Upogebia Pugettensis Class: Multicrustacea, Malacostraca, Eumalacostraca

Home , Clausidium, Orthione griffenis, Upogebia, Upogebia pugettensis, Upogebiidae

Phylum: Arthropoda, Crustacea

Upogebia pugettensis Class: Multicrustacea, Malacostraca, Eumalacostraca

Order: Eucarida, Decapoda, Pleocyemata, Gebiidea The blue mud shrimp Family: Upogebiidae

Taxonomy: Dana described Gebia pu- Antenna: First segment of antennular gettensis in 1852 and this species was later peduncle has a sharp tooth at ventral border redescribed as Upogebia pugettensis and second segment of flagellum has small, (Stevens 1928; Williams 1986). round disto-ventral spine (Wicksten 2011). Mouthparts: The mouth of decapod Description crustaceans comprises six pairs of appendag- Size: The type specimen was 50.8 mm in es including one pair of mandibles (on either length and the illustrated specimen side of the mouth), two pairs of maxillae and (ovigerous female from Coos Bay, Fig. 1) three pairs of maxillipeds. The maxillae and was 90 mm in length. Individuals are often maxillipeds attach posterior to the mouth and larger and reach sizes to 100 mm (range 75 extend to cover the mandibles (Ruppert et al. –112 mm) and northern specimens are larg- 2004). er than those in southern California Carapace: Bears two rows of 11–12 (MacGinitie and MacGinitie 1949; Wicksten teeth laterally (Fig. 1) in addition to a small 2011). distal spines (13 distal spines, 20 lateral teeth Color: Light blue green to deep olive brown on carapace shoulder, see Wicksten 2011). with brown fringes on pleopods and pleon. Carapace with thalassinidean line extending Individual color variable and may depend on from anterior to posterior margin (Wicksten feeding habits (see Fig. 321, Kozloff 1993; 2011). Wicksten 2011). Rostrum: Large, tridentate, obtuse, General Morphology: The body of decapod rough and hairy (Schmitt 1921), the sides crustaceans can be divided into the cepha- bear 3–5 short conical teeth (Wicksten 2011). lothorax (fused head and thorax) and abdo- Rostral tip shorter than antennular peduncle. men. They have a large plate-like carapace Two short processes extending on either side dorsally, beneath which are five pairs of tho- each with 0–2 dorsal teeth (Wicksten 2011). racic appendages (see chelipeds and pere- Teeth: opods) and three pairs of maxillipeds (see Pereopods: Two to five simple walking mouthparts). The abdomen and associat- legs. Second pereopod is not chelate and ed appendages are outstretched and shrimp bears setae on lower segment margins. Car- -like in Upogebiidae (Kuris et al. 2007). For pus has small spines and merus has a single morphology of Upogebia (see Fig. 2, Wil- small spine. Pereopods 3–5 setose, de- liams 1986) and U. pugettensis see Williams crease progressively in size and have dactyls (see Fig. 13, 1986). with spinules (Wicksten 2011). Cephalothorax: Chelipeds: First chelipeds approxi- Eyes: Peduncle cylindrical (Schmitt mately equal and subchelate (Fig. 1). Dactyls 1921), eyestalks short but exceeding lateral curved, with ridged upper surface and lateral rostral process. Corneas terminal and direc- surface with many tubercles (Wicksten 2011). ted antero-laterally (Williams 1986). Fixed finger slender (Williams 1986), with one

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: https://oimb.uoregon.edu/oregon-estuarine-invertebrates and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected]

Hiebert, T.C. 2015. Upogebia pugettensis. 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.

conical tooth. Palm with setose lines, bear- are also more firm and substantial. The most ing setae and small teeth, as well as a sharp noticeable morphological difference between spine at dactyl base. Carpus with lateral these species is the first pair of legs: both of and longitudinal furrow and spine with small which are small, sub-chelate and equal in U. teeth (4–10), two distal spines and, a larger pugettensis. Furthermore, its rostrum is hairy marginal spine. The upper margin of the and has a laterally compressed and slender merus is curved and bears spines and small tip of the short fixed finger of the chela teeth (5–6). Ischium bears a single small (Wicksten 2011). spine (Wicksten 2011). Characteristics defining the Callianas- Abdomen (Pleon): Abdomen elongate and sidae are described by Sakai (1999) and broad, not reflexed, extended, symmetrical Campos et al. (2009). There are three spe- and externally segmented. Bears four pairs cies locally, Neotrypaea californiensis, N. of fan-like pleopods (Fig. 1). gigas and N. biffari (Kuris et al. 2007). Telson & Uropods: Telson wide anteriorly Neotrypaea californiensis can be distinguis- and uropod length exceeds that of the tel- hed from the other two species by the lack of son. Uropods bear dorsal ribs and marginal a prominent rostrum (present in N. gigas) and spines (Wicksten 2011). Fan-like tail formed eyestalks that are acute and diverging tips of by telson, uropods adapted for swimming. the eyestalks (rather than short, blunt and not Sexual Dimorphism: The first pleopod is diverging in N. biffari) (see Campos et al. absent in males and is slender, bi- 2009). Neotrypaea gigas is larger (to 125– articulating and simple in female U. pu- 150 mm) than the other two, and relatively gettensis (Williams 1986). rare in sandy sublittoral habitats. Its rostrum is sharp, with prominent medial tooth (not Upogebiidae-specific Character found in N. californiensis), and its first chela Burrow: Upogebia pugettensis builds U or closes without a gap. It is more common in its Y-shaped burrows that are firm, permanent southern distribution, south of Point and simple with little branching (unlike Ne- Conception (Barnard et al. 1980; Kuris et al. otrypaea californiensis). Burrows extend 2007; Wicksten 2011). Neotrypaea gigas and vertically about 46 cm, then horizontally 0.6– N. californiensis also differ in the morphology 1.2 m and up to the surface (MacGinitie of the second pereopod: In N. californiensis 1930; Ricketts and Calvin 1971). Often, the the propodus and dactyl are of equal length entrance will have a gravel plug if the tide is and in N. gigas, the propodus is curved and out (Stevens 1928). The walls are smooth wider than the dactyl (Kuris et al. 2007). and mucus lined (MacGinitie and MacGinitie Recent examination of these two species 1949). For figure see MacGinitie (1930). using morphological and molecular data Possible Misidentifications suggests that the key characters for Upogebiidae is described by differentiating species is the length of Williams (1986) and Campos et al. (2009) eyestalks and shape of the distal outer edges and the single local species, Upogebia (Pernet et al. 2010). pugettensis (the blue mud shrimp) often co- Ecological Information occurs with N. californiensis. Upogebia Range: Type locality is Puget Sound, Wash- pugettensis is easy to recognize because it ington. Known range includes Alaska to Mor- is larger and its color (bluish and never red row Bay, California (see Fig. 1, Williams 1986; or pink) is strikingly different. Its burrows Wicksten 2011). Southern populations of U.

pugettensis become replaced by the conge- the Pacific coast from Asia in the 1980s and is ner, U. macginitieorum (Kuris et al. 2007). thought to have caused the 2002 collapse of Local Distribution: Oregon estuaries and U. pugettensis on the Pacific coast (e.g. sloughs including Alsea, Nestucca, Netarts, Willapa Bay, Washington, see Fig. 1, Yaquina, Coos Bay. Dumbauld et al. 2011; Williams and Boyko Habitat: Estuarine mudflats (in areas 2012). This parasite was discovered in without Zostera, Stevens 1928), in mud or Yaquina Bay, Oregon in 1999 and was the sandy mud, often with some gravel. In first parasite known from the gills of Upogebia adaptation to living in an environment that is in western North America (Markham 2004; relatively low in oxygen, N. californiensis Chapman et al. 2012). Larvae of the and U. pugettensis exhibit low metabolic commensal phoronid, Phoronis pallida, exhibit rates and can both survive periods of increased swimming speed and settlement anoxia. Upogebia pugettensis has a higher behavior in the presence of Upogebia- metabolic rate and cannot survive periods of conditioned seawater (Santagata 2004). anoxia as well as N. californiensis Abundance: Can be locally common (Kuris et (Thompson and Pritchard 1969a; Barnard et al. 2007). In Willapa Bay, Washington, the al. 1980; Zebe 1982). density of N. californiensis (up to 450 shrimp Salinity: Collected at salinities of 30. A per m2) was always higher than that of the strong hyperosmotic regulator, their lower other locally occurring ghost shrimp, U. pu- lethal limit is 3.5 (Thompson and Pritchard gettensis (up to 100 shrimp per m2) Dumbauld 1969b; Barnard et al. 1980). et al. 1996). Temperature: Life-History Information Tidal Level: Intertidal to shallow subtidal, Reproduction: Each burrow inhabited by one near shore (Wicksten 2011). Mid to lower pair (Barnard et al. 1980). Ovigerous females intertidal of bays (Stevens 1928; Kuris et al. found December and February (Elkhorn 2007) and usually lower than N. cali- Slough California, MacGinitie and MacGinitie forniensis. Occasionally small individuals 1949), October through May (Willapa Bay, occur quite high in the intertidal (Ricketts Washington, Dumbauld et al. 1996) and early and Calvin 1971). April (South Slough, Coos Bay, Oregon). Associates: The blue mud shrimp, Eggs carried under abdomen on pleopods Upogebia pugettensis, is found overlapping (Fig. 2). Neotrypaea californiensis reach the range of N. californiensis, though it is sexual maturity at 2 years and produce 3,900 generally found in lower intertidal burrows eggs while U. pugettensis produces 7,100 and in muddier sediments. Common (Dumbauld et al. 1996). commensals in ghost shrimp burrows Larva: The larvae of U. pugettensis are found include a polynoid worm Hesperonoe, in plankton samples from February to June pinnotherid crabs (Scleroplax glanulata), and their morphology was described by Hart copepods (Hemicyclops, Clausidium), the (1937). Larval development in U. pugettensis shrimp Betaeus harrimani, the bopyrid proceeds via three zoea stages and, a final isopod lone cornuta, the goby Clevelandia, megalopa stage, each marked by a molt (Hart the echiuroid worm Urechis caupo, and the 1937; Puls 2001). Upogebia pugettensis zo- clam Cryptomya californica (MacGinitie ea have rostrum shorter than antennules (1/3 1934; Kuris et al. 2007; Campos et al. 2009; antennule length, compare to Callianassidae), Wicksten 2011). The parasitic bopyrid five abdominal segments, swimming setae on isopod, Orthione griffenis, was introduced to

exopods and maxillipeds, and triangular tel- for U. pugettensis and its commensal bivalve son with indentation at posterior margin with Cryptomya californica at three phytoplankton five setae, unlike N. californiensis, which densities (low, medium and high).. They have a medial tooth at telson posterior (see found that U. pugettensis removed 57, 53 and paguroid zoeae Fig. 53.2, Harvey et al. 40% of phytoplankton drawn into the burrow, 2014: Hart 1937; Puls 2001). Larval size while C. californica removed 12, 19 and 39%, (measured from tip of rostrum to tip of tel- respectively. son) proceeds from 3.7 mm (Zoea I, Fig. 3), Predators: Adults are used by humans for to 4.4 mm (Zoea II), to 5.4 mm (Zoea III) fish bait, individuals avoid predation by re- (Puls 2001). Megalopae are shrimp-like in treating to burrow. Juveniles and larvae are morphology with blunt rostrum, pereopods eaten in the plankton (e.g. fish). and maxillipeds resembling adults. The tel- Behavior: Ghost shrimp species son becomes rectangular and has rounded (Neotrypaea, Upogebia) are known to be eco- uropods laterally. Pleopods on abdominal system engineers with the ability to regulate segments 2–5 have setae (Puls 2001). and change community (macro and microbial Upogebia pugettensis larvae recruit in spring communities) structure by burrowing and de- (April-June) (Willapa Bay, Washington, posit feeding (Dumbauld and Wyllie- Dumbauld et al. 1996). Echeverria 2003; Bertics and Ziebis 2009). Juvenile: Sexual dimorphism occurs before Their presence and behavior effects biogeo- maturation in U. pugettensis (unlike N. cali- chemical composition including sediment forniensis), where claw size occurs almost grain size, nutrient exchange (D’Andrea and immediately (<1 year old), when individuals DeWitt 2009) and organic composition. Biotur- are 4–5 mm in (carapace) length (Dumbauld bation (Kristensen et al. 2012) turns over and et al. 1996). re-suspends sediment, which can increase Longevity: Moderately long lived (Ricketts erosion and sediment instability, having a and Calvin 1971). negative effect on algae and seagrasses that Growth Rate: Growth occurs in conjunction require light for photosynthesis (e.g. Zostera, with molting. In pre-molting periods the epi- Dumbauld and Wyllie-Echeverria 2003) and dermis separates from the old cuticle and a suspension feeders (e.g. oysters, Dumbauld dramatic increase in epidermal cell growth et al. 1996). In turn, seagrasses tend to solid- occurs. Post-molt individuals will have soft ify sediment and are not suitable habitats for shells until a thin membranous layer is de- ghost shrimp species (Berkenbusch et al. posited and the cuticle gradually hard- 2007). In controlled experiments, the pres- ens. During a molt decapods have the abil- ence of U. pugettensis lowers abundances of ity to regenerate limbs that were previously other estuarine crustaceans and polychaetes autotomized (Kuris et al. 2007). The growth (Posey et al. 1991). rate for U. pugettensis is approximately 4–5 Bibliography mm (carapace length) per year (Dumbauld et al. 1996). 1. BARNARD, L. J., D. E. BOWERS, AND E. Food: Detritivore, obtaining food by filtering C. HADERLIE. 1980. Macrura and water through the burrow as it sits near the Anomura, p. 577-593. In: Intertidal inverte- entrance. Individuals make a brates of California. R. H. Morris, D. P. Ab- “basket" (MacGinitie 1930) with its first and bott, and E. C. Haderlie (eds.). Stanford second pereopods, which have long setae. University Press, Stanford, CA. Griffen et al. (2004) estimated filtration rates 2. BERKENBUSCH, K., A. A. ROWDEN,

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iforniensis (Crustacea, Thalassinidea). Comparative Biochemistry and Physiolo- gy B-Biochemistry & Molecular Biology. 72:613-617. Updated 2015 T.C. Hiebert