"Palaeoecology of Middle Palaeozoic "heliolitids" (Tabulata) and their convergence with Coenothecalia" - a project of the National Science Centre of Poland
Principal investigator. Scope of work: field work; laboratory work, palaeontological and sedimentological investigations; interpretation of research results; synthesis of research results.
Co-investigator. Scope of work: field work; palaeontological, biological and regional geology consultation.
Description
Fig. 1. A, B - Heliolites porosus (Heliolitina, Devonian). A - surface of the corallum; B - a transverse thin section; C - Heliopora coerulea (Coenothecalia, Recent), surface of the corallum. The analogous structure of the skeleton comprised of auto- and siphonopores in both species is visible.
"Heliolitids" are a group of colonial corals which died out at the end of Middle Devonian, c.a.
380 million years ago. They belong to tabulates (superorder Tabulata), a group which went extinct by
the end of Paleozoic, and its evolutionary legacy most likely disappeared. "Heliolitids" are unique
among them, because their skeletons share many similarities with some modern corals, especially the
so-called "blue coral", the species Heliopora coerulea, an octocoral (order Coenothecalia) commonly
occurring on the Pacific reefs. Even though the affinity between them is very remote, their skeletons
have a very similar, seemingly identical architecture. It is probably an effect of convergence -
developing similar structures by different organisms as a result of adaptation to similar environments.
The aim of the proposed research is to determine the environmental constraints on life and
development of "heliolitids", as well as to find out how fare does the convergence between them and
their analogues among the Coenothecalia (such as the blue coral) reach. An additional objective of the
project will be the analysis of the distribution of "heliolitids" in the Middle Devonian Rheic ocean. It
will help reconstruct the configuration of this ocean, whose paleogeography remains a matter of
discussion.
In order to achieve that, field works will be undertaken in the Silurian and Givetian outcrops in
Poland, Czech Republic and Gotland. The gathered coral fossils will be assigned to the lowest possible
taxonomic unit (e.g. species). Then their growth forms and the history of colony development will be
analysed. Additionally, a microscopic analysis will be performed, of the rocks from which the fossils
came. It will allow the determination of such parameters of the environment, in which the corals lived,
as water dynamics, depth and the inventory of associatied organisms. The results of the
abovementioned studies will be compared with the data on modern corals belonging to Coenothecalia,
which live on the reefs of Ryukyu Islands in the northern Pacific.
Determining the environments of life and adaptations of the "heliolitids" will allow for a
comparison of these long extinct organisms to modern, endangered species of corals. Thanks to that, it
will be possible to determine, how far does the convergence between them reach, which in turn will
shed a new light on the adaptation processes of corals from very remotely related groups, living in
analogous environments of life.
Differentiation of heliolitid growth forms. Scale bar = 1cm.
Aims of the project
Analysis of heliolitid life strategies and environmental adaptations
Comparison of heliolitid paleoecology and environmental preferences of present-day octocorals of the genus Heliopora
Evaluating which traits of heliolitid corals were controlled extrinsically (environmentally) and intrinsically (genetically); observation of evolutionary trends
Reconstruction of paleogeographic distribution of heliolitid corals within the Rheic domain
Research areas
Sweden
Sweden - Gotland; Silurian
Belgium
South Belgium - synklinorium Dinant; Devonian
Poland
Poland - Grzegorzowice-Skały, Dziewki; Devonian
Publications
Heliolitid corals and their competitors: a case study from the Wellin patch reefs, Middle Devonian, Belgium
May 2021
Jan J. Król, Julien Denayer, Paweł Wolniewicz, Mikołaj K. Zapalski
Abstract:
Wellin patch reefs are small Upper Eifelian build-ups within the fine-grained argillaceous limestone of the Hanonet Formation. Whereas the reefs themselves are not well exposed, their fossil assemblage is accessible in the hills near the town of Wellin, approximately 40 km SE of Dinant in Belgium. It is especially rich in massive stromatoporoids, heliolitids and other tabulate corals. They exhibit predominantly domical and bulbous morphologies. This paper focuses primarily on the palaeoautoecology of the heliolitid corals and their relationships with other organisms. Cases of mutual overgrowth between heliolitids, other corals and stromatoporids suggest a high degree of competition for space on the reefs, possibly related to the scarcity of hard substrates. Coral and stromatoporoid growth forms, as well as the prevalence of micritic matrix, point to a relatively low energy environment. However, abundant growth interruption surfaces, sediment intercalations and rejuvenations of corals suggest episodically increased hydrodynamic regime and sediment supply. It is inferred that the patch reefs developed in a relatively shallow environment, where the reefal assemblage was regularly affected by storms. Heliolitids exhibited high sediment tolerance and relied on passive sediment removal for survival. They also could regenerate effectively and commonly overgrew their epibionts, after the colony’s growth was hampered by the sediment. This is recorded in extremely abundant growth interruption surfaces, which allow the analysis of the impact of sediment influxes on the heliolitid corals.
A record of overgrowth competition between a Heliolites porosus colony and a stromatoporoid in a thin section photograph. Sediment inclusions associated with the growth interruption surfaces are indicated with arrows. Scale bar = 5 mm.
A sequence of encrustations and overgrowths of Heliolites porosus and corals Platyaxum sp. (p), Temnophyllum? sp. (te), and sponges Taleastroma sp. (ta) and Clathrocoilona sp. (ca). A thin section, scale bar = 5 mm.
Heliolites porosus settled on a stromatoporoid Clathrocoilona sp. (ca) and a coral Alveolites sp. (a), encrusted by Actinostroma sp. (ac) and a cheatetid (ch). A polished slab, scale bar = 1 cm.
A domical colony of Heliolites porosus rotated and rejuvenated. The arrow indicates a bioclaustration. A polished slab, scale bar = 1 cm.
Types 1-4 of growth interruption surfaces, as defined by Miller & West (1997), with the proportional abundance in the studied material.
Deducing photosymbiosis in extinct heliolitid corals
January 2024
Jan J. Król, Błażej Berkowski, Julien Denayer, Mikołaj K. Zapalski
Abstract:
The matter of photosymbiosis in Paleozoic corals remains unresolved as it is not possible to directly check for the presence of algal symbionts in fossil corals. However, present-day photosymbiotic corals are characterised by a number of features that can be evaluated in fossil corals as well, such as large, highly integrated colonies, growth banding, and platy growth forms in mesophotic conditions. The present study aims to evaluate these features in heliolitid corals. Heliolitids were relatively highly integrated, compared to other Paleozoic corals and could produce large colonies, over 1 m in diameter. In this study, heliolitid corals from different outcrops from the Silurian (~444 to 419 ma) of Gotland (Sweden) were analysed, and additional Devonian (~419 to 359 ma) specimens from Belgium, Poland and Morocco, featuring cyclic growth banding, were also included. Overall, over 60% of studied specimens from Lower Visby Formation in Ireviken and over 80% from Eke Formation in Lau Käldu are platy or tabular. Those outcrops represent environments that were mesophotic. Specimens from the shallower Upper Visby Formation in Ygne are dominated by branching forms instead. The apparent light-related skeletal plasticity, high colony integration, growth banding, and the absence of heliolitids in deep water environments implies that some of these corals were most likely photosymbiotic.
Colony surfaces of the studied heliolitid species from Gotland. A - Stelliporella parvistella, B - Propora tubulata, C - Heliolites interstinctus, D - H. daintreei, E - H. megastoma, F - H. spongodes. Scale bars are 2 mm.
Platy heliolitids. A-B - Heliolites interstinctus,C-D - Stelliporella parvistella. Scale bars are 1 cm.
Irregular colony of Stelliporella parvistella. A - side view, B - top view. Scale bars are 1 cm.
A polished slab of Heliolites porosus with cyclic growth banding.
Heliolitid growth forms and their relative abundances (%) in the outcrops.
Conference presentations
Paleoecology of heliolitid corals from Wellin bioherm (Eifelian, Dinant Synclinorium, preliminary results)
13th International Symposium on Fossil Cnidaria and Porifera Modena 2019
Jan J. Król, Julien Denayer
Abstract:
Heliolitid tabulate corals from Wellin have been studied in respect to their growth patterns and colony development. The Wellin bioherm occurs within the uppermost Eifelian sediments, cropping out in southern parts of the Dinant Synclinorium in Belgium. It is located c.a. 40 km SE from Dinant. It is a part of the Wellin member of the Upper Eifelian – Lower Givetian Hanonet Formation, known for its abundant fauna of corals, stromatoporoids, brachiopods, and bryozoans.
Heliolitid corals are very abundant. They are small to medium sized (ranging between 3.4 cm and 17.2 cm in diameter). They mostly represent bulbous, domal, and irregular growth forms. Rejuvenations and following radical changes of the growth direction often occur. Growth interruption surfaces are very common within the coralla. They constitute dark layers that in cross-section show how the shape of the colony changed as it grew. These surfaces can be clean, but are often accompanied by sediment or encrusting organisms – mainly stromatoporoid sponges and alveolitid tabulates. Stromatoporoid encrusters commonly co-occur with syringoporid tabulate symbionts, growing inside the sponge body. In some cases the encrusters were later overgrown by the heliolitid, following its recovery after the growth interruption.
The abundant growth interruption surfaces consisting of sediment, as well as the common rejuvenations and changes of the growth direction, point to episodically increased sediment input. According to previous studies, it is very likely that at least some representatives of the genus Heliolites could be able to survive temporary sediment cover. The higher sediment input could be caused by the increased water energy e.g. during storms. The abundant growth interruption surfaces were also observed in the Devonian heliolitids from Morocco, Poland, and other localities in Belgium.
The interactions between heliolitids and their encrusters, settling on the surface of the colony during growth interruption, could suggest a competition for space between benthic organisms. The capability of heliolitids to survive sediment-derived growth interruption and to overgrow their epibionts, imply that these corals were very resistant. Heliolitids competing with stromatoporoids and overgrowing each other are also known from the Givetian of Morocco.
The continuation of the study of heliolitids from Wellin will include detailed taxonomy and further paleoecological analysis. The paleoecology of this group of tabulates is not very well understood, as the research of heliolitids in the past was often more focused on their taxonomy.
Skeletons of Palaeozoic Heliolites (Tabulata) and modern Heliopora (Octocorallia): a case of ecological convergence?
14th International Coral Reef Symposium, Bremen 2021
Jan J. Król, Nina Yasuda
Abstract:
Heliolites was a widespread genus of heliolitid corals, a group belonging to Tabulata. They were unique among tabulates - unlike the others, they developed septa (in hexameric symmetry) and formed highly integrated, plocoid colonies, rare in the Palaeozoic. Their ecology remains, however, poorly understood.
Heliolites featured a skeletal structure almost identical to that of modern octocorals of the genus Heliopora. However, the possibility of these genera being related has been ruled out based on different symmetry, mineralogy, and microstructure of their skeletons. Therefore, in this study, we test the hypothesis of ecological convergence as the reason for the close similarity between Palaeozoic heliolitids and Recent helioporids.
Our analyses of morphology and growth patterns of Middle Devonian Heliolites from Morocco, Belgium, and Poland show that this genus was very resilient. It was able to survive under the conditions of increased sediment influx, high energy events, scarcity of hard substrate for settlement, and strong competition for space on the reef. Additionally, it thrived under the greenhouse climate of the Middle Devonian, commonly considered to be one of the warmest periods in all of the Phanerozoic.
The observations on modern Heliopora coerulea from the Shiraho reef (offshore Ishigaki Island, Japan) show that it occurs abundantly in places characterized by increased temperatures and sediment input in the shallow, protected, low-energy setting of the backreef, but also in the forereef, under increased hydrodynamics and lower temperatures. It shows great skeletal plasticity, which allows it to adapt to different environmental conditions. Interactions with other benthic organisms are also common, particularly with other corals and with Polychaeta.
The resilience and adaptability of both Heliolites and Heliopora seem to be the common link. The simple skeletons provided their plocoid colonies with increased flexibility and versatility. In both genera the gross morphology and the corallite spacing can vary, apparently depending on external factors. More in-depth analysis of the convergence patterns between Heliolites and Heliopora will help us to better understand the adaptive capabilities of such resilient corals, which could resist high temperatures and generally unfavourable conditions. This should prove especially useful during present-day climate change resulting in the retreat of the scleractinians as the main reef builders.
Heliolitids - the best Paleozoic analogues of present-day reef corals?
Mediterranean Geosciences Union Annual Meeting, Marrakech 2022
Jan J. Król
Abstract:
Similarly to present-day reefs, the coral communities of Middle Paleozoic had to live through major climate shifts. Understanding how they dealt with such crises could help us predict the future of coral reefs endangered by current climate change. Unfortunately, most of the Paleozoic corals were not all that similar to their modern counterparts. Highly-integrated colonies of heliolitids however, might share the most similarities with modern reef corals, both in terms of structure and ecology.
Heliolitid coralla were collected from Silurian and Devonian outcrops of Gotland, Morocco, Belgium and Poland. Polished slabs and thin sections of the specimens were studied in respect to their growth patterns, taphonomy, inter-species interactions, associated biota and surrounding microfacies.
Studied specimens exhibit colony-wide responses to unfavorable conditions, e.g. sediment influxes or infestations. These are recorded in abundant growth-interruption surfaces in the coralla. Their high abundance suggests that the colonies were easily affected by environmental stressors, but the corals were able to regenerate from such events. Some species (e.g. Steliporella parvistella, Heliolites interstinctus, Propora tubulata) were also capable of skeletal plasticity – like the ability to form branching colonies under increased sediment input or platy colonies in low-light environments. Additionally, cyclic growth banding is present in some of the specimens, which together with the platy growth forms, suggests the presence of algal symbionts.
These characteristics of heliolitids are very similar to some of the modern hermatypic corals, especially Heliopora coerulea, which also shares with them an uncanny resemblance in the skeletal structure.
Heliolitids present themselves as an evolutionarily advanced group of corals for their time. They exhibited high colony integration, remarkable regenerative capabilities and most likely were photosymbiotic. On the other hand, much like modern corals, they were susceptible to shifts in environmental conditions.
These characteristics of heliolitids as a group living in the time periods of extreme climate change, makes them great analogues of modern corals. Studying their development provides us with insights into how hermatypic corals might react to increasingly more extreme climate change in the future.
Conclusions
Heliolitids formed highly integrated colonies, evolutionarily advanced in comparison to other Paleozoic corals
They exhibited good regenerative capabilities
They preferred relatively shallow-water environments
Most likely heliolitids were photosymbiotic
They were susceptible to shifts in environmental conditions
They actively participated in competition for space and commonly constituted substrate for epibionts
The preferences and adaptations listed above are very similar to the features of Heliopora coerulea, which suggests that their similar skeletal structure is in fact an example of convergent evolution
Heliolitids constitute good analogues for modern reef corals - the knowledge of their paleoecology could help in prognosing the future of coral reefs in the times of climate warming