Cumella Vulgaris Class: Multicrustacea, Malacostraca, Eumalacostraca

Home , Campylaspis, Cumacea, Diastylidae, Eudorella, Leuconidae, Multicrustacea

Phylum: Arthropoda, Crustacea

Cumella vulgaris Class: Multicrustacea, Malacostraca, Eumalacostraca

Order: Peracarida, Cumacea The ghost shrimp Family: Nannastacidae

Description chamber (Watling 2007). Size: In the original description by Hart Eyes: Conspicuous and circular in fe- (1930) from Vancouver Island, an ovigerous males (Gonor et al. 1979) (Fig. 1). In males a female was 2.5 mm in length and a male, single central sessile eye, with seven equal 3.0 mm long (Hart 1930). The illustrated lenses, is more prominent (Gonor et al. 1979) specimens (from Coos Bay) include a fe- (Fig. 4). male, 2.5 mm long, and a young male, 2.1 Antennae: Female antennule is rather mm in length. Cumella vulgaris is one of the stout, not easily visible, and with rudimentary smallest cumacean species (Sars 1900). inner flagellum (Nannastacidae, Fage 1951). Color: Males are dark brown except for ligh- The second antenna in females is with two ter distal segments and appendages. The large plumose setae (Hart 1930) (not figured). female carapace and sixth pleonite are dark Mouthparts: Mandibles are not unique brown and the rest of the body is light brown and the bases are not massive (Fage 1951) or white (Gonor et al. 1979). (not figured). General Morphology: Cumaceans are eas- Carapace: Female carapace is large ily recognizable by a large and inflated cara- and deep, with a smooth mid-dorsal carina pace and a (relatively) slender, flexible thor- (ridge) with a depression on each side (on ax and abdomen (Kozloff 1993; Gerken and posterior margin). A deep antennal notch is Martin 2014) (Fig. 1). Their bodies can be present, with an acute antero-lateral angle divided into these three major regions: the (Fig. 1). The male carapace is slender, the cephalon (head) that is covered by a cara- antennal notch is not as deep as in females, pace and includes the first five pairs of ap- and the dorsal carina is almost absent (Fig. pendages (antennae, mandibles, maxillae, 3). collectively the mouthparts). Posterior to Rostrum: Two pseudorostral lobes the cephalon is the pereon (thorax), usually (together called a pseudorostrum), or exten- consisting of five thoracic somites, followed sions of the carapace, extend anteriorly but by the pleon (abdomen) with consistently do not fuse in front of the head in cumaceans six pleonites. The fifth pleonite is usually the (Watling 2007). The pseudorostrum in female longest and the pleonites are lacking pleo- C. vulgaris is relatively short, minutely serrate pods in female individuals. The cumacean anteriorly and strongly pronounced (Fig. 1). family Nannastacidae are characterized by In males, the pseudorostral projection is shor- the lack of a free telsons and uropod endo- ter (Sars 1900) (Fig. 3). pods that are uniarticulate (Watling 2007). Pereon: Consists of five thoracic somites, (For general morphology of C. vulgaris, see each with paired appendages (pereopods) also Plate 229B, Watling 2007.) (Figs. 1–3). Cephalon: A carapace covers the cephalon Pereopods: The first pereopods in fe- and first three thoracic somites and is ex- males are with bases serrate on the outer dis- panded on either side to form a branchial tal margin. The dactyl and propodus are

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. and L. Rasmuson. 2015. Cumella vulgaris. 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 Biol- ogy, Charleston, OR.

equal to the carpus in length. The second uropods, a strong carapace carina, and exo- pereopods are stout, and the dactyl is twice podites that appear only on the first two pairs as long as the propodus. The exopodites of pereopods. Males have a compound eye, are present on the first two pairs of are slim, lack a strong carapace carina and pereopods only (Cumella, Lie 1969). The have a very long second antenna. Males also last three pereopods are stout (Fig. 1). The have four pereopod exopodites and some first four pereopod bases in males are more uropod distinctions. dilated than in females and exopodites are Possible Misidentifications present on the first four pereopods (absent Cumaceans are very small (range 1 on fifth) (Figs. 2, 3). mm–1 cm) shrimp-like crustaceans. Their Pleon: Long and narrow in males and stou- heads and thorax are fused to form a ter in females. Consists of six articles or carapace, the abdomen is tubular and the pleonites, and lacking pleopods (Figs. 1, 3). uropods are slender and biramous. There are Pleopods: All female cumaceans 1500 species worldwide, approximately 50 of lack pleopods (Fig.1) and males in the family which occur on the Pacific coast of the United Nannastacidae also lack pleopods (Watling States (Watling 2007; Gerken and Martin 2007) (Fig. 3). 2014). Cumaceans belong to the Telson: Telson short, not freely articulated Malacostraca, and are characterized by a and fused to sixth abdominal article carapace that covers the first three or four (Nannastacidae, Watling 2007) (Figs. 1, 3). thoracic somites. They also have an anterior Uropods: The uropod peduncles in females extension (pseudolobes), a telson that is have inner margin with only one spine on present or reduced and fused with the last the inner distal angle (Gonor et al. 1979) pleonite, eyes that are united dorsally, a (Fig. 6). The uropod endopod is uniarticu- second antenna that is without an exopod and late (compare to biarticulate endopod in Nip- pleopods that are absent in females and can poleucon hinumensis), larger than exopod, be absent or reduced in males (Watling denticulate on inner margin, with two stout 2007). spines, and one strong apical spine. The The superorder Peracarida includes exopod is with two articles (as in all cu- cumaceans, mysids, isopods, tanaids and maceans), is ½ the width of the endopod, amphipods. Cumaceans can be separated and with one slender apical spine (Fig. 6) from mysids by their single compound eye (Gonor et al. 1979). The uropods of males (particularly in the males), as mysids have are slim and the peduncle is denticulate, lon- large stalked eyes. Mysids have a carapace ger than rami (Fage 1951), and with three which covers the entire thorax, while distal spines. The endopod is with only a cumaceans have several posterior segments single article (Nannastacidae, Watling exposed (e.g. Figs. 1, 3). Euphausiids belong 1979), while the exopod is with two articles to the superorder Eucarida (along with (Fig. 5). decapods) and are pelagic and marine, but Sexual Dimorphism: Quite strong sexual might occasionally be found in estuaries. dimorphism is observed in C. vulgaris. Fe- They have biramous thoracic appendages males are generally shorter and stouter than (cumacean pereopods are uniramous, with males and mature individuals have a brood some thoracic exopodites). Additionally, pouch. The female eye lacks the obvious euphausiids have strong pleopods for large lenses found in males (Fig. 4). Female swimming and cumacean pleopods, when specimens have a broader carapace and

present, are small. Campylaspis hartae has a carapace with The four local cumacean families can large ridges, but no bumps, and C. rubroma- be divided into those with a freely articulated culata has a carapace with a series of bumps telson and those without, the former com- or tubercles and shallow ridges (Watling prise the Lampropidae and Diastylidae, 2007). while the latter comprise the Leuconidae The Leuconidae (like the Nannastaci- and Nannastacidae (Watling 2007). Cuma- dae) lack an independent telson. However, cean families that lack an articulated telson they always have a biarticulate uropod endo- are consistently monophyletic on molecular pod, not a uniramous one as in Nannastaci- phylogenies and are likely derived within the dae. Members of the Leuconidae often have Cumacea (Haye et al. 2004). However, up to two pairs of male pleopods (there are morphological characters used to differenti- none in Nannastacidae) and leuconid males ate cumacean families (e.g. number of pleo- have exopodites on all five pairs of pereopods pods in males) may be homoplasious (see (rarely on three). Leuconid females have ex- Haye et al. 2004). opodites on four (rarely on three) pairs of The family Nannastacidae, in which pereopods (Watling 1979). Thus, numbers of Cumella occurs, lack an independent telson, pereopodal exopodites in both sexes are too the males have no pleopods and the alike in the families Leuconidae and Nan- endopod of the uropod is uniarticulate. nastacidae to serve as dependable determin- Pereopodal exopodites in the ing characters. Of the Leuconidae, the gene- Nannastacidae are as follows: males have ra Eudorella, and Nippoleucon (see N. five (rarely four or three) pairs and females hinumensis, this guide) occur on the Pacific have three (rarely four or zero) pairs Coast (each with one local species). (Watling 1979). Cumella vulgaris is the only The Lampropidae and Diastylidae have species in this genus locally. However, C. a freely articulated telsons and the former pygmaea, the European species is very like family has three or more terminal setae on the C. vulgaris in color and size. The female of telson while the latter has 0–2. The Lam- C. pygmaea is stouter than C. vulgaris, with propidae includes six local species in the gen- a less inflated carapace and with a dentate era Hemilamprops and Mesolamprops (each crest on the carina. The male of C. pyg- with one local species) and the Lamprops maea is similar to that of C. vulgaris, except (four local species, see L. quadriplicata, this that its pedigerous segments are more une- guide). In the Diastylidae there are five local ven (Hart 1930). The only other genus of species in three genera including Anchico- Nannastacidae from our area is Campylas- lurus and Diastylopsis (one local species pis. In this genus, both males and females each) and Diastylis (three local species) have exopodites only on the first pair of (Watling 2007). pereopods (Lie 1969). The females have a Ecological Information bulbous carapace with rounded anterolateral Range: Type locality is Puget Sound (Hart angles, unlike Cumella, which has an un- 1930), known range from central California to inflated carapace and an acute anterolateral Oregon (Watling 2007). angles. Campylaspis species have a cara- Local Distribution: Known in occur in Coos pace that extends posteriorly and overhangs and Yaquina bays. the first few pereonites. Campylaspis canali- Habitat: Cumacean species choose sub- culata has a smooth carapace and females strates mostly based on food availability. Cu- with a marginal anterior-posterior groove.

mella vulgaris prefers fine sand (grains with Lamprops and Pseudocuma, Gerken and diameter less than 160 µm) and dislikes dry Martin 2014). Little is known about the devel- sand (in lab experiments). Males can be opment of C. vulgaris, specifically. In found in sand with grains of 200 µm diame- Manocuma stellifera, an Atlantic intertidal ter and smaller (Wieser 1956) and are also cumacean, mating occurs at night in plankton found on the water surface, near shore (Hart (Gnewuch and Croker 1973; Watling 1979), 1930). Cumella vulgaris actively avoids during the short swarming period. Females habitats with fast currents (McCauley et al. molt 12–96 hours before oviposition (in the 1977). Applications of the insecticide Sevin lab). Eggs are probably fertilized as they are caused significant decreases (90%) in C. released into the marsupium, where they are vulgaris abundance (Simenstad and Cordell carried to a manca stage. Some other 1989). intertidal species have two breeding Salinity: Collected at salinities of 30 (in Co- generations per year, one in summer and in os Bay). fall (see Corey 1969, 1976 in Watling 1979). Temperature: Larva: Cumacean development proceeds Tidal Level: Intertidal and usually found be- from an egg to two manca stages, a subadult low +1.5 meters MLLW down to -0.6 meters and, finally, an adult. The manca stage re- Wieser 1956). Individuals found on water sembles the adult, but is defined by a lack of surface and in standing water at low tide the fifth pair of pleopods (see Fig. 41.1F, (Hart 1930). Subtidal populations are Gerken and Martin 2014). The mancae of M. reported as deep as 10 m (Jones 1961). stellifera molt three times and the young leave Associates: the marsupium, molt several more times into Abundance: The most common cumacean subadult morphology, with mature gonads in Puget Sound and San Juan Islands, and secondary sexual characteristics present Washington (Wieser 1956) and a common (see Corey 1969, 1976 in Watling 1979). intertidal species in central California. In Juvenile: Coos Bay, it was the second most abundant Longevity: In Atlantic intertidal cumaceans, crustacean (by numbers) found in a North longevity varies with reproductive time of Bend study site (Gonor et al. 1979). With year: an early summer generation may live Nippoleucon hinumensis, it was found at up five months, while late summer and fall to 5,600 individuals per square meter in broods will overwinter and live 12 and nine South Slough of Coos Bay (personal months, respectively (see Corey in Watling communication, M. Posey, OIMB). In 1979). Willapa Bay, C. vulgaris was the third most Growth Rate: Cumacean growth occurs in abundant organism in Neotrypaea beds, conjunction with molting where the exoskele- ninth most abundant in Zostera beds, ton is shed and replaced. Post-molt individu- seventh most abundant in oyster beds and als will have soft shells as the cuticle gradual- twelfth most abundant in bare mud and sand ly hardens. During a molt, arthropods have (Ferraro and Cole 2007). the ability to regenerate limbs that were previously autotomized (Kuris et al. 2007). Life-History Information Food: A deposit feeder in fine sand and mud. Reproduction: Development in cumaceans In coarse sand (>150 µm), it is an epistrate is direct, where eggs hatch within a marsupi- feeder that scrapes food off individual grains um, and development is thought to be simi- (Watling 1979; Kozloff 1993). Cumaceans lar among cumacean genera (e.g. Leucon, feed while buried and swim to new site when

one site has been exploited. Cumella vulgar- recently introduced green crab, Carcinus is aggregates to feed (Watling 1979). maenas, in central California. Marine Biol- Predators: Cumella vulgaris have been ob- ogy. 122:239-247. served in gut contents of the three-spined 7. HART, J. F. L. 1930. Some Cumacea of sticklebacks (Gasterosteus aculeatus) and the Vancouver Island region. Contributions Northern anchovies (Engraulis mordax) to Canadian Biology. 6:23-40. (Rasmuson and Morgan 2013). The 8. HAYE, P. A., I. KORNFIELD, and L. WAT- introduced European green crab (Carcinus LING. 2004. Molecular insights into Cuma- maenas) significantly reduced C. vulgaris cean family relationships (Crustacea, Cu- populations (Grosholz and Ruiz 1995). macea). Molecular Phylogenetics and Shorebirds feed on C. vulgaris, but have no Evolution. 30:798-809. notable impact on abundance (Wilson 9. JONES, M. L. 1961. A quantitative evalua- 1991). tion of the benthic fauna off Point Rich- Behavior: Females and juveniles are capa- mond, California. University of California ble of swimming at speeds of 0.25 to 1.5 cm Publications in Zoology. 67:219-320. per second, while males are capable of 10. KING, A. R. 1977. Acute effects of sedi- higher swimming speeds (~5 cm s-1, King mentation on Cumella vulgaris hart 1930 1977). (Cumacea). Thesis (M.S.)--Oregon State University,1977. Bibliography 11. KOZLOFF, E. N. 1993. Seashore life of 1. FAGE, L. 1951. Cumacés. Faune de the northern Pacific coast: an illustrated France:1-136. guide to northern California, Oregon, 2. FERRARO, S. P., and F. A. COLE. 2007. Washington, and British Columbia. Univer- Benthic macrofauna-habitat associations sity of Washington Press, Seattle. in Willapa Bay, Washington, USA. Estua- 12. KURIS, A. M., P. S. SADEGHIAN, J. T. rine Coastal and Shelf Science. 71:491- CARLTON, and E. CAMPOS. 2007. De- 507. capoda, p. 632-656. In: The Light and 3. GERKEN, S., and J. W. MARTIN. 2014. Smith manual: intertidal invertebrates from Cumacea, p. 216-218. In: Atlas of crusta- central California to Oregon. J. T. Carlton cean larvae. J. W. Martin, J. Olesen, and (ed.). University of California Press, Berke- J. T. Høeg (eds.). Johns Hopkins Univer- ley, CA. sity Press, Baltimore, MD. 13. LIE, U. 1969. Cumacea from Puget Sound 4. GNEWUCH, W. T., and R. A. CROKER. and off the northwestern coast of Wash- 1973. Macrofauna of northern New Eng- ington with descriptions of two new spe- land marine sand. 1. Biology of Manco- cies. Crustaceana. 17:19-30. cuma stellifera (Zimmer, 1943) 14. MCCAULEY, J. E., R. A. PARR, and D. R. (Crustacea, Cumacea). Canadian Jour- HANCOCK. 1977. Benthic infauna and nal of Zoology. 51:1011-1020. maintenance dredging: case study. Water 5. GONOR, J. J., D. R. STREHLOW, and Research. 11:233-242. G. E. JOHNSON. 1979. Ecological as- 15. RASMUSON, L. K., and S. G. MORGAN. sessments at the North Bend airport ex- 2013. Fish predation after weakly synchro- tension site. School of Oceanography, nized larval release in a coastal upwelling Oregon State University, Salem, OR. system. Marine Ecology Progress Series. 6. GROSHOLZ, E. D., and G. M. RUIZ. 490:185-198. 1995. Spread and potential impact of the 16. SARS, G. O. 1900. An account of the

crustacea of Norway. III: Cumacea. Ber- gen Museum. 17. SIMENSTAD, C. A., and J. R. COR- DELL. 1989. Effects of Sevin application on littoral flat meiofauna: preliminary sampling in Willapa Bay, June-July 1988. Seattle, Wash. : Fisheries Research In- stitute, University of Washington School of Fisheries, Seattle, Wash. 18. WATLING, L. 1979. Marine flora and fauna of the northeastern United States: Crustacea, Cumacea. In: NOAA Tech- nical Report NMFS Circular. Dept. of Commerce, National Oceanic and At- mospheric Administration, National Ma- rine Fisheries Service ;, Washington. Updated 2015 T.C. Hiebert and L. Rasmuson