| Opabinia | |
|---|---|
 | |
| Fossil specimen (holotype) on display at theSmithsonian in Washington, D.C. | |
Scientific classification | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Arthropoda |
| Family: | †Opabiniidae |
| Genus: | †Opabinia Walcott, 1912 |
| Species: | †O. regalis |
| Binomial name | |
| †Opabinia regalis Walcott, 1912 | |
Opabinia regalis is anextinct,stem group marinearthropod found in theMiddle CambrianBurgess ShaleLagerstätte (505 million years ago) ofBritish Columbia.[1]Opabinia was a soft-bodied animal, measuring up to 7 cm in body length, and had a segmented trunk with flaps along its sides and a fan-shaped tail. The head showed unusual features: fiveeyes, a mouth under the head and facing backwards, and a clawedproboscis that most likely passed food to its mouth.Opabinia lived on the seafloor, using the proboscis to seek out small, soft food.[2] Fewer than twenty good specimens have been described; 3 specimens ofOpabinia are known from the GreaterPhyllopod bed, where they constitute less than 0.1% of the community.[3]
When the first thorough examination ofOpabinia in 1975 revealed its unusual features, it was thought to be unrelated to any knownphylum,[4] or perhaps a relative of arthropod andannelid ancestors.[2] However, later studies since late 1990s consistently support its affinity as a member of basal arthropods, alongside the closely relatedradiodonts (Anomalocaris and relatives) andgilled lobopodians (Kerygmachela andPambdelurion).[5][6][7][8][1][9][10]
In the 1970s, there was an ongoing debate about whethermulti-celled animals appeared suddenly during the Early Cambrian, in an event called theCambrian explosion, or had arisen earlier but without leaving fossils. At firstOpabinia was regarded as strong evidence for the "explosive" hypothesis.[4] Later the discovery of a whole series of similarlobopodian animals, some with closer resemblances to arthropods, and the development of the idea ofstem groups, suggested that the Early Cambrian was a time of relatively fastevolution, but one that could be understood without assuming any unique evolutionary processes.[11]
In 1911,Charles Doolittle Walcott found in theBurgess Shale nine almost complete fossils ofOpabinia regalis and a few of what he classified asOpabinia ? media, and published a description of all of these in 1912.[12] The generic name is derived fromOpabin pass betweenMount Hungabee andMount Biddle, southeast ofLake O'Hara, British Columbia,Canada.[12] In 1966–1967,Harry B. Whittington found another good specimen,[13] and in 1975 he published a detailed description based on very thoroughdissection of some specimens and photographs of these specimens lit from a variety of angles. Whittington's analysis did not coverOpabinia ? media; Walcott's specimens of this species could not be identified in his collection.[2] In 1960 Russianpaleontologists described specimens they found in theNorilsk region ofSiberia and labelledOpabinia norilica,[14] but these fossils were poorly preserved, and Whittington did not feel they provided enough information to be classified as members of thegenusOpabinia.[2]
All the recognizedOpabinia specimens found so far come from the "Phyllopod bed" of the Burgess Shale, in theCanadian Rockies of British Columbia.[2]
In 1997, Briggs and Nedin reported fromSouth AustraliaEmu Bay Shale a new specimen ofMyoscolex that was much better preserved than previous specimens, leading them to conclude that it was a close relative ofOpabinia[15]—although this interpretation was later questioned by Dzik, who instead concluded thatMyoscolex was anannelid worm.[16]
Opabinia looked so strange that the audience at the first presentation of Whittington's analysis laughed.[2] The length ofOpabinia regalis from head (excluding proboscis) to tail end ranged between 4 centimetres (1.6 in) and 7 centimetres (2.8 in).[2] One of the most distinctive characters ofOpabinia is the hollowproboscis, whose total length was about one-third that of the body, and projected down from under the head. The proboscis wasstriated like avacuum cleaner's hose and flexible, and it ended with a claw-like structure whose terminal edges bore 5 spines that projected inwards and forwards. The bilateral symmetry and lateral (instead of vertical as reconstructed by Whittington 1975[2]) arrangement of the claw suggest it represents a pair of fused frontal appendages, comparable to those ofradiodonts andgilled lobopodians.[17][5][18][19] The head bore five stalked eyes: two near the front and fairly close to the middle of the head, pointing upwards and forwards; two larger eyes with longer stalks near the rear and outer edges of the head, pointing upwards and sideways; and a single eye between the larger pair of stalked eyes, pointing upwards. It has been assumed that the eyes were allcompound, like otherarthropods' lateral eyes,[2] but this reconstruction, which is not backed up by any evidence,[2] is "somewhat fanciful".[20] The mouth was under the head, behind the proboscis, and pointedbackwards, so that thedigestive tract formed a U-bend on its way towards the rear of the animal. The proboscis appears to have been sufficiently long and flexible to reach the mouth.[2]
The main part of the body was typically about 5 millimetres (0.20 in) wide and had 15 segments, on each of which there were pairs of flaps (lobes) pointing downwards and outwards. The flaps overlapped so that the front of each was covered by the rear edge of the one ahead of it. The body ended with what looked like a single conical segment bearing three pairs of overlapping tail fan blades that pointed up and out, forming a tail like a V-shaped double fan.[2]
Interpretations of other features ofOpabinia fossils differ. Since the animals did not havemineralized armor nor even tough organicexoskeletons like those of other arthropods, their bodies were flattened as they were buried and fossilized, and smaller or internal features appear as markings within the outlines of the fossils.[2][5]
![Various interpretations on the flap and gill structures of Opabinia regalis A: Whittington (1975),[2] B: Bergström (1986),[17] C: Budd (1996),[5] D: Zhang & Briggs (2007),[7] E: Budd & Daley (2011)[8]](/mt/?noimg=&dark=on&url=https%3a%2f%2fen.wikipedia.org%2fwiki%2fFile%3a20210809_Opabinia_regalis_flap_gill_interpretation.png)
![Opabinia cross-section based on Budd and Daley (2011)[8]](/mt/?noimg=&dark=on&url=https%3a%2f%2fen.wikipedia.org%2fwiki%2fFile%3a20210807_Opabinia_regalis_trunk_cross_section.png)
Whittington (1975) interpreted the gills as paired extensions attached dorsally to the bases of all but the first flaps on each side, and thought that these gills were flat underneath, had overlapping layers on top.[2] Bergström (1986) revealed the "overlapping layers" were rows of individual blades, interpreted the flaps as part of dorsal coverings (tergite) over the upper surface of the body, with blades attached underneath each of them.[17][8]Budd (1996) thought the gill blades attached along the front edges on the dorsal side of all except the first flaps. He also found marks inside the flaps' front edges that he interpreted as internal channels connecting the gills to the interior of the body, much as Whittington interpreted the mark along the proboscis as an internal channel.[5] Zhang andBriggs (2007) however, interpreted all flaps have posterior spacing where the gill blades attached.[7] Budd and Daley (2011) reject the reconstruction by Zhang & Briggs, showing the flaps have complete posterior edges as in previous reconstructions. They mostly follow the reconstruction by Budd (1996) with modifications on some details (e.g. the first flap pair also have gills; the attachment point of gill blades located more posteriorly than previously thought).[8]
Whittington (1975) found evidence of near-triangular features along the body, and concluded that they were internal structures, most likely sideways extensions of the gut (diverticula).[2] Chenet al. (1994) interpreted them as contained within the lobes along the sides.[21] Budd (1996) thought the "triangles" were too wide to fit withinOpabinia's slender body, and thatcross-section views showed they were attached separately from and lower than the lobes, and extended below the body.[5] He later found specimens that appeared to preserve the legs' exterior cuticle. He therefore interpreted the "triangles" as short, fleshy, conical legs (lobopods). He also found small mineralized patches at the tips of some, and interpreted these as claws. Under this reconstruction, the gill-bearing flap and lobopod were homologized to the outer gill branch and inner leg branch of arthropodbiramous limbs seen inMarrella,trilobites, andcrustaceans.[5] Zhang and Briggs (2007) analyzed the chemical composition of the "triangles", and concluded that they had the same composition as the gut, and therefore agreed with Whittington that they were part of the digestive system. Instead they regardedOpabinia's lobe+gill arrangement as an early form of the arthropod limbs before it split into a biramous structure.[7] However, this similar chemical composition is not only associated with the digestive tract; Budd and Daley (2011) suggest that it represents mineralization forming within fluid-filled cavities within the body, which is consistent with hollow lobopods as seen in unequivocal lobopodian fossils. They also clarify that the gut diverticula ofOpabinia are series of circular gut glands individualized from the "triangles". While they agreed on the absence of terminal claws, the presence of lobopods inOpabinia remain as a plausible interpretation.[8]
The way in which theBurgess Shale animals were buried, by a mudslide or a sediment-laden current that acted as a sandstorm, suggests they lived on the surface of the seafloor.[2]Opabinia probably used its proboscis to search the sediment for food particles and pass them to its mouth.[2] Since there is no sign of anything that might function as jaws, its food was presumably small and soft.[2] The paired gutdiverticula may increase the efficiency of food digestion and intake of nutrition.[22] Whittington (1975) believing thatOpabinia had no legs, thought that it crawled on its lobes and that it could also have swum slowly by flapping the lobes, especially if it timed the movements to create awave with the metachronal movement of its lobes.[2] On the other hand, he thought the body was not flexible enough to allow fish-like undulations of the whole body.[2]
| |||||||||
| Summarized phylogeny betweenOpabinia and otherEcdysozoan taxa.[10] |
Considering how paleontologists' reconstructions ofOpabinia differ, it is not surprising that the animal's classification was highly debated during the 20th century.[5]Charles Doolittle Walcott, the originaldescriber, considered it to be ananostracancrustacean in 1912.[12] The idea was followed byG. Evelyn Hutchinson in 1930, providing the first reconstruction ofOpabinia as an anostracan swimming upside down.[23]Alberto Simonetta provided a new reconstruction ofOpabinia in 1970 very different to those of Hutchinson's, with lots ofarthropod features (e.g. ,dorsal exoskeleton and jointed limbs) which are reminiscent ofYohoia andLeanchoilia.[24]Leif Størmer, following earlier work byPercy Raymond, thought thatOpabinia belonged to the so-called "trilobitoids" (trilobites and similar taxa). After his thorough analysisHarry B. Whittington concluded thatOpabinia was not arthropod in 1975, as he found no evidence for arthropodan jointed limbs, and that nothing like the flexible, probably fluid-filled, proboscis was known in arthropods.[2] Although he leftOpabinia's classification above thefamily level open, the annulated but not articulated body and the unusual lateral flaps with gills persuaded him that it may have been a representative of the ancestral stock from the origin ofannelids and arthropods,[2] two distinct animal phyla (Lophotrochozoan andEcdysozoan, respectively) which were still thought to be close relatives (united underArticulata) at that time.[25]
In 1985,Derek Briggs and Whittington published a major redescription ofAnomalocaris, also from the Burgess Shale.[26] Soon after that, Swedish palaeontologistJan Bergström, noting in 1986 the similarity ofAnomalocaris andOpabinia, suggested that the two animals were related, as they shared numerous features (e.g., lateral flaps, gill blades, stalked eyes, and specialized frontal appendages). He classified them as primitive arthropods, although he considered that arthropods are nota single phylum.[17]
In 1996,Graham Budd found what he considered evidence of short, un-jointed legs inOpabinia.[5] His examination of the gilled lobopodianKerygmachela from theSirius Passetlagerstätte, about518 million years ago and over 10M years older than the Burgess Shale, convinced him that this specimen had similar legs.[27][28] He considered the legs of these twogenera very similar to those of the Burgess Shale lobopodianAysheaia and the modernonychophorans (velvet worms), which are regarded as the bearers of numerousancestral traits shared by the ancestors of arthropods. After examining several sets of features shared by these and similar lobopodians he drew up a "broad-scale reconstruction of the arthropodstem-group",i.e., of arthropods and what he considered to be their evolutionary basal members.[5] One striking feature of this family tree is that moderntardigrades (water bears) may beOpabinia's closest livingevolutionary relatives.[5] On the other hand, Houet al. (1995, 2006) suggestedOpabinia is a member of unusualcycloneuralian worms withconvergent arthropod features.[29][18]
Although Zhang and Briggs (2007) disagreed with Budd's diagnosis thatOpabinia's "triangles" were legs, the resemblance they saw betweenOpabinia's lobe+gill arrangement and arthropods'biramous limbs led them to conclude thatOpabinia was very closely related to arthropods. In fact they presented a family tree very similar to Budd's except that theirs did not mention tardigrades.[7] Regardless of the different morphological interpretations, all major restudies since 1980s similarly concluded that the resemblance betweenOpabinia and arthropods (e.g., stalked eyes, dorsal segmentation, posterior mouth, fused appendages, gill-like limb branches) are taxonomically significant.[17][5][7][8][1]
Since the 2010s, the suggested close relationship betweenOpabinia and tardigrades/cycloneuralians is no longer supported, while the affinity ofOpabinia as a stem-group arthropod alongsideRadiodonta (a clade that includesAnomalocaris and its relatives[30]) andgilled lobopodians is widely accepted,[1][9][10] as consistently shown by multiple phylogenetic analyses,[31][32][33][34][35][36][37][38][39][40][41][42] as well as new discoveries such as the presence of arthropod-like gut glands[8][22] and the intermediate taxonKylinxia.[40]
In2022, Paleontologists described a similar looking animal which was discovered in Cambrian-aged rocks ofUtah.[43] The fossil was namedUtaurora comosa, and was found within theWheeler Shale.[43] The stem-arthropod was actually first described in 2008, but at the time it was originally considered a specimen ofAnomalocaris.[44] This discovery could suggest there were other animals that looked likeOpabinia, and its family may have been more diverse.[43]
Opabinia made it clear how little was known about soft-bodied animals, which do not usually leave fossils.[2] When Whittington described it in the mid-1970s, there was already a vigorous debate about the early evolution ofanimals.Preston Cloud argued in 1948 and 1968 that the process was "explosive",[45] and in the early 1970sNiles Eldredge andStephen Jay Gould developed their theory ofpunctuated equilibrium, which views evolution as long intervals of near-stasis "punctuated" by short periods of rapid change.[46] On the other hand, around the same time Wyatt Durham andMartin Glaessner both argued that the animal kingdom had a longProterozoic history that was hidden by the lack of fossils.[47][48] Whittington (1975) concluded thatOpabinia, and othertaxa such asMarrella andYohoia, cannot be accommodated in modern groups. This was one of the primary reasons why Gould in his book on theBurgess Shale,Wonderful Life, considered that Early Cambrian life was much more disparate and "experimental" than any later set of animals and that theCambrian explosion was a truly dramatic event, possibly driven by unusualevolutionary mechanisms.[4] He regardedOpabinia as so important to understanding this phenomenon that he wanted to call his bookHomage to Opabinia.[49]
However, other discoveries and analyses soon followed, revealing similar-looking animals such asAnomalocaris from the Burgess Shale andKerygmachela fromSirius Passet.[17][27] Another Burgess Shale animal,Aysheaia, was considered very similar to modernOnychophora,[50] which are regarded as close relatives of arthropods.[51] Paleontologists defined a group calledlobopodians to include fossil panarthropods that are thought to be close relatives of onychophorans, tardigrades and arthropods but lack jointed limbs. This group was later widely accepted as a paraphyletic grade that led to the origin of extant panarthropod phyla.
While this discussion about specific fossils such asOpabinia andAnomalocaris was going on in the late 20th century, the concept ofstem groups was introduced to cover evolutionary "aunts" and "cousins". Acrown group is a group of closely related living animals plus their last common ancestor plus all its descendants. A stem group contains offshoots from members of the lineage earlier than the last common ancestor of the crown group; it is arelative concept, for exampletardigrades are living animals that form a crown group in their own right, but Budd (1996) regarded them also as being a stem group relative to the arthropods.[5][52] Viewing strange-looking organisms likeOpabinia in this way makes it possible to see that, while the Cambrian explosion was unusual, it can be understood in terms of normal evolutionary processes.[11]
.png)
.jpg)
