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Reptamsassia n. gen. (Amsassiaceae n. fam.; calcareous algae) from the Lower Ordovician (Floian) of western Newfoundland, and the earliest symbiotic intergrowth of modular species

Published online by Cambridge University Press:  28 January 2022

Dong-Jin Lee*
Affiliation:
College of Earth Sciences, Jilin University, Changchun, 130061, China
Robert J. Elias
Affiliation:
Department of Earth Sciences, University of Manitoba, Winnipeg, MB R3T 2N2, Canada
Brian R. Pratt
Affiliation:
Department of Geological Sciences, University of Saskatchewan, Saskatoon, SK S7N 5E2, Canada
*
*Corresponding author

Abstract

Modular coral-like fossils occur in thrombolitic reefal beds at two stratigraphic levels within the Lower Ordovician (Floian) Barbace Cove Member of the Boat Harbour Formation, in the St. George Group of western Newfoundland. They are here assigned to Reptamsassia n. gen.; R. divergens n. gen. n. sp. is present at both levels, whereas a comparatively small-module species, R. minuta n. gen. n. sp., is confined to the upper level. Reptamsassia n. gen. resembles the Ordovician genus Amsassia in its phacelocerioid structure, back-to-back walls of adjoining modules, module increase by longitudinal fission involving infoldings of the wall, tabula-like structures that are continuous with the vertical module wall, and calices with concave-up bottoms. The new genus is differentiated by its encrusting habit, modules with highly variable growth directions and shapes throughout skeletal growth, and modules that may separate slightly or diverge from one another following fission. Together, Amsassia and Reptamsassia n. gen. are considered to represent a distinct group of calcareous algae, the Amsassiaceae n. fam., which possibly belongs to the green algae. The Early Ordovician origination of Amsassia followed by Reptamsassia n. gen. contributed to the beginning of the rise in diversity on a global scale and in reefal settings during the Great Ordovician Biodiversification Event. Reptamsassia minuta n. gen. n. sp. was an obligate symbiont that colonized living areas on its host, R. divergens n. gen. n. sp., with isolated modules of R. divergens n. gen. n. sp. able to persist in the resulting intergrowth with R. minuta n. gen. n. sp. This is the earliest known symbiotic intergrowth of macroscopic modular species, exemplifying the development of ecologic specialization and ecosystem complexity in Early Ordovician reefs.

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Articles
Copyright
Copyright © The Author(s), 2022. Published by Cambridge University Press on behalf of The Paleontological Society
Figure 0

Figure 1. (1) Map of Newfoundland. (2) Map of western Newfoundland showing autochthonous lower Paleozoic platform carbonates (based on Knight et al., 2008, fig. 1), which are present from Port au Port Peninsula to the northern tip of Great Northern Peninsula. Samples of Amsassia described by Lee et al. (2021) were collected at Green Head, Eddies Cove West, and Boat Head; samples of Reptamsassia n. gen. for the present study were collected at Isthmus Bay (for detailed map see Knight and James, 1987, fig. 3d).

Figure 1

Figure 2. Generalized stratigraphic section of the St. George Group in western Newfoundland (based on Knight et al., 2008, fig. 2; with updated chronostratigraphy in Boyce et al., 2013, fig. 2). The North American Skullrockian through Blackhillsian stages comprise the Ibexian Series. Besides the contact between the St. George Group and the overlying Table Head Group, there are two significant regional disconformities within the Boat Harbour Formation (shown with wavy lines). The middle member and Barbace Cove Member of the Boat Harbour Formation are labeled, but other members are not. Lithologic symbols: rectangles = limestone; right trapezoids = interbedded limestone and dolostone. Stratigraphic positions of Amsassia and Reptamsassia n. gen. are marked with arrows (geographic locations are shown in Fig. 1.2): A, Amsassia terranovensis and Amsassia? sp. A, ~26–33 m above base of Watts Bight Formation at Green Head; B, Amsassia diversa and Amsassia? sp. B, ~18 m below top of middle member in Boat Harbour Formation at Boat Head; C, Amsassia diversa, ~13.5 m below top of middle member in Boat Harbour Formation at Eddies Cove West; D, Reptamsassia divergens n. gen. n. sp., ~4 m above base of Barbace Cove Member in Boat Harbour Formation at Isthmus Bay (locality B86); E, Reptamsassia divergens n. gen. n. sp. and R. minuta n. gen. n. sp., ~15 m above base of Barbace Cove Member in Boat Harbour Formation at Isthmus Bay (locality B97).

Figure 2

Figure 3. (1–3) Representative dolomitic limestone fabrics of thrombolite mounds containing Reptamsassia n. gen. in the Barbace Cove Member (Floian) of the Boat Harbour Formation, Isthmus Bay, Port au Port Peninsula, western Newfoundland; back-lit photographs of thin sections oriented horizontally (1; locality B86) and vertically (2, 3; locality B97) with respect to bedding; scale bar (shown in 1) = 10 mm. (1) Rudstone composed of peloids, rare bioclasts, a fragment of nautiloid (lower center), a turriform gastropod (right), and intraclasts of biolithite, including one containing Reptamsassia n. gen. (top center). (2) Boundstone composed of Reptamsassia n. gen. (center and right) encased in microbialite, flanked by wackestone composed of sparse biomicrite and locally pelsparite, with burrows conforming to Palaeophycus and Planolites where backfilled. (3) Boundstone composed of Reptamsassia n. gen. in upright growth position, mostly enveloped by a thin microbialite coating and flanked by lime mudstone consisting of burrowed micrite and locally pelsparite. (4) Direct association of Reptamsassia n. gen. species; back-lit photograph of thin section. Transverse view of large-module species R. divergens n. gen. n. sp. (left; GSC 142673) and the small-module species R. minuta n. gen. n. sp. (right; GSC 142674) from floatstone/rudstone at locality B97; scale bar = 5 mm.

Figure 3

Figure 4. Thin-section photomicrographs showing transverse (1, 2), longitudinal (3, 4), and oblique longitudinal (5) views of intergrowths of large-module species Reptamsassia divergens n. gen. n. sp. and the small-module species R. minuta n. gen. n. sp. from boundstone at locality B97. (1) Reptamsassia minuta n. gen. n. sp. (middle of photograph) associated with R. divergens n. gen. n. sp. (left and right sides); R. minuta encloses isolated modules of R. divergens (arrows; outlined area enlarged in Fig. 5.1); GSC 142675a (holotype of R. minuta n. gen. n. sp.); scale bar = 5 mm. (2) Reptamsassia minuta n. gen. n. sp. (right side) associated with R. divergens n. gen. n. sp. (left side); isolated modules of R. divergens (arrows) are surrounded by R. minuta; GSC 142675b (holotype of R. minuta n. gen. n. sp.); scale bar = 2 mm. (3) Reptamsassia divergens n. gen. n. sp. overgrown by R. minuta n. gen. n. sp.; isolated modules of R. divergens (arrows) are intergrown with R. minuta; GSC 142676; scale bar = 2 mm. (4) Reptamsassia minuta n. gen. n. sp. overgrowing and intergrown with R. divergens n. gen. n. sp.; tabula-like structures, associated with constrictions of modules, are developed at same level in adjacent modules of R. minuta (between arrows); GSC 142677a (paratype of R. minuta n. gen. n. sp.); scale bar = 2 mm. (5) Reptamsassia divergens n. gen. n. sp. (bottom) partially overgrown by R. minuta n. gen. n. sp. (center), which it subsequently overgrew (top); patch of microbial clotted micrite (A) with continuous module growth on both sides (B); GSC 142677b (paratype of R. minuta n. gen. n. sp.); scale bar = 2 mm.

Figure 4

Figure 5. Thin-section photomicrographs showing transverse (1) and longitudinal (2) views of intergrowths of large-module species Reptamsassia divergens n. gen. n. sp. and the small-module species R. minuta n. gen. n. sp. from boundstone at locality B97; scale bars = 1 mm. (1) Enlargement of outlined area in Figure 4.1, showing isolated modules of R. divergens n. gen. n. sp. (arrows) enclosed in R. minuta n. gen. n. sp.; GSC 142675a (holotype of R. minuta n. gen. n. sp.). (2) Reptamsassia divergens n. gen. n. sp. (lower left) overgrown by R. minuta n. gen. n. sp., showing isolated modules of R. divergens (arrows) intergrown with R. minuta, and terminal growth surface (upper left); GSC 142678.

Figure 5

Figure 6. Thin-section photomicrographs showing transverse (2, 3, 5–7) and both transverse and longitudinal (1, 4) views of Reptamsassia divergens n. gen. n. sp. from localities B86 (4, 6, 7) and B97 (1–3, 5). (1) Skeleton showing phacelocerioid structure and variable growth directions of modules, ranging from upward to lateral and reptant; some small areas of sediment accumulation or microbial encrustation (left of center, lower right of center) seem to have been accommodated by diverted growth of modules without mortality; note module undergoing bipartite fission (arrow); holotype GSC 142679; scale bar = 5 mm. (2) Skeleton showing variable growth directions of modules; paratype GSC 142680; scale bar = 5 mm. (3) Skeleton showing partial growth interruptions associated with peloids and micritic clots likely of microbial origin (arrows); GSC 142681; scale bar = 5 mm. (4) An intraclast consisting of skeleton encrusted by microbialite; longitudinal fission resulting in pairs of modules (arrows); deep calices with concave-up bottoms (outlined area; enlarged in Fig. 8.3); GSC 142682; scale bar = 2 mm. (5) Cerioid area showing low size and shape variability of densely packed, polygonal to subpolygonal modules; GSC 142683; scale bar = 1 mm. (6) Loosely packed area showing high size variability of subcircular to circular modules, some flanked by micrite (dark colored); outlined area enlarged in 7; GSC 142684; scale bar = 0.5 mm. (7) Enlargement of outlined area in 6, showing back-to-back walls between adjoining modules (pairs of white arrows); GSC 142684; scale bar = 0.5 mm.

Figure 6

Figure 7. Thin-section photomicrographs showing transverse views of Reptamsassia divergens n. gen. n. sp. from localities B86 (1, 2, 6) and B97 (3–5, 7, 8). (1) Modules irregularly shaped where they encrust substrate during initial stage of skeletal growth (black arrow) and at base of skeleton during subsequent lateral expansion (white arrows); outlined area enlarged in 2; GSC 142685; scale bar = 2 mm. (2) Enlargement of outlined area in 1, showing variably sized modules, some flanked by micrite, in loosely packed area; module shapes subcircular to circular, or irregular due to interference from adjacent modules; note module undergoing bipartite fission involving infolding of wall (arrow); GSC 142685; scale bar = 0.5 mm. (3–6) Modules undergoing bipartite fission (white arrows) and probable quadripartite fission (black arrow in 5) involving infolding(s) of wall; GSC 142686a, GSC 142687, GSC 142686b, and GSC 142688, respectively; scale bars = 0.5 mm. (7) Module undergoing tripartite fission; GSC 142686a; scale bar = 0.5 mm. (8) Module probably undergoing tripartite fission involving infoldings of wall; GSC 142689; scale bar = 0.5 mm.

Figure 7

Figure 8. Thin-section photomicrographs showing longitudinal views of Reptamsassia divergens n. gen. n. sp. from localities B86 (3, 4) and B97 (1, 2, 5–7). (1) Longitudinal fission resulting in modules that separate slightly; GSC 142690; scale bar = 0.5 mm. (2) Deep calice with concave-up bottom (white arrow) similar in shape to tabula-like structures within modules (black arrows); GSC 142691; scale bar = 1 mm. (3) Enlargement of outlined area in Figure 6.4, showing longitudinal fission resulting in modules that separate slightly (left side); note deep calices of modules; GSC 142682; scale bar = 1 mm. (4) Deep calices with concave-up bottoms; GSC 142692; scale bar = 1 mm. (5, 6) Longitudinal fission resulting in modules that diverge (arrows); GSC 142693 and GSC 142694, respectively; scale bars = 1 mm and 0.5 mm, respectively. (7) Tabula-like structures forming bases of regenerated modules, usually following constriction of modules (e.g., white arrows), commonly developed at same levels in adjacent modules (black arrows); GSC 142695; scale bar = 2 mm.

Figure 8

Table 1. Diameter (mm) of modules of Reptamsassia divergens n. gen. n. sp., based on 12 skeletons from locality B86 and 11 (including holotype and paratype) from locality B97. In transverse thin-sections, diameter was measured internally in the largest 10% of modules; module diameter = (long axis + short axis) ÷ 2 (see Scrutton, 1981). Abbreviations: Min. = minimum; Max. = maximum; Avg. = average; S.D. = standard deviation; N = number of modules.

Figure 9

Figure 9. Thin-section photomicrographs showing transverse views of Reptamsassia minuta n. gen. n. sp. from locality B97. (1) Mostly polygonal to subpolygonal modules showing variable shapes and sizes, with one module undergoing bipartite fission involving infolding of wall (white arrow); adjoining modules have back-to-back walls (black arrows); holotype GSC 142675a; scale bar = 0.5 mm. (2) Shape and size variability of subcircular to circular and irregularly shaped modules, some flanked by sediment (dark area left of center); holotype GSC 142675c; scale bar = 0.5 mm. (3) Module undergoing bipartite fission (arrow) where modules are relatively densely packed; holotype GSC 142675b; scale bar = 0.2 mm. (4, 5) Modules undergoing bipartite fission involving infolding(s) of wall (arrows); holotype GSC 142675c and GSC 142696a, respectively; scale bars = 0.2 mm. (6) Module undergoing tripartite fission involving infoldings of wall (arrows); holotype GSC 142675b; scale bar = 0.2 mm. (7) Module probably undergoing quadripartite fission involving infoldings of wall (arrows); GSC 142697; scale bar = 0.2 mm.

Figure 10

Figure 10. Thin-section photomicrographs showing longitudinal views of Reptamsassia minuta n. gen. n. sp. from locality B97. (1–3) Longitudinal fission resulting in modules that separate slightly or diverge (white arrows); note calice of a module (black arrow in 1); GSC 142696b (1, 2) and GSC 142698 (3); scale bars = 0.2 mm. (4, 5) Modules having moderately deep calices (arrows); holotype GSC 142675a and paratype GSC 142677a, respectively; scale bars = 0.2 mm and 0.5 mm, respectively.

Figure 11

Table 2. Diameter (mm) of modules of Reptamsassia minuta n. gen. n. sp., based on five skeletons (including holotype). In transverse thin-sections, diameter was measured internally in the largest 10% of modules; module diameter = (long axis + short axis) ÷ 2 (see Scrutton, 1981). Abbreviations: Min. = minimum; Max. = maximum; Avg. = average; S.D. = standard deviation; N = number of modules.