The Burgess Shale

Morania confluens

3D animation of Morania confluens (being grazed by Wiwaxia corrugata).

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Class: Cyanophyceae (Order: Nostocales?)
Remarks:

Walcott (1919) considered Morania to be related to the modern cyanobacteria Nostoc. No revisions to the affinities of this cyanobacterium have been published since.

Species name: Morania confluens
Described by: Walcott
Description date: 1919
Etymology:

Morania – from Moraine Lake (1,885 m), in Banff National Park.

confluens – from the Latin fluere, “flow or stream,” and the prefix con, “together.” The name refers to the abundance of this species.

Type Specimens: Syntypes–USNM35378-35390, 35398 (M. confluens); USMN 35391, 35392 (M. costellifera);USNM35393 (M. elongata);USNM35394 (M. fragmenta);USNM35395 – 35397, 35401 (M.? globosa);USNM57718 (M. parasitica);USNM35402 (M.? reticulata) in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: M. costellifera Walcott, 1919; M. elongata Walcott, 1919; M. fragmenta Walcott, 1919; M.? globosa Walcott, 1919; M. parasitica Walcott, 1919; M.? reticulata Walcott, 1919, all from the Walcott Quarry.

Other deposits: M.? antiqua Fenton and Fenton, 1937 from the middle Proterozoic Altyn Limestone of Montana and the Little Dal Group, Mackenzie Mountains (see Hofmann and Aitken, 1979).

Age & Localities:

Period:
Middle Cambrian, Bathyuriscus-Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Walcott Quarry on Fossil Ridge.

History of Research:

Brief history of research:

Walcott described Morania, erecting eight species, in a 1919 paper along with Burgess Shale algae, comparing the genus to the extant cyanobacteria Nostoc. Walcott included thin sections and details of the microstructures of M. confluens showing that it was formed of tangled strings of pyrite. Satterthwait (1976) studied specimens of M. confluens from the Geological Survey of Canada collections as part of her PhD thesis and broadly agreed with Walcott’s original interpretations, in particular regarding a position within the Nostocaceae. Sattertwhait’s work has not been published but she suggested that many species erected by Walcott might not be valid and could represent parts of more complex algae. Mankiewicz (1992) re-observed Walcott’s thin sections and confirmed the presence of Morania in several samples. Rigby (1986) identified M.? frondosa Walcott 1919, as a sponge and reassigned it to a new genus (see Crumillospongia frondosa).

Description:

Morphology:

Morania ranges in shape from spherical to sheet-like. The sheet-like form M. confluens is by far the most common species. Specimens typically range in length between 1 to more than 13 centimeters. The sheets are characteristically perforated, with holes up to 3 centimeters in diameter. The shape, size, number and distribution of holes are highly variable. Thin sections show that the microstructure of M. confluens is represented by a tangle mass of filaments called trichomes. These filaments have a beadlike structure with little spheroids of pyrite ranging 3 to 7 micrometers in diameter, and originally interpreted by Walcott as defining cellular structures.

Abundance:

Estimating the abundance of Morania is difficult since some bedding planes have large tangled masses of this cyanobacterium, and many could represent fragments of the same colony. Morania is very common in the Walcott Quarry and represents 4.9% of the community (Caron and Jackson, 2008).

Maximum Size:
130 mm

Ecology:

Ecological Interpretations:

Caron and Jackson (2006) suggested that Morania covered large areas of the benthos and might have provided a stable substrate and food source for benthic animals, in particular for a number of grazers, like Odontogriphus and Wiwaxia.

References:

CARON, J.-B. AND D. A. JACKSON. 2006. Taphonomy of the Greater Phyllopod Bed Community, Burgess Shale. Palaios, 21: 451-465.

CARON, J.-B. AND D. A. JACKSON. 2008. Paleoecology of the Greater Phyllopod Bed community, Burgess Shale. Palaeogeography, Palaeoclimatology, Palaeoecology, 258: 222-256.

HOFMANN, H. J. AND J. D. AITKEN. 1979. Precambrian biota from the Little Dal Group, Mackenzie Mountains, northwestern Canada. Canadian Journal of Earth Sciences, 16: 150-166.

MANKIEWICZ, C. 1992. Obruchevella and other microfossils in the Burgess Shale: preservation and affinity. Journal of Paleontology, 66(5): 717-729.

SATTERTHWAIT, D. F. 1976. Paleobiology and Paleoecology of Middle Cambrian Algae from Western North America. Unpublished PhD thesis, California, Los Angeles, 120 p.

WALCOTT, C. 1919. Cambrian Geology and Paleontology IV. Middle Cambrian Algae. Smithsonian Miscellaneous Collections, 67(5): 217-260.

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Marpolia spissa

3D animation of Marpolia spissa.

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Class: Cyanophyceae (Order: Oscillatoriales?)
Remarks:

Walcott (1919) considered this species to be a cyanobacterium, but Walton suggested a relationship to red algae instead (Walton, 1923). More recent studies concurred with Walcott’s original interpretation (Conway Morris and Robison, 1988).

Species name: Marpolia spissa
Described by: Walcott
Description date: 1919
Etymology:

Marpolia – from Mount Marpole (2,997 m), a peak located near the Burgess Shale, northwest of Emerald Lake in Yoho National Park.

spissa – from the Latin spissus, “crowded,” in reference to the bush-like aspect of this cyanobacteria.

Type Specimens: Lectotype –USNM35403 (M. spissa); holotype –USNM35412 (M. aequalis) in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: M. aequalis Walcott 1919 from the Trilobite Beds on Mount Stephen (known from a single specimen).

Other deposits: Marpolia (possibly represented by different species) is common in various Cambrian exceptional fossil deposits, in particular from the Middle Cambrian Spence Shale and Wheeler Formation in Utah (Conway Morris and Robison, 1988) and the Middle Cambrian Kaili Formation in China (Yang et al., 2001).

Age & Localities:

Period:
Middle Cambrian, Bathyuriscus-Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Walcott Quarry on Fossil Ridge, the Tulip Beds (S7) on Mount Stephen, and other smaller localities on Mount Field, Mount Stephen and Monarch Cirque.

History of Research:

Brief history of research:

Walcott described Marpolia in 1919 and named two species from the Burgess Shale, M. spissa from the Walcott Quarry and M. aequalis from the Trilobite Beds. M. spissa was compared to the modern Oscillatorialesin an unpublished thesis (Satterthwait, 1976), an interpretation followed by Conway Morris and Robison (1988) based on the study of fossil material from various Utah deposits. M. spissa is commonly found in thin sections (Mankiewicz, 1992) and can be isolated by acid maceration (Butterfield, 1990). A recent taphonomic study demonstrated that the preservation style of Marpolia is similar to other Burgess Shale organisms (Butterfield et al., 2007).

Description:

Morphology:

Marpolia forms dense tufts up to 5 cm in length composed of numerous filaments. Filaments tend to branch near the base of the tuft. Each filament averages about 40 microns in width. Filaments are composed of an outer sheath and one to four strands of inner cells. Each cell is about 2 microns in length. M. aequalis has a central stem and stronger branching structures than M. spissa.

Abundance:

Estimating the abundance of Marpolia is difficult since some bedding planes have large tangled masses of this cyanobacterium, and many could represent fragments of the same colony. M. spissa is rare and represents only 0.07% of the Walcott Quarry community (Caron and Jackson, 2008).

Maximum Size:
50 mm

Ecology:

Ecological Interpretations:

The absence of an attachment structure suggests that Marpolia may have been free-living, floating in large masses (i.e., planktonic). It may have attached to other floating objects as free-living cyanobacteria do today. It is also possible that the lack of attachment structure is taphonomic (a structure that is lost during deposition), due to detachment from the sediment during transport (caused by having been swept up in mud flows) prior to burial.

References:

BUTTERFIELD, N. J. 1990. Organic preservation of non-mineralizing organisms and the taphonomy of the Burgess Shale. Paleobiology, 16: 272-286.

BUTTERFIELD, N. J., U. BALTHASAR AND L. WILSON. 2007. Fossil diagenesis in the Burgess Shale. Palaeontology, 50: 537-543.

CARON, J.-B. AND D. A. JACKSON. 2008. Paleoecology of the Greater Phyllopod Bed community, Burgess Shale. Palaeogeography, Palaeoclimatology, Palaeoecology, 258: 222-256.

CONWAY MORRIS, S. AND R. A. ROBISON. 1988. More soft-bodied animals from the Middle Cambrian of Utah and British Columbia. University of Kansas Paleontological Contributions, 122 p.

MANKIEWICZ, C. 1992. Obruchevella and other microfossils in the Burgess Shale: preservation and affinity. Journal of Paleontology, 66: 717-729.

SATTERTHWAIT, D. F. 1976. Paleobiology and Paleoecology of Middle Cambrian Algae from Western North America. Unpublished PhD thesis, California, Los Angeles, 120 p.

WALCOTT, C. D. 1919. Middle Cambrian Algae. Cambrian Geology and Paleontology IV. Smithsonian Miscellaneous Collections, 67(5): 217-260.

WALTON, J. 1923. On the structure of a Middle Cambrian alga from British Columbia (Marpolia spissa Walcott). Proceedings of the Cambridge Philosophical Society-Biological Sciences, 1: 59-62.

YANG, R., J. MAO, Y. ZHAO, X. CHEN AND X. YANG. 2001. Branching macroalgal fossils of the Early-Middle Cambrian Kaili Formation from Taijiang, Guizhou Province, China. Acta Geologica Sinica, 75: 433-440.

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