Habelia Temporal range: | |
---|---|
Life restoration | |
Fossils of Habelia optata | |
Scientific classification | |
Kingdom: | |
Phylum: | |
(unranked): | |
Order: | |
Family: | †Habeliidae |
Genus: | †Habelia Walcott, 1912 |
Species: | †H. optata |
Binomial name | |
†Habelia optata Walcott, 1912 | |
Habelia is a genus of extinct arthropod from the Middle Cambrian, thought to be one of the earliest known relatives of chelicerates. Its fossils have been found in the Burgess Shale in British Columbia, Canada. Fifty-four specimens of Habelia are known from the Greater Phyllopod bed, where they comprise 0.1% of the community. [1]
The body is divided into two main segments. The cephalon ("head")/prosoma with seven segments/8 somites, and the trunk with 12 segments, making for a total of 19 body segments. The body is around 8.5 to 34 millimetres (0.33 to 1.34 in) in length excluding the long telson that terminates the body. The head, which is 25% wider than it is long, makes up about 30% of the total body length, and has a bulging shape, with triangular genal spines projecting outward at its edges. It housed the stomach of the animal. The head bore a pair of spherical compound eyes, which were accommodated by notches in the head shield. A hypostome-labrum complex was present, with the mouth openling vertically close to the front of the head. The first pair of appendages was very short and probably flexible, this was followed by 5 pairs of appendages that bore large gnathobases (modified basipod/base of the appendages used to process food), increasing in size posteriorly which bore staggered rows of strongly sclerotized teeth on their inner edge. The endopods of these limbs increased in size posteriorly, and had seven segments/podomeres, with the terminating segment being a claw, with podomeres 5 and 6 having borne a brush of inward facing setae. The exopods of these limbs were elongate and thin rami, increasing in size posteriorly, each composed of 7 or more podomeres, which bore setae at the segment joints. [2]
There is one pair of biramous limbs on the margin between the head and trunk, which is morphologically similar to the trunk limbs. The exopods of the biramous trunk limbs are paddle-shaped and fringed with thin lamellae/setae, while the endopods are similar to those of the cephalon in being 7 segmented and clawed, with two smaller claws on segment 6. The basipods of these limbs were cylindrical and broad, and did not function as gnathobases. The trunk is subdivided into the anterior thorax with 5 segments and the posterior post-thorax with 7 segments. Some specimens bear three pairs of upward pointing dorsal spines along the first three trunk segments. This may have been a sexually dimorphic feature. The 5 segments of the thorax each bear biramous limbs. The segments of the post-thorax bore pairs of exopods decreasing in size posteriorly, but there are no visible endopods, meaning that they were either reduced or entirely absent. The body ended with a long rod-shaped telson. [2]
While previously enigmatic, a 2017 redescription found that it formed a clade (Habeliida) with Sanctacaris, Utahcaris, Wisangocaris and Messorocaris as a stem-group to Chelicerata, with Habelia being placed in the monotypic family Habellidae within Habeliida. [2]
Habelia is thought to have been a durophagous predator, using its gnathobases to feed on hard shelled organisms. [2]
Naraoiidae is a family, of extinct, soft-shelled trilobite-like arthropods, that belongs to the order Nektaspida. Species included in the Naraoiidae are known from the second half of the Lower Cambrian to the end of the Upper Silurian. The total number of collection sites is limited and distributed over a vast period of time: Maotianshan Shale and Balang Formation (China), Burgess Shale and Bertie Formation (Canada), the Šárka Formation, Emu Bay Shale (Australia), Idaho and Utah (USA). This is probably due to the rare occurrence of the right circumstances for soft tissue preservation, needed for these non-calcified exoskeletons.
Marrella is an extinct genus of marrellomorph arthropod known from the Middle Cambrian of North America and Asia. It is the most common animal represented in the Burgess Shale of British Columbia, Canada, with tens of thousands of specimens collected. Much rarer remains are also known from deposits in China.
Sanctacaris is a Middle Cambrian arthropod from the Burgess Shale of British Columbia. It was most famously regarded as a stem-group chelicerate, a group which includes horseshoe crabs, spiders and scorpions, although subsequent phylogenetic studies have not always supported this conclusion. Its chelicerate affinities regain support in later observations, alongside the reassignment of Habelia optata as a sanctacaridid-related basal chelicerate. It has been placed as a member of the extinct family Sanctacarididae alongside Wisangocaris and Utahcaris.
Sidneyia is an extinct arthropod known from fossils found from the Early to the Mid Cambrian of China and the Mid Cambrian Burgess Shale of British Columbia, Canada.
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.
Misszhouia is a genus of small to average sized marine trilobite-like arthropods within the Naraoiidae family, that lived during the early Cambrian period. The species are M. longicaudata, from the Maotianshan Shales, described in 1985, and M. canadensis, from the Burgess Shale and described in 2018, although later species may belongs to genus Naraoia instead.
Alalcomenaeus is one of the most widespread and longest-surviving arthropod genera of the Early and Middle Cambrian. Known from over 300 specimens in the Burgess Shale and the Chengjiang biota. It is a member of the family Leanchoiliidae in the group Megacheira.
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 Cambrian to the upper Ordovician.
Chuandianella ovata is an extinct bivalved arthropod that lived during Cambrian Stage 3 of the Early Cambrian. It is the only species classified under the genus Chuandianella. Its fossils were recovered from the Chengjiang Biota in Yunnan, China.
Haikoucaris is a genus of megacheiran arthropod that contains the single species Haikoucaris ercaiensis. It was discovered in the Cambrian Chengjiang biota of China.
Isoxys is a genus of extinct bivalved Cambrian arthropod; the various species of which are thought to have been freely swimming predators. It had a pair of large spherical eyes, and two large frontal appendages used to grasp prey.
Molaria is a genus of Cambrian arthropod, the type species M. spinifera is known from the Middle Cambrian Burgess Shale. 144 specimens of Molaria are known from the Greater Phyllopod bed, where they comprise 0.27% of the community. A second species M. steini was described from the Sirius Passet in Greenland in 2017.
Eoredlichia is an extinct genus of trilobite of average to large size. It lived during the early Cambrian in the Chengjiang fauna of Yunnan, China, and in Australia and Thailand. Eoredlichia is compounded of the Greek ἠώς and Redlichia, a later but related genus, so it means "early Redlichia". The species epithet intermedia means intermediate, indicating it is morphologically intermediate between other species. Eofallotaspis gives rise to Lemdadella, and thence to Eoredlichia and the other Redlichiidae.
Tegopelte gigas is a species of large soft-bodied arthropod known from two specimens found in the Middle Cambrian Burgess Shale of British Columbia, Canada.
Retifacies abnormalis is an extinct arthropod that lived in the lower Cambrian. Its fossil remains have been found in the Maotianshan Shales of Yunnan, China. It is a member of the Artiopoda, and closely related to Pygmaclypeatus.
Cheloniellon is a monotypic genus of cheloniellid arthropod, known only by one species, Cheloniellon calmani, discovered from the Lower Devonian Hunsrück Slate of Germany.
Erratus is an extinct genus of marine arthropod from the Cambrian of China. Its type and only species is Erratus sperare. Erratus is likely one of the most basal known arthropods, and its discovery has helped scientists understand the early evolution of arthropod trunk appendages. Some of the stem-arthropods like radiodonts did not have legs, instead they had flap like appendages that helped them swim. Erratus on the other hand had not only flaps but also a set of primitive legs. It also supported the theory that the gills of aquatic arthropods probably evolved into the wings and lungs of terrestrial arthropods later in the Paleozoic.
Jugatacaris is an extinct genus of bivalved arthropod known from a single species, Jugatacaris agilis found in the Cambrian Stage 3 aged Chengjiang biota of Yunnan, China. The carapace is around 28 to 37 millimetres in length, with a pronounced ridge at the top of the carapace separating the two valves, which formed a fin-like structure raised above the carapace. The head has a pair of stalked eyes, as well as a dumbbell shaped medial eye between them. The head also bore a pair of mandibles as well as at least one and possibly two pairs of antennules. The trunk had up to 65 segments, each with biramous appendages. The appendages had thin endopods with 30 podomeres, each bearing a spiny endite, with the endopods ending with a terminal claw. The appendages also had overlapping flap-like exopods, which are elongated, being at maximum eight times as long as they are wide, which on their posterior edge are covered with setae. The trunk ended with a forked tail. It was likely an actively swimming filter feeder, using its constantly beating appendages to sift food from the water column, which was then passed forward along the U-shaped food groove between the appendage pairs towards the mouth. While initially placed as a crustaceanomorph, later studies considered to be a member of Hymenocarina, which contains numerous other similar bivalved Cambrian arthropods.
Surusicaris is an extinct genus of bivalved arthropod, known from the Cambrian Burgess Shale of British Columbia, Canada. It is considered to be closely related to Isoxys, and like it has spined grasping frontal appendages.