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A paleoecological study of stromatoporoid endobionts was carried out to discern the relationships between symbiotic rugosans and their stromatoporoid hosts. The earliest endobiotic rugosan symbiont Palaeophyllum sp. in Baltica has only been found in the stromatoporoid Ecclimadictyon astrolaxum from Saaremaa, Estonia. The rugosans are vertically ori...
Citations
... The Silurian of Baltica has a rich record of symbiosis (Vinn and Wilson, 2016). There are records of possible polychaetes (Mõtus and Vinn, 2009;Vinn and Mõtus, 2014b), rugosans (Nestor, 1966;Kershaw, 1987;Vinn and Mõtus, 2014a), syringoporids (Nestor, 1966;Kershaw, 1987), and cornulitids (Vinn and Wilson, 2010a, b) in tabulate corals and stromatoporoids from the Silurian of Baltica. ...
Еhe large collection of thin sections of stromatoporoids and corals from the Silurian of Ukraine, Moldova, Belarus, and Komi Republic revealed severalincidences of skeletal intergrowth between stromatoporoids/corals and the other invertebrates. The stomatoporoids formed symbiotic associations with soft-bodied worms (Helicosalpinx and Cahetosalpinx), calcareous tentaculitoid tubeworms (microconchids, Cornulites, Conchicolites), and rugosans. Tabulate corals formed symbiotic associations with cornulitids. The studied stromatoporoid based associations are dominated by bioclaustrations of worms without mineral skeleton. Most likely non-mineralized invertebrates benefitted more from endobiotic life mode than invertebrates with mineralizedskeletons as the latter already had a protection on their own against predators. There was almost no difference in the number of symbiont taxa per host stromatoporoidspeciesindicating that all studied stromatoporoids were rather similar in their tolerance towards different endobionts.
... They were especially abundant in Silurian biostrome stromatoporoids (Kershaw 1987). The rugosans were usually vertically oriented inside the stromatoporoid skeletons (Vinn and Mõtus 2014a). Often numerous rugosans had their corallites open at the upper, external surface of stromatoporoids, but many could be completely embedded within the stromatoporoids (Vinn and Mõtus 2014a). ...
... The rugosans were usually vertically oriented inside the stromatoporoid skeletons (Vinn and Mõtus 2014a). Often numerous rugosans had their corallites open at the upper, external surface of stromatoporoids, but many could be completely embedded within the stromatoporoids (Vinn and Mõtus 2014a). Stromatoporoid hosts were presumably beneficial for rugosans as elevated stable substrates on a sea floor that offered a higher tier for feeding (Vinn and Mõtus 2014a). ...
... Often numerous rugosans had their corallites open at the upper, external surface of stromatoporoids, but many could be completely embedded within the stromatoporoids (Vinn and Mõtus 2014a). Stromatoporoid hosts were presumably beneficial for rugosans as elevated stable substrates on a sea floor that offered a higher tier for feeding (Vinn and Mõtus 2014a). It is possible that numerous rugosans in a single stromatoporoid may have decreased the feeding efficiency of the host by occupying part of its feeding surface. ...
Parasitic associations involving colonial animals are farily evenly distributed through the post-Cambrian Phanerozoic and have a long evolutionary history. Parasitism may have played an important role in the evolution of colonial animals. In the Paleozoic, the majority of marine symbioses involved colonial animals, and it is likely that colonial animals were also important hosts of parasites in the Mesozoic and Cenozoic, but further studies are needed. In the Paleozoic, stromatoporoids and corals were the most common hosts to various invertebrate parasites. Corals continued to be important hosts to parasites in the Mesozoic and Cenozoic. In addition, colonial animals themselves often infest or otherwise live in association with other organisms and can be parasites; however, colonial animals are more often hosts than parasites, and this has been so throughout the Phanerozoic. The stratigraphic distribution of parasitic associations in colonial animals is divided into two separate blocks: Paleozoic (Ordovician to Permian) parasitic associations of colonial animals form the first block and Mesozoic to Recent parasitic associations of colonial animals form the second block. This division of parasitic associations corresponds well to the Sepkoski Paleozoic and Modern faunas and therefore these subdivisions are termed as the Paleozoic and the Modern parasitic associations of colonial animals.
... To summarise: in all three examples illustrated, excellent material brought about a minimum of meaningful information in spite of the lack of systematic identifications. Vinn and Mõtus (2014) could be cited as representing an advancement relative to the trivial examples discussed above. Unfortunately, this is only apparent. ...
... Palaeophyllum is diagnosed as a phaceloid colonial genus with long major septa, reduced minor septa, an absent dissepimentarium, and a wide stereozone present on the periphery (Hill 1981, p. F138). Only one of those characters, the reduced minor septa can be observed in the specimens of Vinn and Mõtus (2014). The most important diagnostic character, a phaceloid colonial growth form, is not suggested by the authors and cannot be attributed to their illustrated specimens. ...
... The solitary growth forms of these specimens are reflected in: 1) the absence of offsetting specimens, despite many mature corallites being sectioned in the transverse, longitudinal, and oblique directions; 2) sections of very young specimens located discretely away from the mature ones, at distances excluding their connection not only by offsetting but also by attachment; 3) the growth directions of neighbouring corallites excluding their direct connection by offsetting. Moreover, either an incomplete dissepimentarium or concave tabularium occurs in Vinn and Mõtus' (2014, fig. 6) specimens, i.e., the very characters lacking in Palaeophyllum. ...
Starting from a subjective viewpoint on the decreasing interest in invertebrate fossil taxonomy, this essay discusses its importance in palaeobiological studies exemplified with cases from the palaeobiogeography and palaeoecology of rugose corals, and aims at provoking a discussion on the topic. The possible causes of this negative declining trend include inherent problems of palaeontological taxonomy, and changing systems in science and higher education.
... Paleozoic stromatoporoids were one of the most abundant organisms in reef complexes and associated facies from the Ordovician to the Late Devonian (Kershaw, 2015;Stearn, 2015;Kershaw et al., 2018). They lived in warm, shallow, tropical to subtropical marine environments, exhibiting a variety of growth forms (Stock et al., 2015; and are commonly found associated with other organisms, including many cases of intergrowth with other organisms such as tabulate and rugose corals, brachiopods, bryozoans, and worm tubes in reef environments (e.g., Kershaw, 1987;Young and Noble, 1989;Zhen and West, 1997;Lin and Webby, 1998;Nestor et al., 2010;Da Silva et al., 2011;Vinn and Wilson, 2012;Vinn and Mõtus, 2014;Stearn, 2015;Lee et al., 2016;Kershaw et al., 2018). ...
... Many associations between stromatoporoids and other organisms may be interpreted as spatial competition with, or predation by, the associated other organisms; some cases have been considered to be symbiotic interactions on the basis of modification of the adjacent skeletal structure of the host stromatoporoid (Kershaw et al., 2018). Most intergrowth associations are known from Silurian and Devonian strata (e.g., Mori, 1970;Kershaw, 1987;Young and Noble, 1989;Nestor et al., 2010;Da Silva et al., 2011;Vinn and Wilson, 2012;Vinn and Mõtus, 2014;Vinn et al., 2015;Vinn, 2016a, b), with a few recorded from Ordovician rocks (e.g., Lin and Webby, 1998;Lee et al., 2016). ...
... (4-9) Schematic drawings to illustrate the process of ecological interactions between two stromatoporoids in (1). Journal of Paleontology 94(1):1-10 adaptation to seek shelter from adverse environmental conditions (e.g., competition, predation, or depositional environments; Webby and Kershaw, 2015;Kershaw et al., 2018) or enhanced substrate stability (Vinn and Mõtus, 2014;Lee et al., 2016;Vinn et al., 2017). It has been proposed that stromatoporoids with well-developed laminae probably provided more favorable substrates than other stromatoporoids for the settlement of tabulate corals (Mori, 1970). ...
The earliest known interpreted spatial competition between two species of stromatoporoids, Clathrodictyon cf. C . mammillatum (Schmidt, 1858) and Labechia sp. is found in the Upper Ordovician Xiazhen Formation at Zhuzhai, South China. The interaction between these taxa was initiated by settlement of Labechia sp. on the surface of Clathrodictyon cf. C . mammillatum . Distortions of the intraskeletal elements of stromatoporoids represented by abnormally large, wide cysts and thick cyst plates in Labechia sp. are observed, along with zigzag crumpled distorted laminae and antagonistic behavior of the skeleton in Clathrodictyon cf. C . mammillatum , indicating syn-vivo interactions. The growth of Labechia sp. was terminated by the overgrowth of Clathrodictyon cf. C . mammillatum , possibly reflecting the ecological superiority of Clathrodictyon cf. C . mammillatum over Labechia sp. The observations are interpreted as competitive interaction between stromatoporoids that was most likely facultative, thus most likely occurring by chance, but the interaction allows assessment of different growth behaviors of the stromatoporoid species. Analysis of the interaction provides evidence to improve understanding of the paleoecology and growth behaviors of early stromatoporoids.
... It seems possible that space maintenance around the rugose polyp was similar to such phenomena in the Recent. Cases similar to that described here, where a rugose coral gains space are known from the Ordovician (Vinn et al. 2016(Vinn et al. , 2017a, Silurian (Sorauf & Kissling 2012;Vinn & Mõtus 2014) and Carboniferous (Pickett 2016). On the other hand, the presence of sweeper tentacles or similar structures in tabulate corals is equivocal, as preserved polyps show rather uniform tentacles (Copper 1985); in contrast, tabulates settling on crinoids damaged the host's tissue by either sweeper tentacles or digestive filaments (Berkowski & Zapalski 2014). ...
An exceptionally well‐preserved, unusual biostrome composed of the framebuilding cateniform tabulate coral Halysites catenularius (Linnaeus, 1767) bears an assemblage of the relatively large solitary cystiphyllid rugosan Cystiphyllum visbyense Wedekind, 1927. The corallites of solitary cystiphyllids are embedded within the ranks of the halysitid colonies, which developed on a soft, muddy substrate and in relatively turbid water. The cystiphyllid larvae successively settled mostly on the ranks of halysitid colonies and on colonies of the tiny phaceloid rugose coral Nanophyllum ramosum Johannessen, 1995, whereas calice‐in‐calice recruitment was not successful for these cystiphyllid corals. Further growth of C. visbyense was supported by rhizoid structures, which were most frequently developed on the cardinal (convex) side of the corallite. The process of formation of the rhizoid structures is here discussed and explained in detail, showing that they were formed by the extension of the basal ectodermal tissue of the polyp. The cystiphyllids, which settled on the walls of living corallites of halysitid colonies, used sweeper tentacles to kill the smaller polyps of the colony to maintain the space around them and expand. Hence, they ultimately used the halysitid colonies only as a hard substrate to stabilize their position on the soft muddy sediment.
... The number of rugosan symbionts in a single stromatoporoid can vary from one to tens of specimens depending on the taxa (Nestor et al. 2010). Some rugosans may have reduced the feeding efficiency of the host (Kershaw 1987;Vinn & Mõtus 2014). Rugosans symbionts possibly benefitted from the stable substrate and a higher tier provided by the sponge. ...
There are at least 47 different symbiotic pairs of taxa and 16 symbiotic associations in the Silurian of North America. Crinoids are most common host species and they hosted variety of epibiotic and endobiotic symbionts, including Tremichnus, platyceratid gastropods, brachiopods, microconchids, cornulitids, cyclostome bryozoans and favositid tabulates. Eighteen symbiotic pairs contain at least one colonial partner. Stromatoporoids hosted the most diverse fauna of endobiotic symbionts, including cornulitids, lingulids, Chaetosalpinx, Heliocosalpinx and rugosans. Among 16 symbiotic associations of Silurian of North America, 8 are common between North America and Baltica. North American symbiotic associations involving stromatoporoid hosts are the most similar to their Baltic equivalents.
... Many examples of different fossil associations were described from the Silurian stromatoporoid outcrops of Sweden (Gotland) and Saaremaa (Estonia) by Kershaw (1987), Kershaw and Mõtus (2016) and Vinn and Wilson (2016 and references therein). Stromatoporoid/Rugosa symbioses were reported from the Silurian of Baltoscandia (Estonia) by Vinn et al. (2015b), Vinn and Mõtus (2014) and Vinn and Toom (2017). Some new associations, such as crinoid/stromatoporoid (MGUWr-6621s; Fig. 9F) and brachiopod/bryozoan (MGUWr-6622s; Fig. 9G) and brachiopod/coral (Fig. 9H), which have not been reported so far from Baltoscandia, also were found in the Mokrzeszów Quarry. ...
... Lower Palaeozoic hardgrounds were encrusted mainly by rugose and tabulate corals, bryozoans, crinoids and microconchids (Vinn and Wilson, 2012a, b;Taylor and Wilson, 2003;Vinn et al., 2015a). Different fossil associations were reported from outcrops in Baltoscandia and the Baltic region by several authors (see Vinn and Wilson, 2016 and references therein), as well as coral/stromatoporoid symbiosis (e.g., Vinn and Mõtus, 2014). Just recently, Zatoń et al. (2016) described the first microconchid-encrusted corals from Estonia (Saaremaa). ...
... Endosymbiotic intergrowth of organisms with stromatoporoids has often been reported from the Paleozoic (Vinn, 2016), involving tabulate and rugose corals (Mistiaen, 1984;Kershaw, 1987;Young and Noble, 1989;Nestor et al., 2010;Da Silva et al., 2011;Vinn and Wilson, 2012;Vinn and Mõtus, 2014a;Vinn et al., 2015), brachiopods (Tapanila and Holmer, 2006), cyanobacteria (Webby, 1991;Nestor et al., 2010), bryozoans (Webby and Zhen, 1997;Nestor et al., 2010), worms (Zhen and West, 1997;Vinn et al., 2013;Vinn and Mõtus, 2014b) and cornulitids (Dixon, 2010;Vinn and Wilson, 2010). Coral-stromatoporoid intergrowths mainly involved tabulate corals living within the coenostea of stromatoporoids, and have been described from many Paleozoic shallow-water facies (Kershaw, 1987;Young and Noble, 1989). ...
... It is speculated that the survivorship of Bajgolia in such environments may have been better on elevated substrates that permitted feeding in a higher tier (cf. Vinn and Mõtus, 2014a) or in clear water, such as on the growth surfaces of stromatoporoids which rose above the surrounding seafloor. It is possible that juvenile stromatoporoids were less effective at removing intruders than mature ones, and therefore were more susceptible to settlement of Bajgolia larvae (cf. ...
... Laminae of the host stromatoporoid commonly curved upward or downward at the contact with corallites, indicating that growth of the host stromatoporoid was affected by the presence of Bajgolia. Such responses may have been related to the occupation of some of the host's feeding surface by endosymbionts, as noted by Vinn and Mõtus (2014a). In the case of upward curvature, the corallites of Bajgolia provided surfaces that enabled the stromatoporoid to rise to higher levels, which may have been beneficial. ...
One of the earliest endosymbiotic associations with stromatoporoids occurs in the Late Ordovician Xiazhen Formation of southeastern China. Bajgolia, an auloporid tabulate coral characterized by dichotomous branching due to longitudinal fission, is represented by free-living as well as endobiontic forms in various lithofacies representing a wide range of environments. Only two of 11 stromatoporoid genera (Clathrodictyon and Ecclimadictyon) hosted Bajgolia, mainly in reef and related facies. Bajgolia–stromatoporoid associations occur occasionally in the lower part of the formation, but eventually become persistent in the upper part. Such associations were initiated by larval settlement of the coral on the growth surface of the stromatoporoid. Growth of Bajgolia usually kept pace with its host, but the coral's ability to change growth direction and grow faster prevented its envelopment and termination by the stromatoporoid, allowing the establishment and recurrence of an ongoing endosymbiotic relationship between the two organisms. Endobiontic Bajgolia was able to survive with its corallites protruding from the host; in some cases, the growth form of the stromatoporoid changed in response to the coral. The relationships between Bajgolia and stromatoporoids were probably commensal, but there is also evidence for mutualism and/or parasitism. Bajgolia–stromatoporoid associations represent an important stage in the development of complex ecological relationships and community structure, prior to the common and widespread syringoporid (“caunopore tubes”)–stromatoporoid associations in the Siluro-Devonian.
... On the other hand, calcareous rigid skeletons of rugosans may have reinforced the skeletons of stromatoporoids. Symbiotic rugosans occur in at least in 17 species and 11 genera of stromatoporoids (Nestor, 1966;Soto and Méndez Bedia, 1985;Kershaw, 1987;May, 2005;Vinn and Mõtus, 2014b;Vinn et al., 2015). There are at least 18 species and 14 genera of rugosans in Paleozoic stromatoporoids (Nestor, 1966;Soto and Méndez Bedia, 1985;Kershaw, 1987;May, 2005;Vinn andMõtus, 2014a, 2014b;Vinn et al., 2015). ...
... There are at least 18 species and 14 genera of rugosans in Paleozoic stromatoporoids (Nestor, 1966;Soto and Méndez Bedia, 1985;Kershaw, 1987;May, 2005;Vinn andMõtus, 2014a, 2014b;Vinn et al., 2015). Solitary rugose corals often form symbiotic associations with the stromatoporoids (Nestor, 1966;Kershaw, 1987;Vinn and Mõtus, 2014b;Vinn et al., 2015). Colonial rugose coral and stromatoporoid intergrowth also occur in the Silurian (Kershaw, 1987;May, 2005). ...
... Solitary (Figure 2(b)) and occasionally ramose colonial rugose corals can also be found embedded within stromatoporoids in Silurian and Devonian deposits (Kershaw, 1987;Stel and Stoep, 1982;Soto and Méndez-Bedia, 1985;Nestor et al., 2010;Vinn and Mõtus, 2014a). Individual stromatoporoids sometimes host both rugose corals and caunopores (Figure 2(a)), but it is unclear whether the rugose coral symbionts were parasites of the host stromatoporoids. ...
... Individual stromatoporoids sometimes host both rugose corals and caunopores (Figure 2(a)), but it is unclear whether the rugose coral symbionts were parasites of the host stromatoporoids. Vinn and Mõtus (2014a) postulated benefits to the corals in terms of being able to feed at a higher tier in the water column and having a more stable substrate than usual, while the stromatoporoid hosts may have incurred advantages in being protected against predation by the nematocysts of the corals. However, the latter would have been balanced by loss of some of their feeding surface area. ...
Colonial species occur in a wide range of aquatic invertebrates, some having excellent fossil records, notably corals, bryozoans and graptolite hemichordates. In contrast to unitary animals, colonial animals grow by adding repetitive modules known as zooids. The ability of colonies to endure partial mortality and the typically plastic growth of benthic colonial species facilitates the formation of macrosymbiotic associations, some of which may be parasitic. However, as with unitary fossils, it is notoriously difficult to identify whether the symbioses are parasitisms (+/-) or mutualisms (+/+). Intergrowths between host colonies of stromatoporoid sponges, corals or bryozoans, and skeletal or soft-bodied symbionts are particularly common in Ordovician-Devonian shallow-water deposits. Soft-bodied symbionts in such intergrowths are represented by moulds in the host skeletons, a process of preservation termed bioclaustration. As yet, however, there is a lack of convincing data showing that any of these symbionts were parasites. By comparison with modern analogues, some fossil galls provide more convincing examples of parasitism, and the destructive effects of borings into the skeletons of benthic colonies also argue in favour of parasitism. Pelagic graptoloid hemichordates from the Early Palaeozoic occasionally contain cysts or tubes that have been attributed to parasites on the grounds that they would have adversely affected the hydrodynamics of the floating colonies. Future studies should test for parasitism by comparing the sizes of colonies hosting symbionts with those lacking symbionts.
