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A new species of Lepeophtheirus (Copepoda; Caligidae) parasitic on three Kelpfish species (Clinidae) from the Southern California coast.

A new copepod species, Lepeophtheirus schaadti n. sp., is established based on female and male specimens obtained from the Giant Kelpfish, Heterostichus rostratus Girard, 1854, and Striped Kelpfish, Gibbonsia metzi Hubbs, 1927, captured at Inner Cabrillo Beach in southern California, U.S.A. In addition, comparisons with copepod specimens identified by Wilson (1935) as L. parviventris Wilson, 1905 from the Spotted Kelpfish, Gibbonsia elegans (Cooper, 1864), in Newport Bay, California, revealed they are conspecific with L. schaadti n. sp. The new species differs from its congeners by a combination of characters that include: female with a genital complex that is more than half the length of the cephalothoracic shield and with posterolateral lobes, an abdomen that is composed of one somite and is less than one-quarter the length of the genital complex, a maxillulary dentiform process bearing a thin ridge on the inner tine and lacking a basal knob, no myxal process on the maxilliped, apically rounded tines on the sternal furca, the spine on the first exopodal segment of leg 3 inserted distally on the basal swelling, a 3-segmented leg 4 exopod, and a broad inner lobe of leg 5 that does not extend beyond the posterior margin of the genital complex; and male with three accessory claws on the antennal endopod and no myxal process on the maxilliped. L. schaadti n. sp. represents the first account of an ectoparasitic species from the Striped Kelpfish and Spotted Kelpfish, as well as the fourth ectoparasitic species reported from the Giant Kelpfish.

Members of the copepod family Caligidae Burmeister, 1835, commonly known as sea lice, are predominantly external parasites of marine fishes (Dojiri and Ho 2013). Among the 30 valid caligid genera, Lepeophtheirus von Nordmann, 1832 is one of the more speciose genera, with 121 valid species and 2 recognized subspecies (Boxshall and Walter 2016). Lepeophtheirus parasitizes marine teleosts worldwide, but is more diverse in temperate latitudes (Kabata 1979). One species, Lepeophtheirus salmonis (Kroyer, 1837) sensu lato, is the most pathogenic ectoparasite on farmed salmonids in the northern hemisphere (Johnson et al. 2004; Costello 2006). Presently, 17 species of Lepeophtheirus have been reported from 16 fish families along the California coast, from San Francisco Bay in the north to La Jolla in the south of the state (Table 1). Recent samples of Giant Kelpfish (Heterostichus rostratus Girard, 1854) and Striped Kelpfish (Gibbonsia metzi Hubbs, 1927) (Clinidae Swainson, 1839) collected within the Port of Los Angeles during Cabrillo Marine Aquarium's Inner Cabrillo Beach Survey (ICBS) were infected with an unidentified species of Lepeophtheirus. The ICBS is a long-term, tri-annual survey designed to monitor the abundance and diversity of the invertebrates and fishes living in subtidal eelgrass beds off Inner Cabrillo Beach. Subsequent examination of the unidentified Lepeophtheirus specimens revealed they were not conspecific with Lepeophtheirus parviventris Wilson, 1905, a species previously reported by Wilson (1935) from the Giant Kelpfish in Newport Bay, California. Indeed, the Lepeophtheirus specimens from Inner Cabrillo Beach, as well as those of Wilson (1935), represent an undescribed species, which is described in detail herein.

Materials and Methods

Nearly all copepod specimens of the new taxon were obtained from Heterostichus rostratus samples that were collected in beach seines at three stations along Inner Cabrillo Beach during the 2011-2014ICBS. Only two copepod specimens were obtained from one individual of Gibbonsia metzi captured in a winter 2011 ICBS. Copepod samples were preserved in 70% ethanol upon removal from the host. Copepod specimens were later soaked in lactophenol prior to examination using an Olympus SZX10 dissection microscope and an Olympus BX53 compound microscope equipped with differential interference contrast optics. Selected specimens were also measured intact using an ocular micrometer and/or dissected and examined according to the wooden slide procedure of Humes and Gooding (1964). In the description, length measurements are provided first, followed by width measurements; all measurements given are expressed as the mean followed by the range in parentheses. Pencil drawings of the copepod body and appendages were made with the aid of a drawing tube. Drawings were subsequently inked in with Sakura Pigma Micron[TM] pens on 110 g/[m.sup.2] tracing paper, digitized with a CanoScan LiDE 500F scanner, and assembled into figure plates using Adobe Photoshop. Morphological terminology follows Huys and Boxshall (1991) and Dojiri and Ho (2013). Fish names and classifications conform to Page et al. (2013). Type material and voucher specimens of the new taxon are deposited at the Crustacea Department of the Natural History Museum of Los Angeles County (LACM), Los Angeles, California, U.S.A., and Cabrillo Marine Aquarium (CMA), San Pedro, California, U.S.A.

Type material and voucher specimens of L. parviventris deposited by Wilson (1905, 1908, 1924, 1935) in the National Museum of Natural History (USNM), Smithsonian Institution, Washington, D.C., were also examined for comparative purposes: syntypes comprising 22 females, 15 males, and 11 juveniles (USNM 42064), ex Gadus macrocephalus Tilesius, 1810 (Gadidae Rafinesque, 1810), ChignikBay, Alaska, 1903; 2 females and 1 male (USNM 69798), ex skin of Gibbonsia evides (Jordan & Gilbert, 1883) (=Gibbonsia elegans (Cooper, 1864)) (as Heterostichus rostratus), Newport Bay, California, 1934; 5 females and 1 male (USNM 38566), ex Sebastes rubrivinctus (Jordan & Gilbert, 1880) (as Sebastodes rubrivinctus (Jordan & Gilbert, 1880)) (Scorpaenidae Risso, 1827), Station 4417, off Santa Barbara Island, 29 fathoms, April 12, 1904; 1 female (USNM 53491), exHalichoeres semicinctus (Ayres, 1859) (as Iridio semicinctus (Ayres, 1859)) (Labridae Cuvier, 1816), southern California, April 12, 1913.

Results

Lepeophtheirus schaadti n. sp. (Figs. 1-6)

Type material. Holotype female (LACM CR-2011-3), allotype male (LACM CR-2011-4), and 1 male and 2 female paratypes (CMA 2017.04.0002), ex Heterostichus rostratus (222 mm SL), Station 2 (33[degrees] 42' 38.3" N, 118[degrees]16'58.5"W), off Inner Cabrillo Beach, San Pedro, California, U.S.A., February 5, 2011.

Other material examined. From Station 1 (33[degrees]42'42.6"N, 118[degrees]16'59.1"W), off Inner Cabrillo Beach, San Pedro, California, U.S.A.: 10 females (7 with an unidentified species of Udonella Johnston, 1835 (Monogenea) attached to the genital complex) and 4 males (3 with Udonella sp. attached to the genital complex) (CMA 2017.04.0006), ex H. rostratus (330 mm SL), October 10, 2014; 2 females (each with Udonella sp. attached to the genital complex and egg sacs) (CMA 2017.04.0005), ex H. rostratus (169 mm SL), October 11, 2014; 3 females (LACM MBPC 17851), ex H. rostratus (193 mm SL), October 11, 2014; 1 female and 1 male (LACM MBPC 17852), ex H. rostratus (197 mm SL), October 11, 2014. From Station 2 (33[degrees]42'38.3"N, 118[degrees]16'58.5"W), off Inner Cabrillo Beach, San Pedro, California, U.S.A.: 1 female and 1 male (CMA 2017.04.0014), ex Gibbonsia metzi (110 mm SL), February 5, 2011; 1 female (CMA 2017.04.0003), ex H. rostratus (170 mm SL), February 5, 2011; 1 female and 1 male (CMA 2017.04.0004), ex H. rostratus (120 mm SL), February 5, 2011; 1 female and 1 male (CMA 2017.04.0010), ex H. rostratus (86 mm SL), October 28, 2011; 1 female (CMA 2017.04.0011), ex H. rostratus (182 mm SL), October 28, 2011; 2 females and 2 males (CMA 2017.04.0012), ex H. rostratus (163 mm SL), October 28, 2011; 1 female and 2 males (CMA 2017.04.0013), ex H. rostratus (178 mm SL), October 28, 2011; 4 females (CMA 2017.04.0008), ex H. rostratus (200 mm SL), October 17, 2013; 1 female (CMA 2017.04.0009), ex H. rostratus (245 mm SL), October 17, 2013. From Station 3 (33[degrees]42'35.5"N, 118[degrees]16'51.3"W), off Inner Cabrillo Beach, San Pedro, California, U.S.A.: 1 female (dissected and mounted on glass slide) and 3 males (1 male partially dissected and mounted on glass slide) (CMA 2017.04.0007), ex H. rostratus (260 mm SL), June 10,2013.

Description of adult female. Body (Figure 1A) 4.08 (3.85-4.25) mm long (excluding caudal setae) (n=7). Cephalothoracic shield subcircular, nearly as long as wide [2.19 (2.05-2.35) x 2.11 (1.98-2.33) mm], with well-developed paired frontal plates, posterior margin of thoracic zone extending beyond posterior limit of lateral zone, and hyaline membrane along frontal and lateral rims. Free fourth pedigerous somite about three times wider than long [209 (170230) x 609 (560-645) [micro]m] and indistinctly separated from genital complex. Genital complex large, more than half the length of cephalothoracic shield, marginally wider than long [1.41 (1.30-1.55) x 1.52 (1.38-1.78) mm], with nearly parallel lateral margins and protruded posterolateral corners. Abdomen (Fig. 1B) composed of 1 somite, 247 (220-280) x 416 (370-460) [micro]m, widest anteriorly, and indistinctly separated from genital complex. Caudal ramus (Fig. 1C) slightly longer than wide [111 (105-120) x 99 (90-110) [micro]m], with 6 plumose setae (seta I absent) and short row of setules along inner margin. Egg sacs (Fig. 1A) uniseriate.

Antennule (Fig. 1D) 2-segmented. Proximal segment longer than distal segment, bearing 1 tiny semispherical knob and 1 bifid process on posterodistal corner and 27 setae (25 hirsute, 2 naked) along anterior margin. Distal segment cylindrical, bearing 12 setae (2 setae near posterodistal corner share a common base) and 2 aesthetascs.

Antenna (Fig. 2A) 3-segmented, comprising coxa, basis and 1-segmented endopod incorporating distal claw. Coxa with long, apically rounded process on posterolateral corner. Basis stout, with corrugated surface on protruded, inner distal corner and 1 large, outer distal adhesion pad on dorsal surface. Endopod long, uncinate, bearing 2 naked setae.

Postantennal process (Fig. 2A) with small bump midway on anterior margin of basal section, pair of setulose papillae on base, 1 setulose papilla posterior to base, and recurved, apically rounded hook.

Mandible (Fig. 2B) modified into elongate stylet bearing distolateral hyaline membrane and 12 distomedial teeth (1 blunt, 11 sharp).

Maxillule (Fig. 2A) composed of trisetose papilla and bifid dentiform process. Sclerite anterior to papilla with posteriorly-directed triangular process. Tines on dentiform process subequal, with thin ridge on inner tine.

Postoral process (Fig. 2A) small, triangular.

Maxilla (Fig. 2C), brachiform, 2-segmented, composed of elongate, unarmed syncoxa and slender basis. Basis with large flabellum and long apical calamus and shorter apical canna; calamus furnished with finely serrated membranes; canna with finely serrated posterior margin.

Maxilliped (Fig. 2D) large, subchelate, 3-segmented, comprising long protopod (corpus) and subchela consisting of free endopodal segment (shaft) and claw. Protopod with 2 large patches of denticles near inner margin and small patch of denticles on distolateral corner. Shaft urnarmed. Claw with long, naked basal seta and 2 thin ridges and fine striations distally.

Tines of sternal furca (Fig. 2E) longer than box, slightly divergent, and apically rounded; shallow T-shaped depression present, situated anterior to base of box.

Legs 1 to 3 (Figs. 3A-B and 4A) biramous; leg 4 (Fig. 4C) uniramous. Armature formula of legs 1-4 is shown in Table 2.

Leg 1 (Fig. 3A) intercoxal sclerite naked and elongate. Protopod with 1 outer and 1 inner plumose setae, 1 proximolateral setulose papilla, and 1 mid-lateral pore. First exopodal segment with 1 small, naked outer spine and inner row of setules. Second exopodal segment with 4 apical elements (3 spines, 1 seta), 3 inner plumose setae, tiny inflated process near apical margin, and pectinate membrane at base of each apical spine; outer apical spine with row of tiny denticles on anterior and posterior sides (denticles on posterior side not drawn); middle and inner apical spines each with serrations on anterior and posterior sides (serrations on posterior side not drawn) and an accessory process; apical seta plumose, shorter than outer apical spine. Endopod digitiform, bearing 2 elements apically.

Leg 2 (Fig. 3B) intercoxal sclerite subquadrate, with large hyaline membrane along distal margin. Coxa with 1 inner plumose seta and 2 pores on anterior surface. Basis with 1 outer short, plumose seta, 1 minute pore near outer margin, 1 inner sensillum, and large hyaline membrane along inner margin. Exopod 3-segmented, with large hyaline membrane covering dorsal surface of ramus. First segment with 1 inner plumose seta, inner row of setules, and pectinate membrane at base of large outer spine; latter with sclerotized flange along outer margin and fine serrations along inner margin. Second segment with 1 inner plumose seta, inner row of setules, 1 outer serrate spine, and 1 minute pore on anterior surface. Third segment with inner row of setules, 5 inner plumose setae, 3 outer spines, and 1 minute pore near lateral margin; proximal outer spine with serrated margins; middle outer spine with hyaline membrane along both margins; outer distal spine with hyaline membrane along outer margin and row of setules along inner margin. Endopod 3-segmented. First segment with 1 inner plumose seta and row of setules on distolateral corner. Second segment with 2 inner plumose setae, row of setules along inner and outer margins, and 1 minute pore on anterior surface. Third segment with 6 plumose setae and short row of setules along proximolateral and proximomedial margins.

Leg 3 (Fig. 4A) protopod large, modified to form apron, with 1 outer plumose seta situated near base of exopod, 1 inner plumose seta near large intercoxal sclerite, 1 proximolateral corrugated pad on dorsal surface, 3 marginal membranes, minute pores scattered on ventral surface, and 2 unequal sensilla along posterior margin. Exopod (Fig. 4B) 3-segmented. First segment with 1 inner plumose seta, 1 apical spine reflexed over second segment and furnished with sclerotized flange along outer margin, and 1 minute pore, several sensilla and sclerotized flange on outer basal swelling. Second segment with 1 outer naked spine, 1 inner plumose seta, 1 minute pore, and setules along lateral and medial margins. Third segment with 4 plumose setae, 3 naked spines, and setules along proximal margins. Endopod 2-segmented. First segment with 1 inner plumose seta and outer row of setules. Second segment with 6 plumose setae and setules along outer and inner margins.

Leg 4 (Fig. 4C) protopod with 1 distolateral plumose seta. First exopodal segment with pectinate membrane at base of small, outer naked spine and serrations and several sensilla along outer margin. Second exopodal segment similar to first segment but with much larger outer spine furnished with pectinate margins. Third exopodal segment with 3 apical pectinate spines, pectinate membrane at base of each spine, and tiny serrations along outer margin; spines progressively increase in length from outer to inner apical margin.

Leg 5 (Fig. 4D) vestigial, comprised of small setiferous papilla and broad trisetose lobe on posteroventral surface of genital complex.

Leg 6 (not figured) rudimentary, represented by unarmed genital operculum at gonopore opening.

Description of adult male. Body (Fig. 5A) 2.64 (2.53-2.78) mm long (excluding caudal setae) (n=4). Cephalothoracic shield slightly longer than wide [1.73 (1.65-1.83) x 1.62 (1.58-1.65) mm], ornamented as in female. Free fourth pedigerous somite wider than long [170 (160-180) x 406 (385-420) [micro]m]. Genital complex wider than long [448 (420-470) x 514 (490-540) [micro]m]. Abdomen composed of 1 somite, 210 (210-210) x 268 (260-275) [micro]m], narrowed at junction with genital complex. Caudal ramus longer than wide [120 (110-130) x 110 (100-115) [micro]m], armed as in female.

All limbs as in female, except for the following. Antennule (Fig. 5B) with 29 setae (27 hirsute, 2 naked) on proximal segment. Antenna (Fig. 5C-E) 3-segmented, comprising coxa, basis, and 1-segmented endopod incorporating distal claw. Coxa with large corrugated pad along outer margin on posterior side and fine striations on inner distal margin on anterior side. Basis with 1 large and 1 small corrugated pad on posterior side and 3 unequal corrugated pads on anterior side. Endopod forming robust terminal claw with sclerotized flange on posterior side and bearing 2 naked setae and 3 accessory claws. Maxillule (Fig. 5F) with hyaline digitiform process medial to bifid dentiform process. Postoral process (Fig. 5F) elongate and corrugated. Maxilliped (Fig. 6A) lacking small patch of denticles on distolateral corner of protopod and fine apical striations on claw. Weakly sclerotized adhesion pad (Fig. 6B) present, situated anterior to sternal furca. Leg 5 (Fig. 6C) lobate, bearing 2 plumose and 2 unipinnate setae. Leg 6 (Fig. 6C) forming genital operculum, armed distally with 1 pinnate and 2 plumose setae.

Variability. Female specimen from H. rostratus captured at Station 3 without row of setules along inner margin of caudal rami (Fig. 1B) and with one apically bifurcate seta on distal endopodal segment of right leg 3 (Fig. 6D).

Attachment site. Body surface.

Prevalence and mean intensity. From a total of 655 Giant Kelpfish that were inspected for Lepeophtherius infections between June 2011 and February 2013, 233 L. schaadti n. sp. were removed from 86 fish (prevalence = 13.1%; mean intensity = 2.71). By contrast, from a total of 2716 Striped Kelpfish captured within the same time period at Inner Cabrillo Beach, only two L. schaadti n. sp. were recovered from one fish (prevalence = 0.04%; mean intensity = 2).

Etymology. This species is named in honor of Mike Schaadt, the Director of the Cabrillo Marine Aquarium.

Remarks. Examination of Wilson's (1908, 1924) Lepeophtherius specimens from the Flag Rockfish, Sebastes rubrivinctus, and Rock Wrasse, Halichoeres semicinctus, captured in California waters revealed they are not conspecific with L. parviventris. More importantly, examination of Wilson's (1935) Lepeophtherius specimens from Newport Bay, California, revealed they are conspecific with L. schaadti n. sp. and the host was the Spotted Kelpfish, Gibbonsia elegans, rather than the Giant Kelpfish, as indicated on the vial label.

L. schaadti n. sp. resembles L. elegans Gusev, 1951, L. hexagrammi Gusev, 1951, and L. hospitalis Fraser, 1920 by having in the female a genital complex that is at least half the length of the cephalothoracic shield (including frontal plates) and with small, rounded posterolateral lobes, a 1-segmented abdomen that is less than one-quarter the length of the genital complex, a maxillule with two large tines on the dentiform process, a maxilliped without a myxal process, a pair of non-bifid tines on the sternal furca, the spine on the first exopodal segment of leg 3 inserted distally on the basal swelling, a 3-segmented leg 4 exopod, and the inner lobe of leg 5 not protruding beyond the posterior margin of the genital complex.

L. elegans can be distinguished from L. schaadti n. sp. by having a smooth inner distal corner on the basis of the female antenna, pointed tines and no ridge on the inner tine of the dentiform process of the female maxillule, no denticles on the outer distal corner of the protopod of the female maxilliped, pointed tines on the female sternal furca, a subtriangular inner lobe on the female leg 5, no accessory claws on the endopod of the male antenna, and a large, cone-shaped myxal process on the protopod of the male maxilliped.

L. hexagrammi can be differentiated from L. schaadti n. sp. by the presence of a smooth inner distal corner on the basis of the female antenna, a broader and less recurved hook on the postantennal process of both sexes, both an outer basal knob and a ridge on both tines on the dentiform process of the female maxillule, more tapered tines on the female sternal furca, one accessory claw on the endopod of the male antenna, and a large, cone-shaped myxal process on the protopod of the male maxilliped.

L. hospitalis can be discerned from L. schaadti n. sp. by having a pointed and less recurved hook on the postantennal process of both sexes, a basal semispherical knob on the dentiform process of the maxillule of both sexes, broad flanges on the pointed tines of the female sternal furca, a subtriangular inner lobe on the female leg 5, and two accessory claws on the endopod of the male antenna.

Discussion and Conclusions

The discovery of L. schaadti n. sp. represents the first account of an ectoparasitic species from the Striped Kelpfish and Spotted Kelpfish, as well as the fourth ectoparasitic species reported from the Giant Kelpfish. The copepods Chondracanthus heterostichi Ho, 1972 and C. horridus Heller, 1865 (Chondracanthidae Milne Edwards, 1840) and the leech Heptacyclus cabrilloi Burreson, Kalman Passarelli & Kim, 2012 (Piscicolidae Johnston, 1865) were previously recorded from the Giant Kelpfish (Wilson 1935; Ho 1972b; Burreson et al. 2012). It must be noted, however, that Wilson's record of C. horridus on the Giant Kelpfish requires verification, as C. horridus was originally described from the Black Goby, Gobius niger Linnaeus, 1758 (as Gobius jozo Linnaeus, 1758) (Gobiidae Cuvier, 1816), from the Mediterranean Sea (Heller 1865).

In this study, 13.1% of the Giant Kelpfish were infected with L. schaadti n. sp. as compared to only 0.04% of the Striped Kelpfish. These disparate infection levels suggest that the Giant Kelpfish is a more common host of L. schaadti n. sp. at Inner Cabrillo Beach. It remains to be determined how common L. schaadti n. sp. is throughout the geographical range of its kelpfish hosts, including the Spotted Kelpfish.

From 2011 to 2014, 20 individuals of L. schaadti n. sp. were infected with the hyperparasitic monogene Udonella sp. (Udonellidae Taschenberg, 1879). Nearly all Udonella specimens were attached to the external surface of the copepod's genital complex, with a few on the cephalothorax and egg sacs. In California, Udonella caligorum Johnston, 1835 has been reported from the copepods Trebius caudatus Kroyer, 1838 and T. latifurcatus Wilson, 1921 (Trebiidae Wilson, 1905) parasitic on the Bat Ray, Myliobatis californica Gill, 1865 (Myliobatidae Bonaparte, 1835); on the Curlfin Sole, Pleuronichthys decurrens Jordan & Gilbert, 1881 (Pleuronectidae Rafinesque, 1815); and on the isopod Elthusa vulgaris (Stimpson, 1857) (as Lironeca vulgaris Stimpson, 1857) (Cymothoidae Leach, 1818) parasitic on the Sand Sole, Psettichthys melanostictus Girard, 1854 (Pleuronectidae) (Love and Moser, 1983). Identification of the Udonella material is currently underway and will be dealt with in detail elsewhere.

Acknowledgements

We thank Don Buth (University of California, Los Angeles) for support during parasite collections, Rafael Lemaitre and Chad Walter (Smithsonian Institution) for kindly arranging the loan of copepod material, and Mas Dojiri (City of Los Angeles) for guidance during the initial phase of this study. We also thank all the Cabrillo Marine Aquarium staff and volunteers for help with collection of kelpfishes during the Inner Cabrillo Beach Survey.

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Julianne Kalman Passarelli (1) and Danny Tang (2)

(1) Cabrillo Marine Aquarium, 3720 Stephen M. White Drive, San Pedro, CA 90731, USA

(2) Laboratory, Monitoring, and Compliance Division, Orange County Sanitation District, 10844 Ellis Ave, Fountain Valley, CA 92708, USA

Corresponding author: julianne.passarelli@lacity.org

Caption: Fig 1. Lepeophtheirus schaadti n. sp., adult female. A) Habitus, dorsal; B) Abdomen and caudal rami, dorsal; C) Right caudal ramus, dorsal; D) Right antennule (arrowheads indicate naked setae on proximal segment), ventral. Scale bars: 1.00 mm for A; 200 [micro]m for B; 50 [micro]m for C; 100 [micro]m for D.

Caption: Fig 2. Lepeophtheirus schaadti n. sp., adult female. A) Right antenna (A2) (ap = adhesion pad), postantennal process (PAP), maxillule (MX1) and postoral process (POP), ventral; B) Left mandible, posterior; C) Right maxilla, anterior; D) Right maxilliped, anterior; E) Sternal furca, ventral. Scale bars: 100 [micro]m for A, E; 50 [micro]m for B; 150 [micro]m for C, D.

Caption: Fig 3. Lepeophtheirus schaadti n. sp., adult female. A) Right leg 1 with detail of endopod and apical spines on second exopodal segment, anterior; B) Left leg 2 (mm = marginal membrane) with detail of outer spine on first and second exopodal segments and proximalmost outer spine on third exopodal segment, anterior. Scale bars: 200 M-m for A, B.

Caption: Fig 4. Lepeophtheirus schaadti n. sp., adult female. A) Left leg 3, ventral; B) Left leg 3 exopod, ventral; C) Left leg 4 with detail of serrations along outer margin of first exopodal segment and fine teeth along outer margin of spine on second exopodal segment, anterior; D) Left leg 5 (P5), spermatophore (S) and egg sac (ES), ventral. Scale bars: 200 [micro]m for A, C, D; 50 [micro]m for B.

Caption: Fig 5. Lepeophtheirus schaadti n. sp., adult male. A) Habitus, dorsal; B) Left antennule, ventral; C) Left antenna, posteromedial; D) Left antenna, anterolateral; E) Distal endopodal segment of left antenna, posterior; F) Left maxillule (MX1) and postoral process (POP), ventral. Scale bars: 1.00 mm for A; 100 [micro]m for B, C, D, F; 50 [micro]m for E.

Caption: Fig 6. Lepeophtheirus schaadti n. sp., adult male (A-C) and adult female (D). A) Left maxilliped, anterior; B) Sternal furca (SF) and adhesion pad (AP), ventral; C) Left legs 5 (P5) and 6 (P6), ventral; D) Distal endopodal segment of right leg 3 (arrow indicates abnormal seta), ventral. Scale bars: 100 [micro]m for A, B, C; 50 [micro]m for D.
Table 1. Fish hosts and locality records for species of
Lepeophtheirus reported from California, U.S.A.

Copepod species         Host family       Host species

Lepeophtheirus sp.      Kyphosidae        Medialuna
                                          californiensis
                                          (Steindachner. 1876)

                        Labridae          Oxyjulis californica
                                          (Gunther. 1861)

                        Pomacentridae     Hypsypops rubicundus
                                          (Girard. 1854) (as
                                          Hypsypops rubicunda)

                        Sciaenidae        Genyonemus lineatus
                                          (Ayres. 1855)

                        Sciaenidae        Menticirrhus
                                          undulatus (Girard.
                                          1854)

                        Scorpaenidae      Sebastes paucispinis
                                          Ayres. 1854

Lepeophtheirus          Paralichthyidae   Paralichthys
bifidus Fraser. 1920                      californiens (Ayres.
                                          1859)

                        Pleuronectidae    Pleuronichthys
                                          guttulatus Girard.
                                          1856 (as Hypsopsetta
                                          guttulata)

Lepeophtheirus          Embiotocidae      Cymatogaster
bifurcatus Wilson.                        aggregata Gibbons.
1905                                      1854

                        Embiotocidae      Phanerodon/meatus
                                          Girard. 1854

                        Paralichthyidae   Paralichthys
                                          californiens (Ayres.
                                          1859)

                        Pleuronectidae    Pleuronichthys
                                          verticalis Jordan &
                                          Gilbert. 1880

                        Pleuronectidae    Psettichthys
                                          melanostictus Girard.
                                          1854

Lepeophtheirus          Scorpaenidae      Scorpaena guttata
brachyurus Heller.                        Girard. 1854
1865

Lepeophtheirus          Serranidae        Paralabrax clathratus
constrictus Wilson.                       (Girard. 1854)
1908

                        Serranidae        Paralabrax
                                          maculatofasciatus
                                          (Steindachner. 1868)

                        Serranidae        Paralabrax nebulifer
                                          (Girard. 1854)

Lepeophtheirus          Sciaenidae        Atractoscion nobilis
longiabdominis                            (Ayres. 1860) (as
Shiino. 1960                              Cynoscion nobilis)

Lepeophtheirus          Polyp rionidae    Stereolepis gigas
longipes Wilson 1905                      Ayres. 1859

                        Scorpaenidae      Sebastes setriceps
                                          (Jordan & Gilbert.
                                          1880)

                        Serranidae        Paralabrax clathratus
                                          (Girard. 1854)

Lepeophtheirus          Molidae           Mola mola (Linnaeus.
nordmanni (Milne                          1758)
Edwards. 1840)

Lepeophtheims           Clinidae          Heterostichus
patviventris Wilson,                      rostratas Girard
1905                                      1854b

                        Labridae          Halichoeres
                                          semicinctus (Ayres,
                                          1859) (as Iridio
                                          semicinctus)

                        Scorpaenidae      Sebastes rubrivinctus
                                          (Jordan & Gilbert,
                                          1880) (as Sebastodes
                                          rubrivinctus)

                        Urotrygonidae     Urobatis halleri
                                          (Cooper, 1863) (as
                                          Urolophus halleri)

Lepeophtheims parvus    Embiotocidae      Cymatogaster
Wilson, 1908                              aggregata Gibbons,
                                          1854

                        Embiotocidae      Damalichthys vacca
                                          (Girard, 1855)

                        Labridae          Semicossyphus pulcher
                                          (Ayres, 1854) (as
                                          Pimelometopon pulcher
                                          and P. pulchrum)

Lepeophtheims paulus    Scorpaenidae      Sebastes serriceps
Cressey, 1969                             (Jordan & Gilbert,
                                          1880) (as Sebastodes
                                          serriceps)

Lepeophtheims           Hexagrammidae     Ophiodon elongatus
pravipes Wilson, 1912                     Girard 1854

                        Scorpaenidae      Scorpaena guttata
                                          Girard, 1854

Lepeophtheims           Batrachoididae    Porichthys notatus
remiopsis Dojiri,                         Girard, 1854
1979

                        Paralichthyidae   Hippoglossina stomata
                                          Eigenmann &
                                          Eigenmann, 1890

                        Paralichthyidae   Xystreurys liolepis
                                          Jordan & Gilbert,
                                          1880

                        Pleuronectidae    Parophtys vetulus
                                          Girard, 1854

                        Pleuronectidae    Pleuronichthys
                                          verticalis Jordan &
                                          Gilbert,
                                          1880

                        Cottidae          Chitonotus pugetensis
                                          (Steindachner, 1876)

Lepeophtheims           Paralichthyidae   Citharichthys
rotundipes Dojiri,                        stigmaeus Jordan &
1979                                      Gilbert, 1882

                        Scorpaenidae      Scorpaen a guttata
                                          Girard, 1854

Lepeophtheirus          Salmonidae        Oncorhynchus
salmonis oncorhynchi                      tshawytscha (Walbaum,
Skern-Mauritzen,                          1792)
Torrissen and Glover,
2014

Lepeophtheirus spatha   Paralichthyidae   Paralichthys
Dojiri and Brantley,                      californiens (Ayres,
1991                                      1859)

Lepeophtheirus          Sciaenidae        Atractoscion nobilis
thompsoni Baird, 1850                     (Ayres, 1860) (as
                                          Cynoscion nobilis)

Copepod species         Host family       Locality

Lepeophtheirus sp.      Kyphosidae        Off La Jolla

                        Labridae          Off La Jolla

                        Pomacentridae     Off La Jolla

                        Sciaenidae        Southern California

                        Sciaenidae        Southern California

                        Scorpaenidae      Southern California

Lepeophtheirus          Paralichthyidae   Anaheim Bay
bifidus Fraser. 1920
                        Pleuronectidae    Anaheim Bay

Lepeophtheirus          Embiotocidae      Southern California
bifurcatus Wilson.
1905                    Embiotocidae      Southern California

                        Paralichthyidae   Santa Monica Bay

                        Pleuronectidae    Santa Monica Bay

                        Pleuronectidae    San Francisco Bay

Lepeophtheirus          Scorpaenidae      Off La Jolla
brachyurus Heller.
1865

Lepeophtheirus          Serranidae        Southern California
constrictus Wilson.
1908                    Serranidae        Off La Jolla

                        Serranidae        Southern California

Lepeophtheirus          Sciaenidae        Off La Jolla
longiabdominis
Shiino. 1960

Lepeophtheirus          Polyp rionidae    Off La Jolla
longipes Wilson 1905
                        Scorpaenidae      Off La Jolla

                        Serranidae        Catalina Island

Lepeophtheirus          Molidae           Santa Catalina Island
nordmanni (Milne                          Southern California
Edwards. 1840)                            Monterey Bay

Lepeophtheims           Clinidae          Newport Bay
patviventris Wilson,
1905                    Labridae          Southern California

                        Scorpaenidae      Off Santa Barbara Island

                        Urotrygonidae     California

Lepeophtheims parvus    Embiotocidae      Southern California
Wilson, 1908
                        Embiotocidae      Southern California

                        Labridae          Off San Diego

                                          Off La Jolla

Lepeophtheims paulus    Scorpaenidae      Off La Jolla
Cressey, 1969

Lepeophtheims           Hexagrammidae     Off San Diego
pravipes Wilson, 1912
                        Scorpaenidae      Off Pacific Grove

Lepeophtheims           Batrachoididae    Off Huntington Beach
remiopsis Dojiri,
1979                    Paralichthyidae   Off Huntington Beach
                                          Santa Monica Bay

                        Paralichthyidae   Santa Monica Bay

                        Pleuronectidae    Off Huntington Beach

                                          Santa Monica Bay

                        Pleuronectidae    Off Huntington Beach

                                          Santa Monica Bay

                        Cottidae          Off Huntington Beach

Lepeophtheims           Paralichthyidae   Off Huntington Beach
rotundipes Dojiri,
1979                    Scorpaenidae      Off Huntington Beach

                                          Santa Monica Bay

Lepeophtheirus          Salmonidae        Monterey Bay
salmonis oncorhynchi
Skern-Mauritzen,
Torrissen and Glover,
2014

Lepeophtheirus spatha   Paralichthyidae   Santa Monica Bay
Dojiri and Brantley,
1991

Lepeophtheirus          Sciaenidae        Off La Jolla
thompsoni Baird, 1850

Copepod species         Host family       Reference

Lepeophtheirus sp.      Kyphosidae        Hobson (1971)

                        Labridae          Hobson (1971)

                        Pomacentridae     Hobson (1971)

                        Sciaenidae        Love and Moser (1983)

                        Sciaenidae        Love and Moser (1983)

                        Scorpaenidae      Love and Moser (1983)

Lepeophtheirus          Paralichthyidae   Ho (1972a)
bifidus Fraser. 1920
                        Pleuronectidae    Ho (1975)

Lepeophtheirus          Embiotocidae      Love and Moser (1983)
bifurcatus Wilson.
1905
                        Embiotocidae      Love and Moser (1983)

                        Paralichthyidae   Kaiman (2006)

                        Pleuronectidae    Kaiman (2006)

                        Pleuronectidae    Wilson (1908)

Lepeophtheirus          Scorpaenidae      Wilson (1908)
brachyurus Heller.
1865

Lepeophtheirus          Serranidae        Love and Moser (1983)
constrictus Wilson.
1908
                        Serranidae        Wilson (1908)

                        Serranidae        Love and Moser (1983)

Lepeophtheirus          Sciaenidae        Shiino (1960)
longiabdominis
Shiino. 1960

Lepeophtheirus          Polyp rionidae    Wilson (1908)
longipes Wilson 1905

                        Scorpaenidae      Hobson (1971)

                        Serranidae        Wilson (1921)

Lepeophtheirus          Molidae           Wilson (1908)
nordmanni (Milne
Edwards. 1840)
                                          Wilson (1908) (a)

                                          Wilson (1935)

Lepeophtheims           Clinidae          Wilson (1935) (b)
patviventris Wilson,
1905
                        Labridae          Wilson (1924) (c)

                        Scorpaenidae      Wilson (1908) (c)

                        Urotrygonidae     Wilson (1924) (d)

Lepeophtheims parvus    Embiotocidae      Love and Moser(1983)
Wilson, 1908
                        Embiotocidae      Love and Moser(1983)

                        Labridae          Wilson (1908); Shiino
                                          (1963)

                                          Hobson (1971)

Lepeophtheims paulus    Scorpaenidae      Cressey (1969)
Cressey, 1969

Lepeophtheims           Hexagrammidae     Shiino (1965) (e)
pravipes Wilson, 1912
                        Scorpaenidae      Wilson (1935)

Lepeophtheims           Batrachoididae    Dojiri (1979)
remiopsis Dojiri,
1979
                        Paralichthyidae   Dojiri (1979)
                                          Kaiman (2006)

                        Paralichthyidae   Kaiman (2006)

                        Pleuronectidae    Dojiri (1979)

                                          Kaiman (2006)

                        Pleuronectidae    Dojiri (1979)

                                          Kaiman (2006)

                        Cottidae          Dojiri (1979)

Lepeophtheims           Paralichthyidae   Dojiri (1979)
rotundipes Dojiri,
1979
                        Scorpaenidae      Dojiri (1979)

                                          Kaiman (2006)

Lepeophtheirus          Salmonidae        Wilson (1908) (f)
salmonis oncorhynchi
Skern-Mauritzen,
Torrissen and Glover,
2014

Lepeophtheirus spatha   Paralichthyidae   Dojiri and Brantley
Dojiri and Brantley,                      (1991); Kaiman (2006)
1991

Lepeophtheirus          Sciaenidae        Wilson (1908)
thompsoni Baird, 1850

(a) Reported as L. insignis.

(b) Specimens were reported as L. parviventris, but examination of
these specimens revealed they are L. schaadti n. sp. Furthermore, the
same specimens were reported from Heterostichus rostratas, but
Gibbonsia evides (=Gibbonsia elegans) was handwritten on the vial
label.

(c) Specimens were reported as L. parviventris, but examination of
these specimens revealed they are not conspecific with L.
parviventris.

(d) Reported from U. hallen held in an aquarium at the marine station
of the University of Southern California, at Venice, California.

(e) Reported as L. trifidus.

(f) Reported asi. salmonis.

Table 2. Armature on legs 1-4 (Roman numerals = spines; Arabic
numerals = setae).

          Coxa   Basis   Exopod             Endopod

Leg 1 *   0-0    1-1     I-0;0,ffl + 1,3    vestigial
Leg 2     0-1    1-0     I-1; I-1; II,I,5   0-1; 0-2; 6
Leg 3 *   0-1    1-0     I-1; I-1; II,I,4   0-1; 6
Leg 4 *   0-0    1-0     I-0; I-0; II,I,0   absent

* Although the coxa and basis are fused to form a protopod in this
leg, these segments are treated separately in this Table.
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