 84% of stations were significantly enriched by As and Zn, respectively. Copper, Cd, and Mo were moderately enriched in all functional sectors, but the greenspace sector had minimal enrichment by these elements. Some areas in the industrial sector had significant to very high enrichment of Cd. The EF value indicated Co, Cr, Mn, and Ni were minimally enriched in all the stations.Figure 2Box plot of the (a) enrichment factor (EF), and (b) geo accumulation index (Igeo) for the dust samples in the studied area.Full size imageThe highest average values of Igeo were obtained in the order of Pb  > Sb  > As  > Zn. PTMs included Co, Cr, Ni, Fe, and Mn were categorized as unpolluted and Cd, Cu, and Mo were in the category of unpolluted to moderately polluted. In the industrial zone, the windowsill dust was extremely polluted with Sb and Pb. The sequence of contamination intensity with Pb, Zn and Sb according to land use was: industrial  > commercial  > residential  > greenspace. The highest concentration of arsenic in the study area belongs to the industrial area.To evaluate the pollution level based on land use, PLI and mCd indices were utilized (Fig. 3). These cumulative indices showed that the dust in Qom city is considerably contaminated with PTMs. According to the PLI index, all the stations were categorized as polluted sites. The PLIzone values were in the order of industrial (3.77)  > commercial (2.05)  > residential (1.67)  > green space (1.38). This pattern was also repeated with the mCd index. The mCd for the industrial sector ranged from 6.98 (high contamination level) to 39.60 (ultra-high contamination level). In the commercial sector, fifty percent of dust samples were classified as having a high degree of contamination. All the greenspace stations were in the moderate pollution category. This shows the possible effect of tree density in diminishing the risk of dust pollution to the receptors.Figure 3Pollution level indexes (a) mCd and (b) PLI, based on four functional areas.Full size imageSpatial distribution of PTMsThe As, Cd, Cu, Sb, Pb, and Zn content in 100% of the dust samples exceeded the background value. Spatial distribution maps were generated for the hotspot PTMs (As, Sb, Pb, Cd, Cu, Mo and Zn) by applying the inverse distance weighted (IDW) interpolation method (ArcGIS 10.3). Figure 4 demonstrates that PTMs dispersions were slightly influenced by the prevailing wind direction (from the west), suggesting they came from the point- or area- sources. On the other hand, the K–S test showed that the overall distribution of PTMs was not normal in the studied region. This might signify the influence of industrial activities and the presence of multiple sources of dust.Figure 4Spatial distribution maps of seven PTMs in windowsill dusts of Qom, Iran. This map was constructed using ArcGIS version 10.3. (https://www.esri.com/en-us/arcgis/products/arcgisdesktop/overview).Full size imageThe highest pollution load of PTMs belonged to the industrial section. The level of pollution gradually decreased from Shokouhieh to Mahmoudabad industrial zones. The reason is related to more active industries, a closed environment, and more construction existing in Shokouhieh industrial town than in Mahmoudabad industrial town.There is a clear decreasing trend from the central part to southern (downtown area) and southwestern (suburb area) parts of the city. In fact, these parts are diffusely populated and the southwestern part is almost new with lots of barren lands. Copper, Mo, and Cd show high concentrations toward the central part of the city. Educational, cultural and commercial activities are mainly located in the central part of the city. Also, historical and religious districts in the city center are accompanied by a huge influx of tourists throughout the year. For this reason, the central part of the city has various public transportations such as bus stands and taxi stations, and is dominated by a high load of motorcycles.In the eastern part of the city, some hotspots can be observed (Fig. 4). This part includes an important transportation system (like highways and a complex interchange) where exhaust traffic emissions might be a probable source of As, Sb, Pb, and Zn. Unlike Pb and Zn, several peaks of As are scattered in the western part, suggesting an area source might exist in the region. It is noteworthy that the western area is densely populated with lots of residential buildings. Bisht et al. (2022)35 also observed hotspots of As in the residential area of Dehradun, India.PTM potential sourcesTo evaluate the relationship between PTMs in dust samples, the Spearman correlation and PCA were developed (Fig. 5) and more details are given in Table S7. Statistical analysis can help to identify the potential source of contamination in urban dust. The Spearman correlation was significant at p  residential ≈ greenspace. The five PTMs with the highest overall HI are ranked as follows: Pb  > As  > Cr  > Mn  > Sb (Fig. 7). The HI values in all the sections were lower than the permissible level (1.00), except for Pb. In the industrial section, Pb recorded the highest HI value for children (HI = 1.73) which exceeded the acceptable value. The HI values were 10 times higher for children than adults indicating they are more susceptible to PTMs in the dust.The dominant pathway for noncancerous risk was ingestion followed by dermal contact and inhalation. The trend is in line with previous research25,51,52. However, for Co and Mn, the descending order was different as follows ingestion  > inhalation  > dermal contact. The highest contribution of HQinh and HQderm to HI was measured for Co (34.0%) and Cd (29.0%), respectively.In this study, the carcinogenic risk from windowsill dust was estimated for the carcinogens including Cd, Co, Cr and Ni, Pb, and As through the possible routes (Fig. 8, Table S9). The contribution of PTMs to CR decreased in the order of Cr (3.24E−05)  > As (2.05E−05)  > Pb (2.52E−06)  > Co (6.91E−09)  > Ni (1.72E−09)  > Cd (2.58E−10). The average CR values for target PTMs through inhalation ranged from 7.9E−10 to 1.7E−07, which remained in the safety zone (CR  inhalation  > dermal (Fig. 8). While the contribution of Cr to carcinogenic risk was higher through inhalation than ingestion. The reports concluded that the primary exposure route of Cr is inhalation54. Considering the predominant forms of Cr in the environment, CrVI is more toxic than CrIII. Exposure to CrVI can cause immunological diseases, dental effects and carcinogenic effects (lung cancer, nose and nasal sinus cancer, suspected laryngeal and stomach cancers)54,55.The result of health risk from target PTMs in windowsills of Qom indicates significant chronic exposure to Pb can take place for children in the industrial zone. The ingestion route is the most probable pathway for children due to their hand–to–mouth behavior56. Lead can bio-accumulate in the body without any obvious symptoms of toxicity56. The total CR values for Pb, Cr and As in different land-use types were in the range of tolerable carcinogenic risk (1 × 10−4  More

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Notwithstanding constraints on the amount of hard data, according to our integrated analysis, the developmental environment and ecology of reef communities have an important impact on the appearance of reefs.Analysis of environmental conditions for reef developmentSettings of reef developmentThe F/F extinction event in Late Devonian caused the complete recession of the reef-building communities based on stromatoporoid-coral assemblages7,17. The Carboniferous is generally considered to be a sub-optimal period for the development of framed reefs. After the biological mass extinction, microorganisms and algae rebuilt new reef-building ecosystems18,19. Some short-term biological frame reefs developed with low diversity, limited reef-building organisms, small sizes, and restricted distribution20. Harsh climate and marine conditions occurred in the Mississippian, including extensive marine hypoxia, repeated glacial and interglacial climate changes, and frequent changes of sea level and seawater surface temperature, potentially hindering the recovery of Early Carboniferous metazoan reefs7,21.Metazoans gradually began to participate in reef building in Early Viséan. A large number of biogenic structures formed by corals and bryozoans began to appear, including a small number of sponge reefs/mounds in the middle and late stage of Viséan. The richness and biodiversity of the Mississippian post-zoobenthic reefs flourished in the late Viséan during which corals, bryozoans, sponges, calcareous microorganisms, and some calcareous algae became the main builders3 and large-scale reefs could also be seen in some areas although most of the Viséan metazoan reefs were tabular or laminar. Thus, the metazoan skeletal reefs in the middle to late Viséan were considered to have been resurrected due to relatively warm climatic conditions and higher sea levels after a period of complete disintegration at the end of the Devonian and recession at the beginning of the Carboniferous7.Consequently, the coral reefs in the study area were the products of shallow benthic communities thriving in relatively favourable conditions of Late Viséan-Serpukhovian, which was common for reef development at that time7. Thus, it is expected that more synchronous reefs would to be identified in southern China, or even in the study area in the future.Paleogeography of reefsLangping is located in Dian-Qian-Gui Basin22 regionally (Fig. 2), in the eastern end of Tethys tectonic domain and at the interjunction of Tethys and Pacific structure globally. The Carboniferous Dian-Qian-Gui Basin is adjacent to the Tethys Basin. During the Early Carboniferous, the continent of Gondwana was close to the equator but was separated from the northern continent by the Tethys, where the tropical currents flowed freely from east to west. The benthic warm-water organisms were distributed widely with high abundance and diversity on both sides of the shallow shelf of Tethys.Figure 2Paleogeographic map of southern China in Viséan-Serpukhovian (modified from Feng23, Yao8, and Maillet24). This figure was obtained from articles by Feng23, Yao8 and Maillet24 respectively. The author modified the picture with CorelDRAW (version 2022, and the URL link: https://www.coreldraw.com/cn/). QG Qian-Gui Basin, DQGX Dian-Qian-Gui-Xiang platform.Full size imageViséan-Serpukhovian ecosystems experienced dramatic climate changes and widespread glaciation25. However, the Viséan was also a key layer for a variety of biological structures, with abundant coral reefs and a high diversity of shallow benthic communities, peaking in the late Viséan. Newly discovered post-faunal reefs in Tianlin were mainly formed in the late Viséan-Serpukhovian period, which coincided with frequent sea level fluctuations and possible glacial changes. It seems counterintuitive that tropical coral communities developed during glacial period. However, recent studies suggest that the persistent warm ocean currents on the platform helped some coral species survive from Carboniferous glacial events24. While other areas of symbiotic reefs were poorly developed, Tianlin may provide an ecological sanctuary for corals associated with ocean currents26.Sedimentation of reef developmentAccording to the regional geological structure, the slope model for Langping paleocarbonate platform was obviously different from that of steep slope platform margin, which could be directly affected by waves. Langping palaeo-platform could be regarded as one of the small blocks (block fault barrier) separated from a large platform (continental margin sea basin)13. The relative positions of these blocks were crucial for the emergence and growth of reefs.In situ development of mud-crystalline tuffs and muddy tuffs with weak hydrodynamic conditions is common in the Langping area, and evaporites are poorly developed. There were patch reefs and reef layers in different sizes in the wide intraclast beach, where obviously developed reef beach complexes were rare. The fragments of carbonate base broken by storm in the clastic beach haven’t been observed. The study area is considered to be gentle-slope open platform27,28 based on sedimentary characteristics. It suggests that the study area was far away from the margin of steep-slope platform that directly affected by waves, and more consistent with less energetic internal environments of gentle-slope platform.On the vast platform of Langping gentle slop, deep water lead to low water energy. While in the coastal area, the water energy was relatively strong, thus coarse-grained bioclastic beach and a small amount of point reef could be developed. The beaches were irregular-shaped due to long term transportation and reformation effects of waves and water flow, showing low and gentle slope angles. Dispersed reef-beach complexes at the platform margin slightly impacted inner-platform seawater and the water flows smoothly29.Therefore, it can be assumed that Langping reefs developed along the intertidal shallows of the terrace. The seawater around Langping carbonate platform in Late Viséan-Serpukhovian was relatively shallow while the water flow was strong. Remains of crinoids, brachiopods, a few foraminifera, and solitary corals were likely broken by strong currents, and deposited in situ with a small amount of gravels and lime-mud (Fig. 3). The clastic beach was unstable, suggesting large-scale wave-resistant structures could not be formed quickly30 due to insufficient cohesive and consolidating organisms. In addition, the circumferential impact of water in extensive terraces leads to mud-lime deposition, which is detrimental to most benthic organisms. However, bondstone was more likely to be formed by some binding algaes in the platform (Fig. 4). Therefore, neither the surrounding or the inner region of the platform could provide favourable living conditions for coral reefs to develop over for a long term. The gently sloping terrace environment of Lanping resulted in significant differences in growth size, wave resistance and reef-building capacity between corals in the study area and those on the edge of the steeply sloping terrace.Figure 3Clastic beaches in the Langping. Various clastic beaches developed in the study area. Diverse composition, fragmentation degree and sorting of the clast indicate different water conditions of formation. This figure is modified by the author from field photos with CorelDRAW (version 2022, and the URL link: https://www.coreldraw.com/cn/).Full size imageFigure 4Algal bondstone in the Langping. Bondstone formed by various algaes living in still water. Morphology of bondstone correlates water environment and deposition of mud. Vast algal bondstones indicate deep water and high deposition rate. This figure is modified by the author from field photos with CorelDRAW (version 2022, and the URL link: https://www.coreldraw.com/cn/).Full size imageAt the same time, the warm climate of the late Viséan-Serpukhovian, the good circulation of seawater around the Langping platform, and the abundant supply of oxygen and nutrients were a series of favourable conditions that facilitated the growth of reef-building corals, which led to uplifts being formed on clastic beach, including patch reefs and reef layers with certain sizes. These uplifts impeded waves and provided a protected nearshore environment, though they were much smaller than those developed at the steep-slope platform margin. The inhabitants on the beach could not resist strong waves. These rises were therefore known as reef-beach complexes and could only persist where waves and currents were mild28. They were essentially different from the framework coral reefs which developed on steep-slope platform margin that reflected changed hydrodynamic conditions, nutrient sources, reef sizes, and growth rates.Another potentially favourable factor in the study area could be the deeper water area in the gentle-slope sedimentary environment, which could provide more stable conditions and reduce the damaging effects of global glacial events and large scale sea level fluctuations on reef-building communities25. The frequent fault activities in Dian-Qian-Gui Basin caused the rise and fall of equivalent sea level. More influence of sea-level fluctuations and hydrodynamic conditions would be exerted on Langping platform due to its small size. Furthermore, reef growth promoted by reef-building communities would be frequently disturbed. The sediments displaying evidence of multicycle sedimentation, different components, and diversely fragile clasts in the study area provided direct evidence of frequently changing environment.Alternatively, the sedimentary environment of Langping platform provided conditions favorable for reef-building communities to develop and reefs to grow rapidly. These factors directly or indirectly determined the ecology of reef-building communities and the general appearance of reef development in the study area.Overall, the environmental factor is the primary factor affecting the overall development trend of reefs.Inferred ecological characteristics of reef communitiesResponse of reef-building corals to hydrodynamic conditionsHydrodynamic conditions are very important factors for reef development, which directly determine the abundance and distribution of each reef-building population and are key factors influencing sedimentation and reef growth, and was particularly evident at Langping. Evidence from the fossils suggested the reef-building corals were also changed in response (Table 1). The hydrodynamic condition changes during the development of reefs are inferred based on analysis of the vertical sediments and microfacies changes of coral reefs in the study area31. How these ancient reef-building corals adapted to hydrodynamic conditions was reconstructed combining the evolution of reef-building communities with the study.Table.1 General situation of reef-building coral population in Langping.Full size tableThe Xiadong coral reef started with colonization and expansion of Diphyphyllum on the bioclastic hard substrates32,33. They grew vertically into upright clusters (Fig. 5A) and were insensitive to more sediment in a relatively calm, turbid water environment34. The relatively dense clumped Siphonodendron and massive Lithostrotion (Fig. 5B) were better suited to the turbulent water environment, becoming dominant over time, with Diphyphyllum subordinate with the continuous increase of the water energy, as indicated by the characteristics of sediment particles from fine to coarse. After flourishing for a period, the Siphonodendron–Lithostrotion assemblage eventually waned, likely due to the failure to adapt to the increasing hydrodynamic conditions. Diphyphyllum had persisted combined with Syringopora, to maintain the growth of the reef. However, this assemblage subsequently declined as a result of strong hydrodynamic conditions and finally died out in response to continuous falling of sea level. Consequently, the reefs stopped developing.Figure 5Sketch of coral cluster with upright growing morphology. Most reef-building corals in Langping grow vertically into cluster colonies. This type of morphology is very favourable for corals to get more living space and is important to reef-building. (A) Cluster coral individuals grow uprightly with certain distance between each other. (B) Polygonal columnar coral individuals grow closely to resist strong water flow. This figure is made by the author with CorelDRAW (version 2022, and the URL link: https://www.coreldraw.com/cn/).Full size imageThe Longjiangdong multi-layer reef was composed of three relatively independent, flat reef layers, suggesting three distinct periods of reef development. Diverse species were identified in the reef, with colonial coral Diphyphyllum contributing greatly to reef growth. Diphyphyllum clusters colonized in patchy form on substrates composed of bioclasts or lithic gravels (Fig. 6A). The first reef-building process was brief, ending under high-energy water conditions after a period of growing (Fig. 6B). Subsequently the hydrodynamic conditions became weaker and favorable. Then Diphyphyllum once again flourished. Diphyphyllum clumps in the unit grew closely together in strong currents, with larger and more sparse individuals than in the lower units. A relatively low energy environment was formed between the Diphyphyllum clusters (Fig. 6C). Subsequently, Diphyphyllum could only grow in a limited area of suitability due to the disturbance of high-energy water brought about by short-term sea-level rise and fall. Afterwards, the environment became more favourable and Diphyphyllum expanded rapidly. As a result, the upper unit of Longjiangdong coral reef was formed, in which Diphyphyllum individuals were slightly larger than those in the first two units. Finally, because the kinetic energy of the water continued to weaken, the plaster deposition forced the whole coral reef to stop growing (Fig. 6D).Figure 6Micrographs of sediments in different positions of reef. (A) Calcareous bioclastic limestone, with biological particles accounting for about 70% of the debris. Abundant and diverse organisms indicate a medium-energy environment of the subtidal zone. Samples were taken from the bioclastic beach at the reef base. (B) Slightly larger bioclastics but lower biologic content than that in (A) suggest an increasing water energy. (C) Various bioclastic particles account for about 80% of the clastic particles contained in the calcareous bio-granular rock. The obviously small benthos indicate a low-energy environment in the subtidal zone barriered by the Diphyphyllum clusters. (D) Bioclastic grainstone is mainly composed of marl, with fine clastic particles (about 30%) and bedding. Low biomass indicates a low-energy environment of the subtidal zone. This figure is modified by the author with CorelDRAW (version 2022, and the URL link: https://www.coreldraw.com/cn/). Meaning of the letters in the figure: C crinoids, BF brachiopods, F foraminifera, B bryozoan, P pelletoid, MF mollusk shell fragment.Full size imageLongjiangdong patch reef started to develop in a relatively deep water environment. Diphyphyllum initially colonized and expanded in favorable conditions with the increase of water energy. Then the reef-builders transitioned from a single coral species to an assemblage of Diphyphyllum–Caninia–Lithostrotionella. These three coral species grew independently and contribute almost equally to the structure of the reef. However, the structure and function of the coral community were not yet stable enough. It was easily influenced by the weakening hydrodynamics and the increasing sedimentation, resulting in only small patch reefs.The Xinzhai layer reef was initialized by colonization and expansion of Lonsdaleia on bioclastic beach. Large coral clusters were formed in the presence of turbulent water. With the weakening of hydrodynamic conditions, an unknown branchlike organism and Antheria communities continued to develop separately in this area. Slender branchlike organisms expanded rapidly in these low-energy water environments until they were replaced by some individual corals as hydrodynamic energy increased. Each builder was short-lived in this layer reef, departing from the reef just at the beginning of colonization and expansion, due to rapidly changed hydrodynamic conditions.The evolution of reef-building corals in these four reefs indicated that both the coral assemblages and coral individuals would constantly adapt to the changing hydrodynamic conditions in Langping as sea level rose and fell. Although this was a reactive adjustment of coral populations in response to long-term environmental impacts, it was clearly positive for the building and development of coral reefs.Impact of disturbance on reef communitiesDisturbance is a relatively discontinuous event, which is ubiquitous in nature. It may indirectly affect the composition and population structure of reef communities by changing the environmental conditions, thus affect the structure and function of reef communities, even the evolution of the reef35. The major disturbances evident in these Mississippian framework reefs were associated with frequent changes of water flow, and drastic changes of climate and weather. These seem to be most obvious in the Langping platform due to its small size, with more frequent environmental influence evident on the reef communities in the study area.The most direct effect of disturbance events on coral reefs is the disruption of continuously evolving reef communities, which is common in coral reef studies. After the interruption caused by disturbances, some communities gradually recover due to the absence of continuous disturbance, or the dominant biota may be substituted by invading communities. The winner after interruption is decided by random factors to a large extent, in a ‘Competitive lottery’36. The conditions for the emergence of ‘Competitive lottery’ also include the need for species in a community to have similar abilities to invade discontinuities and to tolerate environmental conditions.Certainly, low-intensity disturbance does not necessarily produce discontinuity, but medium-intensity disturbance without discontinuity could directly impact on community species diversity. According to the ‘Moderate disturbance hypothesis’, moderate disturbance is conducive to a higher level of community diversity37. In environmental conditions with moderate intensity of disturbance, most species will not disappear entirely. The dominant pioneer species will also be restrained by disturbance to a certain extent, so large number of species can coexist, attaining the highest diversity35.The reef-builders in Langping are diverse compared with the Late Carboniferous reefs in Ziyun County10, which also developed in Dian-Qian-Gui Basin. More than 4 reef-building corals are identified in Xiadong reef, while 4 and 3 are in Xinzhai layer reef, Longjiangdong patch reef respectively. These reef-building corals, mostly Diphyphyllum, Lithostrotion, Siphonodendron and Lonsdaleia, were distributed irregularly in the reefs. Their ecological niche and function were likely similar and none of them was obviously dominant in the community (Fig. 7). This is in line with ‘Competitive lottery’ theory and the ‘Moderate disturbance hypothesis’.Figure 7Different species occupied the discontinuity surface irregularly. (A) Different reef-builders colonized and grew on the same hard substrate. (B) and (C) show detailed morphology of colony corals of (A). (D) Colony corals and a large number of individual corals grew together in a limited area, indicating equal colonization on the newly formed discontinuity surface. This figure is modified by the author from field photos with CorelDRAW (version 2022, and the URL link: https://www.coreldraw.com/cn/).Full size imageThe stability of a classical reef ecosystem includes the ability to withstand external disturbances and the ability to return to its original state once the disturbance is removed37,38. It is generally accepted that communities with high diversity are always more stable although ecosystem stability is not absolutely correlated to biodiversity35.There have been no reef-building corals with strong resistance and rapid recovery ability in the communities in Langping. None of these corals succeeded in developing into dominant species that can build reef shelves, which made the reefs in Langping mostly appear in the form of small patch reefs or reef layers. However, formation of the large reef in Xiadong Village, patch reef in Longjiangdong Village, and layer reef in Xinzhai Village were all related to their relatively high diversity of reef-building corals. Compared with the situation where only one reef-building organism dominated the Bianping large coral reef, Wengdao large phylloid algal reef and Ivanovia cf. manchurica patch reefs in Ziyun County10, Guizhou province, the different coral assemblages in Langping area could effectively adapt to changing hydrodynamic conditions and maintain reef growth.Species diversity increased by disturbance stabilized the ecosystemas shown during the construction of coral reefs in Langping.Effects of non-reef-builders on reef-building coralsBesides reef-building corals, there were a large number of reef-dwellers and off-reef organisms in the study area. Reef-dwellers referred to the species that didn’t directly contribute to reef growth in the community, mainly including various benthos and algaes39. Off-reef organisms are not part of the reef-building community, but also play an important role in participating in energy flow and providing organic matter to the reef ecosystem40.Common reef-dwelling organisms include crinoids, brachiopods, gastropods, various algae, foraminifera, bryozoans and individual corals. Crinoids were overwhelmingly dominant in numbers in the reef samples studied here.Carboniferous echinodermata in Guangxi Province reached its peak in Middle-to-Late Mississippian. In terms of amount and distribution, thick limestone with echinodermata debris in the carbonate platform were often dominated by crinoids41,42,43,44,45,46. The large number of crinoids in Langping excluded other metazoans and restricted the development of benthic reef-builders in Late Viséan-Serpukhovian in Langping, leading to poorly developed reef-building communities.Microorganisms and algaes had limited success in stablishing on the moving clastic beach in frequently disturbed water. There has not been obvious evidence of extensive “algal turf” in the coastal area of Langping platform. Only a few corals bonded by algal mats were observed47 (Fig. 8). In addition to their significant contribution to primary productivity, macroalgae were considered to play an important role in two aspects of coral reef ecosystems. One was to promote reef construction by its own binding and consolidation48,49. The other was to create a good condition for zoobenthos larvae to dwell and develop, thereby improving species diversity50. The limited productivity of algae in Langping constrained coral reef trophic inputs, which may then have limited populations of dependent metazoans. As a result, algaes and other metazoans were unable to achieve a variety of reef-building patterns, such as bonding, bounding, entanglement51,52. The reef framework in the study area was not stable in the presence of strong water flow, and the biological communities could not deal with frequent environmental changes, which were directly related to poor development of calcareous algae.Figure 8Micrographs of microbes and algaes. (A) Encrustations (indicated by black arrows) with distinct thickness around coral clusters formed by microbe and algal mats through bonding mud. The encrustations were formed before the clastic deposition (indicated by white arrows), showing the corals were living then. Microbes and algaes inside of the dense coral clusters had little impact on corals. (B) Single polarized micrograph showed clear and smooth boundaries of coral individuals without encrustation or drilling hole made by microbes or algaes. Few corals surrounded by bonding algaes could be observed in Langping, indicating that algaes were poorly developed between coral clusters. This figure is modified by the author from field photos with CorelDRAW (version 2022, and the URL link: https://www.coreldraw.com/cn/).Full size imageInfluence of coral morphology on reef developmentThe accumulation of reef structure had obvious impact on communities. Large reef structures could support abundance and diverse biota by modifying local environments and creating diverse conditions. Consequently the reef-building communities thrived between disturbances, stabilizing reef construction. In terms of large reef, the framework-building corals would play a key role in reef construction regardless of which kind of patterns was adopted. Therefore, reef-building corals with large size, rapid growth vertically, and strong resistance would become the biggest contributors to reef frame construction.The main reef-building corals in Langping were composed of Diphyphyllum, Lithostrotion, Siphonodendron, and Lonsdaleia, etc., being the dominant builders. These corals were similar in morphology such as cluster colony, thick and strong skeleton, and densely packed individuals (Fig. 9), which enabled them to resist water flow. At the same time, the upright colonies were adaptable to relatively calm water, being insensitive to mud deposition. The ecological characteristics of the Langping corals matched the gently sloping environment, the deep water environment and the rapidly changing energy of the currents. These cluster corals were able to colonize hard substrates and expand rapidly, thus altering the surrounding environment. The visible carbonate uplifts were formed with a large amount of benthos grouped into reef-building communities. These distinct uplifts constructed by coral clusters in different water conditions are composed of coral reefs of different sizes and appearance in the study area.Figure 9Main reef-building corals in the study area. (A) Diphyphyllum, (B) Lithostrotion, (C) Siphonodendron, (D) Lonsdaleia. (A) Rapidly grew clusters of main reef-building corals. The strong individuals are packed tightly when growing to support each other. This figure is modified by the author from field photos with CorelDRAW (version 2022, and the URL link: https://www.coreldraw.com/cn/).Full size imageThe complex and diverse local environments formed by large coral reefs can significantly increase benthic populations and improve reef species diversity. As a result, the nutrient flow in the community becomes complicated, and nutrients could be recycled effectively by reducing loss caused by water flow. Therefore, the overall productivity of large coral reef communities was always high. Complex trophic structure satisfied most of the benthos in the community with sufficient nutrients and inorganic salts.The morphology of reef-building corals in Langping enabled them to become predominant species in various water environments, which promoted the continued domed growth of coral reefs and facilitates the development of reef-building communities that form a variety of reefs. It suggests that the morphology of reef-building corals was a key prerequisite for reef development.In conclusion, coral reef communities are always constrained and influenced by environmental conditions. However, the ecology of the inhabitants is also an important factor in the formation of coral reefs. More