Neotrypaea Californiensis Class: Multicrustacea, Malacostraca, Eumalacostraca

Phylum: Arthropoda, Crustacea

Neotrypaea californiensis Class: Multicrustacea, Malacostraca, Eumalacostraca

Order: Eucarida, Decapoda, Pleocyemata, Axiidea The ghost shrimp Family: Callianassidae, Callianassinae

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

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. 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, Charles- ton, OR.

and unequal (Fig. 1). The large cheliped is 1949). They burrow using their first to third broad, serrate and with an obvious gap in legs, aided by mouthparts (MacGinitie 1934; dactyls. The merus has a conspicuous ven- Kozloff 1993) and begin digging backward tral lobe, the carpus is almost square and before turning and removing excess loose longer than the palm, and with laterally in- sediment from the burrow to the surface (see curved dorsal margin (Campos et al. 2009). MacGinitie 1934 for figure). The dactyl has a recurved hook distally Possible Misidentifications (Wicksten 2011) (Fig. 1). Propodi are of Thalassinidea is a former infraorder nearly equal length (McGinitie 1934). containing Callianassidae and Upogebiidae Second chelipeds are both chelate with and, although shown to be non-monophyletic propodi and dactyls near equal in width (Sakai 2004), most mud and ghost shrimps (Figs. 1, 3). Female and immature are often referred to collectively as thalassi- individuals have hand longer than carpus nids. (Wicksten 2011). Upogebiidae is described in Williams Abdomen (Pleon): Abdomen elongate (1986) and Campos et al. 2009 and, locally, (longer than cephalothorax), not reflexed but consists of a single species, Upogebia extended, symmetrical and externally seg- pugettensis, the blue mud shrimp, often co- mented. It bears three pairs of fan-like pleo- occurs with N. californiensis. Upogebia pods (Fig. 1). First and second pleopods pugettensis is easy to recognize because it is are vestigal and absent in males. Third to larger and its color (bluish and never red or fifth are leaf-like (Fig. 1). In females, the first pink) is strikingly different. Its burrows are pleopods are uniramous and the second are also more firm and substantial. The most biramous (Wicksten 2011). noticeable morphological difference between Telson & Uropods: Telson nearly rectangu- the species is the first pair of legs: both of lar, forming a well-developed fan-shape with which are small, sub-chelate and equal in U. uropods, which are equal in length to the tel- pugettensis. Furthermore, its rostrum is hairy son (Fig. 1). Telson composed of two dorsal and has a laterally compressed and slender ribs and posterior marginal tooth. Exopod tip of the short fixed finger of the chela (outer ramus) also with dorsal ribs. (Wicksten 2011). Sexual Dimorphism: Pleopod (see Abdo- Characteristics defining the Callianas- men (Pleon)) and cheliped (see Chelipeds) sidae are described in Sakai 1999 and and morphology differs between males and fe- Campos et al. 2009. There are three species males. Females are also commonly seen locally, Neotrypaea californiensis, N. gigas with conspicuous bright orange egg masses and N. buffari (Kuris et al. 2007). Neotrypaea attached to their pleopods. californiensis can be distinguished from the Callianassidae-specific character other two species by the lack of a prominent Burrow: Neotrypaea californiensis build and rostrum (present in N. gigas) and eyestalks inhabit large, sloppy and permanent burrows that are acute and diverging tips of the with branching side tunnels (Y-shape, Jen- eyestalks (rather than short, blunt and not sen 1995; Puls 2001). Individuals dig tire- diverging in N. biffari) (see Campos et al. lessly, turning over acres of northwest oyster 2009). Neotrypaea gigas is larger (to 125– beds (Ricketts and Calvin 1971, see Behav- 150 mm) than the other two, and relatively ior). Burrows can be to 0.76–1.00 m deep rare in sandy sublittoral habitats. Its rostrum (MacGinitie 1934; MacGinitie and MacGinitie is sharp, with prominent medial tooth (whichN.

californiensis does not possess), and its first osmotic conformer, lower lethal limit 8.75– chela closes without a gap. It is more 10.5 (Thompson and Pritchard 1969b) and common in its southern distribution, south of the upper limit is 43.75 (Barnard et al. 1980). Point Conception (Barnard et al. 1980; Kuris Temperature: et al. 2007; Wicksten 2011). Neotrypaea Tidal Level: High intertidal. Collected at me- gigas and N. californiensis also differ in the dium high and upper to mid-intertidal zones morphology of the second pereopod: In N. (0.0–1.2 m, Ricketts and Calvin 1971). californiensis the propodus and dactyl are of Associates: The blue mud shrimp, Upogebia equal length and in N. gigas, the propodus is pugettensis, is found overlapping the range of curved and wider than the dactyl (Kuris et al. N. californiensis, though it is generally lower 2007). Recent examination of these two intertidally and in muddier sediments. species using morphological and molecular Common commensals in ghost shrimp data suggests that the key characters for burrows include a polynoid worm differentiating species is the length of Hesperonoe, pinnotherid crabs (Scleroplax eyestalks and shape of the distal outer glanulata), copepods (Hemicyclops, edges (Pernet et al. 2010). Clausidium), the shrimp Betaeus harrimani, the bopyrid isopod lone cornuta, the goby Ecological Information Clevelandia, the echiuroid worm Urechis Range: Type region is California, with pro- caupo, and the clam Cryptomya californica posed locality San Francisco or Monterey (MacGinitie 1934; Kuris et al. 2007; Campos Bay (Wicksten 2011), but type material has 2009; Wicksten 2011). been lost. Known range includes Alaska to Abundance: Common in Oregon's estuarine Tiajuana River, California and Point Abreo- mudflats. In Wallapa Bay, Washington, the jos, Baja California, Mexico (Campos et al. density of N. californiensis (up to 450 shrimp 2009). per m2) was always higher than that of the Local Distribution: Distribution in many Or- other locally occurring ghost shrimp, U. pu- egon estuaries including Coos Bay, Alsea gettensis (up to 100 shrimp per m2) Dumbauld River (Gaumer et al. 1973b), Nestucca es- et al. 1996). tuary (Gaumer et al. 1973a), Netarts Bay (Gaumer et al. 1974), Umpqua estuary Life-History Information (Umpqua Estuary 1978), Tillamook Bay Reproduction: Continuously reproductive in (Gaumer 1973b) and Yaquina Bay (Gaumer central California, especially June and July et al. 1974). (MacGinitie 1934; Ricketts and Calvin 1971). Habitat: Mud or sand. Individuals can sur- A breeding season in late spring and summer vive anoxia for nearly six days (Garth and is known to occur in Yaquina Bay, Oregon Abbott 1980). In adaptation to living in an (Puls 2001). Neotrypaea californiensis environment that is relatively low in oxygen, reach sexual maturity at 2 years and produce N. californiensis and U. pugettensis exhibit 3,900 eggs while U. pugettensis produces low metabolic rates and can both survive pe- 7,100 (Dumbauld et al. 1996). Individuals riods of anoxia. Upogebia pugettensis has a ovigerous from April through August (Willapa higher metabolic rate and N. californiensis is Bay, Washington, Dumbauld et al. 1996). able to survive longer during periods of an- Larva: Larval development in N. californiensis oxia (Thompson and Pritchard 1969a; Zebe proceeds via several zoea (five total) and, a 1982). final, megalopa stage, each marked by a molt Salinity: Collected at salinities of 30. An (Puls 2001). Neotrypaea californiensis zoea

have rostrum longer than antennules Adults are also eaten by bottom feeding fish. (compare to Upogebia pugettensis), ab- Green and white sturgeon collected from dominal segments with dorsal and/or lateral Willapa Bay, Washington and the Columbia spines and telson that is broad and flat with River estuary had N. californiensis within their medial tooth at posterior (see paguroid zoe- guts (Dumbauld et al. 2008). Foraging gray ae Fig. 53.2, Harvey et al. 2014: Fig. 11, whales in British Columbia (Clayoquot Sound) McCrow 1972; Puls 2001). Larval size also ingest adult benthic N. californiensis (measured from tip of rostrum to tip of tel- (Dunham and Duffus 2001). son) proceeds from 2.8–3.6 mm (Zoea I) to Behavior: Ghost shrimp species (e.g. Ne- 6.8–7.5 mm (Zoea V) (Puls 2001). Megalo- otrypaea, Upogebia) are known to be ecosys- pae are shrimp-like in morphology with long tem engineers with the ability to regulate and pereopods, resembling the adult. First and change community (macro and microbial second pereopods chelate or subchelate communities) structure by burrowing and de- (Puls 2001). Larvae are flushed into open posit feeding (Dumbauld and Wyllie- ocean by tides, where they spend most of Echeverria 2003; Bertics and Ziebis 2009). larval period in the plankton and exchange Their presence and behavior effects biogeo- between neighboring bays is common chemical composition including sediment (McCrow 1972; Johnson and Gonor 1982). grain size, nutrient exchange and organic Larvae recruit back to the estuary to settle composition. Bioturbation (Kristensen et al. from August to October (Willapa Bay, Wash- 2012) turns over and re-suspends sediment, ington, Dumbalud et al. 1996) and preferen- which can increase erosion and sediment in- tially settle on mud substrate (rather than stability, having a negative effect on algae shell, Feldman et al. 1997). and seagrasses that require light for photo- Juvenile: Sexual dimorphism and matura- synthesis (e.g. Zostera, Dumbauld and Wyllie- tion in claw size occurs at 2 years of age, Echeverria 2003) and suspension feeders when individuals are 9–10 mm carapace (e.g. oysters, Dumbauld et al. 1996; Feldman length (Dumbauld et al. 1996). et al. 2000). All sediment to 76 cm deep is Longevity: turned over completely in 240 days Growth Rate: Growth occurs in conjunction (MacGinitie 1934). In turn, seagrasses tend with molting where the exoskeleton is shed to solidify sediment and are not suitable habi- and replaced. Post-molt individuals will tats for ghost shrimp species (Berkenbusch et have soft shells as the cuticle gradually al. 2007). Outside of their burrows, N. cali- hardens (Ruppert et al. 2004). The growth forniensis specimens are fragile and lay rather rate for N. californiensis is approximately 2– helpless on the sediment surface (Kozloff 3 mm (carapace length) per year (Dumbauld 1993). They can swim for short distances and et al. 1996). move quickly backwards by flapping the fan- Food: Detritivore, obtains food by ingesting like posterior (MacGinitie 1934). mud as it burrows the top (richest) layer Bibliography (MacGinitie 1934; MacGinitie and MacGinitie 1949) and also filter feeds by pumping water 1. BARNARD, L. J., D. E. BOWERS, AND E. through burrow (Powell 1974). C. HADERLIE. 1980. Macrura and Predators: Adults are used by humans for Anomura, p. 577-593. In: Intertidal inverte- fish bait and individuals avoid predation by brates of California. R. H. Morris, D. P. Ab- retreating to burrow. Juveniles and larvae bott, and E. C. Haderlie (eds.). Stanford are eaten in the plankton (e.g. by fish). University Press, Stanford, CA.

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