Marrella
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''Marrella'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of marrellomorph
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
known from the middle
Cambrian The Cambrian Period ( ; sometimes symbolized 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 years ago ...
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 fo ...
of
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, for ...
. It is the most common animal represented in the Burgess Shale, with tens of thousands of specimens collected. Much rarer remains are also known from deposits in China.


History

''Marrella'' was the first fossil collected by
Charles Doolittle Walcott Charles Doolittle Walcott (March 31, 1850February 9, 1927) was an American paleontologist, administrator of the Smithsonian Institution from 1907 to 1927, and director of the United States Geological Survey. Wonderful Life (book) by Stephen Jay G ...
from the Burgess Shale, in 1909. Walcott described ''Marrella'' informally as a "lace crab" and described it more formally as an odd
trilobite Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the ...
. It was later reassigned to the now defunct
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 differently ...
Trilobitoidea in the ''
Treatise on Invertebrate Paleontology The ''Treatise on Invertebrate Paleontology'' (or ''TIP'') published by the Geological Society of America and the University of Kansas Press, is a definitive multi-authored work of some 50 volumes, written by more than 300 paleontologists, and ...
''. In 1971,
Whittington Whittington may refer to: Places * Whittington, Victoria, Australia * Whittington, Illinois, United States England * Old Whittington, Derbyshire * New Whittington, Derbyshire * Whittington Moor, Derbyshire * Whittington, Gloucestershire * Whit ...
undertook a thorough redescription of the animal and, on the basis of its legs,
gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they ar ...
s and head appendages, concluded that it was neither a trilobite, nor a
chelicerate The subphylum Chelicerata (from New Latin, , ) constitutes one of the major subdivisions of the phylum Arthropoda. It contains the sea spiders, horseshoe crabs, and arachnids (including harvestmen, scorpions, spiders, solifuges, ticks, and mite ...
, nor a
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapoda, decapods, ostracoda, seed shrimp, branchiopoda, branchiopods, argulidae, fish lice, krill, remipedes, isopoda, isopods, barnacles, copepods, ...
. ''Marrella'' is one of several unique arthropod-like organisms found in the Burgess Shale. Other examples are '' Opabinia'' and ''
Yohoia ''Yohoia'' is an extinct genus of megacheiran arthropod from the Cambrian period that has been found as fossils in the Burgess Shale formation of British Columbia, Canada. It has been placed among the arachnomorpha, a group of arthropods that in ...
''. The unusual and varied characteristics of these creatures were startling at the time of discovery. The fossils, when described, helped to demonstrate that the soft-bodied Burgess fauna was more complex and diverse than had previously been anticipated.


Morphology

''Marrella'' itself is was small animal, with known specimens ranging from 2.4 mm to 24.5 mm in length. The head shield had two pairs of long posteriorly curved projections/spines, the posterior pair of which had a serrated margin. On the underside of the head was a pair of long and sweeping antennae, along with a pair of short and stout paddle-like swimming appendages, which were covered with
setae In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. ...
. ''Marrella'' has a body composed of 24–26 body segments ( tagma), each with a pair of branched (
biramous The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments (called podomeres) are of Latin origin, and may be confused with terms for bones: ''coxa'' (meaning hip, plu ...
) appendages. The lower branch of each appendage (the endopod) is elongate and leg-like, while the upper branch (the exopod) is a long, feathery gill. There is a tiny, button-like
telson The telson () is the posterior-most division of the body of an arthropod. Depending on the definition, the telson is either considered to be the final segment of the arthropod body, or an additional division that is not a true segment on acco ...
at the end of the
thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the c ...
.García-Bellido, Diego & Collins, Desmond. (2006). A new study of Marrella splendens (Arthropoda, Marrellomorpha) from the Middle Cambrian Burgess Shale, British Columbia, Canada. Canadian Journal of Earth Sciences. 43. 721-742. 10.1139/e06-012. A 1998 paper suggested that striations present on the front projection of well-preserved specimens of ''Marrella'' represented a
diffraction grating In optics, a diffraction grating is an optical component with a periodic structure that diffracts light into several beams travelling in different directions (i.e., different diffraction angles). The emerging coloration is a form of structur ...
pattern, that in life would have resulted in an
iridescent Iridescence (also known as goniochromism) is the phenomenon of certain surfaces that appear to gradually change color as the angle of view or the angle of illumination changes. Examples of iridescence include soap bubbles, feathers, butterfl ...
sheen. Dark stains are often present at the posterior regions of specimens, probably representing extruded waste matter or
hemolymph Hemolymph, or haemolymph, is a fluid, analogous to the blood in vertebrates, that circulates in the interior of the arthropod (invertebrate) body, remaining in direct contact with the animal's tissues. It is composed of a fluid plasma in which ...
.


Ecology

''Marrella'' is likely to have been an active swimmer that swam close to the seafloor (nektobenthic) with its swimming appendages used in a
backstroke Backstroke or back crawl is one of the four swimming styles used in competitive events regulated by FINA, and the only one of these styles swum on the back. This swimming style has the advantage of easy breathing, but the disadvantage of swimm ...
motion, with the large spines acting as stabilizers, as well as possibly also having a defensive function. They have been suggested to be
filter feeders Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feedin ...
, with food particles sifted out of the water column by the posterior appendages during swimming before being passed forward by the appendages towards the mouth.


Taxonomy

''Marrella'' is placed within the Marrellida clade of the Marrellomorpha, a group of arthropods with uncertain affinities known from the
Cambrian The Cambrian Period ( ; sometimes symbolized 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 years ago ...
to
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
. Within the Marrelliida, is it placed as most basal known member of the group.


Occurrence

Marrella is the most abundant genus in the
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 fo ...
. Most ''Marrella'' specimens herald from the 'Marrella bed', a thin horizon, but it is common in most other outcrops of the shale. Over 25,000 specimens have been collected;5028 specimens of ''Marrella'' are known from the Greater
Phyllopod bed The Phyllopod bed, designated by USNM locality number 35k, is the most famous fossil-bearing member of the Burgess Shale fossil ''Lagerstätte''. It was quarried by Charles Walcott from 1911–1917 (and later named Walcott Quarry), and was t ...
, where they comprise 9.56% of the community. A few dozen specimens of an indeterminate species of ''Marrella'' have been reported from the Kaili Formation of Yunnan, China, dating to the Wuliuan stage of the Cambrian. A single fragmentary specimen of an indeterminate species is also known from the
Balang Formation The Balang formation, is a Cambrian Period geological formation, which outcrops in western Hunan and eastern Guizhou in southern China. It contains Burgess Shale-type soft-bodied fossils. It is intermediate in age between the Chengjiang and Kai ...
of Yunnan, China, dating to
Cambrian Stage 4 Cambrian Stage 4 is the still unnamed fourth stage of the Cambrian and the upper stage of Cambrian Series 2. It follows Cambrian Stage 3 and lies below the Wuliuan. The lower boundary has not been formally defined by the International Commission ...
. Both deposits are earlier than the Burgess Shale.


References


External links

* * {{Taxonbar, from=Q12015622 Marrellomorpha Burgess Shale fossils Cambrian arthropods of North America Cambrian genus extinctions