The Burgess Shale

Actaeus armatus

A very rare arthropod with great appendages and large eyes

Actaeus armatus (USNM 155597) – Holotype. Complete specimen shown in dorsal oblique view. Specimen length = 66 mm. Specimen dry – polarized light (left), wet – direct light (right). Walcott Quarry.

© Smithsonian Institution – National Museum of Natural History. Photos: Jean-Bernard Caron



Kingdom: Animalia
Phylum: Arthropoda
Class: Unranked clade Megacheira? (stem group arthropods)

Actaeus is thought to be closely related to Leanchoilia (Bruton and Whittington, 1981). If confirmed, Actaeus would belong to a clade of “great appendage” arthropods, or Megacheira (Hou and Bergström, 1997; Wills et al., 1998; Cotton and Braddy, 2004). The phylogenetic placement of the megacheirans is uncertain and they are considered to be either stem-lineage chelicerates (Chen et al., 2004; Edgecombe, 2010) or upper stem-lineage euarthropods (Budd, 2002).

Species name: Actaeus armatus
Described by: Simonetta
Description date: 1970

Actaeus – from the Greek Actaeus, the first king of Attica. Attica is an historical region of Greece which includes Athens today.

armatus – from the Greek armos, “joint,” and the Latin atus, “provided with.” The name refers to the large frontal appendages of this species.

Type Specimens: Holotype –USNM155597 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none.

Other deposits: none.

Age & Localities:

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:

Simonetta (1970) described this organism on the basis of a single specimen originally collected by Walcott. Simonetta and Delle Cave (1975) provided the first reconstruction of this animal in a general monograph about Burgess Shale arthropods. The taxon was redescribed by Whittington (1981), and compared to Leanchoilia (Bruton and Whittington, 1983).



The single specimen reaches about 6.6 cm in length, excluding the anterior great appendages. The head has a trapezoidal head shield, a pair of large anterolateral pedunculate eyes, one pair of frontal appendages (the “great appendage”) and three pairs of smaller biramous appendages. The distal podomere of the great appendage has claws and two pairs of slim, flexible branches attached to the next podomere. The outer branch of the biramous limbs has a lobate shape fringed by lamellae and the inner branch is jointed and tapers distally. The trunk is composed of eleven segments, with the first eight showing clear lobes fringed by lamellae. The last segment terminates with a triangular and apparently smooth plate.


A single specimen is known of this species.

Maximum Size:
66 mm


Life habits: Mobile, Nektobenthic
Feeding strategies: Carnivorous
Ecological Interpretations:

Like similar and better known forms such as Leanchoilia and Alalcomenaeus the primary mode of locomotion of Actaeus was probably swimming, powered both by the wave-like fanning of its lateral flaps and flicks of its tail. The outer leg branches also served as gills. The long filaments of the great appendages were probably sensory and suggest an active predatory habit, consistent with the large stalked eyes. Actaeus most likely hunted organisms that lived either in or on the sediment.



BRUTON, D. L. AND H. B. WHITTINGTON. 1983. Emeraldella and Leanchoilia, two arthropods from the Burgess Shale, Middle Cambrian, British Columbia. Philosophical Transactions of the Royal Society of London, Series B, 300: 553-582.

BUDD, G. E. 2002. A palaeontological solution to the arthropod head problem. Nature, 417: 271-275.

CHEN, J. Y., D. WALOSZEK AND A. MAAS. 2004. A new ‘great-appendage’ arthropod from the Lower Cambrian of China and homology of chelicerate chelicerae and raptorial antero-ventral appendages. Lethaia, 37: 3-20.

COTTON, T. J. AND S. J. BRADDY. 2004. The phylogeny of arachnomorph arthropods and the origin of the Chelicerata. Transactions of the Royal Society of Edinburgh: Earth Sciences, 94: 169-193.

EDGECOMBE, G. D. 2010. Arthropod phylogeny: An overview from the perspectives of morphology, molecular data and the fossil record. Arthropod Structure & Development, 39: 74-87.

HOU, X. AND J. BERGSTRÖM. 1997. Arthropods of the Lower Cambrian Chengjiang fauna, southwest China. Fossils and Strata, 45: 1-116.

SIMONETTA, A. M. 1970. Studies on non trilobite arthropods of the Burgess Shale (Middle Cambrian). Palaeontographica Italica, 66: 35-45.

SIMONETTA, A. M. AND L. DELLE CAVE. 1975. The cambrian non Trilobite Arthropods from the Burgess Shale of British Columbia. A study of their comparative morphology taxonomy and evolutionary significance. Palaeontographia Italica, 69 1-37.

WHITTINGTON, H. B. 1981. Rare arthropods from the Burgess Shale, Middle Cambrian, British Columbia. Philosophical Transactions of the Royal Society of London Series B: Biological Sciences, 292: 329-357.

WILLS, M. A., D. E. G. BRIGGS, R. A. FORTEY, M. WILKINSON AND P. H. A. SNEATH. 1998. An arthropod phylogeny based on fossil and recent taxa, p. 33-105. In G. D. Edgecombe (ed.), Arthropod fossils and phylogeny. Columbia University Press, New York.

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