The Burgess Shale

Balhuticaris voltae

A large bivalved arthropod with an arch-like carapace

Balhuticaris voltae, holotype ROMIP 66238


Kingdom: Animalia
Phylum: Arthropoda
Higher Taxonomic Assignment: Hymenocarines, Family: Odaraiidae
Species name: Balhuticaris voltae

Hymenocarines were early arthropods with bivalved carapaces and mandibles, forming the bulk of the first mandibulates (represented today by myriapods, crustaceans and insects) (Aria and Caron 2017; Vannier et al. 2018). In many hymenocarines, including Balhuticaris, determining the exact number and types of appendages in their head remains difficult, which hinders a detailed understanding of the evolutionary relationships inside this group. Balhuticaris most probably belongs to the family Odaraiidae, a group of hymenocarines with highly multisegmented bodies, reduced or absent antennae and highly multisegmented legs.

Described by: Izquierdo-López & Caron
Description date: 2022

Balhuticarisfrom the mythological creature Balhut, a giant aquatic animal in some Persian cosmologies, and the latin caris, meaning “crab” or “shrimp”, and voltae- from the Catalan word volta, an arch-like structure.

Type Specimens: Holotype ROMIP66238
Other species:

Burgess Shale and vicinity: None
Other deposits: None

Age & Localities:

Middle Cambrian, Wuliuan Stage, upper part of the Burgess Shale Formation (Burgess Shale) (around 505 million years old)
Principal localities:

Marble Canyon, Tokumm Creek

History of Research:

Brief history of research:

Balhuticaris has been found from both the Marble Canyon and the Tokumm Creek localities of the Burgess Shale during several expeditions between 2012 to 2022. The different specimens of Balhuticaris were originally not recognized as belonging to the same organism. Instead, these were identified as different undescribed euarthropods or potential radiodonts (Nanglu et al. 2020). Balhuticaris was formally described in 2022 (Izquierdo-López and Caron 2022).



Balhuticaris is a large bivalved arthropod that can reach up to 25 cm in length. The carapace only covers the first quarter of the total body length. It has a dome-like shape. In frontal view, the carapace looks like an arch: each valve extends towards the ventral side of the animal, surpassing the length of the legs. The dorsal side of the carapace extends towards the posterior side of the animal, giving the valves a “bean-like” shape in lateral view. The head bears a pair of well-developed, pedunculate, bilobate eyes. The head also bears one pair of short antennulae and a sclerotized structure that may represent a head sclerite. The body is highly multisegmented, with approximately 110 segments posterior to the head. Approximately the first ten segments are longer, and bear legs that become smaller towards the head. All segments bear a pair of legs, each subdivided into two branches (biramous): a walking leg (endopod) and a paddle-like flap (exopod). The endopod is thin and subdivided into around 14 segments. The exopod is ovoid, almost as long as the endopod. The last segment is longer than the rest, and has a flattened triangular shape. This segment bears two paddle-like legs (caudal rami). Each of these is subdivided into three segments, bears three spines on their outer edge and elongated filaments (setae) on their posterior edge.


Balhuticaris is rare, only known from a dozen specimens from the Marble Canyon and Tokumm Creek sites.

Maximum Size:
About 25 cm


Life habits: Epibenthic, Nektobenthic
Feeding strategies: Suspension feeder, Carnivorous
Ecological Interpretations:

Balhuticaris is the largest bivalved arthropod to date, surpassing in length Tuzoia (Vannier et al. 2007) and Nereocaris exilis (Legg et al. 2012), and rivalling other arthropods from the Burgess Shale, such as radiodonts, including the largest complete Anomalocaris (Briggs 1975) and Cambroraster (Moysiuk and Caron 2019), but smaller than the estimated 50 cm long Titanokorys (Caron and Moysiuk 2021). The general anatomy of Balhuticaris, including its elongated body and large segmented caudal rami, indicates that it was probably a good swimmer. It was hypothesized that it could be swimming upside-down (Izquierdo-López and Caron 2022), similar to its relatives Fibulacaris and Odaraia (Briggs 1981; Izquierdo-López and Caron 2019). Balhuticaris’ feeding could have ranged from suspension-feeder to predator (Izquierdo-López and Caron 2022), similar to some of the largest fairy shrimps today (Fryer 1966).


  • ARIA, C. and CARON, J. B. 2017. Burgess Shale fossils illustrate the origin of the mandibulate body plan. Nature, 545: 89–92.
  • BRIGGS, D. E. G. 1975. Anomalocaris, the largest known Cambrian arthropod. Palaeontology, 22: 631–664.
  • BRIGGS, D. E. G. 1981. The arthropod Odaraia alata Walcott, middle Cambrian, Burgess Shale, British Columbia. Philosophical Transactions of the Royal Society of London. B, Biological Sciences, 291: 541–582.
  • CARON, J.-B. and MOYSIUK, J. 2021. A giant nektobenthic radiodont from the Burgess Shale and the significance of hurdiid carapace diversity. Royal Society Open Science, 8: 210664.
  • FRYER, G. 1966. Branchinecta gigas Lynch, a non‐filter‐feeding raptatory anostracan, with notes on the feeding habits of certain other anostracans. Proceedings of the Linnean Society of London, 177: 19–34.
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  • IZQUIERDO-LÓPEZ, A. and CARON, J. B. 2021. A Burgess Shale mandibulate arthropod with a pygidium: a case of convergent evolution. Papers in Palaeontology, 7: 1877–1894.
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  • NANGLU, K., CARON, J. and GAINES, R. 2020. The Burgess Shale paleocommunity with new insights from Marble Canyon, British Columbia. Paleobiology, 46(1): 58–81.
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