Balhuticaris Temporal range: Cambrian, | |
---|---|
Life restoration | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Order: | † Hymenocarina |
Genus: | † Balhuticaris Izquierdo-López & Caron, 2022 |
Type species | |
†Balhuticaris voltae Izquierdo-López & Caron, 2022 |
Balhuticaris is a genus of extinct bivalved (referring to the carapace) hymenocarine arthropod that lived in the Cambrian aged Burgess Shale in what is now British Columbia around 506 million years ago. This extremely multisegmented (with over 100 segments) arthropod is the largest member of the group, and it was even one of the largest animals of the Cambrian, with individuals reaching lengths of 245 mm (9 in). Fossils of this animal suggests that gigantism occurred in more groups of Arthropoda than had been previously thought. [1] It also presents the possibility that bivalved arthropods were very diverse, and filled in a lot of ecological niches. [1] [2] [3]
The hymenocarines were an order of primitive mandibulates, the arthropod group that includes crustaceans, insects, myriapods and their relatives, that lived throughout the Cambrian period. [2] This group was extremely diverse and attained a wide variety of ecological niches and body plans. [2] [3] Several dozen species are known from deposits of Cambrian, and ranged in size from smaller species like Fibulacaris nereidis reaching a length of 2 cm (0.79 in) long, [2] to larger ones like B. voltae. [1]
This arthropod was described in 2022 based on 11 specimens found in the Burgess Shale between 2014 and 2018, more specifically in the Marble canyon locality. By 2020, scientists realized that these fossils represented a new species. Because of how they were preserved the fossils were found two dimensional in several carbonaceous films. The holotype specimen and several others are nearly or fully complete with possible neural and other soft tissues having been preserved. Balhuticaris is named after Balhūt, a giant fish from Persian cosmography, as well as the Latin caris ("crab"). The specific epithet voltae is derived from the Catalan volta, meaning vault, referring to the shape of the carapace when seen from the front. [1]
Balhuticaris was the largest bivalved arthropod in the fossil record, beating the previous holders of this title Nereocaris exilis and Tuzoia. This animal's body was very long, and had extreme segmentation compared to other Cambrian arthropods, with over 100 distinct segments. In total this creature had about 110 pairs of biramous limbs, the most of any Cambrian-aged arthropod. Covering the head of this creature was a large carapace that resembles an arch or other curved structure. This structure only covers the frontmost part of this arthropod but it does extend ventrally beyond its appendages. [1]
In several studies performed, Balhuticaris was found to be a member of the Hymenocarina. More specifically it was found to be most closely related to the genus Odaraia and its relatives. Although they are normally regarded as pancrustaceans, this study found the Hymenocarina to occupy a more basal branch of the mandibulates. [1]
This cladogram shows the position of B. voltae in relation to other arthropods by López et al., 2022. [1]
This hymenocarine most likely engaged in a fast-paced nektonic (free swimming) lifestyle. Its large size means that it was safe from most of the other predatory fauna of its environment. Many features of the fossils evidence a free swimming, pelagic lifestyle. Examples being the presence of a tripartite caudal rami, a feature only found in hymenocarines, [1] [2] and that the carapace goes ventrally beyond the legs, which would have heavily impaired this arthropods ability to crawl on the ocean floor. Its eyes also have a similar shape seen in modern pelagic crustaceans. What this arthropod ate has been a difficult question to answer due to the lack of cephalic appendages in the fossils. Modern day arthropods of a similar size like lobsters, stomatopods, and giant isopods are mainly scavengers or predators. B. voltae however does not possess features that would suggest this, like chelate limbs and gnathobases. [1] [4] Suspension and deposit feeding can also be readily ruled out due to lack of features needed for these lifestyles in B. voltae. Currently it is thought to have suctioned in prey in water currents through a ventral groove. The animal probably swam while it fed, similar to leptostracan and anostracan crustaceans. [1] [5] [6] It is possible that this animal swam in an upside down, or in an inverted position. [1] This is not unheard of, as many other free-swimming arthropods like anostracans, [6] pelagic trilobites like the Telephinids, [7] xiphosurans, [1] and other odaraiid hymenocarines, like Odaraia and Fibulacaris swam in inverted positions. [1] [2]
The Burgess Shale is a middle Cambrian aged Lagerstätte that lies in British Columbia in Canada. [8] This site was the first of its kind to have been discovered and provided great insights into the soft bodied fauna of the early Paleozoic. [9] Dozens of creatures have been preserved at this site including lobopodians, stem-group and total-group Arthropoda, worms, primitive chordates, echinoderms, sponges, as well as other animal groups. [10] [11] This animal was one of the largest of its time, with only the giant radiodonts like Anomalocaris surpassing it in size. [1] More specifically, this animal was found in Marble Canyon. This Lagerstätte produced new taxa including other hymenocarines like Tokummia , Fibulacaris and Pakucaris , large hurdiid radiodonts like Cambroraster and Titanokorys , as well as other arthropods like the megacheiran genus Yawunik , the isoxyid genus Surusicaris , and the basal chelicerate Mollisonia plenovenatrix . Worms have also been found at this site, like the annelid Kootenayscolex . Well-preserved specimens of primitive chordate Metaspriggina are also known from there. [1] [12] [13] [14] [15] [16] [17] [18] [19] [20]
Canadaspis is an extinct genus of bivalved Cambrian arthropod, known from North America and China. They are thought to have been benthic feeders that moved mainly by walking and possibly used its biramous appendages to stir mud in search of food. They have been placed within the Hymenocarina, which includes other bivalved Cambrian arthropods.
Waptia is an extinct genus of arthropod from the Middle Cambrian of North America. It grew to a length of 6.65 cm (3 in), and had a large bivalved carapace and a segmented body terminating into a pair of tail flaps. It was an active swimmer and likely a predator of soft-bodied prey. It is also one of the oldest animals with direct evidence of brood care. Waptia fieldensis is the only species classified under the genus Waptia, and is known from the Burgess Shale Lagerstätte of British Columbia, Canada. Specimens of Waptia are also known from the Spence Shale of Utah, United States.
Odaraia is a genus of bivalved arthropod from the Middle Cambrian. Its fossils, which reach 15 centimetres (5.9 in) in length, have been found in the Burgess Shale in British Columbia, Canada.
Megacheira is an extinct class of predatory arthropods defined by their possession of spined "great appendages". Their taxonomic position is controversial, with studies either considering them stem-group euarthropods, or stem-group chelicerates. The homology of the great appendages to the cephalic appendages of other arthropods is also controversial. Uncontested members of the group were present in marine environments worldwide from the lower to middle Cambrian.
Plenocaris plena is a genus of extinct bivalved hymenocarine arthropod that lived in the Cambrian aged Burgess Shale and Chengjiang. Originally described as a species of Yohoia by Walcott in 1912, it was placed into its own genus in 1974.
Clypecaris is genus of bivalved Cambrian arthropod known from the Chengjiang biota of Yunnan, China. The genus was initially described for the type species C. pteroidea by Hou, 1999. A second species C. serrata was described by Yang et al. in 2016. The species are primarily distinguished by the presence of a serrated edge on the front of the carapace of C. serrata. C. serrata is noted for the modification of an anterior pair of limbs into spined grasping appendages, indicating a predatory lifestyle. It is unknown whether a similar structure was present in C. pteroidea.Clypecaris is considered to likely be a member or a close relative of Hymenocarina, and is closely related to Perspicaris. As well as to Ercaicunia.
Ercaicunia is genus of bivalved Cambrian arthropod from the Chengjiang biota of Yunnan, China. It contains a single species, E. multinodosa that was described by Luo et al. in 1999. The total length of the body ranges from 8 to 11 millimetres. The bivalved carapace covered about a third of the total body-length, and has up to six serrations on its forward edge. The head has a pair of large uniramous antennae, as well as a smaller pair of secondary antennae, as well as pair of mandibles and maxillae. The trunk has 16 pairs of biramous appendages. Specimens were CT scanned in 2019, which suggested it to be a stem-group crustacean. Other subsequent studies have recovered it as a member of Hymenocarina, which contains other bivalved Cambrian arthropods.
Haikoucaris is a genus of megacheiran arthropod that contains the single species Haikoucaris ercaiensis. It was discovered in the Cambrian Chengjiang biota of China.
Pectocaris is an extinct genus of bivalved arthropods from the Cambrian Maotianshan Shales, Yunnan Province of China. There are currently four known species within the genus.
Radiodonta is an extinct order of stem-group arthropods that was successful worldwide during the Cambrian period. Radiodonts are distinguished by their distinctive frontal appendages, which are morphologically diverse and used for a variety of functions. Radiodonts included the earliest large predators known, but they also included sediment sifters and filter feeders. Some of the most famous species of radiodonts are the Cambrian taxa Anomalocaris canadensis, Hurdia victoria, Peytoia nathorsti, Titanokorys gainesi, Cambroraster falcatus and Amplectobelua symbrachiata. The later surviving members include the subfamily Aegirocassisinae from the Early Ordovician of Morocco and the Early Devonian member Schinderhannes bartelsi from Germany.
Marble Canyon is a canyon surrounding Tokumm Creek just above its confluence with the Vermilion River, at the north end of Kootenay National Park in the Canadian Rockies of British Columbia, Canada. South of Marble Canyon on Highway 93 is Numa Falls on the Vermillion River.
Cucumericrus ("cucumber-leg") is an extinct genus of stem-arthropod. The type and only species is Cucumericrus decoratus, with fossils discovered from the Maotianshan Shales of Yunnan, China.
Hurdiidae is an extinct cosmopolitan family of radiodonts, a group of stem-group arthropods, which lived during the Paleozoic Era. It is the most long-lived radiodont clade, lasting from the Cambrian period to the Devonian period.
Tokummia is a genus of fossil hymenocarine arthropod, known only by one species, Tokummiakatalepsis, from the middle Cambrian Burgess Shale as found in a quarry in Marble Canyon in Canada.
Hymenocarina is an order of extinct arthropods known from the Cambrian. They possess bivalved carapaces, typically with exposed posteriors. Members of the group are morphologically diverse and had a variety of ecologies, including as filter feeders and as predators. Recent research has generally considered them to be stem or crown group members of Mandibulata, due to the presence of mandibles in some species.
Titanokorys is a genus of extinct hurdiid (peytoiid) radiodont that existed during the mid Cambrian. It is the largest member of its family from the Cambrian, with a body length of 50 cm (20 in) long, making it one of the largest animals of the time. It bears a resemblance to the related genus Cambroraster. Fossils of T. gainesi were first found within the Marble Canyon locality within the Burgess Shale in 2018. The fossils were not named until 2021 because they were assumed to be giant specimens of Cambroraster.
Fibulacaris is a monotypic genus of fossil arthropod known only by one species, Fibulacaris nereidis, discovered from the Cambrian Burgess Shale of Canada. It was characterized by a bivalved carapace with an inverted rostrum, sandwiching the slender body with stalked eyes and homonomous appendages. It was probably an actively swimming filter feeder and possibly swam upside-down like some branchiopod crustaceans and horseshoe crabs. Phylogenetic analysis suggest it was a relative or member of Hymenocarina, which contains other bivalved arthropods.
Nereocaris is an extinct genus of bivalved hymenocarine arthropod that lived in the Cambrian aged Burgess Shale in what is now British Columbia around 506 million years ago. Two species are known.
Isoxyids are members of the order Isoxyida and the family Isoxyidae, a group of basal arthropods that existed during the Cambrian period. It contains two genera, Isoxys, with 20 species found worldwide, and Surusicaris known from a single species found in the Burgess Shale of Canada. They are distinguished by their bivalved carapaces and pair of upward curving grasping frontal appendages.
Cordaticaris is a genus of extinct hurdiid (peytoiid) radiodont that lived in what is now northern China during the middle Cambrian period. This animal was described in 2020 based on remains found in the Zhangxia Formation, located in the Shandong Province. It is differentiated from other members of its family by its unique heart-shaped frontal sclerite, and its frontal appendages bearing nine endites and seven more elongated subequal endites. This animal was important as it was the first Miaolingian aged hurdiid known from rock layers outside of laurentia, allowing paleontologists to get a better grasp of this families geographic range in life.