Naraoia
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''Naraoia'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of small to average size (about 2-4½ cm long) marine
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s within the family
Naraoiidae Naraoiidae is a family, of extinct, soft-shelled trilobite-like arthropods, that belongs to the order Nectaspida. Species included in the Naraoiidae are known from the second half of the Lower Cambrian to the end of the Upper Silurian. The tota ...
, that lived from the early
Cambrian The Cambrian Period ( ; sometimes symbolized C with bar, Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million ...
to the late
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleozo ...
period. The species are characterized by a large alimentary system and sideways oriented antennas.


Etymology

The name is derived from Narao, the name of a group of small lakes in Cataract Brook canyon, above Hector on the Canadian Pacific Railway,
British Columbia British Columbia (commonly abbreviated as BC) is the westernmost province of Canada, situated between the Pacific Ocean and the Rocky Mountains. It has a diverse geography, with rugged landscapes that include rocky coastlines, sandy beaches, ...
, Canada.Walcott, C.D. Middle Cambrian Branchiopoda, Malacostraca, Trilobita, and Merostomata. In: Cambrian Geology and Paleontology II. Smithsonian. 1914. s:Cambrian Geology and Paleontology/Volume 2/Middle Cambrian Branchiopoda, Malacostraca, Trilobita, and Merostomata


History of the classification

When the fossil was first discovered in
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...
's
Burgess Shale The Burgess Shale is a fossil-bearing deposit exposed in the Canadian Rockies of British Columbia, Canada. It is famous for the exceptional preservation of the soft parts of its fossils. At old (middle Cambrian), it is one of the earliest foss ...
, it was believed to be a
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
, such was the difference between this and other trilobites. Its continuous shield hid most of its structure, interfering with proper classification. When
Harry B. Whittington Harry Blackmore Whittington FRS (24 March 1916 – 20 June 2010) was a British palaeontologist who made a major contribution to the study of fossils of the Burgess Shale and other Cambrian fauna. His works are largely responsible for the conce ...
began dissecting some specimens (''Naraoia'' was among the most populous of the Burgess Shale animals), he discovered that the legs (and gills) of the beasts were very similar, if not identical to those of trilobites, thus the current placement of ''Naraoia'' in
class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differentl ...
Trilobita Trilobites (; meaning "three lobes") are extinction, extinct marine arthropods that form the class (biology), class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil reco ...
. ''Misszhouia longicaudata'' was formerly considered a member of the genus ''Naraoia'', originally known as ''N. longicaudata'', until separated in 1997.


Description

''Naraoia'' is almost flat (dorso-ventrally). The upper (or dorsal) side of the body consists of a non-calcified transversely oval or semi-circular headshield (
cephalon Cephalon, Inc. was an American biopharmaceutical company co-founded in 1987 by pharmacologist Frank Baldino, Jr., neuroscientist Michael Lewis, and organic chemist James C. Kauer—all three former scientists with the DuPont Company. Baldino s ...
), and a tailshield (
pygidium The pygidium (plural pygidia) is the posterior body part or shield of crustaceans and some other arthropods, such as insects and the extinct trilobites. In groups other than insects, it contains the anus and, in females, the ovipositor. It is compo ...
) longer than the cephalon, without any body segments in between. The body is narrowed at the articulation between cephalon and pygidium. The long many-segmented antennas are directed sideways. There are no eyes. The gut has a relatively large diameter (14-18% of the width of the body), and next to four pairs of large digestive sacs (or caeca). The cephalon has branched diverticula occupying most of the cephalon (unlike in ''
Misszhouia ''Misszhouia'' is a genus of small to average sized (up to long) marine arthropods within the Naraoiidae family, that lived during the early Cambrian period. The only species presently known is ''Misszhouia longicaudata'' (the genus is mono ...
''). Naraoia had appendages with two branches on a common basis, like ''Misszhouia'' and trilobites. At least the anterior trunk limbs have exopods with large, paddle-shaped distal lobes and short flattened side branches (setae) on the shaft. The endopod (known only in ''N. compacta'') is composed of six podomeres.Chen, J.-Y., G.D. Edgecombe and L. Ramskjöld. Morphological and ecological disparity in naraoiids (Arthropoda) from the Early Cambrian Chengjiang Fauna, China. Records of the Australian Museum 49(1), pp. 1-24. 199

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Distribution

Species of ''Naraoia'' are known from Canada, the United States, South China, and Australia, occurring in deposits ranging from the Lower Cambrian (
Atdabanian Cambrian Stage 3 is the still unnamed third stage of the Cambrian. It succeeds Cambrian Stage 2 and precedes Cambrian Stage 4, although neither its base nor top have been formally defined. The plan is for its lower boundary to correspond approxim ...
) to the late Upper Silurian ( Pridoli).


Ecology

Sediments present in the gut of ''Naraoia'' suggest that it may have been a
deposit feeder Detritivores (also known as detrivores, detritophages, detritus feeders, or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrates, ...
, eating large amounts of soil, like an
earthworm An earthworm is a terrestrial invertebrate that belongs to the phylum Annelida. They exhibit a tube-within-a-tube body plan; they are externally segmented with corresponding internal segmentation; and they usually have setae on all segments. Th ...
. A very large, complex system of gut diverticula and a gut with great holding capacity indicate that the diet of ''Naraoia spinosa'' was low in
nutrition Nutrition is the biochemical and physiological process by which an organism uses food to support its life. It provides organisms with nutrients, which can be metabolized to create energy and chemical structures. Failure to obtain sufficient n ...
. On the other hand, the morphology of the digestive system has also been interpreted as representing a predatory habit. The large, paddle-shaped distal lobes and short lamellar setae on the exopods, the implanting of the antennas to the side, and the angle of the cephalon with the pygidium of up to 90° with which many specimen are found, all agree with a life of burrowing.


Habitat

All naraoiids were probably marine bottom dwellers.


Key to the species


References


External links

*
The Smithsonian Intstitution's ''Naraoia'' page

Genus †Naraoia - Hierarchy - The Taxonomicon
{{Taxonbar, from=Q283508 Nektaspida Burgess Shale animals Maotianshan shales fossils Cambrian first appearances Silurian extinctions Wheeler Shale Cambrian genus extinctions