Archaeomysis Grebnitzkii Class: Multicrustacea, Malacostraca, Eumalacostraca

Phylum: Arthropoda, Crustacea

Archaeomysis grebnitzkii Class: Multicrustacea, Malacostraca, Eumalacostraca

Order: Peracarida, Mysida A mysid or opossum shrimp Family: Mysidae, Gastrosaccinae, Archaeomysini

Taxonomy: Archaeomysis grebnitzkii was covered by a carapace), and a pleon described from a specimen collected from (abdomen). At the posterior end, they have a cod gut contents by Czerniavksy in 1882. telson and uropods. Among the Mysidacea Later, Holmes described the same species specifically, the carapace is attached to the under a different name, Callomysis macula- thorax by anterior segments only and the pos- ta, which was collected from a sandy beach terior dorsal edge is free (Banner 1948) (Fig. (Holmquist 1975). In 1932, Tattersall trans- 1). Mysid eyes are stalked, antennules are ferred C. maculata to A. maculata and biramous, antennae have a long scale (or Holmquist (1975) synonymized Archaeomy- squama), pleopods are often reduced, thorac- sis maculata and Callomysis maculata as A. ic legs bear swimming exopodites and uro- grebnitzkii, a species which exhibited a wide pods are lamellar and form tail fan. Mysids North Pacific range (Hanamura 1997; Moldin are easily distinguished from other Peracardia 2007). These species were previously dif- by the presence of a statocyst on the uropod ferentiated by subtle variation in morphologi- endopods (see Plate 220, Moldin 2007; cal characters that were deemed to be intra- Vicente et al. 2014; Fig. 1, Meland et al. specific (e.g. rostrum shape, third pleopod 2015). exopod segments, telson length, Hamanura Cephalon: (see also Figs. 3–4, Hanamura 1997). 1997). Eyes: Large, movable, stalked, with Description black corneas and somewhat pear shaped. Size: Male body length ranges from 9–15 Eye and eyestalk less than twice as long as mm, and females 13–22 mm (Holmquist broad (Fig. 2). Visual pigment absorbances 1975; Hanamura 1997). The illustrated were measured for A. grebnitzkii at 496 nm, specimens (from Columbia River) are 4.5 which phylogenetically placed this species mm (male, Fig. 1a) and 17 mm (female, Fig. within a shorter-wavelength spectral sensitivi- 1). ties species clade (Porter et al. 2007). Color: Transparent, with stellar melano- Antennae: Almost as long as body and phores (“maculate”) (see Fig. 265, Kozloff with slender multi-articulate flagellum (Fig. 1). 1993). Some mysids can change color to The peduncle has three joints and is longer blend with their background (Tattersall and than its antennal scale, but shorter than the Tattersall 1951). antennular peduncle (Fig. 2). Antennule General Morphology: Mysids are shrimp- peduncle with first joint about equal in length like crustaceans that are occasionally called to remaining joints combined. The second opossum shrimp due to the female marsupi- joint is with two spines on the outer margin um or brood pouch, which is composed of (Tattersall 1951) (Fig. 2). oostegites. Like other Peracarida (e.g. cu- Antennae Scale: (= squama) Extends maceans, isopods, amphipods, tanaids), to distal end of second penducular joint and is their bodies are elongated and composed of 3½ times as long as broad. Sclae with an anterior cephalon and pereon (thorax,

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. Archaeomysis grebnitzkii. 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.

straight outer margin, without setae (Mysida, Mauchline 1980) and all are (Archaeomysis, Banner 1948), with strong biramous. The first is with a uniarticulate terminal spine, and weak or absent distal endopod and multi-articulate exopod (Banner suture. On anterior and inner margins, scale 1948) of 7–9 articles (Holmquist 1975) and is setose only (no spines) (Kozloff 1974) the third is with elongate exopod (Fig. 2). (Archaeomysis, Banner 1948) (Fig. 1a, 4). Mouthparts: For general mouthpart The second pleopod with endopod of 4–7 ar- morphology for the Mysida see Fig. 3, ticles, exopod of 8–9 articles, the third is with Meland et al. 2015. Labrum longer than elongate copulatory exopod of 8–10 articles, broad (Tattersall and Tattersall 1951) and endopod of 5 articles (Holmquist 1975) (Fig. with strong frontal spiniform process 4), and the fourth is with exopod of 5–9 ar- (Archaeomysis, Banner 1948). ticles (Holmquist 1975). The illustrated fourth Carapace: Attached to first two or male pleopod has 7 articles (Fig. 5), endopod three thoracic segments and free dorsally at a simple plate, the fifth pleopod is like the posterior edge (Banner 1948) (Figs. 1, 1a). fourth, but shorter, and with 4–8 exopod ar- Posterior margin with rounded lateral lobes ticles (Holmquist 1975) (see also Fig. 4, Ha- (Archaeomysis, Banner 1948) and no fringe namura 1997). or ornamentation. Carapace pronounced Female pleopods are ''usually degener- anteriorly into a short rostrum (Fig. 2). ate"' (i.e. reduced) (Mysida, Mauchline 1980) Rostrum: Rostrum length shorter and all are biramous (Archaeomysis, than eyestalk (Fig. 2) and can be grooved, Mauchline 1980) with each ramus of one bent down slightly, or rounded (Holmquist small article (see also Fig. 3, Hanamura 1975). ''Shortly produced”, according to ori- 1997). The first pleopod is with an elongate ginal genus description (Archaeomysis, protopodite, with basal and distal tufts of long Holmquist 1975). setae, the endopod is longer than the exopod, Pereon: and more than ½ as long as protopodite (Fig. Pereopods: Pereopods without well- 6). The second pleopods are with short developed gills (Mysida, Banner 1948). First protopodite and exopod, and a longer leg with exopodite, second leg without a endopod (Fig. 7). The third, fourth and fifth lobe-like process on merus. Pereopod 3–8 female pleopods are like the second, but with with carpopropodus (carpus and propodus shorter endopods. fused) of endopod that is divided into many Telson: Telson with distinct apical cleft subjoints: 9–11 in females (Fig. 3), 7–9 in (Mauchline 1980) and margins of cleft are males (not figured). Exopods in both male denticulate (Banner 1948) (Fig. 8). Total and female legs 3–8 has a basal joint with length is 2½ times as long as broad (at base). an acute outer distal corner (Fig. 3). No Eight to nine spines are present on each branchiae are present on thoracic legs margin, the last two spines are long, strong, (Mauchline 1980). and close together (Fig. 8) (See Fig. 7, Pleon: Fifth pleonite with a small medial Meland et al. 2015; see Fig. 5, Hanamura projection and sixth with corresponding ridge 1997). (Fig. 5). In females, the lateral pleura on the Uropods: Uropods biramous, with neither first somites help form the brood pouch (Fig. branch articulate (Banner 1948). Both rami 1). Inconspicuous pleura are present on so- with setae on the distal margin mites 3–5, none on six (Banner 1948). (Archaeomysis, Banner 1948). The endopod Pleopods: Male pleopods variable is longer than the exopod, with statocyst near

base, and two basal spines (Fig. 8). Six of up to ½ inch long, with an inflated, spines are present on the inner edge in shrimp-like carapace (see Cumella vulgaris, males, seven in females (Banner 1948). this guide), a single compound recessed The exopod is truncate, without suture eye (except for some eyeless females of (Mauchline 1980), with 14 (male) to 17 some species), and a flexible, tubular abdo- (female) lateral spines on outer margin men. Mysids characteristically have large, (more than 10, Archaeomysis). No setae on stalked, movable eyes, and well developed the outer exopod margin (Banner 1948). exopodites on their thoracic legs. Mature Statocyst: Light and balance organ on en- females have oostegites forming a dopod of uropod (Fig. 8). It is found in all marsupium. Additionally, northeast Pacific neritic and in common oceanic mysids mysids lack thoracic gills, have reduced (Banner 1948) and distinguishes mysids pleopods in females (and sometimes in from larval decapods (Green 1968). males). They also have a statocyst on the Sexual Dimorphism: Pleopod morphology inner ramus of the uropod. varies between males and females. Mature Mysicadea is divided into two subor- females are also recognizable by the pres- ders, the Mysida and Lophogastrida. The ence of oostegites that form a thoracic brood former suborder comprises coastal and in- pouch with two pairs of lamellae (Mauchline tertidal species while the latter includes 1980). These oostegites arise from the mostly large, pelagic and deep sea mysids. seventh and eighth pereopods (Mauchline These suborders are easily differentiable by 1980) to form marsupium (Fig. 1). the presence of branchial gills, biramous pleopods and the lack of statocysts in Possible Misidentifications Lophogastrida (branchia are absent, pleo- Mysidacea and Euphausiacea, be- pods are reduced and statocysts are con- ing superficially similar in appearance, are spicuous in the Mysida) (Moldin 2007). Ar- often treated together (e.g. Banner 1948; chaeomysis grebnitzkii belongs to the My- Mauchline 1980). (They were formerly sida, lacking gills or branchiae on the tho- combined as the Schizopoda.) Both are racic legs (Fig. 3) and having rather reduced orders of the class Malacostraca, but female pleopods. euphausiids are in the group (i.e. superor- Within the Mysicadea locally, there der) Eucarida with the Decapoda (Martin are 15 species comprising the following 2007). Like the mysids, euphausiids differ genera: Acanthomysis and Neomysis from decapods in having biramous thoracic (comprising three species and four species, legs (Kasaoka 1974). Unlike the mysids, respectively), and Hyperacanthomysis, Al- euphausiids have a carapace that is fused ienacanthomysis, Columbiaemysis, Delta- dorsally with all the thoracic segments. mysis, Exacanthomysis, Hippacanthomysis, The mysid carapace is attached only to the Holmesimysis, and Archaeomysis (each first two or three thoracic segments. Fur- with a single species represented locally) thermore, mysid females have oostegites, (Moldin 2007). but euphausiids do not. Alienacanthomysis macropsis is rec- Other orders of Peracarida include ognizable by elongated eyestalks and Delta- lsopoda, Tanaidacea, and Amphipoda, mysis holmquistae has spines on the lateral which are all fairly easily distinguished telson margins, but only distally, where oth- from Mysidacea. One order that might be er species have spines all over the telson confused is Cumacea, small crustaceans margins or in proximal groups. Aliena-

canthomysis macropsis, has a broadly tri- female oostegites, statocyst on the uropod angular rostrum with long acute lateral car- endopod, and male fourth pleopods bira- apace spines and its telson has a fringe of mous). For differentiating Neomysis spe- small spines. It is abundant in San Fran- cies, see N. mercedis (this guide). Colum- cisco Bay and becomes rarer farther north biaemysis ignota has been described from (Holmquist 1979). It has been reported female specimens only, and only from from Yaquina Bay to lower Columbia River British Columbia. Its antennal scale is long, and in Puget Sound in bays amongst Ulva setose all around, and has a suture. Its and in plankton (Kozloff 1974a). telson is tongue shaped, with spines Archaeomysis japonica was de- becoming dense at the tip, and two long scribed in 1996 by Hanamura et al. and spines at the rounded apex. There are four was previously considered A. grebnitzkii. spines on the lower edge of the statocyst. However, the two species differ in the en- Ecological Information dopod of the third pleopod, which is seg- Range: Type locality is Bering and Comman- mented in A. japonica (males, Hanamura dor Islands in the Bering Sea (Hanamura et al. 1996). 1997). Known range in northern Pacific Archaeomysis grebnitzkii has (Holmquist 1975) from Urup Island and north- spines along the lateral margins of uropod ern Sakhalin to southwest Alaska and the exopods, which is not seen in other spe- Bering Sea, to Washington, Oregon, and cies. Of the species without lateral exopod northern California (see Fig. 25, Hanamura spines, Hippacanthomysis platypoda has a 1997). flattened exopod of fourth pleopod (males). Local Distribution: Locally present in estuar- Holmesimysis costata and E. davisi have ies of Coos Bay, Yaquina Bay, lower- distinctly segmented pleonites, the former Columbia River and also along the open has a broadly triangular telson while the coast. latter has a telson that is sharply triangular. Habitat: Primarily littoral, buried in sand with Holmesimysis was extracted from a mixture of pebbles and boulders on both Acanthomysis (Holmquist 1979). Its open ocean coast and in inland waters. In members have fourth male pleopods with mud and with the alga Zostera as well as with only two segments and on the tip are two Phyllospadix and kelp intermixed (Holmquist spiny peg-like structures (Mauchline 1980). 1975). Burrows in bottom substratum, and Holmesimysis costata, the type species for rises to surface of water at night, especially the genus, has a short, bluntly rounded during the breeding season (Mauchline 1980; antennal scale. Tattersall and Tattersall 1951). Individuals Columbiaemysis, Acanthomysis, very sensitive to oxygen reduction (Green Neomysis and Hyperacanthomysis species 1968; Jawed 1973). have pleonites that are smooth and without Salinity: Quite variable, from fresh water to distinct folds or segments. Neomysis spe- salinities of 34 (Holmquist 1975). However, cies have a pointed distal tip of the anten- salinity figures could refer to surface, and the nal scale (see Fig. 3, Neomysis mercedis, species is an inhabitant of the saltier, bottom this guide) and members of the remaining water (Holmquist 1975). genera have a distal antennal scale tip that Temperature: A wide temperature tolerance, is rounded. There are several Pacific spe- ranging from 8.5 °C or lower to 24 °C. cies of the genus Neomysis (all with point- (Holmquist 1975). ed apex on the antennal scale, two pairs of

Tidal Level: Predominantly intertidal and high latitudes on season, but not on also found in shallow waters close to shore temperature (Mauchline 1980). Brood size in (Holmquist 1975). Moves up and down with the closely related genus Gastrosaccus, can the rising and falling tide. Occurs at extreme vary seasonally, with largest broods produced low tide level (Puget Sound, Washington, in early summer and a female G. vulgaris, of Banner 1948) and subtidal (Coos Bay, Ore- similar in size to A. grebnitzkii (17 mm long), gon) as well as intertidally and at low water had 100 embryos per brood (Mauchline on ocean beaches. Juveniles may be more 1980). Numbers of broods/year is not common higher in the intertidal, while larger definitely known for A. grebnitzkii, but the individuals (i.e. adults) can be found in most shallow-living neritic and littoral mysid deeper habitats, offshore (Llewellyn et al. species usually have three broods year, 1981; Takahashi and Kawaguchi 1995). including the closely related Gastrosaccus at Associates: a comparable latitude (Japan, Mauchline Abundance: The most common mysid of 1980). Chromosome counts for A. grebnitzkii the northeastern Pacific, followed by were 2n=10, plus an extra small chromosome Neomysis mercedis (see this guide) (Holmquist 1975). Sex ratios vary within popu- (Holmquist 1975). The most abundant and lations, but females frequently outnumber common species collected in hyperbenthic males (Mauchline 1980). sledge samples at Bastendorff Beach, Larva: The first nauplioid stage has append- Charlsteson, Oregon with approximately ages that resemble the typical nauplius larva 3200 individuals per square meter (Marin (see Balanus glandula, this guide), but lacks Jarrin and Shanks 2011). an eye or swimming setae. The next post- nauplioid stage has all adult appendages, car- Life-History Information apace and eyes. Both stages are non-motile Reproduction: All development takes place and lecithotrophic. Ultimately, the post- within the female marsupium and is nauplioid molts into a free-living juvenile lecithotrophic and proceeds through three (Vicente et al. 2014). Larval development distinct stages: 1) the embryonic stage; 2) time depends on temperature. In G. vulgaris, the first nauplioid stage (eyeless larva); and developmental duration is 10.9–25 days 3) the post-naplioid (eyed larva) stage to a (Matsudaira et al. in Mauchline 1980). juvenile (see Figs. 38.1–38.2, Vicente et al. Juvenile: Juveniles are simply miniature 2014). Copulation in A. grebnitzkii occurs at adults at post-emergence molt, when they are night and lasts only a few seconds usually 1.5–3.0 mm long (Mauchline 1980). (Mauchline 1980). Sperm is shed into fe- Longevity: Mysids will probably live 12 –18 male brood pouch and the female then lays months in temperate water and over two eggs, which are immediately fertilized years in the Arctic (Tattersall and Tattersall (Mauchline 1980) and 510 x 480 µm (a 13 1951). No longevity rates known for A. mm long female, Hanamura 1997). Early grebnitzkii. In overwintering generations, embryos are spherical or sub-spherical. most individuals are born in autumn, a few in Young develop to a subadult stage in the summer, and fewer in winter. Spring breeding brood pouch, and emerge from external is intensive and females may produce two genital openings of oviducts near the bases broods. Spring generations reproduce in the of the sixth thoracic legs (Mauchline 1980). summer (possibly twice), and usually die by The number of eggs depends on size of autumn (Mauchline 1980). female and embryos and in temperate and Growth Rate: Growth among isopods occurs

in conjunction with molting where the exo- Behavior: Locomotion is by exopods of tho- skeleton is shed and replaced. Post-molt racic legs as pleopods are often reduced, and individuals will have soft shells as the cuticle not locomotory. Motion is rotary and unceas- gradually hardens. During a molt, ing. An "escape mechanism” is also ob- arthropods have the ability to regenerate served in A. grebnitzkii, where a sudden limbs that were previously autonomized downward flex of abdomen and tail fan caus- (Kuris et al. 2007). Mysids generally take es the individual to spring backward and even about one year to attain full growth and are leap out of water (Tattersall and Tattersall sexually mature in considerably less time 1951) or through mud (Mauchline 1980). (Tattersall and Tattersall 1951). Females While this has not been shown in A. usually grow larger than males (Mauchline grebnitzkii, females of some mysid species 1980) and the number of instars is fewer will recapture escaped larvae and return them than for most crustaceans. The first and to their marsupium (Mauchline 1980). Larvae second occur in the marsupium, and 10 or can belong to other individuals or to other more occur after the release of young species. Males will eat escaped larvae (Mauchline 1980). (Mauchline 1980). Mysids avoid bright light Food: Feeds either on living or dead materi- (Tattersall and Tattersall 1951), but are at- al that is picked up by thoracic endopods, or tracted to weak light sources and fishing lures (more commonly) on fine suspended matter (Mauchline 1980). Archaeomysis grebnitzkii filtered by thoracic exopods. Danish did not respond to atmospheric pressure Gastrosaccus species, also bottom dwellers, changes of 0.1 atm (Mauchline 1980). Other eat detritus, algae, copepods and burrowers (e.g. Gastrosaccus) have amphipods (Tattersall and Tattersall 1951). pronounced diel vertical migration where To stir up food for filtering, mysids will bal- individuals remain within the substrate during ance, head down, on antennal scales and the day and are pelagic at night. Burrowing inner flagella of antennulae, and create cur- mysids are reotaxtic and face into water rents with thoracic exopods. They can also current. They may move offshore to avoid "plow" the bottom with their scales and breaking waves and waves can also wash flagellae (Cannon and Manton 1927 in them out of their burrows (Mauchline 1980). Mauchline 1980). South African Bibliography Gastrosaccus species feeds most often at night (Mauchline 1980). 1. BANNER, A. H. 1948. A taxonomic study Predators: Fish are the most common and of the Mysidacea and Euphausiacea most important predators (Tattersall and Tat- (Crustacea) of the northeastern Pacific. tersall 1951; Haertel and Osterberg 1967) as Part I. Mysidacea, from family Lophogas- well birds (e.g. eider duck in Aleutians), tridae through tribe Erythropini. Transac- shrimp, ctenophores, squid, and possibly tions of the Royal Canadian Institute. 26. cetaceans and seals. This species is also 2. GREEN, J. 1968. The biology of estuarine eaten by humans or used as fish bait in animals. University of Washington Press, some parts of the world (e.g. India, Tattersall Seattle. and Tattersall 1951). Archaeomysis 3. HAERTEL, L., and C. OSTERBERG. grebnitzkii was the dominant prey of the bay 1967. Ecology of zooplankton, benthos shrimp, Lissocrangon stylirostris (see and fishes in the Columbia River estuary. description in this guide) at night (Marin Ecology. 48:459-472. Jarrin and Shanks 2008, 2011). 4. HANAMURA, Y. 1997. Review of the tax-

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