1. Walcott, C. D, 1920, Middle Cambrian Spongiae. Cambrian Geology and Paleontology, IV.

BibTeX
@misc{walcott1920middle2,
    author = "Walcott, C. D",
    title = "Middle Cambrian Spongiae. Cambrian Geology and Paleontology, IV",
    year = "1920",
    howpublished = "Smithsonian Miscellaneous Collections, v. 67, p. 261-364",
    note = "talkorigins\_source = {true}; raw\_reference = {Walcott, C. D., 1920, Middle Cambrian Spongiae. Cambrian Geology and Paleontology, IV: Smithsonian Miscellaneous Collections, v. 67, p. 261-364.}"
}

2. Whittington, Harry Blackmore, 1975, The enigmatic animal Opabinia regalis, middle Cambrian, Burgess Shale, British Columbia: Philosophical Transactions of the Royal Society of London. B, Biological Sciences: v. 271, no. 910: p. 1-43.

Abstract

Ten almost complete specimens have been studied in detail; nine coming from C. D. Walcott’s original collection, one from the recent re-investigation. The cuticle is preserved as a thin, dark layer; the external surface was apparently smooth, except for striations on the frontal process and adjacent to the mouth. Dorsally on the short cephalon were five eyes, thought to have been compound, the inner and outer pairs pedunculate, the median not stalked. From the anteroventral slope of the cephalon arose a long, flexible frontal process, divisible into a longer, cylindrical proximal portion, and a shorter, broad distal portion. The latter was divided longitudinally, each half bearing a group of long spines, directed inward and forward. The process probably contained a median, fluid-filled canal. The mouth was situated on the vertical, posteroventral wall of the cephalon, the alimentary canal U-shaped. The cylindrical axial region of the trunk tapered slightly backward, the alimentary canal situated ventrally and extending to the tip. The trunk was divided into a main portion of 15 segments, subequal in length, and a short posterior portion lacking segmentation. The junctions between segments gave a limited flexibility to the body. Each segment of the main portion of the trunk bore a pair of thin lateral lobes, directed downward and outward, overlapping, of maximum width medially, the lobes progressively more strongly prolonged backward. Dorsal to lobes 2-15, a paddle-shaped gill was attached near the base of the lobe. The ventral surface of the gill was flat, the outer, dorsal surface bearing imbricated, thin lamellae. The gills lay between adjacent, overlapping lateral lobes. Internally, in the main portion of the trunk what may have been diverticula of the gut are preserved, extending into the proximal portions of the lateral lobes. The posterior portion of the trunk bore three pairs of thin, lobate blades, directed upward and outward, overlapping in the opposite sense to the lateral lobes, the entire structure forming a tail fan. The dorsal margin of the tip of the axial region of the fan appears to have borne a pair of spines. The body is preserved with thin layers of rock between such parts as left and right eyes of a pair, adjacent lateral lobes, between gills and lobes, and between gill lamellae. The parts of the bodies are shown to have been entombed at varied angles to the horizontal bedding planes, and are greatly compressed. It is therefore considered that individuals were trapped in a cloud of sediment in suspension, moving along the sea bottom, and buried as it settled out. If so, the animal was benthonic in habit. Opabinia regalis may have ploughed shallowly in the bottom mud, propelled by movement of the lateral lobes. The eyes are presumed to have been capable of detecting movements in the surrounding waters, and the frontal process to have been used to explore the mud for food and bring it to the backward-facing mouth. The posterior region of the trunk may have aided in producing water currents over the dorsal surface of the body, or have aided in steering if the animal was capable of swimming. No structures that appear to have been antennae, and no other jointed appendages, have been observed, and the gills are not trilobite-like. O. regalis is not considered to have been a trilobitomorph arthiopod, nor is it regarded as an annelid. It may be descended from segmented animals from which arthropod phyla and/or annelids were derived.

BibTeX
@article{whittington1975the,
    author = "Whittington, Harry Blackmore",
    title = "The enigmatic animal Opabinia regalis, middle Cambrian, Burgess Shale, British Columbia",
    year = "1975",
    journal = "Philosophical Transactions of the Royal Society of London. B, Biological Sciences",
    abstract = "Ten almost complete specimens have been studied in detail; nine coming from C. D. Walcott’s original collection, one from the recent re-investigation. The cuticle is preserved as a thin, dark layer; the external surface was apparently smooth, except for striations on the frontal process and adjacent to the mouth. Dorsally on the short cephalon were five eyes, thought to have been compound, the inner and outer pairs pedunculate, the median not stalked. From the anteroventral slope of the cephalon arose a long, flexible frontal process, divisible into a longer, cylindrical proximal portion, and a shorter, broad distal portion. The latter was divided longitudinally, each half bearing a group of long spines, directed inward and forward. The process probably contained a median, fluid-filled canal. The mouth was situated on the vertical, posteroventral wall of the cephalon, the alimentary canal U-shaped. The cylindrical axial region of the trunk tapered slightly backward, the alimentary canal situated ventrally and extending to the tip. The trunk was divided into a main portion of 15 segments, subequal in length, and a short posterior portion lacking segmentation. The junctions between segments gave a limited flexibility to the body. Each segment of the main portion of the trunk bore a pair of thin lateral lobes, directed downward and outward, overlapping, of maximum width medially, the lobes progressively more strongly prolonged backward. Dorsal to lobes 2-15, a paddle-shaped gill was attached near the base of the lobe. The ventral surface of the gill was flat, the outer, dorsal surface bearing imbricated, thin lamellae. The gills lay between adjacent, overlapping lateral lobes. Internally, in the main portion of the trunk what may have been diverticula of the gut are preserved, extending into the proximal portions of the lateral lobes. The posterior portion of the trunk bore three pairs of thin, lobate blades, directed upward and outward, overlapping in the opposite sense to the lateral lobes, the entire structure forming a tail fan. The dorsal margin of the tip of the axial region of the fan appears to have borne a pair of spines. The body is preserved with thin layers of rock between such parts as left and right eyes of a pair, adjacent lateral lobes, between gills and lobes, and between gill lamellae. The parts of the bodies are shown to have been entombed at varied angles to the horizontal bedding planes, and are greatly compressed. It is therefore considered that individuals were trapped in a cloud of sediment in suspension, moving along the sea bottom, and buried as it settled out. If so, the animal was benthonic in habit. Opabinia regalis may have ploughed shallowly in the bottom mud, propelled by movement of the lateral lobes. The eyes are presumed to have been capable of detecting movements in the surrounding waters, and the frontal process to have been used to explore the mud for food and bring it to the backward-facing mouth. The posterior region of the trunk may have aided in producing water currents over the dorsal surface of the body, or have aided in steering if the animal was capable of swimming. No structures that appear to have been antennae, and no other jointed appendages, have been observed, and the gills are not trilobite-like. O. regalis is not considered to have been a trilobitomorph arthiopod, nor is it regarded as an annelid. It may be descended from segmented animals from which arthropod phyla and/or annelids were derived.",
    url = "https://doi.org/10.1098/rstb.1975.0033",
    doi = "10.1098/rstb.1975.0033",
    number = "910",
    pages = "1-43",
    volume = "271"
}

3. Whittington, Harry B., 1975, Trilobites with appendages from the Middle Cambrian, Burgess Shale, British Columbia: Fossils and Strata: p. 97-136.

BibTeX
@incollection{whittington1975trilobites,
    author = "Whittington, Harry B.",
    title = "Trilobites with appendages from the Middle Cambrian, Burgess Shale, British Columbia",
    year = "1975",
    booktitle = "Fossils and Strata",
    url = "https://doi.org/10.18261/8200049639-1975-06",
    doi = "10.18261/8200049639-1975-06",
    pages = "97-136"
}

4. Briggs, D E G, 1976, The Arthropod Branchiocaris N. Gen., Middle Cambrian, Burgess Shale, British Columbia.

Abstract

A study of new specimens of the bivalved arthropod Protocaris preriosa Resser, 1929 from the Burgess Shale has added much information to the original description, and the species is removed to a new genus Bra11c/1iocaris. Prorocaris marshi Walcott, 1884 and Dioxycaris arge11ra (Walcott, 1886) are also redescribed to facilitate comparison. The cephalic region of Bra11chiocaris preriosa bore a paired antenna anterior to a pair of stout appendages which may have been chelate. Suboval valves, the dorsal margin terminating anteriorly and posteriorly in a small pointed process, covered the anterior of the body. The trunk included about 46 divisions followed by a telson bearing a pair of short pointed processes. The lamellate trunk appendages may have been biramous, with a short, segmented proximal element. The animal probably swam near the seabed by metachronal movements of the lamellate appendages, which may also have functioned in respiration. B. pretiosa shows closest affinities to the branchiopod Crustacea among extant arthropods but is considered unlikely to represent a direct evolutionary predecessor of this group. The trunk of Protocaris marshi included about 65 divisions; the tel son processes were elongate and curved. The appendages are unknown. The carapace of Dioxycaris arge11ta is similar in outline and size to that of Branchiocaris preliosa but the only known specimen is poorly preserved and lacks evidence of the soft-part morphology.

BibTeX
@misc{briggs1976the,
    author = "Briggs, D E G",
    title = "The Arthropod Branchiocaris N. Gen., Middle Cambrian, Burgess Shale, British Columbia",
    year = "1976",
    abstract = "A study of new specimens of the bivalved arthropod Protocaris preriosa Resser, 1929 from the Burgess Shale has added much information to the original description, and the species is removed to a new genus Bra11c/1iocaris. Prorocaris marshi Walcott, 1884 and Dioxycaris arge11ra (Walcott, 1886) are also redescribed to facilitate comparison. The cephalic region of Bra11chiocaris preriosa bore a paired antenna anterior to a pair of stout appendages which may have been chelate. Suboval valves, the dorsal margin terminating anteriorly and posteriorly in a small pointed process, covered the anterior of the body. The trunk included about 46 divisions followed by a telson bearing a pair of short pointed processes. The lamellate trunk appendages may have been biramous, with a short, segmented proximal element. The animal probably swam near the seabed by metachronal movements of the lamellate appendages, which may also have functioned in respiration. B. pretiosa shows closest affinities to the branchiopod Crustacea among extant arthropods but is considered unlikely to represent a direct evolutionary predecessor of this group. The trunk of Protocaris marshi included about 65 divisions; the tel son processes were elongate and curved. The appendages are unknown. The carapace of Dioxycaris arge11ta is similar in outline and size to that of Branchiocaris preliosa but the only known specimen is poorly preserved and lacks evidence of the soft-part morphology.",
    url = "https://doi.org/10.4095/103962",
    doi = "10.4095/103962"
}

5. Whittington, Harry Blackmore, 1977, The Middle Cambrian trilobite Naraoia, Burgess Shale, British Columbia: Philosophical Transactions of the Royal Society of London. B, Biological Sciences: v. 280, no. 974: p. 409-443.

Abstract

The type species of the genus, N.compacta, is described from new preparations and measurements of over 100 specimens from C. D. Walcott’s original collection, and 5 from the recent re-investigation. Photographs and explanatory drawings provide the basis for considerations of mode of preservation, and lead to a new reconstruction. The dorsal exoskeleton was divided by a single articulation into two shields, each moderately convex with a raised axial region, the subcircular anterior shield overlapping for a short distance the longer posterior shield; narrow reflexed doublure on both shields. Dorsal surfaces of shields smooth, without transverse furrows, eyes absent. Axial region of anterior shield widest posteriorly, extending forward to threequarters length of shield, labrum may have been present under anterior portion. Axial region of posterior shield tapered back, reaching close to posterior margin of shield. Alimentary canal may be preserved filled with sediment, and was probably U-shaped anteriorly, broadest beneath anterior portion of axial region, tapering back to tip of posterior shield. Two types of alimentary diverticula preserved as reflective bands on anterior shield; single trunk of lateral diverticula ran transversely at mid-length and ramified beneath lateral region of shield; three pairs of axial diverticula, one per segment, originated behind main trunk of lateral diverticula and ramified in posterior part of axial region. Axial diverticula, one per segment and not ramifying, appear to have been present beneath the axial region of the posterior shield. Paired areas of muscle attachment, preserved as reflective or pyritous areas, are segmentally arranged along the axial region, one pair close together at the anterior extremity. One pair of long, uniramous, multi-jointed antennae was attached beside anterior extremity of axial region, followed by a maximum of 19 pairs of similar biramous appendages, three pairs on the posterior part of the anterior shield, remainder beneath posterior shield. Large triangular coxa strongly spinose on adaxial margin; inner, leg branch of five podomeres and terminal, thorn-like spine; large, spinose endite on proximal podomere. Outer branch arose from abaxial, dorsal margin of coxa, and consisted of slim, tapering shaft with terminal lobe, dorsal margin of shaft bore many long, thin, upward and backwardly directed lamellae. Specimens range in length from 9 to 40 mm, some 40 % of the sample being cast dorsal exoskeletons, the remainder whole animals. About one-fifth of the sample bore a posterolateral spine on the anterior shield, rather than having a rounded angle. This difference was recently used to erect two new species, Naraoia halia and N. pammon; here it is taken as the sole evidence of dimorphism in the single species N. compacta. A second species, N. spinfer, is recognized from two poorly-preserved specimens, characterized by seven pairs of lateral spines and a median posterior spine on the margins of the posterior shield; the axial region is poorly defined and appendages virtually unknown. N. compacta is considered to have been a benthonic predator and scavenger, walking, digging and raking in search of food much as did the trilobite Olenoides serratus, and to have had poor swimming powers. The lamellate outer branch of the appendage is regarded as a gill branch, aerated by currents produced when walking and swimming or drifting. There is no evidence of an abdomen or telson, so that N. compacta is a trilobite-like animal lacking the articulated thorax; it is regarded as representing a separate order of class Trilobita.

BibTeX
@article{whittington1977the,
    author = "Whittington, Harry Blackmore",
    title = "The Middle Cambrian trilobite Naraoia, Burgess Shale, British Columbia",
    year = "1977",
    journal = "Philosophical Transactions of the Royal Society of London. B, Biological Sciences",
    abstract = "The type species of the genus, N.compacta, is described from new preparations and measurements of over 100 specimens from C. D. Walcott’s original collection, and 5 from the recent re-investigation. Photographs and explanatory drawings provide the basis for considerations of mode of preservation, and lead to a new reconstruction. The dorsal exoskeleton was divided by a single articulation into two shields, each moderately convex with a raised axial region, the subcircular anterior shield overlapping for a short distance the longer posterior shield; narrow reflexed doublure on both shields. Dorsal surfaces of shields smooth, without transverse furrows, eyes absent. Axial region of anterior shield widest posteriorly, extending forward to threequarters length of shield, labrum may have been present under anterior portion. Axial region of posterior shield tapered back, reaching close to posterior margin of shield. Alimentary canal may be preserved filled with sediment, and was probably U-shaped anteriorly, broadest beneath anterior portion of axial region, tapering back to tip of posterior shield. Two types of alimentary diverticula preserved as reflective bands on anterior shield; single trunk of lateral diverticula ran transversely at mid-length and ramified beneath lateral region of shield; three pairs of axial diverticula, one per segment, originated behind main trunk of lateral diverticula and ramified in posterior part of axial region. Axial diverticula, one per segment and not ramifying, appear to have been present beneath the axial region of the posterior shield. Paired areas of muscle attachment, preserved as reflective or pyritous areas, are segmentally arranged along the axial region, one pair close together at the anterior extremity. One pair of long, uniramous, multi-jointed antennae was attached beside anterior extremity of axial region, followed by a maximum of 19 pairs of similar biramous appendages, three pairs on the posterior part of the anterior shield, remainder beneath posterior shield. Large triangular coxa strongly spinose on adaxial margin; inner, leg branch of five podomeres and terminal, thorn-like spine; large, spinose endite on proximal podomere. Outer branch arose from abaxial, dorsal margin of coxa, and consisted of slim, tapering shaft with terminal lobe, dorsal margin of shaft bore many long, thin, upward and backwardly directed lamellae. Specimens range in length from 9 to 40 mm, some 40 \% of the sample being cast dorsal exoskeletons, the remainder whole animals. About one-fifth of the sample bore a posterolateral spine on the anterior shield, rather than having a rounded angle. This difference was recently used to erect two new species, Naraoia halia and N. pammon; here it is taken as the sole evidence of dimorphism in the single species N. compacta. A second species, N. spinfer, is recognized from two poorly-preserved specimens, characterized by seven pairs of lateral spines and a median posterior spine on the margins of the posterior shield; the axial region is poorly defined and appendages virtually unknown. N. compacta is considered to have been a benthonic predator and scavenger, walking, digging and raking in search of food much as did the trilobite Olenoides serratus, and to have had poor swimming powers. The lamellate outer branch of the appendage is regarded as a gill branch, aerated by currents produced when walking and swimming or drifting. There is no evidence of an abdomen or telson, so that N. compacta is a trilobite-like animal lacking the articulated thorax; it is regarded as representing a separate order of class Trilobita.",
    url = "https://doi.org/10.1098/rstb.1977.0117",
    doi = "10.1098/rstb.1977.0117",
    number = "974",
    pages = "409-443",
    volume = "280"
}

6. Conway Morris, Simon, 1979, Middle Cambrian polychaetes from the Burgess Shale of British Columbia: Philosophical Transactions of the Royal Society of London. B, Biological Sciences: v. 285, no. 1007: p. 227-274.

Abstract

The Burgess Shale (Middle Cambrian) polychaetes Canadia spinosa Walcott, Burgessochaeta setigera (Walcott) gen. nov. and Peronochaeta dubia (Walcott) gen. nov. are redescribed on the basis of Walcott’s type specimens and on much additional material. Two new polychaetes Insolicorypha psygma gen. et sp. nov. and Stephenoscolex argutus gen. et sp. nov. are described. A poorly preserved specimen of unknown generic affinity is described as type A. The polychaetes are preserved as thin films that adhere to both sides of the split in the rock so that part and counterpart may be available. In C. spinosa, B. setigera and I. psygma, parts of the bodies such as the fascicles of setae are separated by thin layers of sediment that apparently seeped in during turbulent transport in turbidites or mudflows. The bodies therefore lie on two or more planes of microbedding and the factors that control exposure across a specimen are discussed. Aspects of the palaeoecology of the Burgess Shale are reviewed, including the distance the biota was transported prior to burial, the reasons for the exquisite preservation, and the effects of sedimentary compaction. C. spinosa was characterized by broad notosetae that extended across the dorsum, and large fascicles of neurosetae. Lobate branchiae were situated in the inter-ramal spaces. The prostomium bore a pair of elongate tentacles and the straight gut had an eversible unarmed proboscis. Several lines of evidence suggest that C. spinosa was an active benthonic swimmer. B. setigera was peculiar in possessing identical notosetae and neurosetae along the entire body. Long anterior tentacles, possibly of peristomial origin, may have been used in feeding. Indirect evidence indicates that B. setigera inhabited a burrow which it might have excavated with its proboscis. P. dubia may also have burrowed but it had uniramous parapodia bearing simple and acicular setae. The prostomium bore a pair of short appendages. I. psygma had extended neuropodia bearing cirri and elongate setae. The notopodia were reduced and cirri appear to have been wanting. The peculiar prostomium carried a pair of appendages. I. psygma is regarded as a pelagic polychaete. S. argutus possessed uniramous parapodia with simple stout setae. The bilobed prostomium bore at least one pair, and perhaps three pairs, of short appendages. Type A was the largest of the Burgess Shale polychaetes and had prominent setae on at least the anterior section of the body. Type A was a sediment eater but the feeding habits of the other polychaetes are uncertain. Particular attention is given to the influence of decay on the Burgess Shale polychaetes. To place the Burgess Shale polychaetes in some geological perspective other Cambrian worms, including a polychaete from the Spence Shale of Utah, are briefly redescribed and the late Precambrian (Ediacarian) worms Dickinsonia, Spriggina and Marywadea are assessed. Contrary to the findings of other workers, no convincing evidence for placing these latter worms in the polychaetes is forthcoming.

BibTeX
@article{conwaymorris1979middle,
    author = "Conway Morris, Simon",
    title = "Middle Cambrian polychaetes from the Burgess Shale of British Columbia",
    year = "1979",
    journal = "Philosophical Transactions of the Royal Society of London. B, Biological Sciences",
    abstract = "The Burgess Shale (Middle Cambrian) polychaetes Canadia spinosa Walcott, Burgessochaeta setigera (Walcott) gen. nov. and Peronochaeta dubia (Walcott) gen. nov. are redescribed on the basis of Walcott’s type specimens and on much additional material. Two new polychaetes Insolicorypha psygma gen. et sp. nov. and Stephenoscolex argutus gen. et sp. nov. are described. A poorly preserved specimen of unknown generic affinity is described as type A. The polychaetes are preserved as thin films that adhere to both sides of the split in the rock so that part and counterpart may be available. In C. spinosa, B. setigera and I. psygma, parts of the bodies such as the fascicles of setae are separated by thin layers of sediment that apparently seeped in during turbulent transport in turbidites or mudflows. The bodies therefore lie on two or more planes of microbedding and the factors that control exposure across a specimen are discussed. Aspects of the palaeoecology of the Burgess Shale are reviewed, including the distance the biota was transported prior to burial, the reasons for the exquisite preservation, and the effects of sedimentary compaction. C. spinosa was characterized by broad notosetae that extended across the dorsum, and large fascicles of neurosetae. Lobate branchiae were situated in the inter-ramal spaces. The prostomium bore a pair of elongate tentacles and the straight gut had an eversible unarmed proboscis. Several lines of evidence suggest that C. spinosa was an active benthonic swimmer. B. setigera was peculiar in possessing identical notosetae and neurosetae along the entire body. Long anterior tentacles, possibly of peristomial origin, may have been used in feeding. Indirect evidence indicates that B. setigera inhabited a burrow which it might have excavated with its proboscis. P. dubia may also have burrowed but it had uniramous parapodia bearing simple and acicular setae. The prostomium bore a pair of short appendages. I. psygma had extended neuropodia bearing cirri and elongate setae. The notopodia were reduced and cirri appear to have been wanting. The peculiar prostomium carried a pair of appendages. I. psygma is regarded as a pelagic polychaete. S. argutus possessed uniramous parapodia with simple stout setae. The bilobed prostomium bore at least one pair, and perhaps three pairs, of short appendages. Type A was the largest of the Burgess Shale polychaetes and had prominent setae on at least the anterior section of the body. Type A was a sediment eater but the feeding habits of the other polychaetes are uncertain. Particular attention is given to the influence of decay on the Burgess Shale polychaetes. To place the Burgess Shale polychaetes in some geological perspective other Cambrian worms, including a polychaete from the Spence Shale of Utah, are briefly redescribed and the late Precambrian (Ediacarian) worms Dickinsonia, Spriggina and Marywadea are assessed. Contrary to the findings of other workers, no convincing evidence for placing these latter worms in the polychaetes is forthcoming.",
    url = "https://doi.org/10.1098/rstb.1979.0006",
    doi = "10.1098/rstb.1979.0006",
    number = "1007",
    pages = "227-274",
    volume = "285"
}

7. Whittington, H.B., 1980, The significance of the fauna of the Burgess Shale, Middle Cambrian, British Columbia: Proceedings of the Geologists' Association: v. 91, no. 3: p. 127-148.

BibTeX
@article{whittington1980the,
    author = "Whittington, H.B.",
    title = "The significance of the fauna of the Burgess Shale, Middle Cambrian, British Columbia",
    year = "1980",
    journal = "Proceedings of the Geologists' Association",
    url = "https://doi.org/10.1016/s0016-7878(80)80034-4",
    doi = "10.1016/s0016-7878(80)80034-4",
    number = "3",
    pages = "127-148",
    volume = "91"
}

8. Briggs, Derek Ernest Gilmor, 1981, The arthropod Odaraia alata Walcott, Middle Cambrian, Burgess Shale, British Columbia: Philosophical Transactions of the Royal Society of London. B, Biological Sciences: v. 291, no. 1056: p. 541-582.

Abstract

This, the first detailed description, interpretation and reconstruction of Odaraia alata, is based on all 29 known specimens. These include material of Eurysaces pielus Simonetta and Delle Cave, 1975, which is synonymized with O. alata herein. The head bore a pair of large eyes anteriorly and a paired mandible posteriorly. Features between these are poorly defined and the number of limb-bearing cephalic somites is unknown. The carapace was bivalved and essentially tubular in configuration, enclosing most of the body anteriorly. The trunk included up to at least 45 uniform short wide limb-bearing somites. The trunk appendages were biramous (with the possible exception of the first two), with an outer lamellate branch projecting dorsad of a segmented, spinose and apparently sometimes bifurcate inner branch which shows some evidence of variation along the trunk. The telson bore three large flukes, two projecting laterally and one vertically. The evidence suggests that O. alata fed by employing the carapace as a filter chamber within which the appendages, which trapped small pelagic animals, were confined. The arthropod probably swam on its back, using the appendages. Although the flukes did not articulate proximally, the telson appears to have been well adapted as a stabilizing and steering organ. O. alata shows some similarities to the Crustacea, particularly the Branchiopoda, but the preservation of the features of the cephalon is inadequate to allow its affinities to be determined unequivocally. It is classified in the family Odaraidae Simonetta and Delle Cave, 1975, but assignment to a higher taxon within the arthropods is not considered to be justified.

BibTeX
@article{briggs1981the,
    author = "Briggs, Derek Ernest Gilmor",
    title = "The arthropod Odaraia alata Walcott, Middle Cambrian, Burgess Shale, British Columbia",
    year = "1981",
    journal = "Philosophical Transactions of the Royal Society of London. B, Biological Sciences",
    abstract = "This, the first detailed description, interpretation and reconstruction of Odaraia alata, is based on all 29 known specimens. These include material of Eurysaces pielus Simonetta and Delle Cave, 1975, which is synonymized with O. alata herein. The head bore a pair of large eyes anteriorly and a paired mandible posteriorly. Features between these are poorly defined and the number of limb-bearing cephalic somites is unknown. The carapace was bivalved and essentially tubular in configuration, enclosing most of the body anteriorly. The trunk included up to at least 45 uniform short wide limb-bearing somites. The trunk appendages were biramous (with the possible exception of the first two), with an outer lamellate branch projecting dorsad of a segmented, spinose and apparently sometimes bifurcate inner branch which shows some evidence of variation along the trunk. The telson bore three large flukes, two projecting laterally and one vertically. The evidence suggests that O. alata fed by employing the carapace as a filter chamber within which the appendages, which trapped small pelagic animals, were confined. The arthropod probably swam on its back, using the appendages. Although the flukes did not articulate proximally, the telson appears to have been well adapted as a stabilizing and steering organ. O. alata shows some similarities to the Crustacea, particularly the Branchiopoda, but the preservation of the features of the cephalon is inadequate to allow its affinities to be determined unequivocally. It is classified in the family Odaraidae Simonetta and Delle Cave, 1975, but assignment to a higher taxon within the arthropods is not considered to be justified.",
    url = "https://doi.org/10.1098/rstb.1981.0007",
    doi = "10.1098/rstb.1981.0007",
    number = "1056",
    pages = "541-582",
    volume = "291"
}

9. Bruton, D. L., 1981, The arthropod Sidneyia inexpectans, Middle Cambrian, Burgess Shale, British Columbia: Philosophical Transactions of the Royal Society of London. B, Biological Sciences: v. 295, no. 1079: p. 619-653.

Abstract

Old and new specimens of Sidneyia inexpectans have been studied and are accompanied by explanatory drawings and photographs. New reconstructions of the animal are given together with a three-dimensional model. The body consisted of a cephalon with a long backwardly directed doublure, a thorax of nine articulating somites, abdomen with cylindrical exoskeleton of two or three somites and a telson. A caudal fan was formed by a pair of uropods articulating at the posterior margin of the last abdominal somite. The cephalon had stalked eyes and preoral antennae but no walking or grasping appendages. The first four somites of the thorax had paired uniramous, prehensile walking legs attached to the body by broad coxae with spiny gnathobases. The coxae were smaller on the five posterior thoracic somites and the paired appendages were biramous, each bearing a gill supported on a flap attached at its proximal end to the first podomere of the leg. The coxa-body attachment resembles that of modern merostomes and is in advance of trilobites. Evidence suggests that Sidneyia was a bottom-living, carnivorous animal eating larger and harder food than trilobites. Gut contents include ostracodes, hyolithids, small trilobites and phosphatic debris. Sidneyia is the earliest known form which could be an ancestor to merostomes, but its body plan and absence of chelicera distinguishes Sidneyia from this group. The holotype of Amiella ornata Walcott, 1911 is reinterpreted and its synonomy with S. inexpectans is confirmed.

BibTeX
@article{bruton1981the,
    author = "Bruton, D. L.",
    title = "The arthropod Sidneyia inexpectans, Middle Cambrian, Burgess Shale, British Columbia",
    year = "1981",
    journal = "Philosophical Transactions of the Royal Society of London. B, Biological Sciences",
    abstract = "Old and new specimens of Sidneyia inexpectans have been studied and are accompanied by explanatory drawings and photographs. New reconstructions of the animal are given together with a three-dimensional model. The body consisted of a cephalon with a long backwardly directed doublure, a thorax of nine articulating somites, abdomen with cylindrical exoskeleton of two or three somites and a telson. A caudal fan was formed by a pair of uropods articulating at the posterior margin of the last abdominal somite. The cephalon had stalked eyes and preoral antennae but no walking or grasping appendages. The first four somites of the thorax had paired uniramous, prehensile walking legs attached to the body by broad coxae with spiny gnathobases. The coxae were smaller on the five posterior thoracic somites and the paired appendages were biramous, each bearing a gill supported on a flap attached at its proximal end to the first podomere of the leg. The coxa-body attachment resembles that of modern merostomes and is in advance of trilobites. Evidence suggests that Sidneyia was a bottom-living, carnivorous animal eating larger and harder food than trilobites. Gut contents include ostracodes, hyolithids, small trilobites and phosphatic debris. Sidneyia is the earliest known form which could be an ancestor to merostomes, but its body plan and absence of chelicera distinguishes Sidneyia from this group. The holotype of Amiella ornata Walcott, 1911 is reinterpreted and its synonomy with S. inexpectans is confirmed.",
    url = "https://doi.org/10.1098/rstb.1981.0164",
    doi = "10.1098/rstb.1981.0164",
    number = "1079",
    pages = "619-653",
    volume = "295"
}

10. Whittington, Harry Blackmore, 1981, Rare arthropods from the Burgess Shale, Middle Cambrian, British Columbia: Philosophical Transactions of the Royal Society of London. B, Biological Sciences: v. 292, no. 1060: p. 329-357.

Abstract

Six species of arthropods from the Walcott Collection, U.S. National Museum, are described. Molaria spinifera Walcott is known from over 100 specimens, a sample that reveals the morphology fairly fully. Between one and 12 specimens of the other species are known, and yield limited information. M. spinifera had a smooth, convex exoskeleton, not trilobed, the cephalic shield being a quarter-sphere in shape, eight trunk tergites diminishing in size posteriorly and the cylindrical telson having a short ventral spine and a long, jointed posterior spine. The cephalon bore a pair of short, slim antennae and three pairs of biramous appendages. There were eight pairs of similar biramous appendages on the trunk. The biramous appendage had a large basal podomere, a segmented inner walking branch, and a lobate outer branch arising from the basal podomere and bearing marginal lamellae. The sagittal length of cephalon, trunk and telson ranged from 8 to 26 mm, the posterior spine slightly exceeding this length; the smallest specimens are similar to the largest. The animal lacked eyes, and was probably benthic and may have been a scavenger and deposit feeder. Habelia optata Walcott was superficially similar to M. spinfera, the trunk being of 12 tergites; there was no cylindrical telson, but a ridged and barbed spine inserted into the 12th tergite, the spine having a joint at about two-thirds its length. The external surface of the exoskeleton was tuberculate; the pleurae of the tergites curved back increasingly strongly posteriorly, the tips being spinose. The cephalon appears to have borne a slim, short pair of antennae and two pairs of biramous appendages; the proximal portions of the jointed inner branches may have been adapted for grinding food. The first six trunk somites bore biramous limbs, the inner branch being a relatively long walking leg, the outer a lobe having marginal lamellae; on the posterior trunk somites there is no trace of the inner branch, but the outer was present. H. optata lacked eyes and was probably a benthic animal. Only the smooth exoskeleton of a possible second species, H? brevicauda Simonetta, is known, of which the posterior spine is short and bluntly rounded. The new genus and species Sarotrocercus oblita is erected for a few specimens, in which the body is about 1 cm in length, and behind which is a slim spine having a group of spines at the tip. From beneath the anterolateral margin of the cephalic shield a large eye projected, and the cephalon bore also one pair of large, jointed appendages. Behind these were pairs of lobed appendages bearing marginal lamellae, one on the cephalon and one on each of the nine trunk somites. This small species may have drifted and swum in the higher water layers, the occasional carcass lying on the sea bottom having been preserved. The single specimen of Actaeus armatus Simonetta is over 6 cm in length. The exoskeleton of this specimen is divided into cephalic shield with marginal eye lobe, 11 trunk tergites and a triangular terminal plate. The anterior cephalic appendage was Leanchoil-like, the stout proximal portion being curved and ending in a group of claws, the next two podomeres bearing long, slim extensions. The head shield also bore three pairs of biramous appendages, consisting of a small jointed inner branch and a large lobed outer branch with marginal lamellae; appendages like these outer branches are preserved beneath the trunk tergites. Only two specimens are identified as Alalcomenaeus cambricus Simonetta (length 3-4 cm). The exoskeleton is divisible into cephalic shield, trunk of probably 12 tergites, and an ovate terminal plate which has lateral bands. The cephalon has a marginal eye lobe and an anterior appendage which is broad proximally, the long distal portion being slim. The holotype shows a series of lobed appendages, the first three cephalic. Between them project the curved, pointed terminations of inner branches. The second specimen suggests that these lobed appendages bore marginal filaments, and reveals the inner branches as blade-shaped, and spinose on the inward-facing margin. These biramous appendages were present on all the trunk somites, being largest anteriorly. These remarkable appendages suggest a benthic scavenger, able to hold on to, and tear up, a carcass. ‘ Leanchoilia protogonia ’ Simonetta is most probably a composite, a poorly preserved Leanchoilia superlata lying on an unidentified, branching organism. The five species showing appendages extend greatly the known range of variation in morphology of the Burgess Shale arthropods. Affinities are discussed, but familial and higher classification is postponed, pending completion of work on all the arthropods from the shale.

BibTeX
@article{whittington1981rare,
    author = "Whittington, Harry Blackmore",
    title = "Rare arthropods from the Burgess Shale, Middle Cambrian, British Columbia",
    year = "1981",
    journal = "Philosophical Transactions of the Royal Society of London. B, Biological Sciences",
    abstract = "Six species of arthropods from the Walcott Collection, U.S. National Museum, are described. Molaria spinifera Walcott is known from over 100 specimens, a sample that reveals the morphology fairly fully. Between one and 12 specimens of the other species are known, and yield limited information. M. spinifera had a smooth, convex exoskeleton, not trilobed, the cephalic shield being a quarter-sphere in shape, eight trunk tergites diminishing in size posteriorly and the cylindrical telson having a short ventral spine and a long, jointed posterior spine. The cephalon bore a pair of short, slim antennae and three pairs of biramous appendages. There were eight pairs of similar biramous appendages on the trunk. The biramous appendage had a large basal podomere, a segmented inner walking branch, and a lobate outer branch arising from the basal podomere and bearing marginal lamellae. The sagittal length of cephalon, trunk and telson ranged from 8 to 26 mm, the posterior spine slightly exceeding this length; the smallest specimens are similar to the largest. The animal lacked eyes, and was probably benthic and may have been a scavenger and deposit feeder. Habelia optata Walcott was superficially similar to M. spinfera, the trunk being of 12 tergites; there was no cylindrical telson, but a ridged and barbed spine inserted into the 12th tergite, the spine having a joint at about two-thirds its length. The external surface of the exoskeleton was tuberculate; the pleurae of the tergites curved back increasingly strongly posteriorly, the tips being spinose. The cephalon appears to have borne a slim, short pair of antennae and two pairs of biramous appendages; the proximal portions of the jointed inner branches may have been adapted for grinding food. The first six trunk somites bore biramous limbs, the inner branch being a relatively long walking leg, the outer a lobe having marginal lamellae; on the posterior trunk somites there is no trace of the inner branch, but the outer was present. H. optata lacked eyes and was probably a benthic animal. Only the smooth exoskeleton of a possible second species, H? brevicauda Simonetta, is known, of which the posterior spine is short and bluntly rounded. The new genus and species Sarotrocercus oblita is erected for a few specimens, in which the body is about 1 cm in length, and behind which is a slim spine having a group of spines at the tip. From beneath the anterolateral margin of the cephalic shield a large eye projected, and the cephalon bore also one pair of large, jointed appendages. Behind these were pairs of lobed appendages bearing marginal lamellae, one on the cephalon and one on each of the nine trunk somites. This small species may have drifted and swum in the higher water layers, the occasional carcass lying on the sea bottom having been preserved. The single specimen of Actaeus armatus Simonetta is over 6 cm in length. The exoskeleton of this specimen is divided into cephalic shield with marginal eye lobe, 11 trunk tergites and a triangular terminal plate. The anterior cephalic appendage was Leanchoil-like, the stout proximal portion being curved and ending in a group of claws, the next two podomeres bearing long, slim extensions. The head shield also bore three pairs of biramous appendages, consisting of a small jointed inner branch and a large lobed outer branch with marginal lamellae; appendages like these outer branches are preserved beneath the trunk tergites. Only two specimens are identified as Alalcomenaeus cambricus Simonetta (length 3-4 cm). The exoskeleton is divisible into cephalic shield, trunk of probably 12 tergites, and an ovate terminal plate which has lateral bands. The cephalon has a marginal eye lobe and an anterior appendage which is broad proximally, the long distal portion being slim. The holotype shows a series of lobed appendages, the first three cephalic. Between them project the curved, pointed terminations of inner branches. The second specimen suggests that these lobed appendages bore marginal filaments, and reveals the inner branches as blade-shaped, and spinose on the inward-facing margin. These biramous appendages were present on all the trunk somites, being largest anteriorly. These remarkable appendages suggest a benthic scavenger, able to hold on to, and tear up, a carcass. ‘ Leanchoilia protogonia ’ Simonetta is most probably a composite, a poorly preserved Leanchoilia superlata lying on an unidentified, branching organism. The five species showing appendages extend greatly the known range of variation in morphology of the Burgess Shale arthropods. Affinities are discussed, but familial and higher classification is postponed, pending completion of work on all the arthropods from the shale.",
    url = "https://doi.org/10.1098/rstb.1981.0033",
    doi = "10.1098/rstb.1981.0033",
    number = "1060",
    pages = "329-357",
    volume = "292"
}

11. Rigby, J. K, 1986, Sponges of the Burgess Shale (Middle Cambrian) British Columbia.

BibTeX
@misc{rigby1986sponges1,
    author = "Rigby, J. K",
    title = "Sponges of the Burgess Shale (Middle Cambrian) British Columbia",
    year = "1986",
    howpublished = "Palaeontographica Canada, v. 2",
    note = "talkorigins\_source = {true}; raw\_reference = {Rigby, J. K., 1986, Sponges of the Burgess Shale (Middle Cambrian) British Columbia: Palaeontographica Canada, v. 2.}"
}

12. Conway Morris, S., 1995, Enigmatic shells, possibly halkieriid, from the Middle Cambrian Burgess Shale, British Columbia: Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen: v. 195, no. 1-3: p. 319-331.

BibTeX
@article{conwaymorris1995enigmatic,
    author = "Conway Morris, S.",
    title = "Enigmatic shells, possibly halkieriid, from the Middle Cambrian Burgess Shale, British Columbia",
    year = "1995",
    journal = "Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen",
    url = "https://doi.org/10.1127/njgpa/195/1995/319",
    doi = "10.1127/njgpa/195/1995/319",
    number = "1-3",
    pages = "319-331",
    volume = "195"
}