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Symbiotic relationships |
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Types of symbioses exhibited by west-coast shrimps include mutualism, where both participants in the relationship benefit, commensalism, where one participant benefits while the other is not harmed, and parasitism, where one participant benefits at the expense of the other.
The following studies are separated into sub-sections for “true shrimps” (F. Thoridae & F. Crangonidae) and “mud & ghost” shrimps (Infraorder Thalassinidea) |
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"True" shrimps |
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Thorid & Crangonid shrimps
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Research study 1 |
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 A cleaning role has been suggested for shrimps Lebbeus mundus on the west coast. Although this behavior appears not to have been formally reported in the scientific literature, anecdotal descriptions of them cleaning the mouths of dusky rockfishes Sebastes ciliatus and ling cods Ophiodon elongatus are recounted by a crustacean researcher at the University of Washington, and a mutualistic symbiosis is strongly suspected. Jensen 2006 Zootaxa 1383: 23; see also Jensen 2006 Pacific Coast Crabs & Shrimps. Sea Challengers, Monterey, California +87pp. Photograph courtesy the author.
"Cleaner" shrimps Lebbeus mundus 4X
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Research study 2 |
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 Although ling cods are voracious predators and routinely eat small crustaceans, a cleaner shrimp busying itself on sensitive parts of a fish presenting itself for cleaning (skin, lips, mouth lining, gills) would never be in harm’s way. If Lebbeus mundus’ behaviour is the same or similar to that of tropical cleaner shrimps, then it would be removing skin flakes, old mucus, and parasites (e.g., copepods, isopods). The relationship thus benefits both partners. The shrimp obtains food and the fish gets a cleaning. Photograph taken from a video filmed by Neil McDaniel, a well-known B.C. underwater photographer and videographer
Ling cod Ophiodon elongatus with alleged
cleaner shrimp Lebbeus mundus 0.75X |
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Research study 3 |
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There are 6-7 crangonid shrimps along the west coast and most are parasitized to some degree or another by a bopyrid isopod Argeia pugettensis. Researchers at the Hatfield Marine Science Center, Oregon report, for example, that both sexes of the bay shrimp Lissocrangon  stylirostris near Coos Bay, Oregon are about 62% infected. The isopod, usually only one per host, inhabits the gill chamber. The parasite has multiple effects on its host, including sterilising of females, reducing the host’s ability to capture food, and affecting its growth. Only a small percentage of shrimps (0.2%) are both brooding and parasitised, suggesting either that the parasite sterilises the host female or that it reduces its metabolic activity to such an extent that it cannot brood. In either case, the researchers suggest that the parasite could reduce the population size of L. stylirostris to such an extent that it eventually becomes ecologically extinct. Jarrin & Shanks 2008 J Crust Biol 28: 613. Photograph courtesy Laury Perry, Portland, Oregon.
NOTE many species of shrimps have some type of protandrous development, meaning that most or all large individuals will be potentially breeding females
Parasitised shrimps Lissocrangon pugettensis 1X |
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Mud & ghost shrimps (Inraorder Thalassinidea) |
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Research study 1 |
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 Mud- and ghost-shrimp burrows may be co-inhabited by several different commensal species including goby fishes, pea crabs, worms, phoronids, and bivalves. Most notable, perhaps, is the bivalve Cryptomya californica which lives embedded in the burrow walls. This commensal style of life effectively increases the depth at which the clams can live, removing them from contact with predators and from competitors at the sediment-water surface. MacGinitie & MacGinitie 1968 Natural history of marine animals. McGraw-Hill, N.Y.
Cryptomya californica 1.5X |
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Research study 1.1 |
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 The goby Typhlogobius californiensis lives commensally and solely with the most southern-most ghost-shrimp species in California, Neotrypaea biffari. Both participants generally come in pairs, one of each sex. The gobies are blind, pale pink in colour, 2-4cm in length, and spend most of their time in the deep reaches of the burrow system (see photograph on Right). The shrimps feed by straining planktonic material from water currents that they pump through the burrow. As both shrimps may be pumping at the same time at different burrow entrances, the system usually has 3 and sometimes more openings (each less than 1cm in diameter). The burrows are located intertidally, but usually amongst boulders, so they are relatively permanent (see photograph below Left). A researcher at the Kerckoff Marine Laboratory, Corona del Mar, California provides details of burrow life through years of laboratory observation, aided by constraining the occupants between 2 glass plates mounted in a frame (2cm separation). The shrimps’ tasks are to dig, clean, and repair the burrow system, and to maintain a steady water current. This is necessary both for themselves and for the well-being of the gobies, who rely on the shrimps’ pumping for respiratory ventilation and to bring in edible organic bits, ones too large for the shrimps to eat and allowed by them to pass into the burrow. Other than cleaning up organic debris by eating it, the gobies’ contributions to the relationship are few. The author notes that they may sometimes act to repel invading species but, as they are not particularly aggressive and also blind, their contributions relative to those of the shrimps are small. Contact between host and commensal is minimal, limited mostly to pinching or shoving by the shrimps when the gobies get in the way, and to occasional deliveries of food bits to the fish in sediment loads that are otherwise destined to be dumped at one of the exit holes. Interestingly, in a laboratory situation where food is provided and predators excluded, the relationship is not obligate for either organism, but how the fish would do on their own in the field is another question. MacGinitie  1939 Am Midl Nat 21 (2): 489.
NOTE the relationship is usually termed commensal but, as little benefit seems to confer to the shrimp hosts, parasitism is also possible
NOTE at hatching the eyes appear to be normal, but once a fish enters a burrow they become distorted in shape and covered over by layers of skin. Light/dark perception is evident in younger fish, but this diminishes with age as the covering skin becomes thicker
Prime shrimp-goby habitat in
southern California ca. 1938
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Research study 2 |
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An obligate commensal with mud shrimps Upogebia pugettensis in most or all areas1 of the west coast is the phoronid Phoronis pallida. The phoronids live as adults in cemented-sand tubes embedded in the burrow wall (photo on Right), which itself is constructed of sediments bound in mucus (in addition to its binding properties, the mucus acts as a lubricant for the shrimp’s movements; the mucus is secreted from special glands in the hindgut area of the shrimp). The phoronid shown in the photo on the Left occupies the comma-shaped sand-tube originating at the 6 o'clock position. Its feeding apparatus, known as a lophophore, is fully extended.
The author's studies show that competent actinotroch larvae of Phoronis modify their behaviour in the presence of Upogebia-conditioned seawater2 such that swimming rates increase significantly (approximately 3-fold) in both downwards and horizontal directions. Once at the sea bottom the larvae probe the sediments with their sensory oral hoods. The author’s experiments further indicate that ammonia released in urinary excretions from Upogebia is not involved; rather, the chemical cue3 appears to be present in the gut egestia. Also effective at cueing the behaviour are seawater extracts of Upogebia gut tissue and burrow walls. The graph at the lower Right illustrates the dosage-dependency of the response. Santagata 2004 Mar Biol 207: 103.
 NOTE1 collections are made in Bodega Bay, California, Coos Bay, Oregon, and False Bay, Washington
NOTE2 2 liters of seawater in which 2 adult shrimps have been sitting for 2h (at 15oC). "Dose" in the graph represents µl of Upogegia-conditioned seawater . ml-1 fresh seawater
NOTE3 isolation of the substance from Upogegia-conditioned seawater indicates a molecular size of 10-50 kDa. Its activity is eliminated by treatment with arginase and significantly reduced by treatment with lipase, suggesting a peptide/fatty acid chemical composition |
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Research study 3 |
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 Transport of species from one region to another as live bait for recreational fisheries invariably runs the risk of escapees surviving to interbreed with the local population. To assess whether this occurs with ghost shrimps Neotrypaea californiensis being shipped to bait shops in southern California from Oregon and Washington, researchers at California State University, Long Beach analyse 2 mitochondrial DNA markers in populations ranging from False Bay, San Juan Islands, Washington to San Diego, California. Results reveal nonsignificant genetic structure across the range sampled, suggesting a low risk of genetic homogenisation from interbreeding. However, the researchers additionally present evidence of a second putative species of Neotrypaea in southern California whose northern range limit is Pt. Conception (see Clade A in diagram), a finding they suggest that is deserving of further reseach. Finally, and germane to this section on shrimp parasites, they note that about 6% of N. californiensis purchased in bait shops are infected with the host-castrating bopyrid isopod Ione cornuta, and suggest that this risk also merits further study. Pernet et al. 2008 Mar Biol 153: 1127.
NOTE genes analysed are fragments of mitochondrial cytochrome b and cytochrome oxidase I
NOTE samples are from 2 baitshops in Los Angeles County and Orange County, and from 5 field locations north of Pt. Conception and 7 south of Pt. Conception
Distribution of 2 clades of ghost shrimps Neotrypaea
californiensis along the west coast. Shrimps sampled from
2 bait shops in Los Angeles County and Orange County
were originally collected in Oregon and Washington |
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Research study 4 |
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 Large mud shrimps Upogebia pugettensis in Yaquina Bay, Oregon are almost 100% infested with bopyrid1 isopods Orthione griffenis (see histograms on Right). Note that the parasites do not infect shrimps less than about 11mm carapace length. The isopods live in the branchial chambers attached to the gills. Researchers at the Hatfield Marine Science Center in Newport, Oregon are able to correlate mass2 loss in the shrimps with the mass of parasites they carry. All bopyrid isopods, including Orthione, are obligate blood-parasites that feed on the host’s hemolymph and thus cause loss in mass of the host proportional to the parasite’s energetic requirements3. The authors' results show an average weight loss of 8% owing to the presence of the parasites, but the data are quite variable. An additional negative effect of a bopyrid parasite is that it often castrates its (male?) host, thus reducing the host’s fitness to zero but without killing it. However, this effect does not seem to be present in the Yaquina shrimp population. Smith et al. 2008 J Crust Biol 28: 228. Photograph courtesy USDA Agricultural Research Service.
NOTE1 world species of thalassinidean decapods in the genera Upogebia and Neotrypaea are commonly infested with bopyrid isopods. Upogebia spp. alone are known to harbour at least 29 species of the parasite. Markham 2005 Proc Biol Soc Washington 118: 84
NOTE2 about 1-2% of the shrimps also have parasitic clams Neaeromyra rugifera attached to their pleopods, and these may impose an additional “parasitic cost”, for example, interference with ventilation and brood carrying. This might be something worth further investigation
 NOTE3 reproductive-energy requirements in Orthione are huge. A single brood represents about 40% of a female’s mass and is made up, on average, of about 40,000 eggs
Mud shrimp Upogebia pugettensis with parasitic isopod Orthione griffensis removed. Note that the shrimp also has parasitic clams attached to its pleopods 1.6X
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Research study 5 |
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 Later work at the Oregon Institute of Marine Biology on bopyrid isopods Orthione griffenis parasitic on host thalassinid shrimps Upogebia pugettensis, shows that the isopod originates in China and was likely introduced to the west coast as larvae released in ballast water. Records of the species in China pre-date the first records of its presence on the west coast by at least 20yr. In North America Orthione infests mud shrimps U. pugettensis from British Columbia to Point Conception, California. South of the Point it switches to a replacement mud-shrimp species U. macginitieorum. The isopod inhabits the branchial chamber of its host and, in addition to possible effects on gas exhange, it may reduce fecundity of female hosts of up to 100%. Infestation of shrimps may reach 65% in Oregon populations. The authors provide the first evidence that specimens of Orthione from Asia and North America are the same species. Williams & An 2009 Integr Comp Biol 49: 1; for a comparison of O. griffenis on both sides of the Pacific see Williams & An 2009 Integr Comp Biol 49: 114. Photograph courtesy Jason Williams, Hofstra University, New York and Oxford University Press, U.K.
Mud shrimp Upogebia pugettensis with a
female bopyrid isopod Orthione griffenis in
its branchial chamber (reflected upwards) 2X |
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Research study 6 |
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 The bopyrid-isopod parasite Orthione griffensis was introduced to the west coast from Asia sometime in the 1980s. Two decades later, in 2002, the mud-shrimp Upogebia pugettensis population in Willapa Bay, Washington suffered a huge decline in numbers (see graph), and the question arises as to whether the two events are related. Researchers based primarily at the Hatfield Marine Science Center, Oregon provide strong correlative evidence from 20yr of data that this is indeed the case. First, levels of infestation of mud shrimps by Orthione are almost non-existent in Willapa Bay during 1988-97 (<1%), but by 1998-2002 are significantly greater, both in males (5-8%) and in females, notably in the 2yr primary-breeder age-class (30%). Second, the increase in some portions of the Willappa Bay population is exponential from about 1997. Although causing blood loss and castration in both sexes, bopyrids do not necessarily kill their hosts. Fecundity, however, my be significantly decreased, as shown by 68% reproductive losses seen in mud-shrimp populations in Yaquina Bay, Oregon between 2005-09. Although not assessed specifically, the researchers perforce assume that the levels of parasitism seen in Willapa Bay must have affected both recruitment and population dynamics of the mud shrimps. Dumbauld et al. 2011 Estuaries & Coasts 34: 336.
NOTE a former west-coast researcher now at the University of South Carolina, Columbia, has earlier used simulation modeling to explore potential mechanisms for causal links between the invasive parasite and decline of its host. The author’s overall conclusion is that the parasite interferes with its host’s reproduction by increasing the host’s metabolic requirements. Several hypotheses, all testable, are generated by the models. These are: 1) infection should be proportional to degree of exposure to parasite-laden water, and this will vary with tidal height and pumping rate of the shrimp, 2) the metabolic burden will be proportional to parasite size, 3) the greater the metabolic imposition, the greater will be the degree of feminisation of male hosts, and 4) metabolic costs of infection will be most severe in higher intertidal regions, where feeding time is most restricted. The author concludes by stating that, if tested, these hypotheses will greatly contribute to our understanding of the interactive dynamics of Upogebia and Orthione. Griffen 2009 Mar Ecol Progr Ser 391: 73. |
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