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Crustacean Biology Rediscovery of the horseshoe shrimp Lightiella serendipita Jones, 1961 (Cephalocarida: Hutchinsoniellidae) in San Francisco Bay, California, USA, with a key to the worldwide species of Cephalocarida Type Article Author Garcia, Crystal; Woo, Isa; Rogers, D. Christopher; Flanagan, Alison M.; De La Cruz, Susan E. W. Title Rediscovery of the horseshoe shrimp Lightiella serendipita Jones, 1961 (Cephalocarida: Hutchinsoniellidae) in San Francisco Bay, California, USA, with a key to the worldwide species of Cephalocarida Journal title Journal of Crustacean Biology URI http://hdl.handle.net/20.500.12634/629 Rights This work is written by (a) US Government employee(s) and is in the public domain in the US. Download date 30/09/2021 08:05:15 applyparastyle "fig//caption/p[1]" parastyle "FigCapt" applyparastyle "fig" parastyle "Figure" Journal of Crustacean Biology Journal of Crustacean Biology The Crustacean Society Journal of Crustacean Biology (2020) 1–7. doi:10.1093/jcbiol/ruaa044 Downloaded from https://academic.oup.com/jcb/article-abstract/doi/10.1093/jcbiol/ruaa044/5874902 by Zoological Society of San Diego user on 04 August 2020 Rediscovery of the horseshoe shrimp Lightiella serendipita Jones, 1961 (Cephalocarida: Hutchinsoniellidae) in San Francisco Bay, California, USA, with a key to the worldwide species of Cephalocarida Crystal Garcia1,2, , Isa Woo1, , D. Christopher Rogers3, , Alison M. Flanagan1,4, and Susan E.W. De La Cruz1, 1U.S. Geological Survey, Western Ecological Research Center, San Francisco Bay Estuary Field Station, PO Box 158, Moffett Field, CA 94035-0158, USA; 2Current affiliation: ICF International Inc., 2600 Hilltop Dr. Suite C137, Richmond, CA 94086, USA; 3Kansas Biological Survey and Biodiversity Institute, University of Kansas, Higuchi Hall, 2101 Constant Avenue, Lawrence, KS 66047-3759, USA; and 4Current affiliation: Department of Recovery Ecology, Institute for Conservation Research, San Diego Zoo Global, Escondido, CA 92027, USA Correspondence: S.E.W. De La Cruz; e-mail: [email protected] (Received 10 April 2020; accepted 25 June 2020) ABSTRACT Lightiella serendipita Jones, 1961 was first discovered in San Francisco Bay, California in 1953, but it had not been observed since 1988. In 2017, a total of 13 adult L. serendipita specimens were found as part of a study in central San Francisco Bay, nearly doubling the total number of specimens ever collected. We measured vertical distribution of macroinvertebrates and environmental variables, including grain size and chemical composition of sediment sam- ples, to evaluate potential features associated with the habitat of the species. Specimens were generally found in sediments with low organic matter (1.7–3%), high sulfate concentrations (594.6–647 ppm SO4), fine grain size (12.8–36.2% sand, 35.6–58% silt, 22.8–37.6% clay) and were mostly found in deep core sections (4–10 cm). Specimens were also consistently observed in cores containing tube-forming Polychaeta (i.e., Sabaco elongatus (Verrill, 1873) and Capitellidae), suggesting L. serendipita may have a commensal relationship with sedentary poly- chaetes, as do other cephalocaridans such as Lightiella incisa Gooding, 1963. We provide a scanning electron micrograph of L. serendipita and the first complete key to the species in class Cephalocarida to help elucidate the taxonomy of this rare crustacean taxon. The per- ceived absence of L. serendipita in previous surveys of the Bay may be attributable to its rarity; however, additional research is needed to fully understand habitat requirements and popula- tion size of this unique endemic species. Key Words: benthic invertebrates, endemic species, subtidal habitats INTRODUCTION poorly understood (Sanders & Hessler, 1964; McLaughlin, 1976; Saloman, 1978; De Troch et al., 2000; Carcupino et al., 2006; Cephalocaridans (class Cephalocrida), commonly referred to as Addis, 2008). horseshoe shrimps, is a poorly understood group of marine crust- Adult cephalocaridans measure ~ 4 mm in length. The adult aceans, with only 13 species recognized worldwide, all of which body posterior to the cephalon consists of nine limb-bearing are reported as uncommon to rare (Olesen et al., 2011). Only thoracic segments (thoracic limb VIII is absent in Lightiella Jones, three species have been collected from more than one locality 1961), 10 limbless abdominal segments, and a telson (Olesen et al., (Carcupino et al., 2006; Olesen et al., 2011); the rest are considered 2014). No eyes have been observed in either the adult or larval endemic. Due to their rarity, small size, and wholly benthic life stages (Olesen et al., 2011). stages, cephalocaridans remain substantially understudied. Since Cephalocaridans are simultaneous hermaphrodites with prob- cephalocaridans have been observed across wide temperature able self-fertilization (Hessler et al., 1970, 1995). Their immo- ranges, in substrates with and without seagrasses, and in inter- bile sperm suggests that individuals must be in direct contact to tidal as well as deep-sea habitats, their habitat requirements are Published by Oxford University Press on behalf of The Crustacean Society 2020. This work is written by (a) US Government employee(s) and is in the public domain in the US. C. GARCIA ET AL. cross-fertilize (Addis, 2008). Larval development shows a pro- METHODS nounced anamorphic gradual growth characterized by a high number of postembryonic stages (metanauplius and juvenile) Sampling and collection (Olesen et al., 2014). Lightiella serendipita was found in benthic cores that were collected Lightiella (family Hutchinsoniellidae Sanders, 1955) is the lar- as part of a multi-year (USGS, Western Ecological Research gest genus of Cephalocrida, with five described species. Lightiella Center (WERC)) study that evaluated the potential impacts of Downloaded from https://academic.oup.com/jcb/article-abstract/doi/10.1093/jcbiol/ruaa044/5874902 by Zoological Society of San Diego user on 04 August 2020 serendipita (Fig. 1) is the type species for the genus by designation. It maintenance dredging on demersal-fish foraging habitats (De La is the only known cephalocaridan from the eastern North Pacific Cruz et al., 2017). Study sites in six marinas (Table 2) were sam- and is one of two species from the Pacific coast of the Americas. pled as part of this effort. All sites were located on the western Other species of the genus are recorded from the American side of central SFB, a predominately polyhaline zone (Thompson Atlantic and Caribbean Sea, the Mediterranean, and New et al., 2007, 2013), and were relatively shallow (≤ 3.2 m mean Caledonia (Olesen et al., 2011). lower low water) with muddy (silt-clay) sediments. Lightiella serendipita was first documented by Jones & Hand Each study site contained three to seven transects (each ap- (1956) and later described by Jones (1961), who collected seven proximately 120 m in length) placed in a recently dredged area specimens during four sampling events from 1953 to 1957 in as well as an adjacent undredged reference area. A minimum of San Francisco Bay (SFB), California, USA (Fig. 2). The seven six coring locations were placed 20 m apart from each other on specimens included two juveniles, three adults, and two in- each of the transects. Two replicate cylindrical cores (10 cm × dividuals of unknown life stages. Reish et al. (1975) collected 10 cm) were collected at each coring location. A total of 1,362 an unidentified cephalocaridan from Anaheim Bay, southern samples were collected during three sampling periods (August California, which may either represent a range extension of 2016, January 2017, and August 2017; Table 2). In August 2016, L. serendipita or a new species. Nine confirmed adult specimens cores were separated into 2 cm sections from Mooring Road and of L. serendipita were collected three decades after its description Pier 32 sites to evaluate how invertebrate abundance and biomass by Jones (1961) in Berkeley, Coyote Point, and San Leandro, changed with sediment depth. Thereafter, samples were divided California by the U.S. Geological Survey (USGS) San Francisco into two sections (0 to 4 cm and 4 to 10 cm) because preliminary Bay Regional Effects Monitoring Program (Table 1, Fig. 2; analyses indicated that most of the abundance and biomass was in Schemel et al., 1988; 1990). the upper 4 cm of the cores (De La Cruz et al., 2017). One sedi- San Francisco Bay is the largest estuary on the west coast of ment core was collected at each transect to characterize sediment the Americas and contains approximately 90% of the coastal grain size (percent sand, silt, and clay) and chemical composition wetlands remaining in California (Goals Project, 2015). The es- (nitrate, sulfur, organic matter, and sedimentary pH). Sediment tuary is one of the most important ecosystems for sustaining en- samples were processed by A & L Agricultural Laboratories demic or endangered species in California, including several (Modesto, CA, USA). anadromous fishes (Swanson, 2007; California State Coastal Faunal cores were rinsed through a 0.5 mm sieve, and ani- Conservancy, 2010) and the commercially important Dungeness mals retained by the sieve were preserved in a 70% ethanol, 1% crab (Metacarcinus magister (Dana, 1852) (Taso, 1979; California rose bengal solution. All macroinvertebrates were sorted, identi- State Coastal Conservancy, 2010). It also supports almost 500 spe- fied, and enumerated by the USGS San Francisco Bay Estuary cies of invertebrates (Goals Project, 1999). While there are many Invertebrate Ecology
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