Neotrypaea Californiensis Class: Malacostraca Order: Decapoda the Ghost Shrimp Section: Anomura, Paguroidea Family: Callinassidae

Home , Betaeus, Callianassidae, Neotrypaea, Scleroplax, Upogebia pugettensis

Phylum: Arthropoda, Crustacea Neotrypaea californiensis Class: Malacostraca Order: Decapoda The ghost shrimp Section: Anomura, Paguroidea Family: Callinassidae

Taxonomy: Neotrypaea californiensis was Antennae: Antennal angles rounded described as a member of the genus and naked and antennal peduncle shorter Callianassa 1854 by Dana and remained than antennular peduncle (Campos et al. there until an analysis by Manning and Felder 2009). (1991) resulted in the three NE pacific Mouthparts: The mouth of decapod Callianassa species moving to the new genus crustaceans comprises six pairs of Neotrypaea. Tudge et al. (2000) later appendages including one pair of mandibles analyzed 93 characters of adult morphology (on either side of the mouth), two pairs of and found the genus Neotrypaea to be non- maxillae and three pairs of maxillipeds. The monophyletic and Sakai (1999) synonymized maxillae and maxillipeds attach posterior to Neotrypaea and Callianassa. The monophyly the mouth and extend to cover the mandibles of Neotrypaea is still supported by some (Ruppert et al. 2004). The third maxilliped in authors (e.g. Campos et al. 2009) and we N. californiensis is operculiform, with widened follow the most current local intertidal guides, merus extending beyond articulation with which use N. californiensis (Kuris et al. 2007). carpus and ischium (Manning and Felder For complete list of synonymies see Sakai 1991; Campos et al. 2009; Wicksten 2011). (2005). Carapace: Smooth and with lateral grooves (Wicksten 2011). Description Rostrum: Not prominent, rounded Size: Males up to 115 mm and females to and with small blunt tooth (Wicksten 2011) 120 mm in length (Barnard et al. 1980; Puls (Fig. 2). 2001; Wicksten 2011). Teeth: Color: Can be white to cream with patches of Pereopods: Second pereopod pinkish red or orange on the abdomen and flattened, chelate (Fig. 3) and with row of appendages (see Plate 19, Kozloff 1993; setae along posterior margin (Campos et al. Wicksten 2011). The illustrated specimen 2009). Third to fifth pereopods are (from Coos Bay) is pale pink with light orange predominantly used in walking (MacGinitie abdomen. 1934). Third pereopod with triangular carpus General Morphology: The body of decapod and round, small dactyl. Fourth and fifth crustaceans can be divided into the pereopods are slender (Wicksten 2011). cephalothorax (fused head and thorax) and Chelipeds: First chelipeds are abdomen. They have a large plate-like chelate and unequal (Fig. 1). The large carapace dorsally, beneath which are five cheliped is broad, serrate and with an obvious pairs of thoracic appendages (see chelipeds gap in dactyls. The merus has a conspicuous and pereopods) and three pairs of ventral lobe, the carpus is almost square and maxillipeds (see mouthparts). The abdomen longer than the palm, and with laterally and associated appendages are outstretched incurved dorsal margin (Campos et al. 2009). and shrimp-like in Callianassidae (Stevens The dactyl has a recurved hook distally 1928; Kuris et al. 2007). (Wicksten 2011) (Fig. 1). Propodi are of Cephalothorax: nearly equal length (McGinitie 1934). Second Eyes: Eyestalks flattened and with chelipeds are both chelate with propodi and acute tips. Pigmented corneas are mid-dorsal dactyls near equal in width (Figs. 1, 3). within eyestalk (Fig. 2) (Wicksten 2011). Female and immature individuals have hand Eyes triangular and with diverging tip longer than carpus (Wicksten 2011). (Campos et al. 2009).

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12725 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected] Abdomen (Pleon): Abdomen elongate pink) is strikingly different. Its burrows are (longer than cephalothorax), not reflexed but also more firm and substantial. The most extended, symmetrical and externally noticeable morphological difference between segmented. It bears three pairs of fan-like the species is the first pair of legs: both of pleopods (Fig. 1). First and second pleopods which are small, sub-chelate and equal in U. are vestigal and absent in males. Third to pugettensis. Furthermore, its rostrum is hairy fifth are leaf-like (Fig. 1). In females, the first and has a laterally compressed and slender pleopods are uniramous and the second are tip of the short fixed finger of the chela biramous (Wicksten 2011). (Wicksten 2011). Telson & Uropods: Telson nearly Characteristics defining the rectangular, forming a well-developed fan- Callianassidae are described in Sakai 1999 shape with uropods, which are equal in length and and Campos et al. 2009. There are three to the telson (Fig. 1). Telson composed of species locally, Neotrypaea californiensis, N. two dorsal ribs and posterior marginal tooth. gigas and N. buffari (Kuris et al. 2007). Exopod (outer ramus) also with dorsal ribs. Neotrypaea californiensis can be Sexual Dimorphism: Pleopod (see distinguished from the other two species by Abdomen (Pleon)) and cheliped (see the lack of a prominent rostrum (present in N. Chelipeds) morphology differs between gigas) and eyestalks that are acute and males and females. Females are also diverging tips of the eyestalks (rather than commonly seen with conspicuous bright short, blunt and not diverging in N. biffari) orange egg masses attached to their (see Campos et al. 2009). Neotrypaea gigas pleopods. is larger (to 125–150 mm) than the other two, and relatively rare in sandy sublittoral Callianassidae-specific character habitats. Its rostrum is sharp, with prominent Burrow: Neotrypaea californiensis build and medial tooth (whichN. californiensis does not inhabit large, sloppy and permanent burrows possess), and its first chela closes without a with branching side tunnels (Y-shape, Jensen gap. It is more common in its southern 1995; Puls 2001). Individuals dig tirelessly, distribution, south of Point Conception turning over acres of northwest oyster beds (Barnard et al. 1980; Kuris et al. 2007; (Ricketts and Calvin 1971, see Behavior). Wicksten 2011). Neotrypaea gigas and N. Burrows can be to 0.76–1.00 m deep californiensis also differ in the morphology of (MacGinitie 1934; MacGinitie and MacGinitie the second pereopod: In N. californiensis the 1949). They burrow using their first to third propodus and dactyl are of equal length and legs, aided by mouthparts (MacGinitie 1934; in N. gigas, the propodus is curved and wider Kozloff 1993) and begin digging backward than the dactyl (Kuris et al. 2007). Recent before turning and removing excess loose examination of these two species using sediment from the burrow to the surface (see morphological and molecular data suggests MacGinitie 1934 for figure). that the key characters for differentiating species is the length of eyestalks and shape Possible Misidentifications of the distal outer edges (Pernet et al. 2010). Thalassinidea is a former infraorder containing Callianassidae and Upogebiidae Ecological Information and, although shown to be non-monophyletic Range: Type region is California, with (Sakai 2004), most mud and ghost shrimps proposed locality San Francisco or Monterey are often referred to collectively as Bay (Wicksten 2011), but type material has thalassinids. been lost. Known range includes Alaska to Upogebiidae is described in Williams Tiajuana River, California and Point Abreojos, (1986) and Campos et al. 2009 and, locally, Baja California, Mexico (Campos et al. 2009) consists of a single species, Upogebia Local Distribution: Distribution in many pugettensis, the blue mud shrimp, often co- Oregon estuaries including Coos Bay, Alsea occurs with N. californiensis. Upogebia River (Gaumer et al. 1973b), Nestucca pugettensis is easy to recognize because it is estuary (Gaumer et al. 1973a), Netarts Bay larger and its color (bluish and never red or (Gaumer et al. 1974), Umpqua estuary

Hiebert, T.C. 2015. Neotrypaea californiensis. 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. (Umpqua Estuary 1978), Tillamook Bay 7,100 (Dumbauld et al. 1996). Individuals (Gaumer 1973b) and Yaquina Bay (Gaumer ovigerous from April through August (Willapa et al. 1974). Bay, Washington, Dumbauld et al. 1996). Habitat: Mud or sand. Individuals can Larva: Larval development in N. survive anoxia for nearly six days (Garth and californiensis proceeds via several zoea (five Abbott 1980). In adaptation to living in an total) and, a final, megalopa stage, each environment that is relatively low in oxygen, marked by a molt (Puls 2001). Neotrypaea N. californiensis and U. pugettensis exhibit californiensis zoea have rostrum longer than low metabolic rates and can both survive antennules (compare to Upogebia periods of anoxia. Upogebia pugettensis has pugettensis), abdominal segments with dorsal a higher metabolic rate and N. californiensis and/or lateral spines and telson that is broad is able to survive longer during periods of and flat with medial tooth at posterior (see anoxia (Thompson and Pritchard 1969a; Zebe paguroid zoeae Fig. 53.2, Harvey et al. 2014: 1982). Fig. 11, McCrow 1972; Puls 2001). Larval Salinity: Collected at salinities of 30. An size (measured from tip of rostrum to tip of osmotic conformer, lower lethal limit 8.75– telson) proceeds from 2.8–3.6 mm (Zoea I) to 10.5 (Thompson and Pritchard 1969b) and 6.8–7.5 mm (Zoea V) (Puls 2001). the upper limit is 43.75 (Barnard et al. 1980). Megalopae are shrimp-like in morphology with Temperature: long pereopods, resembling the adult. First Tidal Level: High intertidal. Collected at and second pereopods chelate or subchelate medium high and upper to mid-intertidal (Puls 2001). Larvae are flushed into open zones (0.0–1.2 m, Ricketts and Calvin 1971). ocean by tides, where they spend most of Associates: The blue mud shrimp, Upogebia larval period in the plankton and exchange pugettensis, is found overlapping the range of between neighboring bays is common N. californiensis, though it is generally lower (McCrow 1972; Johnson and Gonor 1982). intertidally and in muddier sediments. Larvae recruit back to the estuary to settle Common commensals in ghost shrimp from August to October (Willapa Bay, burrows include a polynoid worm Washington, Dumbalud et al. 1996) and Hesperonoe, pinnotherid crabs (Scleroplax preferentially settle on mud substrate (rather glanulata), copepods (Hemicyclops, than shell, Feldman et al. 1997). Clausidium), the shrimp Betaeus harrimani, Juvenile: Sexual dimorphism and the bopyrid isopod lone cornuta, the goby maturation in claw size occurs at 2 years of Clevelandia, the echiuroid worm Urechis age, when individuals are 9–10 mm carapace caupo, and the clam Cryptomya californica length (Dumbauld et al. 1996). (MacGinitie 1934; Kuris et al. 2007; Campos Longevity: 2009; Wicksten 2011). Growth Rate: Growth occurs in conjunction Abundance: Common in Oregon's estuarine with molting where the exoskeleton is shed mudflats. In Wallapa Bay, Washington, the and replaced. Post-molt individuals will have density of N. californiensis (up to 450 shrimp soft shells as the cuticle gradually hardens per m2) was always higher than that of the (Ruppert et al. 2004). The growth rate for N. other locally occurring ghost shrimp, U. californiensis is approximately 2–3 mm pugettensis (up to 100 shrimp per m2) (carapace length) per year (Dumbauld et al. Dumbauld et al. 1996). 1996). Food: Detritivore, obtains food by ingesting Life-History Information mud as it burrows the top (richest) layer Reproduction: Continuously reproductive in (MacGinitie 1934; MacGinitie and MacGinitie central California, especially June and July 1949) and also filter feeds by pumping water (MacGinitie 1934; Ricketts and Calvin 1971). through burrow (Powell 1974). A breeding season in late spring and summer Predators: Adults are used by humans for is known to occur in Yaquina Bay, Oregon fish bait and individuals avoid predation by (Puls 2001). Neotrypaea californiensis retreating to burrow. Juveniles and larvae are reach sexual maturity at 2 years and produce eaten in the plankton (e.g. by fish). Adults are 3,900 eggs while U. pugettensis produces also eaten by bottom feeding fish. Green and

A publication of the University of Oregon Libraries and the Oregon Institute of Marine Biology Individual species: http://hdl.handle.net/1794/12725 and full 3rd edition: http://hdl.handle.net/1794/18839 Email corrections to: [email protected] white sturgeon collected from Willapa Bay, Experimental Marine Biology and Washington and the Columbia River estuary Ecology. 341:70-84. had N. californiensis within their guts 3. BERTICS, V. J., AND W. ZIEBIS. (Dumbauld et al. 2008). Foraging gray 2009. Biodiversity of benthic microbial whales in British Columbia (Clayoquot Sound) communities in bioturbated coastal also ingest adult benthic N. californiensis sediments is controlled by (Dunham and Duffus 2001). geochemical microniches. Isme Behavior: Ghost shrimp species (e.g. Journal. 3:1269-1285. Neotrypaea, Upogebia) are known to be 4. CAMPOS, E., A. R. DE CAMPOS, ecosystem engineers with the ability to AND I. MANRIQUEZ. 2009. Intertidal regulate and change community (macro and thalassinidean shrimps microbial communities) structure by burrowing (Thalassinidea, Callianassidae and and deposit feeding (Dumbauld and Wyllie- Upogebiidae) of the west coast of Echeverria 2003; Bertics and Ziebis 2009). Baja, California, Mexico: annotated Their presence and behavior effects checklist, key for identification, and biogeochemical composition including symbionts. Crustaceana. 82:1249- sediment grain size, nutrient exchange and 1263. organic composition. Bioturbation (Kristensen 5. DUMBAULD, B. R., D. A. et al. 2012) turns over and re-suspends ARMSTRONG, AND K. L. FELDMAN. sediment, which can increase erosion and 1996. Life-history characteristics of sediment instability, having a negative effect two sympatric thalassinidean shrimps, on algae and seagrasses that require light for Neotrypaea californiensis and photosynthesis (e.g. Zostera, Dumbauld and Upogebia pugettensis, with Wyllie-Echeverria 2003) and suspension implications for oyster culture. Journal feeders (e.g. oysters, Dumbauld et al. 1996; of Crustacean Biology. 16:689-708. Feldman et al. 2000). All sediment to 76 cm 6. DUMBAULD, B. R., D. L. HOLDEN, deep is turned over completely in 240 days AND O. P. LANGNESS. 2008. Do (MacGinitie 1934). In turn, seagrasses tend sturgeon limit burrowing shrimp to solidify sediment and are not suitable populations in Pacific Northwest habitats for ghost shrimp species estuaries? Environmental Biology of (Berkenbusch et al. 2007). Outside of their Fishes. 83:283-296. burrows, N. californiensis specimens are 7. DUMBAULD, B. R., AND S. WYLLIE- fragile and lay rather helpless on the ECHEVERRIA. 2003. The influence of sediment surface (Kozloff 1993). They can burrowing thalassinid shrimps on the swim for short distances and move quickly distribution of intertidal seagrasses in backwards by flapping the fan-like posterior Willapa Bay, Washington, USA. (MacGinitie 1934). Aquatic Botany. 77:27-42. 8. DUNHAM, J. S., AND D. A. DUFFUS. Bibliography 2001. Foraging patterns of gray whales in Central Clayoquot Sound, 1. BARNARD, L. J., D. E. BOWERS, British Columbia, Canada. Marine AND E. C. HADERLIE. 1980. Macrura Ecology Progress Series. 223:299- and Anomura, p. 577-593. In: Intertidal 310. invertebrates of California. R. H. 9. FELDMAN, K. L., D. A. Morris, D. P. Abbott, and E. C. ARMSTRONG, B. R. DUMBAULD, T. Haderlie (eds.). Stanford University H. DEWITT, AND D. C. DOTY. 2000. Press, Stanford, CA. Oysters, crabs, and burrowing shrimp: 2. BERKENBUSCH, K., A. A. ROWDEN, Review of an environmental conflict AND T. E. MYERS. 2007. Interactions over aquatic resources and pesticide between seagrasses and burrowing use in Washington State's (USA) ghost shrimps and their influence on coastal estuaries. Estuaries. 23:141- infaunal assemblages. Journal of 176.

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