Diversity
The phylum
Cycliophora
, only recently described in 1995, consists of at least three species of acoelomate,
bilaterally symmetrical organisms that are obligate commensalists on the mouthparts
of lobsters. There are two formally described species in the phylum,
Symbion pandora
and
Symbion americanus
, with at least one additional, undescribed species known.
Symbion pandora
was first discovered on the mouthparts of Norway lobsters (
Nephrops norvegicus
) in Scandinavian waters, and
Symbion americanus
was described from American lobsters (
Homarus americanus
) in North American waters. The third, undescribed, species may be found on European
lobsters (
Homarus gammarus
), in European waters. These organisms are among the smallest known free-living metazoans,
with females measuring around 350 µm in length, and males only reaching lengths of
30 to 40 µm, and containing (often significantly) fewer than 200 cells in their entire
body. These organisms filter feed on bacteria and food particles that escape from
their host's mouthparts, and despite their small size, exhibit a complex and distinctive
life cycle with multiple stages, including asexual feeding stages that may reproduce
by budding, as well as free-swimming male (Prometheus), female (Pandora), and internally
brooded chordoid larvae, and sessile dwarf males that live attached to the body wall
of females.
Geographic Range
Cycliophorans are only known from marine waters of the Northern hemisphere, with the
same distributions as the lobster species they live on.
- Biogeographic Regions
- nearctic
- palearctic
- atlantic ocean
- mediterranean sea
- Other Geographic Terms
- holarctic
Habitat
Cycliophorans live on the mouthparts of their lobster hosts. Feeding individuals and
chordoid cysts may be found on all six feeding mouthparts, most often on individuals
with a carapace length greater than 35 mm. The numbers of individuals living on a
host increases with size; over a thousand feeding cycliophorans and nearly 200 chordoid
cysts have been found on larger lobsters. Sessile larvae may settle near their female
progenitors, or disperse and colonize a new host. They have been found from the intertidal
zone to depths of 720 m.
- Habitat Regions
- temperate
- saltwater or marine
Systematic and Taxonomic History
The phylum
Cycliophora
was described by Peter Funch and Reinhardt Kristensen in 1995. Due to its recent
description, no alternative names for this phylum or its member species exist. Based
upon morphological characteristics, the authors hypothesized a sister relationship
to the phylum
Entoprocta
, or perhaps
Polyzoa
(then referred to as the superphylum
Bryozoa
), which contains both
Entoprocta
and
Ectoprocta
. Many subsequent molecular and morphological phylogenetic analyses have supported
a sister group relationship between
Entoprocta
and
Cycliophora
, and thus, their inclusion in
Polyzoa
. However, other molecular and combined analyses have suggested a relationship to
the taxon
Gnathifera
, a group which contains rotifers, thorny-headed worms, gnathostomulids, and
Micrognathozoa
(which contains a single species,
Limnognathia maerski
). Though their exact placement remains the subject of debate, either set of relationships
would warrant their inclusion in the superphylum
Lophotrochozoa
.
Physical Description
Cycliophorans have an anterior buccal funnel; oval-shaped trunk; and posterior, acellular
stalk, with an adhesive disc they use to attach themselves to their hosts’ mouthparts.
Sessile stage females are approximately 350 µm long and 100 µm wide. The trunk and
adhesive disc are covered in a layered cuticle (the disc itself may also be comprised
of cuticle). Cycliophorans are acoelomate, with the area between their guts and body
walls filled with mesenchyme. They have a feeding ring around the buccal funnel that
is densely packed with cilia and contractile cells, which form a pair of sphincters
capable of closing the oral area. Two muscle fibers extend dorsally from the base
to the ventral side of the trunk, and are likely used to move the buccal tube during
feeding. The gut, which is entirely ciliated, is U-shaped. A curved esophagus connects
the buccal funnel to a stomach with large gland cells and a narrow lumen. The intestine
leads to a dorsal rectum and anus, located near the buccal funnel.
Cycliophorans exhibit sexual dimorphism. Males only measure 30 to 40 µm in length
with heavily ciliated bodies and a ventral-posterior penis and associated pouch. They
are typically found free swimming or on the body of a female. Their bodies may be
round or more rectangular in shape. Once thought to have greatly reduced body structures,
recent research has shown that males do possess complex musculature, a large cerebral
ganglion and nerve cords, fully developed gonads and mating structures, and sensory
organs, as do females.
- Other Physical Features
- ectothermic
- heterothermic
- bilateral symmetry
- Sexual Dimorphism
- female larger
- sexes shaped differently
Development
The most definitive characteristic of cycliophorans is their complex reproductive
cycle, consisting of an asexual and sexual generation. There are two forms of asexual,
sessile, feeding animals. One form may house “Pandora larvae” and the other form may
house a primary male and female together in a brooding chamber. Sessile animals may
also undergo internal budding, in which they lose their buccal funnels and entire
digestive systems, replacing them with a new bud. The bud arises from embryonic cells
in the posterior region of the animal’s trunk. This process is repeated many times
during the sessile animal’s life. Pandora larvae also undergo internal budding in
order to form a feeding stage.
Asexual feeding individuals may change to sexual reproduction and create a primary
male and sexually mature female instead of asexual offspring; this may be triggered
by an impending molt of the host, as sexual individuals are sessile for a time. In
this case, a primary male, also known as a "Prometheus larva," is released from the
brooding chamber with no sexual organs or gonads, only developing them if it settles
on a chamber housing a female. In this event, the male produces secondary males via
budding. A secondary male has a cuticular, tubular penis and one spermatozoa compartment.
A sexual female will not bud internally; instead, she has one large anterior oocyte.
Fertilization occurs just before or after her release from the brooding chamber; the
exact method of sperm transfer is not known. She will then settle and brood herself
into a chordoid larva, which will engulf all of her tissue, leaving only her cuticle.
The chordoid larvae hatches and disperses, settling on a new host and beginning the
cycle again by budding a new, asexual, feeding cycliophoran.
Reproduction
Cycliophorans reproduce both asexually (producing either Pandora larvae, which settle
and become females, or brooding chambers housing male and female individuals) and
sexually. When they reproduce sexually, asexual males bud and creates multiple, sexual
males, which are free swimming until they settle on a female and transfer sperm to
her.
- Mating System
- polygynandrous (promiscuous)
Individuals reproduce both sexually and asexually. No specific breeding season has
been identified for these animals but the appearance of sexually mature individuals
is linked to the molting cycle of their hosts.
- Key Reproductive Features
- year-round breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- asexual
- fertilization
Cycliophorans exhibit no parental investment beyond production of asexual and sexual
offspring.
- Parental Investment
- no parental involvement
Lifespan/Longevity
There is no data currently available regarding the average lifespan of cycliophorans.
Behavior
Although thousands of individuals may be found on the mouthparts of a single host,
cycliophorans are solitary. They are sessile, using the adhesive discs at the end
of their stalks to attach to their hosts’ mouthparts. Only larvae and males are free-swimming.
Communication and Perception
Cycliophorans possess a relatively well-developed cerebral ganglion and a pair of
longitudinal nerves that proceed ventrolaterally from this structure. Males, females,
Pandora, and Prometheus larvae all possess frontal and lateral head sensilla that
may serve as mechanosensory structures. Electron microscopy of male cycliophorans
has revealed structures that may represent additional tactile sensory organs, such
as frontal palps and dorsal papillae, and a structure in the cerebral ganglion of
one individual that was tentatively identified as a statocyst. The prevalence and
function of these structures requires further investigation and verification. It would
appear likely that cycliophorans must possess some chemosensory ability as well, as
they are able to synchronize their reproductive and developmental behaviors with their
host's molting and feeding cycles. The basis of their interactions with one another
remains unknown.
Food Habits
Cycliophorans are filter feeders during their sessile stage; they do not feed during
free swimming stages. Their circular mouths are surrounded by a ring of compound cilia
that create a feeding current; they typically consume small food particles from their
hosts, or bacteria. It has been hypothesized that they depend solely on particles
generated by their hosts and that the sessile stage is triggered by increased feeding
by the host.
- Primary Diet
- planktivore
- Foraging Behavior
- filter-feeding
Predation
No predators specific to cycliophorans have been identified; however, any predator
of hosts carrying them, such as large demersal fishes, would necessarily consume these
animals as well.
Ecosystem Roles
Cycliophorans are only found living as commensals on (or, in free swimming stages,
near) their lobster hosts.
- American lobsters Homarus americanus (Class Malacostraca , Phylum Arthropoda )
- European lobsters Homarus gammarus (Class Malacostraca , Phylum Arthropoda )
- Norway lobsters Nephrops norvegicus (Class Malacostraca , Phylum Arthropoda )
Economic Importance for Humans: Positive
There are no known positive effects of cycliophorans on humans, outside of the potential
for scientific research.
- Positive Impacts
- research and education
Economic Importance for Humans: Negative
Although these animals live on lobsters, they do not generally adversely affect their
hosts, although it is possible for large numbers of cycliophorans to clog their host's
mouthparts. There are no known adverse effects of cycliophorans on humans.
Conservation Status
Additional Links
Contributors
Jeremy Wright (author), University of Michigan-Ann Arbor, Leila Siciliano Martina (editor), Texas State University.
- 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.
- 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.
- 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.
- 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).
- saltwater or marine
-
mainly lives in oceans, seas, or other bodies of salt water.
- 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.
- coastal
-
the nearshore aquatic habitats near a coast, or shoreline.
- 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.
- polygynandrous
-
the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.
- year-round breeding
-
breeding takes place throughout the year
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- asexual
-
reproduction that is not sexual; that is, reproduction that does not include recombining the genotypes of two parents
- fertilization
-
union of egg and spermatozoan
- internal fertilization
-
fertilization takes place within the female's body
- sessile
-
non-motile; permanently attached at the base.
Attached to substratum and moving little or not at all. Synapomorphy of the Anthozoa
- motile
-
having the capacity to move from one place to another.
- sedentary
-
remains in the same area
- solitary
-
lives alone
- tactile
-
uses touch to communicate
- chemical
-
uses smells or other chemicals to communicate
- filter-feeding
-
a method of feeding where small food particles are filtered from the surrounding water by various mechanisms. Used mainly by aquatic invertebrates, especially plankton, but also by baleen whales.
- planktivore
-
an animal that mainly eats plankton
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