Pikaia gracilens is an extinct, primitive chordate marine animal known from the Middle Cambrian Burgess Shale of British Columbia. Described in 1911 by Charles Doolittle Walcott as an annelid, and in 1979 by Harry B. Whittington and Simon Conway Morris as a chordate, it became "the most famous early chordate fossil", or "famously known as the earliest described Cambrian chordate". It is estimated to have lived during the latter period of the Cambrian explosion. Since its initial discovery, more than a hundred specimens have been recovered.
The body structure resembles that of the lancelet and it swam perhaps much like an eel. A notochord and myomeres (segmented blocks of skeletal muscles) span the entire length of the body, and are considered the defining signatures of chordate characters. Its primitive nature is indicated by the body covering, a cuticle, which is characteristic of invertebrates and some protochordates. A reinterpretation in 2024 found evidence of the gut canal, dorsal nerve cord and myomeres, and suggested that the taxon was previously interpreted upside down.
The exact phylogenetic position is unclear, though recent studies suggest that it is likely a stem-chordate with crown group traits. Previously proposed affinities include those of cephalochordata, craniata, or a stem-chordate not closely related to any extant lineage."Pikaia gracilens" Burgess Shale Fossil Gallery. Virtual Museum of Canada. 2011. Popularly but incorrectly attributed as an ancestor of all vertebrates, or the oldest fish, or the oldest ancestor of humans, it is generally viewed as a basal chordate alongside other Cambrian chordates; it is a close relative of vertebrate ancestors but it is not an ancestor itself.
Discovery
The fossils of Pikaia gracilens was discovered by Charles Walcott from the Burgess Shale member of the Stephen Formation in British Columbia, and described it in 1911. He named it after Pika Peak, a mountain in Alberta, Canada. Based on the obvious and regular segmentation of the body, as is the feature of annelids, Walcott classified it as a polychaete worm and created a new family Pikaidae for it. (Princeton palaeontologist Benjamin Franklin Howell changed the name of the family to Pikaiidae in 1962.) Walcott was aware of the limitation of his classification, as he noted: "I am unable to place it within any of the families of the Polychaeta, owing to the absence of parapodia [paired protrusions on the sides of polychaete worms] on the body segments back of the fifth."
University of Cambridge palaeontologist Harry B. Whittington and his student Simon Conway Morris re-examined the Burgess Shale fauna and noted the anatomical details of Pikaia for the first time. The fossil specimens bears features of notochord and muscle blocks that are fundamental structures of chordates, and not of annelids. In 1977, Conway Morris presented a paper that indicated the possible chordate position, without further explanation. He and Whittington were convinced that the animal was obviously a chordate, as they wrote in Scientific American in 1979:Finally, we find among the Burgess Shale fauna one of the earliest-known invertebrate representatives of our own conspicuous corner of the animal kingdom: the chordate phylum... The chordates are represented in the Burgess Shale by the genus Pikaia and the single species P. gracilens.Conway Morris formally placed P. gracilens among the chordates in a paper in the Annual Review of Ecology and Systematics that same year. However, he provided no structural analyses such as using microscopes to confirm the chordate features. The comparative description only earned a "putative" chordate status. The fossil's chordate nature was received sceptically for several decades. Only in 2012, when detailed analysis was reported by Conway Morris and Jean-Bernard Caron, that the chordate position became generally accepted.
The fossils are found only in a restricted series of horizons in the strata exposed on Fossil Ridge, close to the Yoho National Park. From the same location, other fish-like animal fossils named Metaspriggina were discovered in 1993. Conway Morris identified the animals as another Cambrian chordate. The fossil specimens are preserved in the Smithsonian Institution and the Royal Ontario Museum.
Description Pikaia has a lancelet-like body, tapering at both ends, laterally flat and lacked a well-defined head. It measures an average of about in length. Walcott recorded the longest individuals as in length. Pikaia has a pair of large, antenna-like tentacles on its head that resembles those of invertebrates such as snails. The attachment of the tentacles makes a two-lobed structure of the head. The tentacles may be comparable to those in the present-day hagfish, a jawless chordate. It has a small circular mouth that could be used to eat small food particles in a single bite. There is a series of short appendages on either side of the underside of the head just after the mouth, and their exact nature or function is unknown. The pharynx is associated with six pairs of slits with tiny filaments that could be used for respiratory apparatus. In these ways, it differs from the modern lancelets, which have distinct pharyngeal gill slits on either sides of the pharynx and are used for filter feeding.
A major primitive structure of Pikaia is a cuticle as its body covering. Cuticle is a hard protein layer predominantly found in invertebrates such as arthropods, molluscs, echinoderms and nematodes. Unlike a typical cuticle, the cuticle of Pikaia does not have hard extracellular (exoskeleton) protection, and the entire body is essentially soft-bodied. Although primitive, Pikaia shows the essential prerequisites for vertebrates. When alive, Pikaia was a compressed, leaf-shaped animal with an expanded tail fin; the flattened body is divided into pairs of segmented muscle blocks, seen as faint vertical lines. The muscles lie on either side of a flexible structure resembling a rod that runs from the tip of the head to the tip of the tail.Palmer, D., (2000). The Atlas of the Prehistoric World. London: Marshall Publishing Ltd. p66-67.
Pikaia was an active and free swimmer. It likely swam by throwing its body into a series of S-shaped, zigzag curves, similar to the movement of eels; fish inherited the same swimming movement, but they generally have stiffer backbones. These adaptations may have allowed Pikaia to filter particles from the water as it swam along. Pikaia was probably a slow swimmer, since it lacked the fast-twitch fibers that are associated with rapid swimming in modern chordates.
Reinterpretations
Walcott's original summary of the description of Pikaia reads:Body elongate, slender, and tapering at each end. It is formed of many segments that are defined by strong annular shiny lines. Head small with two large eyes and two tentacles... Back of the head the first five segments carry short parapodia that appear to be divided into two parts. The enteric canal extends from end to end without change in character... This was one of the active, free-swimming annelids that suggest the Nephthydidae of the Polychaeta.Whittington and Conway Morris were the first to realise that Walcott's description and classification were not reliable and mostly inaccurate. They compared the body segments as described by Walcott with living animals and found that they were similar to the muscle bundles of chordates such as the living Amphioxus (Branchiostoma) as well as fishes, and not to superficial segments of annelids. They pictured that the muscles would be essential for swimming in water in wriggling motions. The enteric canal as observed by Walcott was not an ordinary digestive tract, it runs along with a stiff rod that resembles a notochord. They reported in 1979: "Although Pikaia differs from Amphioxus, in several important respects, the conclusion is that it is not a worm but a chordate appears inescapable."
Conway Morris was convinced that the longitudinal rod was a notochord and the segments were muscle blocks that he concluded that Pikaia "is a primitive chordate rather than a polychaete. The earliest fish scales are Upper Cambrian, and Pikaia may not be far removed from the ancestral fish." In 1982, he added further description in his Atlas of the Burgess Shale that Pikaia had one or more fins, but did not specify where they were present.
Pikaia was not popularly known as a chordate fossil or as an ancient chordate until 1989. That year, Harvard University palaeontologist Stephen Jay Gould wrote in his book Wonderful Life: The Burgess Shale and the Nature of History: "Pikaia is not an annelid worm. It is a chordate, a member of our own phylum—in fact, the first recorded member of our immediate ancestry." From this remark Pikaia became generally recognised as a chordate and ancestor of vertebrates.
In 1993, Conway Morris came up with another possible chordate feature. He identified structures that looked like gill slits but gave a cautious remark: "[They] may have been present, but are hard to identify with certainty in the compressed material available. The tiny pores on the side of the pharynx are normally gill slits in living chordates. He also noticed that Pikaia is similar to Amphioxus in most general aspects, with major difference in its notochord not reaching the anterior end.
Not all palaeontologists were convinced of the chordate designation without better analysis. In 2001, Nicholas D. Holland from the Scripps Institution of Oceanography and Junyuan Chen from the Chinese Academy of Sciences criticised the presentation in Wonderful Life, saying that the "reinterpretation [of Pikaia as a chordate] became almost universally accepted after its unqualified and forceful endorsement by Gould"; concluding that "the cephalochordate affinity of Pikaia is at best only weakly indicated by the characters visible in fossils discovered so far." In 2010, an international team of palaeontologists argued that Pikaia has sufficiently invertebrate characters, and that it mostly look like a much younger extinct animal, the Tully monster (Tullimonstrum gregarium), which is still debated as either an invertebrate or a chordate.
Another component of Pikaia fossils that constrains the animal to be accepted as a chordate is its distinct invertebrate character; its preservational mode suggests that it had cuticle. The cuticle as a body covering is uncharacteristic of the vertebrates, but is a dominant feature of invertebrates. The presence of earlier chordates among the Chengjiang, including Haikouichthys and Myllokunmingia, appears to show that cuticle is not necessary for preservation, overruling the taphonomic argument, but the presence of tentacles remains intriguing, and the organism cannot be assigned conclusively, even to the vertebrate stem group. Its anatomy closely resembles the modern creature Branchiostoma.
A fossil species Myoscolex ateles, discovered in 1979 from Cambrian Emu Bay Shale of Kangaroo Island in South Australia, had been debated as among the oldest annelids, or at least other invertebrate groups. Polish palaeontologist Jerzy Dzik in his formal description in 2003 notes that it "closely resembles the slightly geologically younger Pikaia" in having smooth cuticle as well as muscular segmentation, and projections on its backside (ventral chaetae) that look like Pikaia's tentacles. He concluded:In fact, there is little evidence for chordate affinities of Pikaia. Its relationship with Myoscolex [as annelid in his proposition] appears a much better solution. Both were initially identified as polychaetes and this line of inference perhaps deserves confrontation with more recent evidence than that available to the authors who proposed these genera.
Comprehensive description