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Research Article
Status of Exosphaeroma amplicauda (Stimpson, 1857), E. aphrodita (Boone, 1923) and description of three new species (Crustacea, Isopoda, Sphaeromatidae) from the north-eastern Pacific
expand article infoAdam Wall, Niel Bruce§, Regina Wetzer
‡ Natural History Museum of Los Angeles County, Los Angeles, United States of America
§ Museum of Tropical Queensland, Townsville, Queensland, Australia
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Abstract

Exosphaeroma amplicauda (Stimpson, 1857) from the west coast of North America is reviewed and redescribed and revealed to be a group of closely related species. A neotype is designated and the species redescribed based on the neotype and topotypic specimens. Exosphaeroma amplicauda is known only from the coast of California, at Marin, Sonoma and San Mateo Counties. E. aphrodita (Boone, 1923), type locality La Jolla, California and previously considered nomen dubium is taken out of synonymy and re-validated. A further three species: E. paydenae sp. n., E. russellhansoni sp. n., and E. pentcheffi sp. n. are described herein. Sphaeroma octonctum Richardson, 1899 is placed into junior synonymy with Exosphaeroma amplicauda. A key to the Pacific West Coast Exosphaeroma is provided.

Keywords

Isopoda, Sphaeromatidae, Exosphaeroma, Alaska, Washington, California, intertidal

Introduction

The Sphaeromatidae is a large family, currently with 99 accepted genera (WoRMS, World Register of Marine Species, Bruce and Schotte 2013) and nearly 700 species. The phylogenetic relationships of the Sphaeromatidea were reviewed by Wetzer et al. 2013, but no family-wide treatment since the time of Hansen (1905) and the much later key of Harrison and Ellis (1991) are available. The most recent comprehensive treatment for the United States is Richardson’s (1905) monograph, which to a degree was updated by Kensley and Schotte (1989). The number of described species and genera of North American Sphaeromatidae have slowly increased over the 20th century but many species remain poorly known and attributed to inappropriate genera. At last count, marine and freshwater sphaeromatids in North America included 21 genera with a total of 67 species (seven species inquirenda, incertae sedis or both).

The North American western coast lies within the East Pacific biogeographic zone, and the Sphaeromatidae are represented by 37 species in 11 genera, six of these regarded as species inquirenda and incertae sedis (see Appendix 1). While some western coast United States species have been described in detail (e.g. Bruce and Wetzer 2004; Carvacho and Haasmann 1984; Espinosa-Pérez and Hendrickx 2002; Hendrickx and Espinosa-Pérez 1998; Wetzer and Bruce 2007) many others remain poorly described, and unrecognizable by modern standards (see Appendix 1).

One such poorly-known North American species is Exosphaeroma amplicauda (Stimpson, 1857). The original description of Exosphaeroma amplicauda is brief with a single postage-stamp sized (1.5×2.0 cm) figure of the dorsum taken from specimens “found adhering to fragments of star-fishes picked up on the beach of Tomales Bay by Mr. Samuels, 6.4 mm long and deposited at the Smithsonian” (Stimpson 1857). Stimpson (1858) later provided a paragraph-long description without additional details. The species was redescribed by Kussakin (1979) based on material collected from Amchitka Island, Alaska, some 2000 kilometers north of the type locality. Differences between Kussakin’s (1979) description and fresh material of what appeared to be E. amplicauda from California, including the type location, prompted a re-evaluation of the species. Reviewing morphological and molecular data, we realize that there is a ‘species flock’ of five morphologically similar species on the western coast of North America. Such ‘species flocks’ have been reported for other sphaeromatid genera (e.g. Paracassidina – see Bruce 1994; Oxinasphaera – see Bruce 1997) and other families (e.g. Cymothoidae, see Bruce 1986; Cirolanidae, see Bruce 2004; Aegidae, see Bruce 2004, 2009; Serolidae, see Poore 1987), but this is the first such example in the East Pacific.

We redescribe Exosphaeroma amplicauda from the type locality Tomales Bay (central California coast) and E. aphrodita from San Diego, and describe three new closely related species: E. paydenae sp. n., Aleutians; E. russellhansoni sp. n., Puget Sound, E. pentcheffi sp. n., Palos Verdes Peninsula.

Abbreviations

LACM–Natural History Museum of Los Angeles County; USNM–United States National Museum, Smithsonian Institution; BM–British Museum; MCZ–Museum of Comparative Zoology Harvard, ANSP–Academy of Natural Sciences Philadelphia; UAF–University of Alaska, Fairbanks; AM–Amherst College, Massachusetts; PM–Yale Peabody Museum, Connecticut; RS–robust seta/e; PMS–plumose marginal setae; SEM–scanning electron microscopy; SCAMIT–Southern California Association of Marine Invertebrate Taxonomists. Latitudes and longitudes denoted with “~” are approximate and estimated from Google Earth.

Material and methods

Descriptions are based on the male holotype, female allotype, and topotypic paratypes. Specimens examined have been assigned a USNM or LACM catalog/type numbers. Numbers preceded by “RW” are field and station numbers. Species descriptions were prepared using DELTA (Dallwitz et al. 1997). Setal terminology broadly follows Watling (1989).

Specimens prepared for SEM were cleaned for 10–20 seconds in a Branson 1200 ultrasonic cleaner in a weak solution of Branson GP jewelry soap and distilled water. Specimens were then dehydrated with 100% ethanol. Specimens were placed in solutions of pure ethanol and distilled water in the ratios 2:1, 1:1, 1:2, and finally into 100% ethanol (20 minutes per treatment). Once dehydrated and in 100% ethanol, hexamethyldisilzane (HMDS) was used to replace the ethanol in the specimens. Specimens were transferred through ethanol and HMDS solutions in the following ratios 2:1, 1:1, 1:2 and finally into 100% HMDS (20 minutes per treatment). Specimens were transferred from the final 100% HMDS to fresh HMDS and allowed to evaporate overnight. Specimens were mounted on carbon conductive tabs and coated with gold/palladium using an Emitech K550x sputter coater (Quorum Technologies, LTD, Kent, UK) and imaged using a Hitachi S-3000N variable pressure SEM (Hitachi, Troy, MI) at the LACM.

Drawings were made with the aid of a camera lucida and illustrations were electronically “inked” with Adobe Illustrator CS6. Whole body illustrations were made with a Wild M5D stereo dissecting scope. Appendages were illustrated by dissecting off the appendage and placing them in glycerol on a depression slide and then imaged using a Nikon Labophot-2 compound scope.

Specimens were measured by tracing their dorsal surface along their longitudinal axis with the aid of a camera lucida. A scale bar in the same plane as the specimens allowed calculation of total body length. All lengths reported were mesured in this fashion and may slightly overestimate total body length because pereonites and pleonites are expanded in this position. The lengths given in the “Material Examined” are of the largest specimen of each species and sex. Not all specimens were measured. If a length is provided and multiple specimens were present in a lot, the length refers to largest specimen. In all species mature males appear larger than females, but body lengths for mature adults are similar. Males in all species have much broader uropods than females, which contributes to this illusion. Large sexually mature males tend to be rare compared to females and subadults. Gravid females are rare. Smaller non-gravid individuals cannot be sexed. Females of the different species are virtually indistinguishable and cannot be confidently assigned to a species without an accompanying male. It appears that the largest males guard harems. No individual male-female mate guarding was observed (as occurs in Exosphaeroma inornata Dow, 1958 which also occurs on the Pacific west coast). All species described herein occur in aggregates either under rocks or amongst dead barnacle tests.

We provide dorsal and lateral line drawings of all males for each species. We also provide dorsal and lateral SEMs of both males and females of each species.

Taxonomy

Key to the north-eastern Pacific species of Exosphaeroma of the North American West Coast

This key is based on adult ♂ characters. Also note that weak pereon tubercles are visible only with SEM and not necessarily evident with light microscopy – e.g., compare Figures 1 and 21.

1 Pereonites 1–7, pleon, and pleotelson without ornamentation; pleotelson to overall body length ratio 0.21; apex of posterior margin of pleotelson rounded and truncate; uropodal endopods posterior margin evenly rounded; sex ratio nearly 1:1; individual mate guarding Exosphaeroma inornata (Fig. 26)
Pleon with tubercles; pleotelson and uropods long, pleotelson to overall body length ratio 0.30 or greater; posterior margin of pleotelson acuminate; uropodal endopods posterior margin falcate; large adult males rare, one alpha male guarding many females and juveniles (harem guarding) 2
2 Pereonites 5–7 without tubercles; pleon with 1 anterior and 1 posterior weak tubercles on either side of longitudinal axis; pleotelson dorsal surface without ornamentation; appendix masculina straight, distally narrowing, distal apex acute, length 16.0 basal width Exosphaeroma paydenae sp. n. (Figs 5; 8B; 22)
Pereonite 7 with weak or strong median process; pleon with 1 medium tubercle on either side of longitudinal axis; pleotelson dorsal surface with tubercle 3
3 Pereonites 5 and 6 without ornamentation, pereonite 7 with weak median process; pleon with 1 medium tubercle on either side of longitudinal axis; pleotelson dorsal surface with 2 small anterior tubercles; appendix masculina distal end curving mesially, apex weakly hooked mesially, length 11.4 basal width Exosphaeroma russellhansoni sp. n. (Figs 9; 12B; 23)
Pereonite 5 without ornamentation, pereonite 6 with 1 lateral weak tubercle, pereonite 7 with weak median process, and paired weak lateral tubercles; pleon with 1 medium tubercle on either side of longitudinal axis; pleotelson dorsal surface with 1 anterior median strong tubercle and 2 weak medial tubercles; appendix masculina apically narrowly rounded, length 13.0 basal width Exosphaeroma aphrodita (Figs 17; 20B; 25)
Pereonites 5 and 6 with tubercles; pereonite 7 with median process, and tubercles; pleon with 1 posterior strong tubercle, on either side of longitudinal axis; pleotelson dorsal surface with tubercles 4
4 Pereonites 5 and 6 with 1 median weak tubercle, and 1 weak lateral tubercle; pereonite 7 with weak median process and paired lateral tubercles; pleotelson dorsal surface with 2 small anterior tubercles; appendix masculina distal end curving mesially, straightening at distal tip, length 15.4 basal width Exosphaeroma amplicauda (Figs 1; 4B; 21)
Pereonites 5–6 each with 7 longitudinal rows of strong tubercles, pereonite 7 with strong median process with 3 lateral tubercles; pleotelson dorsal surface with 3 strong medial tubercles on either side of the longitudinal axis, with 1 strong medial tubercle between the longitudinal axis and lateral margin, pleotelson covered with numerous, additional, small tubercles; appendix masculina distally narrowing to an acute rounded tip, length 15 basal width Exosphaeroma pentcheffi sp. n. (Figs 13; 16B; 24)

Exosphaeroma Stebbing, 1900

Restricted synonymy: Exosphaeroma Stebbing, 1900: 553. – Bruce 2003: 327.

Type species

Sphaeroma gigas Leach, 1818; by original designation (Stebbing 1900).

Remarks

A diagnosis and comprehensive synonymy was provided by Bruce (2003). The genus occurs in shallow water in all the world oceans and is one of the few sphaeromatid genera extending to southern reaches of the Southern Ocean. Greatest diversity is found in the Southern Hemisphere. The genus has groups of morphologically similar species, including those species close to the type species, and a group of species with a broad rim to the pleotelson ventral margin, while some species have broad uropods and a posteriorly produced pleotelson apex. At present, the relationships between these different species groups remains unassessed.

Exosphaeroma amplicauda (Stimpson, 1857), E. aphrodita and the three new species described herein form a distinct group within the genus Exosphaeroma. This group of species is characterised by a posteriorly produced and somewhat posteriorly depressed pleotelson, with an acute apex, flattened ventrolateral margins, and the posterior margin overriding a shallow exit channel; the uropods are distally wide and the exopod is distally broadly falcate. The dorsum varies from smooth to nodular. Typically mature males of the “amplicauda group” have a large pleotelson and enlarged posterior coxal plates and cannot completely roll up or fold. Some similar species are known from the Southern Hemisphere, including Exosphaeroma alveola Bruce, 2003 (southeastern Australia); E. antikraussi Barnard, 1940, E. kraussi Tattersall, 1913, E. planum Barnard, 1914 and E. varicolor Barnard, 1914 (all South Africa); and E. montis (Hurley & Jansen, 1977) (New Zealand). All other North American Exosphaeroma have an evenly rounded pleotelson, with a narrow ventral margin, and uropods that are not posteriorly wide.

Other Exosphaeroma occurring between Alaska and the Mexican border that are morphologically not closely related to the Pacific west coast species include E. inornata (known from Puget Sound, Washington to central-southern Baja California Norte, Mexico). E. inornata differs from the “amplicauda group” in that E. inornata lacks marked sexual dimorphism. Males mate guard individual females with males clasping and holding females until mating. E. inornata can roll up into perfect balls, and their bodies are unornamented. This distinguishes them clearly from the “amplicauda” clade (E. amplicauda, E. aphrodita, and the three new species described here).

The type specimens of E. rhomburum (USNM 22573) were borrowed and consist of two specimens from Monterey Bay, neither specimen is an adult male. Richardson’s (1899b: 835) original species description only figures the pleotelson, and she did not note whether the description was based on a male or female. We were not able to further evaluate the status of this species.

In collections from the type locality at Tomales Bay in 2009 for E. amplicauda, we found “family groups” with all life stages (gravid and non-gravid females, subadult males, juveniles, and adult males). These family groups consisted of ca. 10–30 individuals, but in which adult, fully mature males are rare, leading us to conclude that males in these species guard harems rather than guard individual females. For every 10 individuals, we found one, sometimes two, large adult males. We found no evidence for multiple male morphs in these collections [e.g., alpha, beta, gamma males in Paracerceis sculpta as described by Shuster (1989)].

Exosphaeroma amplicauda (Stimpson, 1857)

Figures 1, 2, 3, 4, 21, 27, 28

Sphaeroma amplicauda Stimpson, 1857: 510; Richardson 1899a: 835; 1899b: 179; 1900: 222.

Exosphaeroma amplicauda. – Richardson 1905: 288, 289, fig. 301, 302; Gurjanova 1936: 122, fig. 69; Schultz 1969: 131, fig. 190.

Sphaeroma octoncum Richardson 1905: 293.

Exosphaeroma octoncum. – Iverson 1974: 166. – Richardson 1905: 293, fig. 309, 310. – Schotte 2012: online.

Not Exosphaeroma amplicauda. – Gurjanova 1936: 122, fig. 69. – Kussakin 1979: 399, figs 254, 255 [= Exosphaeroma paydenae sp. n.].

Material examined

NEOTYPE (here designated): ♂ (5.1 mm): California, Marin County, Tomales Bay, north end of bay across from Hog Island, boat launch parking lot, 38.201°N, 122.922°W, intertidal, from underside of rocks, fixed and preserved in 95% ethanol, 9 Jan 2009, coll. R. Wetzer & A. Wall. RW09.003.1, LACM CR-2014.1.

Non-type material. 2 ♂ (RW09.003.2, LACM CR-2014.1), 3 ♀ (RW09.003.3, LACM CR-2014.1) [used for SEM], 1 ♂, ~40 ♀ and juveniles: same locality as RW09.003, LACM CR-2014.1. 6 ♂ (8.1 mm), 4 ♀ (7.1 mm), 10 juveniles (RW09.004.1), plus 2 ♂ and 4 ♀ prepared as SEM (RW09.004.2): intertidal, from underside of rocks, “family group”, coll. A. Wall. RW09.004, LACM CR-2014.2. 7 ♂ (5.8 mm), ~20 ♀ (6.8 mm) and juveniles, and 3 ♀ used for SEM: intertidal, from underside of rocks, “family group”, coll. A. Wall. RW09.005, LACM CR-2014.3. 2 ♂ (8.4 mm), ~25 ♀ (7.4 mm), and juveniles: intertidal, from empty Balanus glandula shells, coll. N.D. Pentcheff, RW09.006.1, LACM CR-2014.4. 1 ♂ (7.7 mm), 2 ♀ (6.5 mm), and 2 juveniles: E. side in cove across from Hog Island (Nick’s Cove), ~38.197°N, ~122.935°W, 1 Nov 1971, A.0030, coll. E.W. Iverson & J. Carlton. RW04.020.1, LACM CR-2014.5. California, Monterey County, Monterey Bay, 4 specimens (labeled E. octoncum), all are ♀. Acc. No. 03472, USNM 22574 (part).

Description of male

Body length 1.6 width; pereonites 5–6 each with 1 median weak tubercle, and 1 weak lateral tubercle; pereonite 7 with weak median process and paired lateral tubercles (Figures 1A, B; 21A, D). Pleon with 1 posterior strong tubercle on either side of longitudinal axis (Figures 1A, B; 21A, D). Pleotelson length 0.82 width, dorsal surface with 2 small anterior tubercles; ventrolateral ridge extending posteriorly 0.75 of total length, with long setae (Figures 1A, B; 21A, C, D).

Figure 1.

Exosphaeroma amplicauda male neotype LACM CR-2014.1. A dorsal B lateral.

Antennula peduncle article 1 length 1.7 width, anterior medial margin with 2 palm setae; article 2 length 1.4 width, inferior distal margin with 2 palm setae; article 3 length 3.2 width; flagellum with 9 articles (Figure 2B). Antenna reaching anterior margin of pereonite 2; flagellum with 14 articles (Figure 2A).

Figure 2.

Exosphaeroma amplicauda male neotype LACM CR-2014.1. A left antenna B left antennula, basal article broken C right mandible palp D right mandible E left mandible F left maxillula G left maxilla H left maxilliped.

Left mandible incisor with 3 cusps; lacinia mobilis with 3 cusps; lacinia mobilis spine row comprised of 5 curved, serrate spines (Figures 2E, 27D). Right mandible incisor with 4 cusps; spine row comprised of 6 curved, serrate spines; crushing surfaces strongly ridged (Figures 2D, 27D). Maxillula mesial lobe with 4 circumplumose RS; lateral lobe with 6 long, curved, pectinate RS (Figures 2F; 27E, F, G). Maxilla mesial lobe with 6 plumose RS on gnathal surface; middle lobe with 4 long, curved, pectinate RS; lateral lobe with 4 long, curved, pectinate RS (Figures 2G; 27E, G). Maxilliped endite distal surface with 5 plumose setae, and 2 simple RS; distomesial margin with 1 coupling hook; palp article 2 distal apex with 9 long, simple RS; article 3 distal apex with 12 long, simple RS, lateral distal angle with 1 long, simple RS; article 4 distal apex with 9 long, simple RS, lateral distal angle with 1 long, simple RS; article 5 distal apex with 7 long, simple RS (Figures 2H; 27E, G).

Pereopod 1 (Figure 3A) basis superior margin without palm setae, inferior distal angle with 1 long, simple seta, inferior medial margin setal patch absent; ischium length 2.4 width, superior margin with 4 long, simple setae, inferior distal angle with 1 long, simple seta; merus 0.42 ischium length, superior distal angle with 2 long, simple setae; carpus inferior distal angle with 1 long, simple seta; propodus length 2.5 width, 0.82 ischium length, superior distal angle with 2 long, simple setae, inferior margin with 3 long, simple setae; dactylus length 1.7 width, length 0.33 propodus length, inferior margin covered with scales, distal margin with 4 simple setae (Figure 3A). Pereopod 3 (Figures 3B, 27A) basis superior margin without palm setae, inferior distal angle with 1 long simple seta, inferior proximal margin with setal patch present; ischium length 2.5 width, superior margin with 3 long, simple RS, inferior distal angle with 1 simple RS, and with setal patch absent; merus lobate, length 1.4 width, 0.57 ischium length, superior distal angle with cluster of 4 simple RS, inferior margin covered in setal mat; carpus length 0.71 merus length, 1.2 width, superior margin with 1 long, simple seta on distal angle, inferior margin with setal mat and 1 long, simple seta; propodus weakly curved, length 2.5 width, 2.3 carpus length, superior distal margin with 1 palm seta, inferior margin first 0.67 covered in setal mat; dactylus length 1.3 width, length 0.36 propodus length, inferior margin distal 0.75 covered with scales, distal margin with 3 long, simple setae (Figures 3B, 27A). Pereopod 7 (Figures 3C; 27B, C) basis superior margin with palm setae absent, inferior proximal margin with setal patch absent, inferior distal angle with long, simple setae absent; ischium length 2.9 width, superior margin with 3 long, simple RS; merus lobate, merus length 1.8 width, merus length 0.66 ischium length, superior distal angle with 4 RS, inferior margin with setal mat, inferior distal angle with biserrate setae absent; carpus length 2.5 width, carpus length 1.3 merus length, inferior margin with setal mat, superior distal angle with a cluster of 5 long, biserrate setae, superior distal angle with a cluster of 3 long, simple, RS, distomesial margin with a cluster of 3 long, biserrate setae, inferior distal angle with a cluster of 5 long, biserrate setae, 1 long, simple RS; propodus weakly curved, length 4.0 width, length 1.3 carpus length, inferior margin proximal 0.33 with setal mat, superior distal angle with 2 long, simple setae, inferior margin with 2 long, simple setae, with palm setae absent; dactylus length 1.8 width, dactylus length 0.28 propodus length, margin with scales, distal margin with 3 simple setae (Figures 3C; 27B, C).

Figure 3.

Exosphaeroma amplicauda male neotype LACM CR-2014.1. A left pereopod 1 B left pereopod 3 C left pereopod 7.

Penial process length 2.5 basal width (Figure 21B, C).

Pleopod 1 peduncle length 0.48 width, with a cluster of 3 coupling hooks; endopod mesial margin heavily covered in fine, simple setae; exopod length 1.7 width, ventral surface without fine, simple setae (Figure 4A). Pleopod 2 appendix masculina proximally swollen, distally narrowing, distal end curving mesially, straightening at distal tip, length 15.4 basal width (Figure 4B). Pleopod 3 peduncle with a cluster of 3 coupling hooks, distolateral angle with 1 large, plumose seta (Figure 4C). Pleopod 4 peduncle length 0.48 width, distolateral angle with 1 large, plumose seta; endopod distal apex 1 large, plumose seta; exopod distal margin with 2 plumose setae (Figure 4D). Pleopod 5 exopod proximolateral margin with palm setae absent; exopod with transverse suture entire, endopod with 1 scale patch; exopod with 3 scale patches (Figure 4E). Uropod exopod length 2.3 width; rolled proximolateral margin weakening moving distally; mesial margin without setae; endopod length 2.5 width, extends past exopod, mesial margin without setae (Figures 4F; 21A, C, D).

Figure 4.

Exosphaeroma amplicauda male neotype LACM CR-2014.1. A–E left pleopods 1–5, respectively F right uropod.

Description of female

Body length 2.7 width; pereonites 1–7 without tubercles, pereonite 7 distomesial margin convex (Figure 21E, F). Pleon with 1 posterior tubercle on either side of longitudinal axis (Figure 21E, F). Pleotelson length 2.6 width, dorsal surface with 2 medium tubercles on either side of longitudinal axis; posterior margin of pleotelson acuminate (Figure 21E, F). Uropod exopod proximolateral margin weakly rolled; endopod posterior margin tapering to evenly rounded tip, length 4.8 width, extends past exopod (Figure 21E, F).

Size. Largest ♂ to 8.4 mm, largest ♀ to 7.5 mm.

Color. Without chromatophores: preserved specimen pale buff, whitish.

Remarks

Exosphaeroma amplicauda is most morphologically similar to Exosphaeroma russellhansoni sp. n. but can be distinguished by: pereonites 5 and 6 with one weak median tubercle, and one weak lateral tubercle; pereonite 7 with weak median process and paired lateral tubercles. (Figures 1A, B; 21A, D). Appendix masculina distal end curving mesially, straightening at distal tip, length 15.4 basal width (Figure 4B).

E. russellhansoni sp. n. is characterized by: pereonite 5–6 each without ornamentation, pereonite 7 with weak median process (Figures 9A, B; 23A, D). Appendix masculina distal end curving mesially, apex weakly hooked mesially, length 11.4 basal width (Figure 12B). E. amplicauda is strongly sexually dimorphic; females lack dorsal tubercles on pereonites 1–7. Overall for all species in this ‘species flock’ the males have a larger pleotelson and uropods. Weak pereon tubercles are visible only with SEM and not necessarily evident with light microscopy. Tubercles visible with light microscopy are figured in the line drawings (compare Figures 1 and 21).

We searched all probable museum collections for Stimpson’s type specimens, but to no avail (see Acknowledgements). It is highly likely that the type specimens are lost. The original and subsequent description (Stimpson 1857, 1858) do not allow for definitive identification of the species. There are five morphologically similar species in the northeast Pacific. A neotype is here designated to stabilize the use of the name Exosphaeroma amplicauda (Stimpson, 1857) and conserve Stimpson’s concept for the species.

We borrowed the types of Sphaeroma octonctum Richardson, 1899 (USNM Cat. No. 22574); Richardson (1899) noted that there were five specimens from the type locality, Monterey Bay). We received only four specimens–three had been previously dissected, some with pleopods removed, and only one specimen was entire. None of these specimens are adult males, and these specimens are indistinguishable from female Exosphaeroma amplicauda from Tomales Bay. We place Sphaeroma octonctum into junior synonymy with Exosphaeroma amplicauda.

Distribution

California: Marin, Sonoma, and San Mateo Counties.

Exosphaeroma paydenae sp. n.

Figures 5, 6, 7, 8, 22, 28

Exosphaeroma amplicauda. – Gurjanova 1936: 122, fig. 69. – Kussakin 1979: 399, Figs 254, 255.

Material examined

HOLOTYPE: ♂ (7.8 mm): Alaska, Aleutian Islands, Kiska Harbor, ~52.00°N, ~177.31°E, ca. 1873, beach, low water, USNM 20474, 211(1025), coll. W.H. Dall [Specimen label reads “Alaska, Kyoka Harbor.” per Marilyn Schotte, 15 Nov 2004 USNM 20474 reads “Aleutian Islands, Kiska Harbor” – maybe a transcription error on the label; specimens denoted as USNM 20474 are also possibly collected ca. 1873 similar to USNM 13312.] USNM 1251663.

PARATYPES: Allotype: ♀ (8.6 mm, whole animal figured): same locality as USNM 1251663, USNM 1251664. 1 ♂, 9 ♀, 2 juveniles, plus 1 ♂ and 1 ♀ prepared as SEM: all same locality as USNM 20474. 8 ♂ (8.0 mm), 1 ♂ broken: north coast of Amchitka, ~51.3°N, 179°E, 1 Jan 1873, USNM 13312, 284(1044), coll. W.H. Dall, USNM 1251665.

Description of male

Body length 1.6 width; pereonites 5–7 each without ornamentation (Figures 5A, B; 22A, B). Pleon with 1 anterior weak tubercle on either side of longitudinal axis, 1 posterior weak tubercle on either side of longitudinal axis (Figures 5A, B; 22A). Pleotelson length 0.59 width, dorsal surface without ornamentation; ventrolateral ridge entire, with few setae (Figures 5A, B; 22A, D).

Figure 5.

Exosphaeroma paydenae sp. n., male holotype USNM 20474. A dorsal B lateral.

Antennula peduncle article 1 length 1.5 width, anterior medial margin with palm setae absent; article 2 length 1.1 width, inferior distal margin with palm setae absent; article 3 length 2.6 width; flagellum with 9 articles (Figure 6B). Antenna reaching posterior margin of pereonite 3, peduncle article 1 with fine, simple setae on superior margin; flagellum with 11 articles (Figure 6A).

Figure 6.

Exosphaeroma paydenae sp. n., male holotype USNM 20474. A right antenna B right antennula C right maxilla D right maxillula E right mandible F left mandible G right maxilliped.

Left mandible incisor with 3 cusps; lacinia mobilis with 2 cusps; lacinia mobilis spine row comprised of 6 curved, serrate spines (Figure 6F). Right mandible incisor with 3 cusps; spine row comprised of 7 curved, serrate spines; crushing surfaces strongly ridged (Figure 6E). Maxillula mesial lobe with 4 circumplumose RS, and 2 long, simple setae; lateral lobe with 7 long, curved, pectinate RS, gnathal surface with 1 curved, simple RS (Figure 6D). Maxilla mesial lobe with 1 long, straight RS, and 8 plumose RS on gnathal surface; middle lobe with 5 long, curved, pectinate RS; lateral lobe with 3 long, curved, pectinate RS (Figure 6C). Maxilliped endite distal surface with 8 plumose setae, and 1 simple RS; distomesial margin with 1 coupling hook; palp article 1 with 1 long, simple RS; article 2 distal apex with 12 long, simple RS; article 3 distal apex with 9 long, simple RS, lateral distal angle with 1 long, simple RS; article 4 distal apex with 9 long, simple RS, lateral distal angle with 1 long, simple RS; article 5 distal apex with 9 long, simple RS (Figure 6G).

Pereopod 1 (Figure 7A) basis superior margin without palm setae, inferior distal angle without long, simple setae, inferior medial margin setal patch absent; ischium length 1.9 width, superior margin with 2 long, simple setae, inferior distal angle without long, simple setae; merus 0.58 ischium length, superior distal angle with 3 long, simple setae; carpus inferior distal angle with 2 long, simple setae; propodus length 2.1 width, 0.83 ischium length, superior distal angle with 1 long, simple seta, inferior margin with 3 long, simple setae; dactylus length 1.1 width, length 0.3 propodus length, inferior margin covered with fine scales, distal margin with 2 simple setae (Figure 7A). Pereopod 3 (Figure 7B) basis superior margin without palm setae, inferior proximal margin with setal patch present; ischium length 2.6 width, superior margin with 3 long, simple RS, inferior distal angle with long, simple RS absent, and with setal patch absent; merus weakly lobate, length 1.4 width, 0.53 ischium length, superior distal angle with a cluster of 5 long, simple RS, inferior margin covered in setal mat; carpus length 0.88 merus length, 1.5 width, superior margin with 2 long, simple setae on distal angle, inferior margin with setal mat, and 1 long, simple seta; propodus weakly curved, length 2.9 width, 1.7 carpus length, superior distal margin without palm setae, inferior margin covered in setal mat; dactylus length 1.6 width, length 0.33 propodus length, inferior margin first 0.75 covered with scales, distal margin with 3 long, simple setae (Figure 7B). Pereopod 7 (Figure 7C) basis superior margin with palm setae absent, inferior proximal margin with setal patch, inferior distal angle with long, simple setae absent; ischium length 2.6 width, superior margin with 7 long, simple RS; merus lobate, merus length 1.6 width, merus length 0.56 ischium length, superior distal angle with 9 RS, inferior margin with setal mat, inferior distal angle with 2 biserrate setae; carpus length 1.8 width, carpus length 0.96 merus length, inferior margin with setal mat, superior distal angle with a cluster of 9 long, biserrate setae, superior distal angle with a cluster of 2 long, simple, RS, inferior distal angle with a cluster of 4 long, biserrate setae, inferior distal angle with 1 long, simple RS; propodus weakly curved, length 3.9 width, length 1.8 carpus length, inferior margin first 0.75 with setal mat, inferior distal margin with 2 long, simple setae, and with palm setae absent; dactylus length 2.0 width, dactylus length 0.26 propodus length, inferior margin with fine scales, distal margin with 4 simple setae (Figure 7C).

Figure 7.

Exosphaeroma paydenae sp. n., male holotype USNM 20474. A right pereopod 1 B right pereopod 3 C right pereopod 7.

Penial process length 3.2 basal width (Figure 22C, D).

Pleopod 1 peduncle length 0.46 width, with a cluster of 3 coupling hooks; endopod mesial margin entirely covered with fine, simple setae; exopod length 1.7 width, ventral surface without fine, simple setae (Figure 8A). Pleopod 2 appendix masculina straight, distally narrowing, distal apex acute, length 16.0 basal width (Figure 8B). Pleopod 3 peduncle with a cluster of 3 coupling hooks, distolateral angle with 2 long, simple setae (Figure 8C). Pleopod 4 peduncle distolateral angle with 1 long, palm seta; endopod distal apex without plumose setae; exopod distal margin with 2 simple setae (Figure 8D). Pleopod 5 exopod proximolateral margin with palm setae absent; exopod with transverse suture starting laterally moving mesially, incomplete; exopod with 2 scale patches (Figure 8E). Uropod exopod length 2.5 width; rolled proximolateral margin weakening moving toward lateral, medial margin; mesial margin without setae; endopod length 2.8 width, extends past exopod, mesial margin without setae (Figures 8F; 22A, B, D).

Figure 8.

Exosphaeroma paydenae sp. n., male holotype USNM 20474. A–E right pleopods 1–5, respectively F right uropod.

Description of female

Body length 2.2 width; pereonites 1–7 without tubercles, pereonite 7 distomesial margin weakly convex (Figure 22E, F). Pleon with 1 posterior weak tubercle on either side of longitudinal axis (Figure 22E, F). Pleotelson length 1.8 width, dorsal surface with 2 tubercles on either side of longitudinal axis; posterior margin of pleotelson acuminate (Figure 22E, F). Uropod exopod proximolateral margin rolled weakly; endopod posterior margin tapering to an evenly rounded tip, length 2.9 width, extends past exopod (Figure 22E, F).

Size

Largest ♂ 8.0 mm, largest ♀ 8.6 mm.

Color

Without chromatophores. Preserved specimen pale cream.

Remarks

Exosphaeroma paydenae sp. n., unlike other Exosphaeroma sp. in this ‘species flock’, lacks strong sexual dimorphism. Males have overall larger pleotelson and uropods than females. E. paydenae sp. n. is morphologically most similar to Exosphaeroma russellhansoni sp. n. E. paydenae sp. n. can be identified by: pereonites 1–7 without tubercles; pleon with one anterior weak tubercle on either side of longitudinal axis, one posterior weak tubercle on either side of longitudinal axis; pleotelson dorsal surface without ornamentation (Figures 5A, B; 22A, B).

Exosphaeroma russellhansoni sp. n., in contrast to E. paydenae has only one weak tubercle on either side of longitudinal axis of its pleon; pleotelson dorsum, with 2 small anterior tubercles (Figures 9A, B; 23A, D). Weak pereon tubercles are visible only with SEM and not necessarily evident with light microscopy. Tubercles visible with light microscopy are figured in the line drawings (compare Figures 5 and 22).

Kussakin (1979) provided new figures for what he considered to be specimens of E. amplicauda from Alaska. In his description he wrote “one sample (three specimens) from Alaska was examined from the collections of the Zoological Institute, Academy of Sciences of the USSR.” We here recognize the Alaska specimens as E. paydenae sp. n., which does not overlap in occurrence with species from further south, all described herein.

Distribution

Alaska, Aleutians.

Etymology

This species is named to honor LACM Trustee and long supporter of science at the Natural History Museum of Los Angeles County, Joan Payden. She is thanked for her gracious philanthropy which in part supported ARW as an undergraduate student researcher. ARW’s research experience describing and redescribing the Exosphaeroma along our coast piqued his interest in marine isopods and launched his career in Crustacea at the LACM.

Exosphaeroma russellhansoni sp. n.

Figures 9, 10, 11, 12, 23, 28

Material examined

HOLOTYPE: ♂ (7.0 mm): Washington, Puget Sound, Seattle, Puget Sound Naval Supply Depot, Smith Cove, ~47.5°N, ~122.2°W, under rocks in sand, 11 Aug 1973. A.0030, coll. E.W. Iverson. RW04.010.1, LACM CR-2014.6.

PARATYPES: Allotype gravid ♀ (6.7 mm): same data as holotype, LACM CR-2014.6. 1 ♂ dissected, appendages figured (RW04.010.3), 1 ♂ (RW04.010.4, LACM CR-2014.6.4) and 2 ♀ (6.7 mm RW04.010.5, LACM CR-2014.6.5) prepared as SEM, plus ~70 additional specimens (all life stages RW04.010.6): same locality as RW04.010, LACM CR-2014.6. 3 ♀ (7.2 mm), 1 subadult ♂: south end of San Juan Island, Cattle Point, 48.451°N, 122.967°W, rocky intertidal barnacles from Semibalanus cariosus, fixed and preserved in 95% ethanol, 7 Apr 2004, coll. R. Wetzer & N.D. Pentcheff. RW04.036.1, LACM CR-2014.7. 1 ♂ (7.9 mm), 1 subadult intermolt: northeast of San Juan Island, Reuben Tarte County Park, 48.612°N, 123.098°W, scrapings of vertical rock surface in intertidal, fixed and preserved in 95% ethanol, 9 Apr 2004, coll. R. Wetzer & N.D. Pentcheff. RW04.041.1, LACM CR-2014.8. 6 ♂ (8.3 mm), 4 ♀ (8.7 mm), 2 ♂ dissected for mandibles and figured: San Juan Island, old man’s farm, ~48.6°N, ~122.9°W, under rocks, 30 Jul 1950. USNM Acc. No. 187867, coll. L. Peternick and P. Illg. USNM 1251666. 3 ♂, 1 gravid ♀, 1 subadult: San Juan Islands, False Bay, 1 Aug 1975, transferred to 95% ethanol 5 Oct 2012, coll. R.R. Hessler. RW12.215.1, LACM CR-2014.9.

Description of male

Body length 1.6 width; pereonite 5–6 each without ornamentation, pereonite 7 with weak median process (Figures 9A, B; 23A, D). Pleon with 1 medium tubercle on either side of longitudinal axis (Figures 9A, B; 23A, D). Pleotelson length 0.65 width, dorsal surface with 2 small anterior tubercles; ventrolateral ridge entire, with long setae (Figures 9A, B; 23A, C, D).

Figure 9.

Exosphaeroma russellhansoni sp. n. male holotype LACM CR-2014.6. A dorsal B lateral.

Antennula peduncle article 1 length 1.2 width, anterior medial margin with 1 palm seta; article 2 length 1.2 width, inferior distal margin with 3 palm setae; article 3 length 2.9 width; flagellum with 9 articles (Figure 10B). Antenna reaching medium margin of pereonite 2, peduncle article 1 with numerous fine simple setae on anterior posterior margin; flagellum with 13 articles (Figure 10A).

Figure 10.

Exosphaeroma russellhansoni sp. n., male paratype LACM CR-2014.6. Male paratype RW04.010.3 A right antenna B right antennula, basal article broken C left maxilliped D left maxilla E male paratype left mandible F right maxillula.

Left mandible incisor with 3 cusps; lacinia mobilis with 2 cusps; lacinia mobilis spine row comprised of 6 curved, serrate spines; crushing surfaces strongly ridged, with 2 serrate spines (Figure 10E). Maxillula mesial lobe with 4 circumplumose RS; lateral lobe with 9 long, curved, pectinate RS (Figure 10F). Maxilla mesial lobe with 6 plumose RS on gnathal surface; middle lobe with 4 long, curved, pectinate RS; lateral lobe with 4 long, curved, pectinate RS (Figure 10D). Maxilliped endite distal surface with 11 plumose setae; distomesial margin 2 coupling hooks, and 2 large stout plumose setae, and 1 large simple RS; palp article 2 distal apex with 15 long, simple RS; article 3 distal apex with 17 long, simple RS, lateral distal angle with 1 long, simple RS; article 4 distal apex with 8 long, simple RS, lateral distal angle with 1 long, simple RS; article 5 distal apex with 9 long, simple RS (Figure 10C).

Pereopod 1 (Figure 11A) basis superior margin with 1 palm seta, inferior distal angle with 1 long, simple seta, inferior medial margin setal patch present; ischium length 1.9 width, superior margin with 3 long, simple setae, inferior distal angle with 1 long, simple seta; merus 0.45 ischium length, superior distal angle with 3 long, simple setae; carpus inferior distal angle with 1 long, simple seta; propodus length 2.3 width, 0.93 ischium length, superior distal angle without long, simple setae, inferior margin with 1 long, simple seta; dactylus length 1.5 width, length 0.46 propodus length, inferior margin covered with scales, distal margin with 3 simple setae (Figure 11A). Pereopod 3 (Figure 11B) basis superior margin with 1 palm seta, inferior distal angle with 1 long simple seta, inferior proximal margin with setal patch absent; ischium length 2.4 width, superior margin with 5 long, simple RS, inferior distal angle with 1 simple RS, and with setal patch absent; merus lobate, length 0.95 width, 0.4 ischium length, superior distal angle with a cluster of 5 simple RS, inferior margin covered in setal mat; carpus length 1.3 merus length, 1.4 width, superior margin with 1 long, simple seta on distal angle, inferior margin with setal mat, and long, simple setae absent; propodus weakly curved, length 2.6 width, 1.7 carpus length, superior distal margin with 1 palm seta, inferior margin first 0.67 covered in setal mat; dactylus length 1.2 width, length 0.33 propodus length, inferior margin proximal half with scales, distal margin with 3 long, simple setae (Figure 11B). Pereopod 7 (Figure 11C) basis superior margin with palm setae absent, inferior proximal margin with setal patch absent, inferior distal angle with 1 long, simple seta present; ischium length 3.0 width, superior margin with 10 long, simple RS; merus lobate, merus length 1.7 width, merus length 0.53 ischium length, superior distal angle with 9 RS, inferior margin with setal mat, inferior distal angle with biserrate setae absent; carpus length 2.5 width, carpus length 1.3 merus length, inferior margin with setal mat, superior distal angle with a cluster of 7 long, biserrate setae, superior distal angle with a cluster of 3 long, simple, RS, inferior distal angle with a cluster of 5 long, biserrate setae, inferior distal angle with 1 long, simple RS; propodus weakly curved, length 4.0 width, length 1.2 carpus length, inferior margin with setal mat absent, superior distal angle with 2 long, simple setae, inferior distal margin with 3 long, simple setae, and with palm setae absent; dactylus length 1.6 width, dactylus length 0.24 propodus length, inferior margin with scales, distal margin with 2 simple setae (Figure 11C).

Figure 11.

Exosphaeroma russellhansoni sp. n., male holotype LACM CR-2014.6. A left pereopod 1 B left pereopod 3 C left pereopod 7.

Penial process length 2.7 basal width (Figure 23B).

Pleopod 1 peduncle length 0.41 width, with a cluster of 4 coupling hooks; endopod mesial margin lightly covered in fine, simple setae; exopod length 1.8 width, ventral surface without fine, simple setae (Figure 12A). Pleopod 2 appendix masculina proximally swollen, distally narrowing, distal end curving mesially, apex weakly hooked mesially, length 11.4 basal width (Figure 12B). Pleopod 3 peduncle with a cluster of 3 coupling hooks, distolateral angle with 2 long, simple setae (Figure 12C). Pleopod 4 peduncle distolateral angle with fine setal patch; endopod distal apex 1 large, plumose seta; exopod distal margin without setae (Figure 12D). Pleopod 5 exopod proximolateral margin with palm setae absent; exopod with distal transverse suture starting laterally, incomplete; exopod with 4 scale patches (Figure 12E). Uropod exopod length 2.4 width; rolled proximolateral margin weakening moving toward lateral, medial margin; mesial margin without setae; endopod length 2.7 width, extends past exopod, mesial proximal margin with setal patch (Figures 12F; 23A, C, D).

Figure 12.

Exosphaeroma russellhansoni sp. n., male paratype LACM CR-2014.6. A–E left pleopods 1–5, respectively F left uropod.

Description of female

Body length 2.7 width; pereonites 1–7 without tubercles, pereonite 7 distomesial margin convex (Figure 23E, F). Pleon with 1 weak posterior tubercle on either side of longitudinal axis (Figure 23E, F). Pleotelson length 0.75 width, dorsum without tubercles; posterior margin of pleotelson acuminate (Figure 23E, F). Uropod exopod proximolateral margin rolled weakly; endopod length 3.7 width, extends past exopod (Figure 23E, F).

Size

Largest ♂ 8.3 mm, largest ♀ 8.7 mm.

Color

Without chromatophores. Preserved specimen pale buff, whitish.

Remarks

E. russellhansoni sp. n. is morphologically most similar to E. amplicauda but can be easily distinguished by: pereonite 5–6 each without ornamentation, pereonite 7 with weak median process (Figures 9A, B; 23A, D). Appendix masculina distal end curving mesially, apex weakly hooked mesially, length 11.4 basal width (Figure 12B).

E. amplicauda is distinguished by: pereonites 5 and 6 with one weak median tubercle, and one weak lateral tubercle; pereonite 7 with weak median process and paired lateral tubercles (Figures 1A, B; 21A, D). Appendix masculina distal end curving mesially, straightening at distal tip, length 15.4 basal width (Figure 4B). E. russellhansoni sp. n. is strongly sexually dimorphic; females lacking dorsal tubercles on pereonites 1–7. Weak pereon tubercles are visible only with SEM, but not necessarily evident with light microscopy, and therefore are omitted from line drawings (compare Figures 9 and 23).

Distribution

Washington, Puget Sound and San Juan Island.

Etymology

Named to honor Russell Kenneth Hanson, ARW’s only maternal uncle who has shaped the person Adam is today by so graciously sharing with Adam his insatiable curiosity, life-long pursuit of perfection and tireless work ethic.

Exosphaeroma pentcheffi sp. n.

Figures 13, 14, 15, 16, 24, 28

Material examined

HOLOTYPE ♂ (4.6 mm): California, Los Angeles County, Palos Verdes Peninsula, Pt. Fermin, shore at Paseo del Mar, ~0.5 mi. W of Gaffey Street, 33.71°N, 118.3°W, mid-low intertidal, chipping overhanging rock with hammer and Phragmatopoma tubes on underside of rock, 0.99 m depth, fixed and preserved in 95% ethanol, 27 Mar 2004, coll. R. Wetzer, N.D. Pentcheff, and LMU students. RW04.030.1, LACM CR-2014.10.

PARATYPES: Allotype ♀ (4.6 mm) (whole animal figured): shore at Paseo del Mar, ~0.5 mi. W of Gaffey Street, 33.71°N, 118.3°W, mostly barnacles, some algal turf, medium to high intertidal, paint scraper, fixed and preserved in 95% ethanol, 16 Feb 2004, coll. R. Wetzer. RW04.002.1, LACM CR-2014.11. 1 ♂ accidently destroyed after being imaged (RW04.255, LACM CR-2014.12), 1 ♂, 3 ♀ (RW04.030.2), plus 1 ♀ (4.6 mm) prepared as SEM: RW04.030.3, LACM CR-2014.10. 3♀(RW04.002.2), plus 1 ♂ (RW04.002.3) and 1 ♀ (RW04.002.4) prepared for SEM: shore at Paseo del Mar, ~0.5 mi. W of Gaffey Street, 33.71°N, 118.3°W, mostly barnacles, some algal turf, medium to high intertidal, paint scrapper, fixed and preserved in 95% ethanol, 16 Feb 2004, coll. R. Wetzer. RW04.002, LACM CR-2014.11. 1 ♂ (5.1 mm), 2 ♀: shore at Paseo del Mar, ~0.5 mi. W of Gaffey Street, 33.71°N, 118.3°W, found in bottom of bucket with sea stars, mid- to low intertidal, fixed and preserved in 95% ethanol, 16 Feb 2004. Loyola Marymount University Invertebrate Class, N.D. Pentcheff, coll. E. Pattison and K. Stanley. RW04.003.1, LACM CR-2014.13.

Description of male

Body length 1.8 width; pereonites 5–6 each with 7 longitudinal rows of strong tubercles, pereonite 7 with strong median process with 3 lateral tubercles (Figures 13A, B; 24A, D). Pleon with 1 medium tubercle on posterior margin, on either side of longitudinal axis (Figures 13A, B; 24A, D). Pleotelson length 0.85 width, dorsal surface with 3 strong medial tubercles on either side of the longitudinal axis, with 1 strong medial tubercle between the longitudinal axis and lateral margin, pleotelson covered with numerous, additional, small tubercles; ventrolateral ridge extending posteriorly 0.80 of total length, with long setae (Figures 13A, B; 24A, C, D).

Figure 13.

Exosphaeroma pentcheffi sp. n., male holotype LACM CR-2014.10. A lateral B dorsal.

Antennula peduncle article 1 length 1.4 width, anterior medial margin with 2 palm setae; article 2 length 1.1 width, inferior distal margin with 3 palm setae; article 3 length 3.1 width; flagellum with 9 articles (Figure 14B). Antenna reaching anterior margin of pereonite 2, peduncle article 1 with fine, simple setae on superior margin; flagellum with 11 articles (Figure 14A).

Figure 14.

Exosphaeroma pentcheffi sp. n., male holotype LACM CR-2014.10. A left antenna B left antennula C left mandible D right mandible E left maxilliped F left maxilla G left maxillula.

Left mandible incisor with 4 cusps; lacinia mobilis with 3 cusps; lacinia mobilis spine row comprised of 8 curved, serrate spines, and 1 curved, robust, simple spine (Figure 14C). Right mandible incisor with 3 cusps; spine row comprised of 7 curved, serrate spines; crushing surfaces strongly ridged, with 1 serrate spine (Figure 14D). Maxillula mesial lobe with 4 circumplumose RS; lateral lobe with 10 long, curved, pectinate RS, gnathal surface with 1 curved, simple RS (Figure 14G). Maxilla mesial lobe with 1 long, straight RS, and 8 plumose RS on gnathal surface; middle lobe with 8 long, curved, pectinate RS; lateral lobe with 5 long, curved, pectinate RS (Figure 14F). Maxilliped endite distal surface with 7 plumose setae, and 3 simple RS; distomesial margin with 1 coupling hook, and 3 large stout plumose setae; palp article 2 distal apex with 6 long, simple RS; article 3 distal apex with 8 long, simple RS, lateral distal angle with 1 long, simple RS; article 4 distal apex with 7 long, simple RS; article 5 distal apex with 7 long, simple RS (Figure 14E).

Pereopod 1 (Figure 15A) basis superior margin without palm setae, inferior distal angle with 1 long, simple seta, inferior medial margin setal patch absent; ischium length 2.3 width, superior margin with 3 long, simple setae, inferior distal angle without long, simple setae; merus 0.50 ischium length, superior distal angle with 3 long, simple setae; carpus inferior distal angle with 1 long, simple seta; propodus length 2.7 width, 1.0 ischium length, superior distal angle without long, simple setae, inferior margin with 1 long, simple seta; dactylus length 1.4 width, length 0.36 propodus length, inferior margin without setal scales, distal margin with 4 simple setae (Figure 15A). Pereopod 3 (Figure 15B) basis superior margin with 1 palm seta, inferior distal angle with 2 long simple setae, inferior proximal margin with setal patch present; ischium length 3.1 width, superior margin with 6 long, simple RS, inferior distal angle with 2 long, simple RS, and with setal patch present; merus lobate, length 2.2 width, 0.73 ischium length, superior distal angle with 4 long, simple RS, inferior margin covered in setal mat; carpus superior margin with 1 long, simple seta on distal angle, inferior margin with setal mat, and 3 long, simple setae; propodus weakly curved, length 3.2 width, 1.8 carpus length, superior distal margin with 1 palm seta, inferior margin covered in setal mat; dactylus length 1.6 width, length 0.30 propodus length, inferior margin without scales, distal margin with 4 long, simple setae (Figure 15B). Pereopod 7 (Figure 15C) basis superior margin with 1 palm seta, inferior proximal margin with setal patch, inferior distal angle with long, simple setae absent; ischium length 2.8 width, superior margin with 2 long, simple RS; merus lobate, merus length 2.1 width, merus length 0.63 ischium length, superior distal angle with 4 RS, inferior margin with setal mat, inferior distal angle with biserrate setae absent; carpus length 2.3 width, carpus length 1.1 merus length, inferior margin with setal mat, superior distal angle with a cluster of 8 long, biserrate setae, superior distal angle with a cluster of 2 long, simple, RS, inferior distal angle with a cluster of 3 long, biserrate setae; propodus weakly, curved, length 5.0 width, length 1.6 carpus length, inferior margin first 0.33 with setal mat, superior distal angle with 2 long, simple setae, inferior margin with 2 long, simple setae, and with palm setae absent; dactylus length 1.7 width, dactylus length 0.22 propodus length, inferior margin without fine scales, distal margin with 3 long, simple setae (Figure 15C).

Figure 15.

Exosphaeroma pentcheffi sp. n., male holotype LACM CR-2014.10. A left pereopod 1 B left pereopod 3 C left pereopod 7.

Penial process length 3.0 basal width (Figure 24B).

Pleopod 1 peduncle length 0.56 width, with a cluster of 3 coupling hooks; endopod mesial margin covered in fine, simple setae; exopod length 1.7 width, ventral surface with fine, simple setae (Figure 16A). Pleopod 2 appendix masculina distally narrowing to an acute rounded tip, length 15 basal width (Figure 16B). Pleopod 3 peduncle with a cluster of 3 coupling hooks, distolateral angle with 2 large, simple setae (Figure 16C). Pleopod 4 peduncle length 0.46 width, distolateral angle with 1 large, simple seta; endopod distal apex without plumose setae; exopod distal margin with 2 simple setae (Figure 16D). Pleopod 5 exopod proximolateral margin with palm setae absent; exopod with transverse suture starting laterally moving mesially, incomplete; exopod with 4 scale patches (Figure 16E). Uropod exopod length 2.4 width; rolled proximolateral margin weakening moving toward lateral, distal margin; mesial margin with evenly spaced fine simple setae; endopod length 2.8 width, extends past exopod, distal apex with short, simple setal patch, dorsal surface covered with numerous small tubercles, mesial margin with evenly spaced fine simple setae (Figures 16F; 24A, C, D).

Figure 16.

Exosphaeroma pentcheffi sp. n., male holotype LACM CR-2014.10. A–E left pleopods 1–5, respectively F left uropod.

Description of female

Body length 2.3 width; pereonites 2–6 each with 7 longitudinal rows of strong tubercles, pereonite 7 distomesial margin convex with strong median process, and 3 lateral tubercles (Figure 24E, F). Pleon with 1 posterior strong tubercle on either side of longitudinal axis (Figure 24E, F). Pleotelson length 0.61 width, dorsal surface with 3 strong medial tubercles on either side of the longitudinal axis, with 1 strong medial tubercle between the longitudinal axis and lateral margin, pleotelson covered with numerous, additional, small tubercles (Figure 24E, F). Uropod exopod proximolateral margin rolled; endopod length 3.6 width, extends past exopod, dorsal surface covered with numerous small tubercles, mesial margin without setae (Figure 24E, F).

Size

Largest ♂ 6.8 mm, largest ♀ 4.6 mm.

Colour

No chromatophores: preserved specimen pale buff, whitish.

Remarks

Exosphaeroma pentcheffi sp. n. unlike the other Exosphaeroma species in this ‘species flock’ lacks strong sexual dimorphism and is unique in that females shares the same dorsal ornamentation as males; males differ from females in having slightly stronger tubercles, longer pleotelson and longer uropods. Females of E. pentcheffi sp. n. are the only females of this ‘species flock’ that can reliably be identified at the species level. E. pentcheffi sp. n. males can be identified by: pereonites 5 and 6 having 7 longitudinal rows of strong tubercles, pereonite 7 with a strong median process with 3 lateral tubercles; pleotelson dorsum with 3 strong medial tubercles on either side of the longitudinal axis, with 1 strong medial tubercle between the longitudinal axis and lateral margin, pleotelson covered with numerous, additional, small tubercles (Figures 13A, B; 24A, D, E, F). Weak pereon tubercles are visible only with SEM and not necessarily evident with light microscopy. Tubercles visible with light microscopy are figured in the line drawings (compare Figures 13 and 24).

Distribution

California, Los Angeles County, Palos Verdes Peninsula.

Etymology

This beautiful species is named for N. Dean Pentcheff, expert isopod collector, superb field and dive buddy, travel companion and IT support par excellence. Dean is commended for his reliable patience, support and solid friendship.

Exosphaeroma aphrodita Boone, 1923

Figures 17, 18, 19, 20, 25, 28

Exosphaeroma aphrodita Boone, 1923. – Bruce 2003: 369. – Espinosa and Hendrickx 2006: 238. – Brusca et al. 2007: 537. – Bruce and Schotte 2012: online.

Material examined

LECTOTYPE, here designated: 1 ♂ USNM 1251667 with mandibles dissected: California, San Diego County, La Jolla, Scripps Institute pier pilings, 6 Nov 1915. USNM Acc. No. 53848, #1045-1-4. Identified as Exosphaeroma amplicauda by P.L. Boone.

PARALECTOTYPES: 5 ♂ (USNM 1251667), with mandibles dissected. USNM 53848.

Non-type Material: 1 ♂: Scripps, ~32.87°N, ~117.26°W, littoral in algae, March 1938, coll. Olga Hartman and Loyola e Silva. USNM 1251668. 1 ♂ (RW01.002.1), 1 ♂, 3 ♀, (RW01.002.2) plus 1 ♂ (RW01.002.3) and 3 ♀ (RW01.002.4) prepared as SEM: Scripps Institute of Oceanography, beneath seaward end of Scripps Pier, ~32.87°N, ~117.26°W, to 8 m, among detritus at base of pilings, water temp. 59 °F, SCUBA, fixed and preserved in 95%, 7 Jan 2001, coll. T. Haney. RW01.002, LACM CR-2014.14. 1 ♂ (broken): San Diego, pilings, 1 Jul 1996. USNM Acc. No. 180084, Sta. No. 256. USNM 1251669.

Description of male

Body length 2.0 width; pereonites 5 without ornamentation, pereonite 6 with 1 lateral weak tubercle, pereonite 7 with weak median process, and paired weak lateral tubercles (Figures 17A, B; 25A, C). Pleon with 1 medium tubercle on either side of longitudinal axis (Figures 17A, B; 25A, C). Pleotelson length 0.89 width, dorsal surface with 1 anterior median strong tubercle and 2 weak medial tubercles; ventrolateral ridge extending posteriorly 0.75 of total length, with long setae (Figures 17A, B; 25A, B, C).

Figure 17.

Exosphaeroma aphrodita male lectotype LACM CR-2014.14. A dorsal B lateral.

Antennula peduncle article 1 length 1.5 width, anterior medial margin with palm setae absent; article 2 length 1.3 width, inferior distal margin with 1 palm seta; article 3 length 2.9 width; flagellum with 8 articles (Figure 18B). Antenna reaching posterior margin of pereonite 3, peduncle article 1 superior margin without palm setae; flagellum with 12 articles (Figure 18A).

Figure 18.

Exosphaeroma aphrodita male lectotype LACM CR-2014.14. A left anntenna B left antennula C left mandible D right mandible E left maxilliped F left maxillula G left maxilla.

Left mandible incisor with 2 cusps; lacinia mobilis with 2 cusps; lacinia mobilis spine row comprised of 6 curved, serrate spines (Figure 18C). Right mandible incisor with 4 cusps; spine row comprised of 5 curved, serrate spines; crushing surfaces strongly ridged (Figure 18D). Maxillula mesial lobe with 5 circumplumose RS; lateral lobe with 6 long, curved, pectinate RS (Figure 18F). Maxilla mesial lobe with 3 long, curved RS, and 6 plumose RS on gnathal surface; middle lobe with 8 long, curved, pectinate RS; and 7 long, curved RS; lateral lobe with 3 long, curved, pectinate RS (Figure 18G). Maxilliped endite distal surface with 2 plumose setae; distomesial margin with 1 coupling hook; palp article 2 distal apex with 1 long, simple RS; article 3 distal apex with 3 long, simple RS, lateral distal angle with 1 long, simple RS; article 4 distal apex with 3 long, simple RS, lateral distal angle with 1 long, simple RS; article 5 distal apex with 3 long, simple RS (Figure 18E).

Pereopod 1 (Figure 19A) basis superior margin with 1 palm seta, inferior distal angle with 1 long, simple seta, inferior medial margin setal patch absent; ischium length 2.3 width, superior margin with 3 long, simple setae, inferior distal angle without long, simple setae; merus 0.40 ischium length, superior distal angle with 3 long, simple setae; carpus inferior distal angle with 1 long, simple seta; propodus length 2.1 width, 0.60 ischium length, superior distal angle with 1 long, simple seta, inferior margin with 3 long, simple setae; dactylus length 1.5 width, length 0.55 propodus length, inferior margin distal 0.67 covered with scales, distal margin with 2 simple setae (Figure 19A). Pereopod 3 (Figure 19B) basis superior margin without palm setae, inferior distal angle with 1 long simple seta, inferior proximal margin with setal patch absent; ischium length 3.2 width, superior margin with 3 long, simple RS, inferior distal angle with 1 simple RS, and with setal patch absent; merus lobate, length 1.4 width, 0.56 ischium length, superior distal angle with a cluster of 3 RS, inferior margin covered in setal mat; carpus length 0.90 merus length, 1.9 width, superior margin with 1 long, simple seta on distal angle, inferior margin with setal mat, and 1 long, simple seta; propodus weakly curved, length 3.2 width, 2.0 carpus length, superior distal margin with 1 palm seta, inferior margin first 0.67 covered in setal mat; dactylus length 1.3 width, length 0.45 propodus length, inferior margin with scales, distal margin with 3 long, simple setae (Figure 19B). Pereopod 7 (Figure 19C) basis superior margin with 2 palm setae, inferior proximal margin with setal patch, inferior distal angle with long, simple setae absent; ischium length 3.7 width, superior margin with 1 long, simple RS; merus weakly lobate, merus length 2.0 width, merus length 0.65 ischium length, superior distal angle with 2 RS, inferior margin with setal mat, inferior distal angle with biserrate setae absent; carpus length 2.9 width, carpus length 1.1 merus length, inferior margin with setal mat, superior distal angle with a cluster of 6 long, biserrate setae, superior distal angle with a cluster of 2 long, simple, RS, inferior distal angle with a cluster of 5 long, biserrate setae, inferior distal angle with 1 long, simple RS; propodus weakly curved, length 4.2 width, length 1.3 carpus length, inferior margin with setal patch, superior distal angle with 1 long, simple seta, inferior distal margin with simple, setae absent, and with 1 palm seta; dactylus length 2.0 width, dactylus length 0.27 propodus length, inferior margin with scales starting medially moving distally, distal margin with 2 simple setae (Figure 19C).

Figure 19.

Exosphaeroma aphrodita male lectotype LACM CR-2014.14. A left pereopod 1 B left pereopod 3 C left pereopod 7.

Penial process length 3.1 basal width (Figure 25B, D).

Pleopod 1 peduncle length 0.42 width, with a cluster of 4 coupling hooks; endopod mesial margin lightly covered in fine, simple setae; exopod length 1.6 width, ventral surface without fine, simple setae (Figure 20A). Pleopod 2 appendix masculina apically narrowly rounded, length 13.0 basal width (Figure 20B). Pleopod 3 peduncle with a cluster of 3 coupling hooks, distolateral angle with 3 large, simple setae (Figure 20C). Pleopod 4 peduncle length 0.61 width, distolateral angle with 1 large, simple seta; endopod distal apex 1 large, plumose seta; exopod distal margin with 2 plumose setae (Figure 20D). Pleopod 5 exopod proximolateral margin with 1 palm seta; exopod with transverse suture entire; exopod with 3 scale patches (Figure 20E). Uropod exopod length 2.4 width; rolled proximolateral margin weakening moving toward lateral, medial margin; mesial margin without setae; endopod length 3.1 width, extends past exopod, mesial margin without setae (Figures 20F, G; 25A, B, C).

Figure 20.

Exosphaeroma aphrodita male lectotype LACM CR-2014.14. A–E left pleopods 1–5, respectively F left uropod exopod G left uropod endopod.

Description of female

Body length 2.9 width; pereonites 1–7 without tubercles, pereonite 7 distomesial margin weakly convex (Figure 25E, F). Pleon with 1 weak posterior tubercle on either side of longitudinal axis (Figure 25E, F). Pleotelson length 0.65 width, dorsal surface without visible tubercles; posterior margin of pleotelson acuminate (Figure 25E, F).

Uropod exopod rolled proximolateral margin rolled weakly; endopod length 3.4 width, extends past exopod, mesial margin without setae (Figure 25E, F).

Size

Largest ♂ 8.3 mm, largest ♀ 8.7 mm.

Color

No chromatophores: preserved specimen pale buff, whitish.

Remarks

Exosphaeroma aphrodita can best be identified by: pereonite 5 without ornamentation, pereonite 6 with one lateral weak tubercle, pereonite 7 with weak median process, and paired weak lateral tubercles; pleotelson dorsum with one anterior median strong tubercle and two weak medial tubercles. E. aphrodita is strongly sexually dimorphic; females lack dorsal tubercles on the pereonites. Weak pereon tubercles are visible only with SEM and not necessarily evident with light microscopy. Tubercles visible with light microscopy are figured in the line drawings (compare Figures 17 and 25).

Exosphaeroma aphrodita, considered nomen dubium by Brusca et al. (2007), is here revalidated. Pearl Lee Boone described this species and several other isopods and tanaids without providing figures (Boone 1923, pp. 147–156). The original description states that “the type and additional material were collected at La Jolla, California and are in the collections of the United States National Museum.” We examined all of the USNM material available. We conclude that the species is valid.

Distribution

California, San Diego–La Jolla.

Figure 21.

SEMs of Exosphaeroma amplicauda LACM CR-2014.1.1. A male dorsal B penes C male pleotelson ventral D male lateral E female dorsal F female lateral.

Figure 22.

SEMs of Exosphaeroma paydenae sp. n. paratype USNM 20474. A USNM 20474x male dorsal B male lateral C penes D male pleotelson ventral E USNM 20474xi female lateral F female dorsal.

Figure 23.

SEMs of Exosphaeroma russellhansoni sp. n., paratype. LACM CR-2014.6.4. A male dorsal B penes C male pleotelson ventral D male lateral E LACM CR-2014.6.5 female dorsal F female lateral.

Figure 24.

SEMs of Exosphaeroma pentcheffi sp. n. paratype LACM CR-2014.12. A male dorsal B penes C male pleotelson ventral D male lateral; LACM CR-2014.11 E female dorsal F female lateral.

Figure 25.

SEMs of Exosphaeroma aphrodita LACM CR-2014.14. A male dorsal B male pleotelson ventral C male lateral D penes E female dorsal F female lateral.

Figure 26.

Exosphaeroma inornata male RW05.315, USA, California, Los Angeles County, Palos Verdes Peninsula, San Pedro, Pt. Fermin, shore at Paseo del Mar, ~0.5 mi. W of Gaffey Street, 33.71°N, 118.3°W. A male dorsal B male pleotelson ventral C male lateral D RW05.106, Pacific, Mexico, Baja California Norte, west of El Rosario, south of Bocana el Rosario, north of Punta Baja, 30.013°N, 115.797°W, male mate-guarding female (male uropods were removed).

Figure 27.

SEM images of Exosphaeroma amplicauda LACM CR-2014.1.1. A pereopod 3 dactylus scales B pereopod 7 merus distal setal patch C pereopod 7 carpus distal setal patch D left and right mandibles ventral E left and right maxillipeds, maxillulae, and maxillae dorsal F left maxillula G maxilliped and other mouth parts dorsal.

Figure 28.

SEM images of epistomes. A Exosphaeroma amplicauda LACM CR-2014.1.1 B Exosphaeroma aphrodita LACM CR-2014.14 C Exosphaeroma pentcheffi sp. n. LACM CR-2014.12 D Exosphaeroma russellhansoni sp. n. LACM CR-2014.6.4 E Exosphaeroma paydenae sp. n. USNM 1251663.

Acknowledgements

We thank the many museum staff who so kindly searched all corners of their collections for Stimpson’s specimens–Marilyn Schotte (USNM), Robert VanSyoc and Elizabeth Kools (CalAcad), Paul Callomon (ANSP), Adam Baldinger (MCZ), Paul Clark (BM), Andres Lopez (UAF), Ardis Johnston (MCZ), Eric Lazo-Wasem and Daniel Drew (PM). Vasily Radashevsky and Marina Malyutina (Institute of Marine Biology, Vladivostok) and Victor Petrjashev (Zoological Museum in St.-Petersburg) are thanked for searching for Kussakin’s specimens from Amchitka. Mark Grygier (Lake Biwa Museum, Japan) is thanked for translating Kussakin from Russian. N. Dean Pentcheff contributed to field collections from Washington to Pt. Fermin, and Todd Haney and Ernie Iverson contributed specimens from San Diego and several other locals. LaiShan Mui is thanked for her whole body drawings funded by NSF Systematics grant DEB-0129317, and Giar-Ann Kung is thanked for her help with the operation of the NHM SEM (funded by DBI-0216506). Gary Poore taught ARW the fine art of electronic inking. Phyllis Sun with her digital expertise helped keep drawing progress moving forward, and Jean Pongsai is thanked for her help preparing the final figures. Our gratitude goes to volunteer par excellence, Jim Cline, for his decade long dedication to isopods, help with sample sorting, and his invaluable contributions to the isopod image gallery (http://isopods.nhm.org/images) which these authors and colleagues around the world have come to rely on.

References

  • Baker WH (1926) Species of the isopod family Sphaeromidae, from the eastern, southern, and western coasts of Australia. Transactions of the Royal Society of South Australia 50: 247–279.
  • Barnard KH (1914) Contributions to the crustacean fauna of South Africa. 1. Additions to the marine Isopoda. Annals of the South African Museum 10(7): 197–230.
  • Barnard KH (1940) Contribution to the crustacean fauna of South Africa. XII. Further additions to the Tanaidacea, Isopoda, and Amphipoda, together with keys for the identification of hitherto recorded marine and freshwater species. Annals of the South African Museum 32: 381–543.
  • Boone PL (1923) New marine tanaid and isopod Crustacea from California. Proceedings of the Biological Society of Washington 36: 147–156.
  • Branch ML, Griffiths CL, Kensley B, Sieg J (1991) The benthic Crustacea of subantarctic Marion and Prince Edward Islands: Illustrated keys to the species and results of the 1982–1989 University of Cape Town Surveys. South African Journal of Antarctic Research 21: 3–44.
  • Brandt A, Wägele J-W (1989) Redescriptions of Cymodocella tubicauda Pfeffer, 1887 and Exosphaeroma gigas (Leach, 1818) (Crustacea, Isopoda, Sphaeromatidae). Antarctic Science 1(3): 205–214. doi: 10.1017/S0954102089000325
  • Bruce NL (1986) Revision of the isopod crustacean genus Mothocya Costa, in Hope, 1851 (Cymothoidae: Flabellifera), parasitic on marine fishes. Journal of Natural History 20: 1089–1192. doi: 10.1080/00222938600770781
  • Bruce NL (1994) The Cassidininae Hansen, 1905 (Crustacea: Isopoda: Sphaeromatidae) of Australia. Journal of Natural History 28: 1077–1173. doi: 10.1080/00222939400770571
  • Bruce NL (1997) A new genus of marine isopod (Crustacea: Flabellifera: Sphaeromatidae) from Australia and the Indo-Pacific region. Memoirs of Museum Victoria 56: 145–234.
  • Bruce NL (2003) New genera and species of sphaeromatid isopod crustaceans from Australian marine coastal waters. Memoirs of Museum Victoria 60: 309–369.
  • Bruce NL (2004) Reassessment of the isopod crustacean Aega deshaysiana (Milne Edwards, 1840) (Cymothoidea: Aegidae) – a worldwide complex of 21 species. Zoological Journal of the Linnean Society 142: 135–232.
  • Bruce NL (2004) New Zealand’s cirolanid isopods – highlighting the diversity of these marine garbage cleaners. Aquatic Biodiversity and Biosecurity 7: 10–11. doi: 10.1111/j.1096-3642.2004.00127.x
  • Bruce NL (2009) The marine fauna of New Zealand: Isopoda, Aegidae (Crustacea). NIWA Biodiversity Memoir 122: 1–252.
  • Bruce NL, Schotte M (2012) Exosphaeroma aphrodita Boone, 1923. In: Schotte M, Boyko CB, Bruce NL, Poore GCB, Taiti S, Wilson GDF (Eds) World Marine, Freshwater and Terrestrial Isopod Crustaceans database. Accessed through: World Register of Marine Species. http://www.marinespecies.org/aphia.php?p=taxdetails&id=257017 [on 2012-10-30]
  • Bruce NL, Schotte M (2013) Sphaeromatidae. In: Schotte M, Boyko CB, Bruce NL, Poore GCB, Taiti S, Wilson GDF (Eds) World Marine, Freshwater and Terrestrial Isopod Crustaceans database. Accessed through: World Register of Marine Species. http://www.marinespecies.org/aphia.php?p=taxdetails&id=118277 [on 2013-05-17]
  • Bruce NL, Wetzer R (2004) Paradella tiffany sp. n., a distinctive sphaeromatid isopod (Crustacea: Isopoda: Sphaeromatidae) from Baja California, Mexico. Zootaxa 623: 1–12.
  • Bruce NL, Wetzer R (2008) New Zealand exports: Pseudosphaeroma Chilton, 1909 (Isopoda; Sphaeromatidae), a Southern Hemisphere genus introduced to the Pacific coast of North America. Zootaxa 1908: 51–56.
  • Brusca RC, Coelho VR, Taiti S (2007) Isopoda. In: Carlton JT (Ed.) The Light and Smith Manual: Intertidal Invertebrates From Central California to Oregon, 4th Edition. University of California Press, Berkeley, California, 503–542.
  • Carlton JT, Iverson EW (1981) Biogeography and natural history of Sphaeroma walkeri (Crustacea: Isopoda) and its introduction to San Diego Bay, California. Journal of Natural History 15: 31–48. doi: 10.1080/00222938100770031
  • Carvacho A, Haasmann Y (1984) Isopodos litorales de Oaxaca, Pacifico Mexicano. Cahiers de Biologie marine 25: 15–32.
  • Dallwitz MJ, Paine TA, Zurcher EJ (1997) Users Guide to the DELTA System: A general system for processing taxonomic descriptions. 4.08. CSIRO Division of Entomology, Canberra.
  • Dow TG (1958) Description of a new isopod from California, Exosphaeroma inornata. Bulletin of the Southern California Academy of Sciences 57: 93–97.
  • Espinosa-Pérez MC, Hendrickx ME (2001) A new species of Exosphaeroma Stebbing (Crustacea: Isopoda: Sphaeromatidae) from the Pacific coast of Mexico. Proceedings of the Biological Society of Washington 114: 640–648.
  • Espinosa-Pérez MC, Hendrickx ME (2002) The genus Paracerceis Hansen, 1905 (Isopoda: Sphaeromatidae) in the eastern tropical Pacific, with the description of a new species. Crustaceana 74: 1169–1187. doi: 10.1163/15685400152885165
  • Espinosa-Pérez MC, Hendrickx ME (2006) A comparative analysis of biodiversity and distribution of shallow-water marine isopods (Crustacea: Isopoda) from polar and temperate water in the East Pacific. Belgium Journal of Zoology 136: 219–247.
  • George RY, Strömberg JO (1968) Some new species and new records of marine isopods from San Juan Archipelago, Washington, USA. Crustaceana 14: 225–254. doi: 10.1163/156854068X00827
  • Glynn PW (1968) A new genus and two new species of sphaeromatid isopods from the high intertidal zone at Naos Island, Panama. Proceedings of the Biological Society of Washington 81: 587–604.
  • Gurjanova EF (1936) Beiträge zur Kenntnis der Isopodenfauna des Pazifischen Ozeans. IV. Neue Isopodenarten aus dem Japanischen und Bering-meer. Zoologischer Anzeiger 114: 250–265.
  • Hale HM (1929) The Crustaceans of South Australia. Part 2. Handbooks of the Flora and Fauna of South Australia, issued by the British Science Guild (South Australia Branch). British Science Guild (South Australian Branch), Adelaide, 201–380.
  • Hansen HJ (1905) On the propagation, structure and classification of the family Sphaeromatidae. Quarterly Journal of Microscopical Science 49: 69–135.
  • Harrison K (1984) Some sphaeromatid isopods (Crustacea) from southern and south-western Australia, with description of a new genus and two new species. Records of the Western Australian Museum 11: 259–286.
  • Harrison K, Ellis JP (1991) The genera of the Sphaeromatidae (Crustacea: Isopoda): a key and distribution list. Invertebrate Taxonomy 5: 915–952. doi: 10.1071/IT9910915
  • Harrison K, Holdich DM (1982) New eubrachiate sphaeromatid isopods from Queensland waters. Memoirs of the Queensland Museum 20: 421–446.
  • Harrison K, Holdich DM (1984) Hemibranchiate sphaeromatids (Crustacea: Isopoda) from Queensland, Australia, with a world-wide review of the genera discussed. Zoological Journal of the Linnean Society 81: 275–387. doi: 10.1111/j.1096-3642.1984.tb01175.x
  • Hendrickx ME, Espinosa-Pérez MC (1998) A new species of Cassidinidea Hansen (Isopoda: Sphaeromatidae) and first record of the genus from the eastern tropical Pacific. Proceedings of the Biological Society of Washington 111: 295–302.
  • Hoestlandt H (1977) Description complémentaire de l’isopode flabellifère Gnorimosphaeroma insulare Van Name et synonimie de G. luteum avec cette espèce. Crustaceana 32: 45–54. doi: 10.1163/156854077X00863
  • Hurley DE, Jansen KP (1977) The marine fauna of New Zealand: family Sphaeromatidae (Crustacea: Isopoda: Flabellifera). New Zealand Oceanographic Institute Memoir 63: 1–95.
  • Iverson EW (1974) Range extensions for some California marine isopod crustaceans. Bulletin of the Southern California Academy of Sciences 73: 164–169.
  • Iverson EW (1978) Status of Exosphaeroma inornata Dow and E. media George and Stromberg (Isopoda: Sphaeromatidae) with ecological notes. Journal of the Fisheries Research Board of Canada 35: 1381–1384. doi: 10.1139/f78-218
  • Iverson EW (1982) Revision of the isopod family Sphaeromatidae (Crustacea: Isopoda: Flabellifera) I. Subfamily names with diagnoses and key. Journal of Crustacean Biology 2: 248–254. doi: 10.2307/1548005
  • Jacobs BJM (1987) A taxonomic revision of the European, Mediterranean and NW African species generally placed in Sphaeroma Bosc, 1802 (Isopoda: Flabellifera: Sphaeromatidae). Zoologische Verhandelingen, Leiden 238: 1–71.
  • Kensley B, Schotte M (1989) Guide to the Marine Isopod Crustaceans of the Caribbean. Smithsonian Institution Press, Washington, DC, 308 pp.
  • Kussakin OG (1979) Marine and brackish-water Isopoda of cold and temperate (boreal) waters of the Northern Hemisphere. Part 1. Flabellifera, Valvifera, and Tyloidea). National Academy of Sciences, USSR, Zoology (Opredeliteli po Faune SSR, Akademiya Nauk, SSSR) 122: 1–470.
  • Kussakin OG, Malyutina MV (1993) Sphaeromatidae (Crustacea: Isopoda: Flabellifera) from the South China Sea. Invertebrate Taxonomy 7: 1167–1203. doi: 10.1071/IT9931167
  • Leach WE (1818) Crustacés, Crustacea. In: Cuvier F (Ed.) Dictionnaire des Sciences Naturelles, dans lequel on traite Méthodiquement des Différents êtres de la Nature, considérés soit en eux-mêmes, d’après l’état actuel de nos connaissances, soit relativement à l’utilité qu’en peuvent retirer la Médecine, l’Agriculture, le Commerce et les Arts. Suivi d’une biographie des plus Célèbres Naturalistes. Ouvrage destiné aux médecins, aux agriculteurs, aux commerçans, aux artistes, aux manufacturiers, et à tous ceux qui ont intérêt à connaître les productions de la nature, leurs caractères génériques et spécifiques, leur lieu natal, leurs propiétés et leurs usages. Vol. 12. F.G. Levrault et Le Normant, Strasbourg et Paris, 69–75.
  • Li L (2000) A new species of Dynoides (Crustacea: Isopoda: Sphaeromatidae) from the Cape d’Aguilar Marine Reserve, Hong Kong. Records of the Australian Museum 52: 137–149. doi: 10.3853/j.0067-1975.52.2000.1311
  • Lombardo AC (1988) Paracerceis richardsonae sp. n. di crostacea isopodo (Sphaeromatidae, Eubranchiatae) delle coste pacifico de Messico. Animalia, Catania 15: 5–15.
  • Loyola e Silva J, Masunari JS, Dubiaski-Silva J (1999) Redescrição de Paracerceis sculpta (Holmes, 1904) (Crustacea, Isopoda, Sphaeromatidae) e nova ocorrência em Bombinhas, Santa Caterina, Brazil. Acta Biologica Paraná, Curitiba 28: 109–124.
  • Menzies RJ (1954) A review of the systematics and ecology of the genus Exosphaeroma with the description of a new genus, a new species, and a new subspecies (Crustacea, Isopoda, Sphaeromidae). American Museum Novitates 1683: 1–24.
  • Menzies RJ (1962) The marine isopod fauna of Bahia de San Quintin, Baja California, Mexico. Pacific Naturalist 3: 339–348.
  • Nunomura N (1998) On the genus Gnorimosphaeroma (Crustacea, Isopoda, Sphaeromatidae) in Japan with descriptions of six new species. Bulletin of the Toyama Science Museum 21: 23–54.
  • Nunomura N (1988) Description of Nishimuraia paradoxa gen. et sp. n., and the first record of the genus Paracerceis in Japan (Isopoda, Sphaeromatidae). Bulletin of the Toyama Science Museum 12: 1–7.
  • Poore GCB (1987) Serolina, a new genus for Serolis minuta Beddard (Crustacea: Isopoda: Serolidae) with descriptions of eight new species from eastern Australia. Memoirs of the Museum of Victoria 48: 141–189.
  • Poore GCB, Bruce NL (2012) Global diversity of marine isopods (except Asellota and crustacean symbionts). PLoS ONE 7: e43529. doi: 10.1371/journal.pone.0043529
  • Richardson H (1897) A new genus and species of Sphaeromatidae from Alaskan waters. Proceedings of the Biological Society of Washington 11: 181–183.
  • Richardson H (1899a) Key to the isopods of the Pacific Coast of North America, with descriptions of twenty-two new species. Proceedings of the United States National Museum 21: 815–869. doi: 10.5479/si.00963801.21-1175.815 [without figures of E. amplicauda]
  • Richardson H (1899b) Key to the isopods of the Pacific Coast of North America, with descriptions of twenty-two new species. Annals Magazine of Natural History London 7, IV: 157–187, 260–277, 321–338. [above reprinted]
  • Richardson H (1899c) Key to the isopods of the Pacific Coast of North America, with descriptions of twenty-two new species. Proceedings of the United States National Museum 21: 815–869. doi: 10.5479/si.00963801.21-1175.815
  • Richardson H (1900) Synopses of the North-American invertebrates. VIII. The Isopoda. – Part I. Chelifera, Flabellifera, Valvifera. American Naturalist 34: 207–230. doi: 10.1086/277593
  • Richardson H (1905) A Monograph of the Isopods of North America. Vol. 54. Smithsonian Institution United States National Museum Bulletin, Washington, DC, 1–727.
  • Schotte M (2012) Exosphaeroma octoncum (Richardson, 1897). In: Schotte M, Boyko CB, Bruce NL, Poore GCB, Taiti S, Wilson GDF (Eds) World Marine, Freshwater and Terrestrial Isopod Crustaceans database. Accessed through: World Register of Marine Species. http://www.marinespecies.org/aphia.php?p=taxdetails&id=257031 [on 2012-10-30]
  • Schultz G (1969) How to Know the Marine Isopod Crustaceans. William C. Brown Company Publishers, Dubuque, Illinois, 359 pp.
  • Shuster SM (1989) Male alternative reproductive strategies in a marine isopod crustacean (Paracerceis sculpta): the use of genetic markers to measure differences in fertilization success among alpha, beta, and gamma males. Evolution 43: 1683–1698. doi: 10.2307/2409384
  • Southern California Association of Marine Invertebrate Taxonomists (SCAMIT) (2013) A taxonomic listing of benthic macro- and megainvertebrates from infaunal and epifaunal monitoring and research programs in the Southern California Bight. Edition 8. http://scamit.org
  • Stebbing TRR (1900) On some crustaceans from the Falkland Islands, collected by Mr. Rupert Vallentin. Proceedings of the General Meetings for Scientific Business of the Zoological Society of London 1900: 517–568.
  • Stimpson W (1857) The Crustacea and Echinodermata of the Pacific shores of North America. Boston Journal of Natural History 6: 503–513.
  • Stimpson W (1858) Sphaeroma amplicauda. Proceedings Boston Society of Natural History 6: 89.
  • Stimpson W (1859) Notes on North American Crustacea, No. 1. Annals of the Lyceum of Natural History of New York 7: 49–93, Plate I. doi: 10.1111/j.1749-6632.1862.tb00142.x
  • Tattersall WM (1913) The Schizopoda, Stomatopoda and non-arctic Isopoda of the Scottish National Antarctic Expedition. Transactions of the Royal Society of Edinburgh 49: 865–894. doi: 10.1017/S0080456800017178
  • Van Name WG (1940) A supplement to the American land and fresh-water isopod Crustacea. Bulletin of the American Museum of Natural History 77: 109–142.
  • Watling L (1989) A classification system for crustacean setae based on the homology concept. In: Felgenhauer BE, Watling L, Thistle AB (Eds) Functional Morphology of Feeding and Grooming in Crustacea. AA. Balkema, Rotterdam, 15–26.
  • Wetzer R, Bruce NL (2007) A new species of Paradella Harrison & Holdich, 1982 (Crustacea: Isopoda: Sphaeromatidae) from Baja California, Mexico, with a key to East Pacific species. Zootaxa 1512: 39–49.
  • Wetzer R, Pérez-Losada M, Bruce NL (2013) Phylogenetic relationships of the family Sphaeromatidae Latreille, 1825 (Crustacea: Peracarida: Isopoda) within Sphaeromatidea based on 18S-rDNA molecular data. Zootaxa 3599: 161–177. doi: 10.11646/zootaxa.3599.2.3

Appendix 1

Sphaeromatidae of the western coast of North America (Alaska to Mexico)

Cassidinidea Hansen, 1905

Cassidinidea mexicana Hendrickx & Espinoza-Pérez, 1998. Gulf of California, Mexico.

Dynoides Barnard, 1914

Dynoides crenulatus Carvacho & Haasmann, 1984. Pacific Mexico.

Dynoides dentisinus Shen, 1929. Introduced. Kussakin 1979; Kussakin and Malyutina 1993.

Dynoides elegans (Boone, 1923) [formerly Clianella elegans Li (2000)]. California.

Dynoides saldani Carvacho & Haasmann, 1984. Pacific Mexico (SCAMIT 2013).

Dynamene

Dynamene benedicti Richardson, 1899. Monterey Bay, California. Correct generic placement uncertain; Kussakin 1979; excluded from the genus by Harrison and Holdich (1982); Brusca et al. 2007.

Dynamene dilatata Richardson, 1899; Monterey Bay, California. Correct generic placement uncertain; Kussakin 1979; Brusca et al. 2007.

Dynamene glabra Richardson, 1899. Oregon to California. Correct generic placement uncertain; Kussakin 1979; excluded from the genus by Harrison and Holdich 1982; SCAMIT 2013.

Dynamene sheareri Hatch, 1947. Oregon. Correct generic placement uncertain; redescribed by George and Strömberg (1968); excluded from the genus by Harrison and Holdich 1982; Brusca et al. 2007 (SCAMIT 2013).

Dynamene tuberculosa Richardson, 1899. California. Correct generic placement uncertain; SCAMIT 2013.

Dynamenella Hansen, 1905

Dynamenella conica (Boone, 1923). California. Correct generic placement uncertain.

Exosphaeroma Stebbing, 1900

Exosphaeroma amplicauda (Stimpson, 1857); = Exosphaeroma octoncum (Richardson, 1897). Marin County to Monterey, California. Brusca et al. 2007; SCAMIT 2013.

Exosphaeroma aphrodita Boone, 1923. San Diego, California.

Exosphaeroma bruscai Espinosa-Perez & Hendrickx, 2002. Pacific Mexico.

Exosphaeroma inornata Dow, 1958; = E. media George & Strömberg, 1968. Puget Sound, Washington to Southern California. Iverson 1978, 1982; Brusca et al. 2007.

Exosphaeroma paydenae sp. n. Alaska.

Exosphaeroma pentcheffi sp. n. Los Angeles, California.

Exosphaeroma rhomburum (Richardson, 1899); Monterey Bay, California. Incertae sedis, males not known.

Exosphaeroma russelhansoni sp. n. Washington.

Gnorimosphaeroma Menzies, 1954

Gnorimosphaeroma insulare (Van Name, 1940), Washington. Menzies 1954; Hoestlandt 1977; Brusca et al. 2007.

Gnorimosphaeroma noblei Menzies, 1954. Washington to California. Kussakin 1979; Brusca et al. 2007.

Gnorimosphaeroma oregonense (Dana, 1852). Alaska to northern California. Menzies 1954; Kussakin 1979, Brusca et al. 2007, SCAMIT 2013.

Gnorimosphaeroma rayi Hoestland, 1969. Nunomura 1998; Brusca et al. 2007.

Paracerceis Hansen, 1905

Paracerceis cordata (Richardson, 1899). California. Kussakin 1979; Brusca et al. 2007.

Paracerceis gilliana (Richardson, 1899); California.

Paracerceis granulosa (Richardson, 1899); Cerros Island, California.

Paracerceis richardsoni Lombardo, 1988. Pacific Mexico. Espinosa-Pérez and Hendrickx 2002.

Paracerceis sculpta (Holmes, 1904). San Clemente Island, California. Globally translocated and widespread. Menzies 1962; Harrison and Holdich 1982; Nunomura 1988; Loyola e Silva et al. 1999; Brusca et al. 2007.

Paracerceis spinulosa Espinosa-Perez & Hendrickx, 2002. Sonora, Mexico.

Paradella Harrison & Holdich, 1982

Paradella dianae (Menzies, 1962). Brusca et al. 2007.

Paradella tiffany Bruce & Wetzer, 2004; Baja California, Mexico.

Paradella garsonrum Wetzer & Bruce, 2007; Baja California, Mexico.

Pseudosphaeroma Chilton, 1909

Pseudosphaeroma sp. cf. campbellensis Bruce & Wetzer, 2008. Introduced species San Francisco to Morro Bay, California. Not P. campbellensis of Hurley and Jansen (1977) and Harrison (1984).

Sphaeroma Latreille, 1802

Sphaeroma quoianum Milne Edwards, 1840 [= Sphaeroma pentodon Richardson, 1904]. Introduced to San Francisco and San Diego, California. Baker 1926; Hale 1929; Kussakin 1979; Harrison and Holdich 1984; Brusca et al. 2007; (SCAMIT 2013).

Sphaeroma serratum (Fabricius, 1787). Worldwide. Kussakin 1979; Jacobs 1987.

Sphaeroma walkeri Stebbing, 1905. Introduced to San Diego, California. Carlton and Iverson 1981; Pires 1982; Brusca et al. 2007, Kussakin 1979.

Striella Glynn, 1968

Striella sp. La Paz, Gulf of California (LACM collections, UC Mexus station 43).