Limnoria Tripunctata Phylum: Arthropoda, Crustacea Class: Malacostraca Order: Isopoda; Flabellifera a Gribble Family: Limnoridae

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Limnoria tripunctata Phylum: Arthropoda, Crustacea Class: Malacostraca Order: Isopoda; Flabellifera A gribble Family: Limnoridae

Taxonomy: Limnoria was described in 1813 Antenna 1: First antenna flagellum by Leach and has been placed in a variety of with four articles and peduncle with three (Fig. isopod families since (e.g. Asellidae), until 3). Both antennae are reduced, separated at Harger erected the family Limnoriidae for it, in midline, and positioned in a nearly transverse 1880 (Menzies 1957). It was divided into two line (Fig. 1). subgenera on the basis of boring substrate Antenna 2: Second antenna and associated mouthparts (Cookson 1991). flagellum with five articles (Fig. 4). Limnoria Limnoria were the wood-borers while Mouthparts: Mandibles with file-like Limnoria Phycolimnoria were the algae-borers ridges (right) and rasping surface (left), but (Menzies 1957; Brusca 1980). Thus, lack lacina mobilis and molar processes Limnoria Limnoria tripunctata is sometimes (Brusca 1980). seen, although these subgeneric names are Pereon: rarely used today (Cookson 1991; Brusca et Pereonites: Seven total segments, al. 2007). the first of which is widest (Figs. 1, 2) and coxal plates are present on pereonites 2–7 Description (Brusca 1980). Size: Limnoriids are small and L. tripunctata Pereopods: In mature females, leaf- is no exception, reaching maximum lengths of like ooestegites are present at the base of 2.5 mm. each of first four pairs of legs and forms a Color: Light tan, whitish and often encrusted brood pouch or marsupium (see Fig. 6, with debris. Corophiurn spinicorne, this guide). General Morphology: Isopod bodies are Pleon: dorso-ventrally flattened and can be divided Pleonites: Five free pleonites with into a compact cephalon, with eyes, two fifth somite bearing three tubercles (Fig. 1). antennae and mouthparts, and a pereon Pleopods: (thorax) with eight segments, each bearing Uropods: Uropod branches dissimilar, with similar pereopods (hence the name “iso- short and claw-like exopod and long, apically pod”). Posterior to the pereon is the pleon, or blunt endopod (Fig. 6). abdomen, with six segments, the last of which Pleotelson: Ornamented pleotelson with is fused with the telson (the pleotelson) (see three anterior tubercles (“tri-punctata", Fig. 1) Plate 231, Brusca et al. 2007). The Isopoda and tuberculate posterior and lateral borders can be divided into two groups: ancestral (Fig. 5). (“short-tailed”) groups (i.e. suborders) that Sexual Dimorphism: Mature females are have short telsons and derived (“long-tailed”) conspicuous with a marsupium (see groups with long telsons. Members of the pereopods) and males with modified Flabellifera, to which L. tripunctata belongs, posterior end of the seventh pereonite (see fall into the long-tailed variety (see Fig. 86, Fig. 1, Menzies 1972). Kozloff 1993). Limnoria tripunctata individuals are able to roll into a ball and are Possible Misidentifications easily recognizable by their small size and The order Isopoda contains 10,000 species, wood-boring habits (Brusca 1980). 1/2 of which are marine and comprise 10 Cephalon: Smooth, rounded and modified suborders, with eight present from central for boring (Fig. 1). California to Oregon (see Brusca et al. Rostrum: 2007). Among isopods with elongated Eyes: Lateral and anterior (Fig. 1). telsons (with anuses and uropods that are subterminal), there are several groups (i.e.

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12720 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected] suborders) including the Valvifera, diameter throughout and have smooth walls Anthuridea, Gnathiidea, Epicaridea and (Sleeter and Coull 1973). Bite marks left on Flabellifera. wood by Limnoria species are distinct and The Flabellifera is a large measure 50–80 µm in diameter and tunnels assemblage and contains 3,000 species reach depths of 2 cm (Pitman et al. 1997). with seven families occurring locally, threeof They can completely bury themselves within which are not present north of Point wood in 4–6 days, but remain close to the Conception, California (Brusca et al. 2007). wood surface (Ricketts and Calvin 1952). Limnoriids are wood-boring species that are Salinity: A stenohaline species (Borges et al. characterized by reduced uropods, small 2014) that tolerates salinity and oxygen exopods, and a body that is less that 4 mm fluctuations as individuals occur in warm, in length (see Brusca et al. 2007 for other often salty upper bays. The ideal salinity distinguishing characters). This family range is 30–34 (in the lab, Borges et al. comprises only four local species (70 2009), but individuals tolerate salinities from described worldwide) and all are in the 12 to 48 (Menzies 1972). Other Limnoria genus Limnoria. species (e.g. L. lignorum) can't tolerate low Limnoria algarum, bores in algal salinity (15) or dissolved oxygen content holdfasts, not wood, and is the only species below 1.6 ppm. Limnoria tripunctata can with a simple incisor mandibular process, stand periodic oxygen depletion, however, lacking a file that is present in the other (Menzies 1957) and has been observed at three, wood-boring, species. Limnoria salinities of 12 near San Francisco, California, lignorum has a pleotelson with dorsal surface and can survive at salinities between 10–18. that forms a Y-shaped keel at the base while However, even L. tripunctata cannot survive L. quadripuntcata and L. tripunctata have exposure to freshwater for greater than one pleotelsons with symmetrically arranged day (Menzies 1957). anterior tubercles. The two latter species Temperature: Temperature range from 15° can be differentiated (as their names to 30°C (Beckman and Menzies 1960; suggest) by the number of tubercles present, Menzies 1972; Borges et al. 2009). Highest four in L. quadripuntcata and three in L. reproductive rates were observed between tripunctata (Brusca et al. 2007). 20–25˚C, reproduction is impaired below 10°C and egg production takes twice as long Ecological Information at 15˚C than at 20˚C (Menzies 1957, 1972). Range: Type locality is San Diego, Tidal Level: A shallow water species, L. California. Known range from Atlantic and tripunctata occurs from the water surface to Pacific coasts in temperate and tropical 18 meters deep. Individuals prefer lower waters and capable of interbreeding over depths when surface salinity is low or tidal large geographic distances (Menzies 1972) fluctuation is great. Individuals tend to prefer within a temperature range of 15–30˚C (44˚ to estuary benthos, and commonly occur at the 12° N) (Beckman and Menzies 1960). bases of pilings. Limnoria tripunctata is a well-established Associates: Limnoria burrows can be species in European coastal waters (Borges inhabited by the commensal isopod, et al. 2014). Caecijaera; the sphaeromatid isopod, Local Distribution: Oregon distribution in Gnorimosphaeroma; the amphipod, Chelura; upper bays including Coos, Yaquina, and the copepod Donsiella (Menzies 1957), Tillamook estuaries. none of which are borers. The boring Habitat: Docks and pilings, chiefly in bays mollusk, Teredo, can also co-occur in wood and estuaries, where it burrows into wood, where Limnoria burrows. After 4–6 months whether it is floating or submerged (Johnson submerged (suspended 1–4 meters above and Menzies 1956). The wood serves as the bottom), untreated wood with L. both food and protection. Limnoria tripunctata tripunctata developed a community consisting is even undeterred by creosote preserved of turbellarians, nematodes, the archiannelid wood (Menzies 1951; Ricketts and Calvin Dinophilus; the polychaete Polydora; the 1952; Borges et al. 2014) (see also Food). tanaid, Leptochelia savignyi; copepods and Within the wood, burrows are equal in amphipods (Sleeter and Coull 1973). The

Hiebert, T.C. 2015. Limnoria tripunctata. In: Oregon Estuarine Invertebrates: Rudys' Illustrated Guide to Common Species, 3rd ed. T.C. Hiebert, B.A. Butler and A.L. Shanks (eds.). University of Oregon Libraries and Oregon Institute of Marine Biology, Charleston, OR. heterotrich ciliate Microfolliculina limnoriae, Larva: Since most isopods are direct attaches to the dorsal surface of the developing, they lack a definite larval stage. pleotelson (up to four individuals per single L. Instead this young developmental stage tripunctata). The presence of this obligate resembles small adults (e.g. Fig. 40.1, Boyko ciliate may reduce feeding rate and negatively and Wolff 2014). Most isopods develop from affect swimming, suggesting that this embryo to a manca larva, consisting of three relationship is a form of ectoparasitism stages. Manca larvae are recognizable by (Delgery et al. 2006). lacking the seventh pair of pereopods, but Abundance: The small size of individuals in otherwise resemble small adults. They this genus allows for hundreds to co-occur in usually hatch from the female marsupium at a single square inch of wood. the second stage and the molt from second to third manca produces the seventh pair of Life-History Information pereopods and sexual characteristics (Boyko Reproduction: Most isopods have separate and Wolff 2014). Isopod development and sexes (i.e. dioecious, Brusca and Iverson larval morphology can vary between groups 1985) (although protogynous and protandric (e.g. Gnathiidae, Cryptoniscoidea, species are known, Araujo et al. 2004; Boyko Bopyroidae, Cymothoidae, Oniscoidea) (see and Wolff 2014). Reproduction proceeds by Boyko and Wolff 2014). Parasitic isopods, for copulation and internal fertilization where example, have larvae that are morphologically eggs are deposited within a few hours after dissimilar from adults (Sadro 2001). Isopod copulation and brooded within the female larvae are not common members of the marsupium (Brusca and Iverson 1985). The plankton, with parasitic larvae most likely to biphasic molting of isopods allows for be observed. Occasionally, suspended copulation; the posterior portion of the body benthic juveniles or pelagic species are molts and individuals mate, then the anterior collected in plankton samples, but these can portion, which holds the brood pouch, molts be differentiated from larvae by their larger (Sadro 2001). Embryonic development size (Sadro 2001). Newly hatched Limnoria proceeds within the brood chamber and is larvae do not swim, but develop this trait over direct with individuals hatching as manca time (Ricketts and Calvin 1952). larvae that resemble small adults, with no Juvenile: larval stage (Boyko and Wolff 2014). Longevity: There is some evidence that Limnoria species exhibit low fecundity, individuals leave their burrows and dig iteroparity and direct development (Menzies separate “tombs” into which they settle to die 1972) and females in the genus only carry an (Sleeter and Coull 1973). average of about 9–10 eggs and breeding Growth Rate: Growth among isopods occurs occurs year-round (Ricketts and Calvin 1952). in conjunction with molting where the Adult L. tripunctata occur and copulate as exoskeleton is shed and replaced. Post-molt pairs within tunnels (see Fig. 2, Menzies individuals will have soft shells as the cuticle 1972) and eggs require 2–4 weeks for gradually hardens. During a molt, arthropods development (Borges et al. 2014). Females have the ability to regenerate limbs that were can produce up to three broods per year, and previously autonomized (Kuris et al. 2007), the number of gravid females in a single however, isopods do not autotomize limbs as population is highest when water readily as other groups (Brusca and Iverson temperatures are between 17 and 19˚C 1985). Compared to other arthropods, (Johnson and Menzies 1956). Locally, peak isopods exhibit a unique biphasic molting, in breeding time for L. tripunctata is from April to which the posterior 1/2 of the body molts May (Friday Harbor, WA, Welton and Miller before the anterior 1/2 (Brusca et al. 2007). 1980) and the average number of eggs per Intermolt period is 25 days in L. tripunctata, female is 22 (Welton and Miller 1980). but decreases with time and age of individual Development time from egg deposition to (Ria Formosa, Portugal, Delgery et al. 2006). hatching is 17 days (at 20 °C), 15 days (at See Fig. 4 Johnson and Menzies 1956 for plot 22°C), 13 days (at 26°C), 11 days (at 30°C of seasonal chart of growth rate. but numbers greatly reduced) (Eltringham Food: Limnoria tripunctata is an 1967). economically significant species due to its

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12720 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected] ability to alter wooden structures by burrowing (Crustacea, Isopoda, Oniscidea): and ingesting wood (e.g. Fig. 1 Menzies sexual differentiation and size at 1957). They use wood as their primary onset of sexual maturity. carbon source by producing lignocellulose Invertebrate Reproduction and digesting enzymes (Borges et al. 2014) and, Development. 45:221-230. interestingly, have digestive systems void of 2. BECKMAN, C., and R. J. the microorganisms that aid in digestion of wood and cellulose among other metazoans MENZIES. 1960. The relationship (Boyle and Mitchell 1978; Sleeter et al. 1978). of reproductive temperature and However, epiphytic bacteria ingested with the geographical range of the wood may serve as nutritionally beneficial to marine woodborer Limnoria species with nitrogen-poor wood diets tripuncata. Biological Bulletin. (Zachary and Colwell 1979; Zachary et al. 118:9-16. 1983; Cragg et al. 1999). Digestion occurs 3. BEST, R. J., and J. J. rapidly (8 minutes total, Ricketts and Calvin STACHOWICZ. 2012. Trophic 1952). The wood-boring ability of limnoriids cascades in seagrass meadows has instigated research toward alternative depend on mesograzer variation in structures that deter boring (e.g. Borges et al. feeding rates, predation 2009; Cragg et al. 1999). Limnoria tripunctata is apparently undeterred by creosote susceptibility, and abundance. preserved wood (Menzies 1951) and Marine Ecology Progress Series. populations that ingest creosote wood appear 456:29-42. to possess a microbial gut flora unlike those 4. BORGES, L. M. S., S. M. CRAGG, that ingest untreated wood (Zachary and and S. BUSCH. 2009. A laboratory Colwell 1979; Zachary et al. 1983). Limnoria assay for measuring feeding and tripunctata may represent a resistant strain of mortality of the marine wood borer gribble, which developed in response to Limnoria under forced feeding creosote (Welton and Miller 1980). conditions: A basis for a standard Furthermore, L. tripunctata is also reported to test method. International tunnel into wood treated with copper chrome Biodeterioration & Biodegradation. arsenic (Pitman et al. 1997). Predators: Isopods play a significant role as 63:289-296. intermediate food web links, like amphipods, 5. BORGES, L. M. S., L. M. (e.g. see Americorophium salmonis, this MERCKELBACH, and S. M. guide) that are consumed by more than 20 CRAGG. 2014. Biogeography of species of marine fish (Welton and Miller wood-boring crustaceans (Isopoda: 1980; cabezon, Best and Stachowicz 2012), Limnoriidae) established in whales (Brusca et al. 2007) and other European coastal waters. Plos invertebrates (e.g. polychaete worms, Reish One. 9. 1954; Brusca 1980). 6. BOYKO, C. B., and C. WOLFF. Behavior: Dispersal between wood habitats 2014. Isopoda and Tanaidacea, p. occurs by swimming and crawling in young 210-215. In: Atlas of crustacean adults. In Teredo species, dispersal is by larvae only, adults burrow but do not swim or larvae. J. W. Margtin, J. Olesen, crawl. and J. T. Høeg (eds.). Johns Hopkins University Press, Bibliography Baltimore. 7. BOYLE, P. J., and R. MITCHELL. 1. ARAUJO, P. B., A. F. QUADROS, 1978. Absence of microorganisms M. M. AUGUSTO, and G. BOND- in crustacean digestive tracts. BUCKUP. 2004. Postmarsupial Science. 200:1157-1159. development of Atlantoscia 8. BRUSCA, R. C. 1980. Common floridana (van Name, 1940) intertidal invertebrates of the Gulf

of California. University of Arizona 16. KOZLOFF, E. N. 1993. Seashore Press, Tucson. life of the northern Pacific coast: an 9. BRUSCA, R. C., C. R. COELHO, illustrated guide to northern and S. TAITI. 2007. Isopoda, p. California, Oregon, Washington, 503-541. In: The Light and Smith and British Columbia. University of manual: intertidal invertebrates Washington Press, Seattle. from central California to Oregon. 17. KURIS, A. M., P. S. SADEGHIAN, J. T. Carlton (ed.). University of J. T. CARLTON, and E. CAMPOS. California Press, Berkeley, CA. 2007. Decapoda, p. 632-656. In: 10. BRUSCA, R. C., and E. W. The Light and Smith manual: IVERSON. 1985. A guide to the intertidal invertebrates from central marine isopod crustacea of Pacific California to Oregon. J. T. Carlton Costa Rica. Revista de Biologia (ed.). University of California Press, Tropical. 33:1-77. Berkeley, CA. 11. COOKSON, L. J. 1991. 18. MENZIES, R. J. 1951. Limnoria Australasian species of Limnoriidae and the premature failure of (Crustacea: Isopoda). Memoirs of creosoted marine structures in the Museum of Victoria. 52:137- North America, p. M1-17. In: 262. Report of the Marine Borer 12. CRAGG, S. M., A. J. PITMAN, and Conference. United States Naval S. M. HENDERSON. 1999. Civil Engineers Research and Developments in the understanding Evaluation Lab, Port Hueneme, of the biology of marine wood California. boring crustaceans and in methods 19. —. 1957. The marine borer family of controlling them. International Limnoriidae (Crustacea, Isopoda). Biodeterioration & Biodegradation. Part 1: Northern and Central 43:197-205. America: Systematics, distribution, 13. DELGERY, C. C., S. M. CRAGG, and ecology. Bulletin of the Marine S. BUSCH, and E. A. MORGAN. Science of the Gulf and Caribbean. 2006. Effects of the epibiotic 7:101-200. heterotrich ciliate Mirofolliculina 20. —. 1972. Experimental limnoriae and of moulting on faecal interbreeding between pellet production by the wood- geographically separated boring isopods, Limnoria populations of marine wood-boring tripunctata and Limnoria isopod Limnoria tripunctata with quadripunctata. Journal of preliminary indications of hybrid Experimental Marine Biology and vigor. Marine Biology. 17:149-&. Ecology. 334:165-173. 21. PITMAN, A. J., S. M. CRAGG, and 14. ELTRINGHAM, S. K. 1967. The G. DANIEL. 1997. The attack of effects of temperature on the greenheart (Ocotea rodiaei Mez) development of Limnoria eggs and creosote treated Douglas fir (Isopoda: Crustacea). Journal of (Pseudotsuga menziesii Mirb.) by Applied Ecology. 4:521-529. Limnoria tripunctata Menzies. 15. JOHNSON, M. W., and R. J. Material Und Organismen. 31:281- MENZIES. 1956. The migratory 291. habits of the marine gribble 22. REISH, D. J. 1954. Polychaetous Limnoria tripunctata Menzies in annelids as associates and San Diego Harbor, California. predators of the crustacean Biological Bulletin. 110:54-68. woodborer, Limnoria. Wasmann Journal of Biology. 12:223-226.

23. RICKETTS, E. F., and J. CALVIN. 1952. Between Pacific tides : an account of the habits and habitats of some five hundred of the common, conspicuous seashore invertebrates of the Pacific Coast between Sitka, Alaska, and Northern Mexico. Stanford : Stanford University Press, Stanford. 24. SADRO, S. 2001. Arthropoda: Decapoda, p. 176-178. In: Identification guide to larval marine invertebrates of the Pacific Northwest. A. Shanks (ed.). Oregon State University Press, Corvallis, OR. 25. SLEETER, T. D., P. J. BOYLE, A. M. CUNDELL, and R. MITCHELL. 1978. Relationships between marine microorganisms and wood- boring isopod Limnoria tripunctata. Marine Biology. 45:329-336. 26. SLEETER, T. D., and B. C. COULL. 1973. Invertebrates associated with marine wood boring isopod, Limnoria tripunctata. Oecologia. 13:97-102. 27. WELTON, L. L., and M. A. MILLER. 1980. Isopoda and Tanaidacea: the isopods and allies, p. 536-558. In: Intertidal invertebrates of California. R. H. Morris, D. P. Abbott, and E. C. Haderlie (eds.). Stanford University Press, California. 28. ZACHARY, A., and R. R. COLWELL. 1979. Gut-associated microflora of Limnoria tripunctata in marine creosote-treated wood pilings. Nature. 282:716-717. 29. ZACHARY, A., K. K. PARRISH, and J. D. BULTMAN. 1983. Possible role of marine bacteria in providing the creosote-resistance of Limnoria tripunctata. Marine Biology. 75:1-8.