Neotrypaea californiensis
Encyclopedia
Neotrypaea californiensis (formerly Callianassa californiensis), the Bay ghost shrimp, is a species
of ghost shrimp
that lives on the Pacific coast
of North America
. It is a pale animal which grows to a length of 11.5 cm (4.5 in). One claw is bigger than the other, especially in males, and the enlarged claw is thought to have a function in mating
. N. californiensis is a deposit feeder that lives in extensive burrow systems, and is responsible for high rates of bioturbation
. It adversely affects oyster farms
, and its numbers are controlled in some places by the application of pesticides. It carries out an important role in the ecosystem, and is used by fishermen as bait
.
s, and a pink abdomen
.
Adult N. californiensis have one claw larger than the other, and in the males, the "master claw" can make up as much as 25% of the animal's mass – compared to only 10% in females – with the minor claw making up around 3% of the total body mass in both sexes. The enlarged claw is equally likely to be on the right side or the left side. The male's larger claw is thought to be used in agonistic encounters
or during mating
, and may be the result of sexual selection
.
Eggs are laid in spring or early summer, and the larvae
hatch in summer, living as plankton
. They settle to the sea floor again as post-larvae in the late summer and fall.
in 1854 by James Dwight Dana
as a member of the genus Callianassa
, giving the type locality as "California"; the material Dana studied was probably collected from San Francisco Bay
or Monterey
, but the original specimens have since been lost. In 1991, Raymond Manning
and Darryl Felder transferred the three species in that genus that come from California
and Oregon
into the new genus Neotrypaea
. N. californiensis is distinguished from the other two species of Neotrypaea by the lack of a rostrum
(which is present in Neotrypaea gigas) and the acute and diverging tips of the eyestalk
s (which are short, blunt and not diverging in Neotrypaea biffari).
Both Neotrypaea californiensis and the mud shrimp Upogebia pugettensis live in mudflat
s and sand
y substrates
in the intertidal zone
of estuaries
in western North America. N. californiensis is found from Mutiny Bay, Alaska
to Punta Abreojos
, Mulegé
, Baja California Sur
, Mexico
. Its habitat is also used for the aquaculture
of the Pacific oyster
, Crassostrea gigas. Since the bioturbation
carried out by N. californiensis and U. pugettensis reduces the productivity of the oyster beds, they are considered pests. Their effects may, however, have knock-on effects across the entire ecosystem
, and may buffer it from the hazards of nutrient enrichment
and increase primary
and secondary productivity by increasing the amount of dissolved inorganic nitrogen
.
The burrows made by N. californiensis have many branches, and a number of other animals live in them, including snapping shrimp
of the genus Betaeus
, the copepod
Clausidium vancouverense
, and the crab
Scleroplax granulata. The gut flora
of N. californiensis includes a wide range of bacteria
, comprising around 40% Alphaproteobacteria
, 20% gram-positive bacteria, 20% in the Cryptophaga–Flavobacteria
–Bacteroides
group, and 5% of each of Gammaproteobacteria
and Epsilonproteobacteria
. Predators of N. californiensis include bottom-dwelling fish
and Dungeness crab
s (Metacarcinus magister).
N. californiensis has a negative impact on oyster
production, and as a result, the insecticide
carbaryl
(1-napthyl N-methyl carbamate) is sprayed in some areas (including Willapa Bay
, Washington) to reduce the population of N. californiensis. The addition of shelly debris also reduces numbers of N. californiensis both by preventing the settlement of larva
e, and through predation on the young N. californiensis by young Dungeness crabs in the shelly debris.
N. californiensis is used as fishing bait
, and is frequently transported alive between U.S. state
s, prompting fears that existing population structure
may be obliterated, and that it could introduce the castrating
parasitic
isopod
Ione cornuta outside its native range.
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...
of ghost shrimp
Thalassinidea
Thalassinidea is an infraorder of decapod crustaceans that live in burrows in muddy bottoms of the world's oceans. In Australian English, the littoral thalassinidean Trypaea australiensis is referred to as the yabby , frequently used as bait for estuarine fishing; elsewhere, however, they are...
that lives on the Pacific coast
Pacific Ocean
The Pacific Ocean is the largest of the Earth's oceanic divisions. It extends from the Arctic in the north to the Southern Ocean in the south, bounded by Asia and Australia in the west, and the Americas in the east.At 165.2 million square kilometres in area, this largest division of the World...
of North America
North America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...
. It is a pale animal which grows to a length of 11.5 cm (4.5 in). One claw is bigger than the other, especially in males, and the enlarged claw is thought to have a function in mating
Mating
In biology, mating is the pairing of opposite-sex or hermaphroditic organisms for copulation. In social animals, it also includes the raising of their offspring. Copulation is the union of the sex organs of two sexually reproducing animals for insemination and subsequent internal fertilization...
. N. californiensis is a deposit feeder that lives in extensive burrow systems, and is responsible for high rates of bioturbation
Bioturbation
In oceanography, limnology, pedology, geology , and archaeology, bioturbation is the displacement and mixing of sediment particles and solutes by fauna or flora . The mediators of bioturbation are typically annelid worms , bivalves In oceanography, limnology, pedology, geology (especially...
. It adversely affects oyster farms
Oyster farming
Oyster farming is an aquaculture practice in which oysters are raised for human consumption. Oyster farming most likely developed in tandem with pearl farming, a similar practice in which oysters are farmed for the purpose of developing pearls...
, and its numbers are controlled in some places by the application of pesticides. It carries out an important role in the ecosystem, and is used by fishermen as bait
Fishing bait
Fishing bait is any substance used to attract and catch fish, e.g. on the end of a fishing hook, or inside a fish trap. Traditionally, nightcrawlers, insects, and smaller bait fish have been used for this purpose...
.
Description and life cycle
Neotrypaea californiensis reaches a length of 11.5 centimetres (4.5 in). The body is creamy white, with patches of pale colour (pink, yellow or orange) on the appendageAppendage
In invertebrate biology, an appendage is an external body part, or natural prolongation, that protrudes from an organism's body . It is a general term that covers any of the homologous body parts that may extend from a body segment...
s, and a pink abdomen
Abdomen
In vertebrates such as mammals the abdomen constitutes the part of the body between the thorax and pelvis. The region enclosed by the abdomen is termed the abdominal cavity...
.
Adult N. californiensis have one claw larger than the other, and in the males, the "master claw" can make up as much as 25% of the animal's mass – compared to only 10% in females – with the minor claw making up around 3% of the total body mass in both sexes. The enlarged claw is equally likely to be on the right side or the left side. The male's larger claw is thought to be used in agonistic encounters
Agonistic behaviour
In ethology, agonistic behaviour is any social behaviour related to fighting. Thus it is broader than aggressive behaviour because it includes not only actual aggression but also threats, displays, retreats, placating aggressors, and conciliation. The term was coined by Scott and Fredericson in 1951...
or during mating
Mating
In biology, mating is the pairing of opposite-sex or hermaphroditic organisms for copulation. In social animals, it also includes the raising of their offspring. Copulation is the union of the sex organs of two sexually reproducing animals for insemination and subsequent internal fertilization...
, and may be the result of sexual selection
Sexual selection
Sexual selection, a concept introduced by Charles Darwin in his 1859 book On the Origin of Species, is a significant element of his theory of natural selection...
.
Eggs are laid in spring or early summer, and the larvae
Crustacean larvae
Crustaceans may pass through a number of larval and immature stages between hatching from their eggs and reaching their adult form. Each of the stages is separated by a moult, in which the hard exoskeleton is shed to allow the animal to grow...
hatch in summer, living as plankton
Plankton
Plankton are any drifting organisms that inhabit the pelagic zone of oceans, seas, or bodies of fresh water. That is, plankton are defined by their ecological niche rather than phylogenetic or taxonomic classification...
. They settle to the sea floor again as post-larvae in the late summer and fall.
Taxonomy
N. californiensis was originally describedAlpha taxonomy
Alpha taxonomy is the discipline concerned with finding, describing and naming species of living or fossil organisms. This field is supported by institutions holding collections of these organisms, with relevant data, carefully curated: such institutes include natural history museums, herbaria and...
in 1854 by James Dwight Dana
James Dwight Dana
James Dwight Dana was an American geologist, mineralogist and zoologist. He made pioneering studies of mountain-building, volcanic activity, and the origin and structure of continents and oceans around the world.-Early life and career:...
as a member of the genus Callianassa
Callianassa (genus)
Callianassa is a genus of mud shrimps, in the family Callianassidae. Three of the species in this genus Callianassa is a genus of mud shrimps, in the family Callianassidae. Three of the species in this genus Callianassa is a genus of mud shrimps, in the family Callianassidae. Three of the species...
, giving the type locality as "California"; the material Dana studied was probably collected from San Francisco Bay
San Francisco Bay
San Francisco Bay is a shallow, productive estuary through which water draining from approximately forty percent of California, flowing in the Sacramento and San Joaquin rivers from the Sierra Nevada mountains, enters the Pacific Ocean...
or Monterey
Monterey, California
The City of Monterey in Monterey County is located on Monterey Bay along the Pacific coast in Central California. Monterey lies at an elevation of 26 feet above sea level. As of the 2010 census, the city population was 27,810. Monterey is of historical importance because it was the capital of...
, but the original specimens have since been lost. In 1991, Raymond Manning
Raymond B. Manning
Raymond Brendan Manning was an American carcinologist, specialising in alpha taxonomy and mantis shrimp.-Biography:...
and Darryl Felder transferred the three species in that genus that come from California
California
California is a state located on the West Coast of the United States. It is by far the most populous U.S. state, and the third-largest by land area...
and Oregon
Oregon
Oregon is a state in the Pacific Northwest region of the United States. It is located on the Pacific coast, with Washington to the north, California to the south, Nevada on the southeast and Idaho to the east. The Columbia and Snake rivers delineate much of Oregon's northern and eastern...
into the new genus Neotrypaea
Neotrypaea
Neotrypaea is a genus of ghost shrimp in the family Callianassidae, containing the following five species:*Neotrypaea biffari *Neotrypaea californiensis *Neotrypaea gigas...
. N. californiensis is distinguished from the other two species of Neotrypaea by the lack of a rostrum
Rostrum (anatomy)
The term rostrum is used for a number of unrelated structures in different groups of animals:*In crustaceans, the rostrum is the forward extension of the carapace in front of the eyes....
(which is present in Neotrypaea gigas) and the acute and diverging tips of the eyestalk
Eyestalk
In anatomy, an eyestalk is a protrusion that extends the eye away from the body, giving the eye a better field of view than if it were unextended. It is common in nature and in fiction....
s (which are short, blunt and not diverging in Neotrypaea biffari).
Ecology and human impact
Mudflat
Mudflats or mud flats, also known as tidal flats, are coastal wetlands that form when mud is deposited by tides or rivers. They are found in sheltered areas such as bays, bayous, lagoons, and estuaries. Mudflats may be viewed geologically as exposed layers of bay mud, resulting from deposition of...
s and sand
Sand
Sand is a naturally occurring granular material composed of finely divided rock and mineral particles.The composition of sand is highly variable, depending on the local rock sources and conditions, but the most common constituent of sand in inland continental settings and non-tropical coastal...
y substrates
Substrate (biology)
In biology a substrate is the surface a plant or animal lives upon and grows on. A substrate can include biotic or abiotic materials and animals. For example, encrusting algae that lives on a rock can be substrate for another animal that lives on top of the algae. See also substrate .-External...
in the intertidal zone
Intertidal zone
The intertidal zone is the area that is above water at low tide and under water at high tide . This area can include many different types of habitats, with many types of animals like starfish, sea urchins, and some species of coral...
of estuaries
Estuary
An estuary is a partly enclosed coastal body of water with one or more rivers or streams flowing into it, and with a free connection to the open sea....
in western North America. N. californiensis is found from Mutiny Bay, Alaska
Alaska
Alaska is the largest state in the United States by area. It is situated in the northwest extremity of the North American continent, with Canada to the east, the Arctic Ocean to the north, and the Pacific Ocean to the west and south, with Russia further west across the Bering Strait...
to Punta Abreojos
Punta Abreojos
' is a fishing town in Baja California Sur, Mexico.It is located at the entrance to , a Biosphere Reserve which is frequently visited by pods of grey whales. An isolated hill, high, rises a few miles behind the town, and is a conspicuous landmark for sailors...
, Mulegé
Mulegé (municipality)
Mulegé is the northernmost municipality of the Mexican state of Baja California Sur. It is the second-largest municipality by area in the country, with an area of 32,092.2 km² . In the census of 2010 it had a population of 59,114 inhabitants...
, Baja California Sur
Baja California Sur
Baja California Sur , is one of the 31 states which, with the Federal District, comprise the 32 Federal Entities of Mexico. Before becoming a state on October 8, 1974, the area was known as the South Territory of Baja California. It has an area of , or 3.57% of the land mass of Mexico and comprises...
, Mexico
Mexico
The United Mexican States , commonly known as Mexico , is a federal constitutional republic in North America. It is bordered on the north by the United States; on the south and west by the Pacific Ocean; on the southeast by Guatemala, Belize, and the Caribbean Sea; and on the east by the Gulf of...
. Its habitat is also used for the aquaculture
Aquaculture
Aquaculture, also known as aquafarming, is the farming of aquatic organisms such as fish, crustaceans, molluscs and aquatic plants. Aquaculture involves cultivating freshwater and saltwater populations under controlled conditions, and can be contrasted with commercial fishing, which is the...
of the Pacific oyster
Pacific oyster
The Pacific oyster, Japanese oyster or Miyagi oyster , is an oyster native to the Pacific coast of Asia. It has become an introduced species in North America, Australia, Europe, and New Zealand.- Etymology :...
, Crassostrea gigas. Since the bioturbation
Bioturbation
In oceanography, limnology, pedology, geology , and archaeology, bioturbation is the displacement and mixing of sediment particles and solutes by fauna or flora . The mediators of bioturbation are typically annelid worms , bivalves In oceanography, limnology, pedology, geology (especially...
carried out by N. californiensis and U. pugettensis reduces the productivity of the oyster beds, they are considered pests. Their effects may, however, have knock-on effects across the entire ecosystem
Ecosystem
An ecosystem is a biological environment consisting of all the organisms living in a particular area, as well as all the nonliving , physical components of the environment with which the organisms interact, such as air, soil, water and sunlight....
, and may buffer it from the hazards of nutrient enrichment
Eutrophication
Eutrophication or more precisely hypertrophication, is the movement of a body of water′s trophic status in the direction of increasing plant biomass, by the addition of artificial or natural substances, such as nitrates and phosphates, through fertilizers or sewage, to an aquatic system...
and increase primary
Primary production
400px|thumb|Global oceanic and terrestrial photoautotroph abundance, from September [[1997]] to August 2000. As an estimate of autotroph biomass, it is only a rough indicator of primary production potential, and not an actual estimate of it...
and secondary productivity by increasing the amount of dissolved inorganic nitrogen
Nitrogen
Nitrogen is a chemical element that has the symbol N, atomic number of 7 and atomic mass 14.00674 u. Elemental nitrogen is a colorless, odorless, tasteless, and mostly inert diatomic gas at standard conditions, constituting 78.08% by volume of Earth's atmosphere...
.
The burrows made by N. californiensis have many branches, and a number of other animals live in them, including snapping shrimp
Alpheidae
Alpheidae is a family of caridean snapping shrimp characterized by having asymmetrical claws, the larger of which is typically capable of producing a loud snapping sound. Other common names of these species include pistol shrimp or alpheid shrimp.The family is diverse and worldwide in distribution,...
of the genus Betaeus
Betaeus
Betaeus is a genus of shrimps of the family Alpheidae, containing the following species:*Betaeus australis Stimpson, 1860*Betaeus emarginatus Betaeus is a genus of shrimps of the family Alpheidae, containing the following species:*Betaeus australis Stimpson, 1860*Betaeus emarginatus Betaeus is a...
, the copepod
Copepod
Copepods are a group of small crustaceans found in the sea and nearly every freshwater habitat. Some species are planktonic , some are benthic , and some continental species may live in limno-terrestrial habitats and other wet terrestrial places, such as swamps, under leaf fall in wet forests,...
Clausidium vancouverense
Clausidium vancouverense
Clausidium vancouverense, the red copepod, is a symbiont of the ghost shrimp Neotrypaea californiensis. It is one of six species in the genus Clausidium and is found with its host in the Pacific Ocean from Alaska to Baja California....
, and the crab
Crab
True crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" , or where the reduced abdomen is entirely hidden under the thorax...
Scleroplax granulata. The gut flora
Gut flora
Gut flora consists of microorganisms that live in the digestive tracts of animals and is the largest reservoir of human flora. In this context, gut is synonymous with intestinal, and flora with microbiota and microflora....
of N. californiensis includes a wide range of bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...
, comprising around 40% Alphaproteobacteria
Alphaproteobacteria
Alphaproteobacteria is a class of Proteobacteria. Like all Proteobacteria, they are Gram-negative.-Characteristics:The Alphaproteobacteria comprise most phototrophic genera, but also several genera metabolising C1-compounds , symbionts of plants and animals, and a group of pathogens, the...
, 20% gram-positive bacteria, 20% in the Cryptophaga–Flavobacteria
Flavobacteria
The class Flavobacteria is composed of a single order of environmental bacteria. Flavobacteria are a group of commensal bacteria and opportunistic pathogens. Flavobacterium psychrophilum causes the septicemic diseases rainbow trout fry syndrome and bacterial cold water disease...
–Bacteroides
Bacteroides
Bacteroides is a genus of Gram-negative, bacillus bacteria. Bacteroides species are non-endospore-forming, anaerobes, and may be either motile or non-motile, depending on the species. The DNA base composition is 40-48% GC. Unusual in bacterial organisms, Bacteroides membranes contain sphingolipids...
group, and 5% of each of Gammaproteobacteria
Gammaproteobacteria
Gammaproteobacteria is a class of several medically, ecologically and scientifically important groups of bacteria, such as the Enterobacteriaceae , Vibrionaceae and Pseudomonadaceae. An exceeding number of important pathogens belongs to this class, e.g...
and Epsilonproteobacteria
Epsilonproteobacteria
Epsilonproteobacteria is a class of Proteobacteria. All species of this class are, like all Proteobacteria, gram-negative.The Epsilonproteobacteria consist of few known genera, mainly the curved to spirilloid Wolinella spp., Helicobacter spp., and Campylobacter spp...
. Predators of N. californiensis include bottom-dwelling fish
Fish
Fish are a paraphyletic group of organisms that consist of all gill-bearing aquatic vertebrate animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish, as well as various extinct related groups...
and Dungeness crab
Dungeness crab
The Dungeness crab, Metacarcinus magister , is a species of crab that inhabits eelgrass beds and water bottoms on the west coast of North America. It typically grows to across the carapace and is a popular seafood...
s (Metacarcinus magister).
N. californiensis has a negative impact on oyster
Oyster
The word oyster is used as a common name for a number of distinct groups of bivalve molluscs which live in marine or brackish habitats. The valves are highly calcified....
production, and as a result, the insecticide
Insecticide
An insecticide is a pesticide used against insects. They include ovicides and larvicides used against the eggs and larvae of insects respectively. Insecticides are used in agriculture, medicine, industry and the household. The use of insecticides is believed to be one of the major factors behind...
carbaryl
Carbaryl
Carbaryl is a chemical in the carbamate family used chiefly as an insecticide. It is a white crystalline solid commonly sold under the brand name Sevin, a trademark of the Bayer Company. Union Carbide discovered carbaryl and introduced it commercially in 1958...
(1-napthyl N-methyl carbamate) is sprayed in some areas (including Willapa Bay
Willapa Bay
Willapa Bay is a bay located on the southwest Pacific coast of Washington state in the United States. The Long Beach Peninsula separates Willapa Bay from the greater expanse of the Pacific Ocean. With over of water surface Willapa Bay is the second largest estuary on the United States Pacific coast...
, Washington) to reduce the population of N. californiensis. The addition of shelly debris also reduces numbers of N. californiensis both by preventing the settlement of larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
e, and through predation on the young N. californiensis by young Dungeness crabs in the shelly debris.
N. californiensis is used as fishing bait
Fishing bait
Fishing bait is any substance used to attract and catch fish, e.g. on the end of a fishing hook, or inside a fish trap. Traditionally, nightcrawlers, insects, and smaller bait fish have been used for this purpose...
, and is frequently transported alive between U.S. state
U.S. state
A U.S. state is any one of the 50 federated states of the United States of America that share sovereignty with the federal government. Because of this shared sovereignty, an American is a citizen both of the federal entity and of his or her state of domicile. Four states use the official title of...
s, prompting fears that existing population structure
Population stratification
Population stratification is the presence of a systematic difference in allele frequencies between subpopulations in a population possibly due to different ancestry, especially in the context of association studies...
may be obliterated, and that it could introduce the castrating
Castration
Castration is any action, surgical, chemical, or otherwise, by which a male loses the functions of the testicles or a female loses the functions of the ovaries.-Humans:...
parasitic
Parasitism
Parasitism is a type of symbiotic relationship between organisms of different species where one organism, the parasite, benefits at the expense of the other, the host. Traditionally parasite referred to organisms with lifestages that needed more than one host . These are now called macroparasites...
isopod
Isopoda
Isopods are an order of peracarid crustaceans, including familiar animals such as woodlice and pill bugs. The name Isopoda derives from the Greek roots and...
Ione cornuta outside its native range.