Diversity
The phylum
Chaetognatha
, also known as arrow worms, contains nearly 200 species of mostly planktonic, bilaterally
symmetrical, coelomate, worm-like organisms. The phylum contains two orders,
Phragmophora
and
Aphragmophora
. The main difference between the two is the presence of ventral transverse muscle
bands in
Phragmophora
, which are absent in
Aphragmophora
. Chaetognaths may be found in marine and some estuarine environments throughout the
world. About a fifth of the total species are benthic, some living just above the
deep ocean floor; these are often attached to the substrate by adhesive secretions.
Chaetognaths may range from 1 mm to 12 cm in length and are typically transparent,
although some deep-water species may be orange in color, and phragmophorids may be
opaque, due to their musculature. The common name, arrow worms is derived from their
streamlined appearance, with paired lateral fins and a single caudal (tail) fin, while
their scientific name comes from the hooked set of jaws that protrude lateral to the
mouth. These structures are used in prey capture, with chaetognaths feeding on a number
of crustacean (mainly copepods) and fish (mainly larvae) species, which they track
through daily vertical migrations in the water column (these migrations may also protect
them from predators). Chaetognaths are hermaphroditic, and may undergo reciprocal,
nonreciprocal, or self-fertilization.
Geographic Range
Chaetognaths may be found in marine and some estuarine environments throughout the
world, including polar and tropical regions.
- Biogeographic Regions
- nearctic
- palearctic
- oriental
- ethiopian
- neotropical
- australian
- antarctica
- arctic ocean
- indian ocean
- atlantic ocean
- pacific ocean
- mediterranean sea
- Other Geographic Terms
- holarctic
- cosmopolitan
Habitat
Chaetognaths are mainly planktonic organisms in marine and estuarine environments.
About a fifth of the total species are benthic, some living just above the deep ocean
floor. They are often found in great numbers, particularly in mid-water and neritic
waters, and may be found in rock pools or associated with certain oceanic currents.
- Habitat Regions
- temperate
- tropical
- polar
- saltwater or marine
- Other Habitat Features
- estuarine
Systematic and Taxonomic History
Since first being recorded in 1775, chaetognaths have been classified many ways, including
as
molluscs
,
arthropods
, and
nematodes
. They were elevated to their own phylum by the German zoologist Rudolf Leuckart in
1854, a decision which is supported by numerous morphological and embryological autapomorphies,
indicated below. The monophyly of this phylum is widely accepted, and molecular phylogenetic
analyses of 28S ribosomal DNA have supported the monophyly of its two constituent
orders,
Phragmophora
and
Aphragmophora
(without the traditionally included family
Pterosagittidae
).
Chaetognaths were originally considered
deuterostomes
, based on embryological observations. All molecular phylogenetic analyses performed
thus far, however, strongly support the placement of
Chaetognatha
within
protostome
animals. The precise phylogenetic placement of
Chaetognatha
within
Protostomata
remains somewhat confused today, as molecular analyses have suggested
Chaetognatha
is a sister group to all other protostomes,
Spiralia
, or phylum
Priapulida
(within the superphylum
Ecdysozoa
). Their potential identity as a basal sister group to such a large metazoan group
makes further examination of their phylogenetic placement essential, as it has the
potential to greatly contribute to our understanding of morphological and developmental
characters found in some of the earliest bilaterian organisms.
Physical Description
Chaetognaths range from 1 mm to 12 cm in length and are typically transparent, although
some deep-water species may be orange in color (carotenoid pigmentation), and phragmophorids
may be opaque, due to their musculature. They are bilaterally symmetrical and have
long, streamlined bodies, which may be divided into head, trunk, and tail regions.
They have paired lateral fins and a single tail fin. The mouth is located ventrally
on the head, and is set into a vestibule; this structure is typically associated with
grasping spines or hooks, located laterally to the mouth, as well as teeth, which
are in located at the front of the mouth. Some species have serrated hooks and/or
cuspidate teeth. A hood (anterolateral body wall fold) may be pulled over the head
to enclose the vestibule.
Chaetognaths are covered in a thin, flexible cuticle on top of the epidermis. Epidermal
cells are mainly squamous and have interlocking margins; they may be stratified. Epidermal
cells covering the fins are elongated and the cells lining the vestibule are columnar
rather than squamous. The cuticle is not continuous and, where it is not present,
there are many secretory cells in the epidermis. There is a basement membrane present
between the epidermis and body wall; the body wall is made up of four quadrants of
dorsolateral and ventrolateral longitudinal muscles. The body cavities are most likely
derived from enterocoelic cavities, which form during development. The body cavity
has a tripartite arrangement, with a head cavity (protocoel, reduced in space by the
cephalic musculature), and paired trunk and tail coeloms with dorsal and ventral longitudinal
mesenteries, which correspond to the mesocoel and matcoel, respectively. Transverse
septa separate the body regions. The body fluid has a variety of cells and cell-like
structures, although their functions are largely unknown. The fluid-filled coeloms,
body wall, basement membrane, and cuticle all provide support to the body. They do
not have circulatory, respiratory, or excretory organs; gases are diffused across
the body wall and fluid transport is via cilial action within the body cavities.
A few species of deep sea chaetognaths, including
Eukrohnia fowleri
and
Caecosagitta macrocephala
, are bioluminescent.
- Other Physical Features
- ectothermic
- heterothermic
- bilateral symmetry
- venomous
Development
Chaetognaths are hermaphroditic. Cross-fertilization is most common, although some
species will self-fertilize. Fertilization is typically internal and eggs may be released
into the water, deposited on the sea floor or other substrate, or brooded in pouches
near the tail. Cleavage is radial, holoblastic, and equal, leading to a coeloblastula.
Development is direct and accomplished quickly, typically from zygote to juvenile
within 48 hours.
Reproduction
Chaetognaths may undergo reciprocal, nonreciprocal, or self-fertilization. Some benthic
species have been documented performing a mating "dance," with an individual depositing
balls of sperm onto a mate.
- Mating System
- polygynandrous (promiscuous)
Chaetognaths have paired ovaries located in their trunks and paired testes located
in their tails. Sperm mature before eggs (which makes self-fertilization less likely),
and are stored in coelomic cavities within the tail until they are released in clusters
outside the body via a pair of seminal vesicles. Ovaries have oviducts, which lead
to genital pores located near the trunk-tail junction. In populations of at least
a few species, breeding occurs twice a year, and hatching occurs from April to June
and late September to December (typically fewer hatchlings).
- Key Reproductive Features
- iteroparous
- seasonal breeding
- simultaneous hermaphrodite
- sexual
- fertilization
- oviparous
Outside of a few species (such as members of genus
Eukronhnia
) that brood their young until they are ready to swim, chaetognaths exhibit no parental
investment beyond the production of gametes.
- Parental Investment
- female parental care
-
pre-fertilization
- provisioning
-
pre-hatching/birth
-
protecting
- female
-
protecting
Lifespan/Longevity
Most cold water chaetognaths have a longer life expectancy than those in tropical
waters, two years versus six weeks, respectively.
Behavior
Many species within this phylum are known to undergo daily vertical migrations, rising
to the surface at night to follow prey and sinking during the day, which provides
protection from predators. These worms have ammonia-filled vacuolated cells in their
trunks, which help them to regulate their depth in the water column. Pelagic chaetognaths
move by contracting the longitudinal muscles of their right and left sides alternately,
creating forward, darting motions. Fins do not appear to aid in locomotion, instead
acting as stabilizers.
- Key Behaviors
- natatorial
- diurnal
- nocturnal
- crepuscular
- motile
- sedentary
- solitary
Communication and Perception
Chaetognaths have a central nervous system with a large cerebral ganglion, dorsal
to the pharynx. Additional ganglia, which serve muscles and sensory organs of the
head, arise from this structure. They also have a pair of circumenteric connective
nerves, emerging from the rear of the cerebral ganglion and extending (posterioventrally)
to meet in a ventral ganglion in the epidermis of the trunk. This ganglion controls
swimming motion and also gives rise to many pairs of nerves, which create a subepidermal
nerve plexus.
Chaetognaths have a pair of compound eyes below the epidermis, on the head. They are
made up of five inverted pigment-cup ocelli, one large ocellus directed laterally
and four smaller ones directed medially; this gives these worms a nearly uninterrupted
field of vision. Their eyes do not typically have lenses and likely do not form images,
but are used for light reception and body orientation. The ocelli also contain ciliated
receptor cells.
Chaetognaths have a ciliary loop (corona ciliata) on the dorsal surface of the head-trunk
juncture, made up of two rings of cilial cells that may be chemoreceptive and/or aid
in sperm transfer. They are covered in patches of ciliary fans, which enhance the
detection of vibrations in the water.
- Other Communication Modes
- photic/bioluminescent
- Perception Channels
- visual
- tactile
- vibrations
- chemical
Food Habits
Chaetognaths are carnivorous predators, particularly of
copepods
. They are also known to feed on other crustaceans and small fishes. Benthic species
are typically ambush predators. They use adhesive secretions to affix themselves to
substrate and extend their mouths and vestibules, as well as their associated hooks.
When prey is detected by a worm (by the cilia on the body), the head darts forward
and prey is captured, using the hooks. Prey is swallowed whole. Panktonic species
dart forward in the water column to catch prey within reach, using their grasping
spines to pull prey in. The majority of these worms inject their prey with a neurotoxin
(tetrodotoxin); it has been hypothesized that chaetognaths have a commensal relationship
with bacteria (from genus
Vibro
) in their heads or guts, which actually produce the toxin.
- Primary Diet
-
carnivore
- piscivore
- eats non-insect arthropods
- planktivore
Predation
Chaetognaths are prey to many larger organisms including fishes, whales, other marine
invertebrates, and molluscs.
Ecosystem Roles
A number of tetrodotoxin producing bacteria have been isolated from the guts of chaetognaths;
these are likely responsible for the production of the toxin used by the worms in
prey capture. They may be hosts to parasitic
digeneans
,
nematodes
, and metacestodes; infections may be the result of the worm ingesting infested
copepods
, which serve as intermediate hosts. In turn, they may pass these parasites on to
their predators (particularly fishes). They may also serve as hosts to ectoparasites
such as
copepods
and
dinoflagellates
.
- Vibrio species
- Vibrio alginolyticus
- Hysterothylacium aduncum (Order Ascaridida , Phylum Nematoda )
- Adolescaria progastrica (Order Plagiorchiida , Phylum Platyhelminthes )
- Derogenes sp. (Order Plagiorchiida , Phylum Platyhelminthes )
- Derogenes varicus (Order Plagiorchiida , Phylum Platyhelminthes )
- Didymozoidae (Order Plagiorchiida , Phylum Platyhelminthes )
- Ectenurus virgulus (Order Plagiorchiida , Phylum Platyhelminthes )
- Hemiurus levinseni (Order Plagiorchiida , Phylum Platyhelminthes )
- Lepocreadiid metacercariae (Order Plagiorchiida , Phylum Platyhelminthes )
- Monascus filiformis (Order Plagiorchiida , Phylum Platyhelminthes )
- Parahemiurus merus (Order Plagiorchiida , Phylum Platyhelminthes )
- Tetraphyllidea sp. (Class Cestoda , Phylum Platyhelminthes )
- Microsetella norwegica (Order Harpacticoida , Phylum Arthropoda )
- Corycaeus amazonicus (Order Poecilostomatoida , Phylum Arthropoda )
- Oodinium jordani (Order Blastodinida , Phylum Myzozoa )
Economic Importance for Humans: Positive
Chaetognaths are important to humans not only in terms of scientific research possibilities,
but also as prey items for a variety of fish eaten by humans.
- Positive Impacts
- research and education
Economic Importance for Humans: Negative
Chaetognaths may negatively impact humans if they pass along parasites to fishes eaten
by humans.
Conservation Status
Neither this phylum, nor its constituent species is considered at risk of becoming
threatened or endangered.
Other Comments
In spite of being soft-bodied, chaetognaths are known in the fossil record, dating
back to the early Cambrian Period.
Additional Links
Contributors
Jeremy Wright (author), University of Michigan-Ann Arbor, Leila Siciliano Martina (editor), Animal Diversity Web Staff.
- Nearctic
-
living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Palearctic
-
living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- oriental
-
found in the oriental region of the world. In other words, India and southeast Asia.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Ethiopian
-
living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Neotropical
-
living in the southern part of the New World. In other words, Central and South America.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Australian
-
Living in Australia, New Zealand, Tasmania, New Guinea and associated islands.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Antarctica
-
lives on Antarctica, the southernmost continent which sits astride the southern pole.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Arctic Ocean
-
the body of water between Europe, Asia, and North America which occurs mostly north of the Arctic circle.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Atlantic Ocean
-
the body of water between Africa, Europe, the southern ocean (above 60 degrees south latitude), and the western hemisphere. It is the second largest ocean in the world after the Pacific Ocean.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Pacific Ocean
-
body of water between the southern ocean (above 60 degrees south latitude), Australia, Asia, and the western hemisphere. This is the world's largest ocean, covering about 28% of the world's surface.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- holarctic
-
a distribution that more or less circles the Arctic, so occurring in both the Nearctic and Palearctic biogeographic regions.
Found in northern North America and northern Europe or Asia.
- cosmopolitan
-
having a worldwide distribution. Found on all continents (except maybe Antarctica) and in all biogeographic provinces; or in all the major oceans (Atlantic, Indian, and Pacific.
- temperate
-
that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).
- tropical
-
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
- polar
-
the regions of the earth that surround the north and south poles, from the north pole to 60 degrees north and from the south pole to 60 degrees south.
- saltwater or marine
-
mainly lives in oceans, seas, or other bodies of salt water.
- pelagic
-
An aquatic biome consisting of the open ocean, far from land, does not include sea bottom (benthic zone).
- benthic
-
Referring to an animal that lives on or near the bottom of a body of water. Also an aquatic biome consisting of the ocean bottom below the pelagic and coastal zones. Bottom habitats in the very deepest oceans (below 9000 m) are sometimes referred to as the abyssal zone. see also oceanic vent.
- reef
-
structure produced by the calcium carbonate skeletons of coral polyps (Class Anthozoa). Coral reefs are found in warm, shallow oceans with low nutrient availability. They form the basis for rich communities of other invertebrates, plants, fish, and protists. The polyps live only on the reef surface. Because they depend on symbiotic photosynthetic algae, zooxanthellae, they cannot live where light does not penetrate.
- coastal
-
the nearshore aquatic habitats near a coast, or shoreline.
- abyssal
-
on or near the ocean floor in the deep ocean. Abyssal regions are characterized by complete lack of light, extremely high water pressure, low nutrient availability, and continuous cold (3 degrees C).
- brackish water
-
areas with salty water, usually in coastal marshes and estuaries.
- estuarine
-
an area where a freshwater river meets the ocean and tidal influences result in fluctuations in salinity.
- ectothermic
-
animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature
- heterothermic
-
having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.
- bilateral symmetry
-
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
- venomous
-
an animal which has an organ capable of injecting a poisonous substance into a wound (for example, scorpions, jellyfish, and rattlesnakes).
- polygynandrous
-
the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.
- iteroparous
-
offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).
- seasonal breeding
-
breeding is confined to a particular season
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- fertilization
-
union of egg and spermatozoan
- internal fertilization
-
fertilization takes place within the female's body
- oviparous
-
reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.
- female parental care
-
parental care is carried out by females
- natatorial
-
specialized for swimming
- diurnal
-
- active during the day, 2. lasting for one day.
- nocturnal
-
active during the night
- crepuscular
-
active at dawn and dusk
- motile
-
having the capacity to move from one place to another.
- sedentary
-
remains in the same area
- solitary
-
lives alone
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- chemical
-
uses smells or other chemicals to communicate
- photic/bioluminescent
-
generates and uses light to communicate
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- vibrations
-
movements of a hard surface that are produced by animals as signals to others
- chemical
-
uses smells or other chemicals to communicate
- carnivore
-
an animal that mainly eats meat
- piscivore
-
an animal that mainly eats fish
- planktivore
-
an animal that mainly eats plankton
References
Brands, S. 2009. "Classification: Phylum Chaetognatha " (On-line). Systema Naturae 2000. Accessed April 01, 2013 at http://sn2000.taxonomy.nl .
Brusca, R., G. Brusca. 2003. Invertebrates (2nd Edition) . Sunderland, MA: Sinauer Associates.
Chen, J., D. Huang. 2002. A possible Lower Cambrian chaetognath (arrow worm). Science , 298: 187.
DaPonte, M., A. Gil de Pertierra, M. Palmieri, M. Ostrowski de Nunez. 2008. Monthly occurrence of parasites of the chaetognath Sagitta friderici off Mar del Plata, Argentina. Journal of Plankton Research , 30/5: 567-576. Accessed April 02, 2013 at http://plankt.oxfordjournals.org/content/30/5/567.full.pdf .
Dunn, C., A. Hejnol, D. Matus, K. Pang, W. Browne, S. Smith, E. Seaver, G. Rouse, M. Obst, G. Edgecombe, M. Sørensen, S. Haddock, A. Schmidt-Rhaesa, A. Okusu, R. Kristensen, W. Wheeler, M. Martindale, G. Giribet. 2008. Broad phylogenomic sampling improves resolution of the animal tree of life. Nature , 452: 745-749.
Edgecombe, G., G. Giribet, C. Dunn, A. Hejnol, R. Kristensen, R. Neves, G. Rouse, K. Worsaae, M. Sørensen. 2011. Higher-level metazoan relationships: recent progress and remaining questions. Organisms Diversity and Evolution , 11: 151-172.
Ghirardelli, E. 1968. Some aspects of the biology of chaetognaths. Pp. 271-375 in Advances in Marine Biology , Vol. 6. London, England: Academic Press. Accessed April 01, 2013 at http://books.google.com/books?id=UefCHv9EU5IC&pg=PA271&lpg=PA271&dq=some+aspects+of+the+biology+of+the+chaetognaths&source=bl&ots=-vkegBU-fE&sig=zUEXXI0Dull8LnX8ESE7InZ45Ig&hl=en&sa=X&ei=97VZUZzKFqHQ0wH-xYHIAg&ved=0CBgQ6AEwAA#v=onepage&q=some%20aspects%20of%20the%20biology%20of%20the%20chaetognaths&f=false .
Giribet, G., C. Dunn, G. Edgecombe, G. Rouse. 2007. A modern look at the Animal Tree of Life. Zootaxa , 1668: 61-79.
Goto, T., M. Yoshida. 1985. The mating sequence of the benthic arrowworm Spadella schizoptera . The Biological Bulletin , 169: 328-333. Accessed April 01, 2013 at http://faculty.uml.edu/rhochberg/hochberglab/Courses/InvertZool%202/CoursePDFS/1-Spadella%20mating%20sequence.pdf .
Helmkampf, M., I. Bruchhaus, B. Hausdorf. 2008. Multigene analysis of lophophorate and chaetognath phylogenetic relationships. Molecular Phylogenetics and Evolution , 46/1: 206-214.
Leuckart, R. 1854. Zoologische Untersuchungen . Giessen, Germany: J. Ricker'sche Buchhandlung.
Littlewood, D., M. Telford, R. Bray. 2004. Protostomes and Platyhelminthes : The Worm's Turn. Pp. 209-236 in Assembling the Tree of Life . Oxford, UK: Oxford University Press.
Margulis, L., M. Chapman. 2010. Kingdoms and Domains: An Illustrated Guide to the Phyla of Life on Earth, 4th Edition, reprinted with corrections . Philadelphia, PA: Academic Press.
Matus, D., R. Copley, C. Dunn, A. Hejnol, H. Eccleston, K. Halanych, M. Martindale, M. Telford. 2006. Broad taxon and gene sampling indicate that chaetognaths are protostomes. Current Biology , 16/15: R575-R576.
McLean, N., C. Nielsen. 1989. Oodinium jordani n. sp., a dinoflagellate ( Dinoflagellata : Oodinidae ) ectoparasitic on Sagitta elegans ( Chaetognatha ). Diseases of Aquatic Organisms , 7: 61-66. Accessed April 02, 2013 at http://www.int-res.com/articles/dao/7/d007p061.pdf .
Papillon, D., Y. Perez, X. Caubit, Y. Le Parco. 2004. Identification of Chaetognaths as Protostomes Is Supported by the Analysis of Their Mitochondrial Genome. Molecular Biology and Evolution , 21/11: 2122-2129.
Papillon, D., Y. Perez, X. Caubit, Y. Le Parco. 2006. Systematics of Chaetognatha under the light of molecular data, using duplicated ribosomal 18S DNA sequences. Molecular Phylogenetics and Evolution , 38/3: 621-634.
Paps, J., J. Baguñà , M. Riutort. 2009. Bilaterian Phylogeny: A Broad Sampling of 13 Nuclear Genes Provides a New Lophotrochozoa Phylogeny and Supports a Paraphyletic Basal Acoelomorpha . Molecular Biology and Evolution , 26/10: 2397-2406.
Ramel, G. 2012. "The Phylum Chaetognatha " (On-line). Earthlife. Accessed April 01, 2013 at http://www.earthlife.net/inverts/chaetognatha.html .
Shapiro, L. 2012. " Chaetognatha " (On-line). Encyclopedia of Life. Accessed April 01, 2013 at http://eol.org/pages/1740/overview .
Szaniawski, H. 2005. Cambrian chaetognaths recognized in Burgess Shale fossils. Acta Palaeontologica Polonica , 50/1: 1-8.
Telford, M., P. Holland. 1997. Evolution of 28S ribosomal DNA in chaetognaths: duplicate genes and molecular phylogeny. Journal of Molecular Evolution , 44/2: 135-144.
Telford, M., P. Holland. 1993. The phylogenetic affinities of the chaetognaths: a molecular analysis. Molecular Biology and Evolution , 10/3: 660-676.
Thuesen, E. 2009. " Chaetognatha " (On-line). Accessed April 01, 2013 at http://academic.evergreen.edu/t/thuesene/chaetognaths/chaetognaths.htm .
Thuesen, E., F. Goetz, S. Haddock. 2010. Bioluminescent organs of two deep-sea arrow worms, Eukrohnia fowleri and Caecosagitta macrocephala , with further observations on bioluminescence in chaetognaths. The Biological Bulletin , 219/2: 100-111. Accessed April 01, 2013 at http://www.ncbi.nlm.nih.gov/pubmed/20972255 .
Thuesen, E., K. Kogure. 1989. Bacterial production of tetrodotoxin in four species of Chaetognatha . The Biological Bulletin , 176: 191-194. Accessed April 02, 2013 at http://www.jstor.org/stable/1541587 .
Vannier, J., M. Steiner, E. Renvoise, S. Hu, J. Casanova. 2007. Early Cambrian origin of modern food webs: evidence from predator arrow worms. Proceedings of the Royal society B: Biological Science , 274: 627-633.
Zhang, Z. 2011. Animal biodiversity: An introduction to higher-level classification and taxonomic richness. Zootaxa , 3148: 7-12.
Zo, Z. 1973. Breeding and growth of the chaetognath Sagitta elegans in Bedford Basin. Limnology and Oceanography , 18/5: 750-756. Accessed April 01, 2013 at http://www.aslo.org/lo/toc/vol_18/issue_5/0750.pdf .
Øresland, V. 1986. Parasites of the chaetognath Sagitta setosa in the western English Channel. Marine Biology , 92/1: 87-91. Accessed April 02, 2013 at http://link.springer.com/article/10.1007/BF00392750?LI=true .
2012. " Chaetognatha " (On-line). World Register of Marine Species. Accessed April 01, 2013 at http://www.marinespecies.org/aphia.php?p=taxdetails&id=2081 .
2002. "Arrow Worms — Phylum Chaetognatha " (On-line). Canada's Polar Life. Accessed April 01, 2013 at http://www.polarlife.ca/organisms/inverts/marine_inverts/chaetognaths.htm .