# Nectocaris

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Extinct animal genus

Not to be confused with [Nectocotis](/source/Nectocotis).

Nectocaris Temporal range: Cambrian Stage 3–Wuliuan PreꞒ Ꞓ O S D C P T J K Pg N Specimen ROM 60079 Scientific classification Kingdom: Animalia Clade: Gnathifera Clade: Cucullophora Phylum: Chaetognatha Family: †Nectocarididae Conway Morris, 1976 Genus: †Nectocaris Conway Morris, 1976 Type species Nectocaris pteryx Conway Morris, 1976 Other species N. latus? Luo and Hu, 1999 Synonyms Vetustovermis Glaessner, 1979 Petalilium Luo and Hu, 1999

***Nectocaris*** is a genus of squid-like animal known from the [Cambrian](/source/Cambrian) period. The initial fossils were described from the [Burgess Shale](/source/Burgess_Shale) of Canada. Other similar remains possibly referrable to the genus are known from the [Emu Bay Shale](/source/Emu_Bay_Shale) of Australia and [Chengjiang Biota](/source/Chengjiang_Biota) of China.

*Nectocaris* was a free-swimming, predatory or scavenging organism. This lifestyle is reflected in its binomial name: *Nectocaris* means "swimming shrimp" (from the [Ancient Greek](/source/Ancient_Greek) νηκτόν, **[nekton](https://en.wiktionary.org/wiki/nekton)**, meaning "swimmer" and καρίς, **karis**, "shrimp"). Two morphs are known: a small morph, about an inch long, and a large morph, anatomically identical but around four times longer.[1]

Nectocaridids had controversial affinities before 2025. Some authors have suggested that they represent the earliest known [cephalopods](/source/Cephalopod). However, their morphology is strongly dissimilar to confirmed early cephalopods, and thus their affinities to cephalopods and even to [molluscs](/source/Mollusca) more broadly are rejected by many authors.[2][3] Their affinities to any animal group beyond [Bilateria](/source/Bilateria) were long considered uncertain,[3] until a 2025 study, which interpreted them as relatives of modern [chaetognaths](/source/Chaetognatha) (arrow worms).[4]

## Anatomy

*Nectocaris pteryx* from the Burgess Shale; funnel is visible folded to left of specimen. Image from Smith (2013).[5]

*Nectocaris* had a flattened, spear head-shaped body with a fleshy-flat fin running along the length of each side.[6] The small head had two stalked eyes, a single pair of [tentacles](/source/Tentacle), and a flexible funnel-shaped structure opening out to the underside of the body.[6] The funnel often gets wider away from the head.[6] The funnel has been suggested to represent an eversible (able to be turned inside out) [pharynx](/source/Pharynx).[3] Internally, a long cavity runs along the body axis, which is suggested to represent the digestive tract.[3] The body contains a pair of [gills](/source/Gill); the gills comprise blades emerging from a zig-zag axis. Muscle blocks surrounded the axial cavity, and are now preserved as dark blocks in the lateral body.[1] The fins also show dark blocks, with fine striations superimposed over them. These striations often stand in high relief above the rock surface itself.[1]

## Diversity

Although *Nectocaris* is known from Canada, China and Australia, in rocks spanning some 20 million years, there does not seem to be much diversity; size excepted, all specimens are anatomically very similar. Historically, three genera have been erected for nectocaridid taxa from different localities, but these 'species' – *Petalilium latus* and *Vetustovermis planus* – likely belong to the same genus or even the same species as *N. pteryx*. Within *N. pteryx*, there seem to be two discrete morphs, one large (~10 cm in length), one small (~3 cm long). These perhaps represent [separate male and female forms](/source/Sexual_dimorphism).[1]

## Ecology

Life restoration following the now erroneous cephalopod interpretation

The unusual shape of the nectocaridid funnel has led to its interpretation as an eversible [proboscis](/source/Proboscis). Martin R. Smith and Jean-Bernard Caron have suggested that it was used for [jet propulsion](/source/Jet_propulsion),[6][1] though this has been questioned by other authors.[7] The eyes of *Nectocaris* would have had a similar visual acuity to modern *[Nautilus](/source/Nautilus)* (if they lacked a lens) or [squid](/source/Squid) (if they did not).[1] They are thought to have been freely-swimming [nektonic](/source/Nektonic) organisms,[3] that were either scavengers or predators on soft-bodied animals, using their tentacles to manipulate food items.[6]

## Affinity

The affinity of *Nectocaris* is controversial.[7][3] Martin R. Smith and Jean-Bernard Caron have suggested that nectocaridids represent early cephalopods. In a 2010 publication in Nature, they suggested that the ancestor of modern cephalopods and nectocaridids probably lacked a mineralised shell,[6] while Smith in a later 2013 publication suggested that it may be more plausible that nectocaridids had instead lost a mineralised shell and developed a morphology convergent on modern [coleoids](/source/Coleoidea),[1] and in 2020 Smith described resembling taxon *[Nectocotis](/source/Nectocotis)* from [Ordovician](/source/Ordovician), which is suggested to have had an internal skeletal element.[8] However, other authors contend that the morphology of nectocaridids is contrary to what is known about cephalopod and mollusc evolution, and they cannot be accommodated within these groups,[2][3][7] and can only be confidently placed as members of [Bilateria](/source/Bilateria).[3] A 2025 study based on specimens of the related *[Nektognathus](/source/Nektognathus)* from [Sirius Passet](/source/Sirius_Passet) of Greenland which preserved neural tissue showed the presence of a ventral ganglion (part of the nervous system within the body cavity), a distinctive characteristic found only among [arrow worms](/source/Arrow_worms) (chaetognaths) as well as other extinct Cambrian relatives like the large *[Amiskwia](/source/Amiskwia)* also known from the Burgess Shale, firmly placing nectocaridids in the chaetognath [stem-group](/source/Stem-group). Due to lack of diagnostic features, authors have not included *Nectocotis* to nectocaridid.[9]

## History of study

		- Holotype of *Nectocaris*, an incomplete specimen that gave rise to erroneous reconstructions

		- Original (and obsolete) reconstruction based on a single, incompletely preserved, lateral specimen (left). The author of this reconstruction, based on the material then available, considered *Nectocaris* to bear arthropod and chordate-like features

*Nectocaris* has a long and convoluted history of study. [Charles Doolittle Walcott](/source/Charles_Doolittle_Walcott), the discoverer of the Burgess Shale, had photographed the one specimen he had collected in the 1910s, but never had time to investigate it further. As such, it was not until 1976 that *Nectocaris* was formally described, by [Simon Conway Morris](/source/Simon_Conway_Morris).[10]

Because the genus was originally known from a single, incomplete specimen and with no [counterpart](/source/Counter_slab),[11] Conway Morris was unable to deduce its affinity. It had some features which were reminiscent of [arthropods](/source/Arthropod), but these could well have been convergently derived.[10][12] Its fins were very unlike those of arthropods.[10]

Working from photographs, the Italian palaeontologist Alberto Simonetta believed he could classify *Nectocaris* within the [chordates](/source/Chordate).[13] He focused mainly on the tail and fin morphology, interpreting Conway Morris's 'gut' as a [notochord](/source/Notochord) – a distinctive chordate feature.[13]

The classification of *Nectocaris* was revisited in 2010, when Martin Smith and Jean-Bernard Caron described 91 additional specimens, many of them better preserved than the type. These allowed them to reinterpret *Nectocaris* as a primitive [cephalopod](/source/Cephalopod), with only 2 tentacles instead of the 8 or 10 limbs of modern cephalopods. The structure previous researchers had identified as an oval [carapace](/source/Carapace) or shield behind the eyes[14] was suggested to be a soft funnel, similar to the ones used for propulsion by modern cephalopods. The interpretation would push back the origin of cephalopods by at least 30 million years, much closer to the first appearance of complex animals, in the [Cambrian explosion](/source/Cambrian_explosion), and implied that – against the widespread expectation – cephalopods evolved from non-mineralized ancestors.[6]

Later independent analyses questioned the cephalopod interpretation, stating that it did not square with the established theory of [cephalopod evolution](/source/Cephalopod_evolution), and that nectocaridids should be considered *[incertae sedis](/source/Incertae_sedis)* among [Bilateria](/source/Bilateria).[15][7][3][2]

### *Vetustovermis*

***Vetustovermis*** (from Latin: "very old worm")[16] is a soft-bodied middle Cambrian [animal](/source/Animal), known from a single reported fossil specimen from the South Australian [Emu Bay Shale](/source/Emu_Bay_Shale). It is probably a [junior synonym](/source/Synonym_(taxonomy)) of *Nectocaris pteryx*.[1]

The original description of *Vetustovermis* hedged its bets regarding classification, but tentatively highlighted some similarities with the [annelid](/source/Annelid) worms.[16] It was later considered an [arthropod](/source/Arthropod),[17][18] and in 2010 Smith and Caron, agreeing that *[Petalilium](/source/Petalilium)* was at least a close relative of *Vetustovermis* (but that treating it as a synonym was premature, given the poor preservation of the *Vetustovermis* type), placed it with *Nectocaris* in the [clade](/source/Clade) [Nectocarididae](/source/Nectocarididae).[6]

Early press reports misspelled the genus name as *Vetusto**d**ermis*.

### *Petalilium*

Fossil of *Petalilium*

***Petalilium*** (sometimes misspelled ***Petalium***)[19] is an enigmatic genus of [Cambrian](/source/Cambrian) organism known from the Haikou area,[20] from the Maotianshan mudstone member of the [Chengjiang biota](/source/Chengjiang_biota).[21] The taxon is a junior synonym of *Nectocaris pteryx*.[1]

Fossils of *Petalilium*[a] show a [dorsoventrally](/source/Anatomical_terms_of_location#Directional_terms) flattened body, usually 5 to 6 centimetres, but ranging from 1.5 to 10 cm. It has an ovate trunk region and a large [muscular](/source/Muscle) foot, and a head with stalked eyes and a pair of long tentacles. The trunk region possesses about 50 soft, flexible, [transverse](/source/Anatomical_terms_of_location#Invertebrate_directional_terms) bars, lateral serialised structures of unknown function. The upper part of the body, interpreted as a [mantle](/source/Mantle_(mollusc)), is covered with a random array of spines on the back, while gills project underneath. A complete, tubular gut runs the length of the body.

Whilst it was originally described as a [phyllocarid](/source/Phyllocarida),[17] and a [ctenophore](/source/Ctenophore) affinity has been suggested,[22] neither interpretation is supported by any compelling evidence.[23]

Some of the characters observed in Chen *et al*.'s (2005) study[19] suggested that *Petalilium* may be related to *Nectocaris*.[6]

The 2025 study supported the synonymy of *Nectocaris* and *Petalilium*, but it was considered that this was a separate species (*N. latus*) rather than *N. pteryx*.[4]

## See also

- [paleontology portal](https://en.wikipedia.org/wiki/Portal:Paleontology)

- [Cambrian explosion](/source/Cambrian_explosion)

- [Paleobiota of the Burgess Shale](/source/Paleobiota_of_the_Burgess_Shale)

- [Chengjiang biota](/source/Chengjiang_biota) - [List of Chengjiang Biota species by phylum](/source/List_of_Chengjiang_Biota_species_by_phylum)

## Footnotes

1. **[^](#cite_ref-22)** *Petalilium* was originally described as *Vetustovermis* by Chen *et al*. (2005),[19] but recognised as *Petalilium* by Smith & Caron (2010).[6]

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1. **[^](#cite_ref-Hu2007_24-0)** Hu, S.; Steiner, M.; Zhu, M.; Erdtmann, B.D.; Luo, H.; Chen, L.; Weber, B. (2007). "Diverse pelagic predators from the Chengjiang Lagerstätte and the establishment of modern-style pelagic ecosystems in the early Cambrian". *Palaeogeography, Palaeoclimatology, Palaeoecology*. **254** (1–2): 307–316. [Bibcode](/source/Bibcode_(identifier)):[2007PPP...254..307H](https://ui.adsabs.harvard.edu/abs/2007PPP...254..307H). [doi](/source/Doi_(identifier)):[10.1016/j.palaeo.2007.03.044](https://doi.org/10.1016%2Fj.palaeo.2007.03.044).

## Further reading

- ["*Nectocaris pteryx*"](https://web.archive.org/web/20201112025257/http://burgess-shale.rom.on.ca/en/fossil-gallery/view-species.php?id=87). *Burgess Shale Fossil Gallery*. Virtual Museum of Canada. 2011. Archived from [the original](http://burgess-shale.rom.on.ca/en/fossil-gallery/view-species.php?id=87) on 2020-11-12. Retrieved 2023-01-21. – 3D animations are available and a more detailed consideration of *Nectocaris*

- Switek, Brian (5 July 2011). ["Nectocaris: What the heck is this thing?"](https://www.wired.com/wiredscience/2011/07/nectocaris-what-the-heck-is-this-thing/). *Laelaps*. – Brian Switek discusses the taxonomy and history of *Nectocaris* in his blog

- Moskvitch, Katia (27 May 2010). ["Mystery fossil is ancestor of squid"](https://www.bbc.co.uk/news/10173293). [BBC News](/source/BBC_News). – [BBC News](/source/BBC_News) coverage of the Smith & Caron's (2010) re-description

- Taylor, Christopher (14 January 2011). ["So nice when people agree with you"](http://coo.fieldofscience.com/2011/01/so-nice-when-people-agree-with-you.html). *Catalogue of Organisms*. – a blog article supporting Mazurek & Zatoń's (2011) view

- Conway Morris, S. (1997). *The Crucible of Creation: the Burgess Shale and the rise of animals*. Oxford University Press. [ISBN](/source/ISBN_(identifier)) [0-19-286202-2](https://en.wikipedia.org/wiki/Special:BookSources/0-19-286202-2).

## External links

- Media related to [*Nectocaris*](https://commons.wikimedia.org/wiki/Category:Nectocaris) at Wikimedia Commons

Taxon identifiers Nectocaris Wikidata: Q132992 GBIF: 4632567 IRMNG: 1331953 Open Tree of Life: 4723195 Paleobiology Database: 7555

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Adapted from the Wikipedia article [Nectocaris](https://en.wikipedia.org/wiki/Nectocaris) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Nectocaris?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.