Nephtyidae: Polychaeta) Described from the Eastern Pacific

BULLETIN OF MARINE SCIENCE, 60(2): 350-363, 1997

A REVIEW OF SIX SPECIES OF NEPHTYS (CUVIER, 1817) (NEPHTYIDAE: POLYCHAETA) DESCRIBED FROM THE EASTERN PACIFIC

Lawrence L. Lovell

ABSTRACT Proboscidial characters and the first occurrence of the interramal cirri have been confused or misreported in literature accounts for Eastern Pacific Ocean species of Nephtys Cuvier, 1817. Six species are reviewed: Nephtys caecoides Hartman, 1938; N. californiensis Hart- man, 1938; N. cormUfa Berkeley and Bcrkeley, 1945; N.ferruginea Hartman, 1940; N. parva Clark and Jones, 1955; and N. signifera Hilbig, 1992. Type material is reexamined and com- pared to literature accounts and non-type material. Corrections to the descriptions and the resulting changes in the taxonomic status are presented, and as a result, N. parva is synonymized with N. cornuta. and N. signifera is synonymized with N. ferruginea.

Species within the genus Nephtys have been described worldwide. Important reviews of the genus include Hartman (1938, 1950), Fauchald (1968, 1977), Rainer and Hutchings (1977), and Imajima and Takeda (1987). Early records of the genus Nephtys from the Eastern Pacific Ocean are reported in Johnson (1901), Moore (1908,1909,1911), Treadwell (1914), Chamberlin (1919,1920) and Berkeley (1924). Both Moore (1911) and Treadwell (1914) noted difficulty in determining the actual number of species present in their material. Hartman (1938) published the first comprehensive review of the genus for the NE Pacific. Additional species descriptions and regional information are provided by Hartman (1940, 1968), Berkeley and Berkeley (1945), Clark and Jones (1955), Banse (1972), Fauchald and Hancock (1981), Ellis and Ronaldson (1988), and Hilbig (1992, 1994). In 1993 members of the Southern California Association of Marine Invertebrate Taxonomists (SCAMIT) began reporting Nephtys signifera Hilbig, 1992 from samples collected in southern California during wastewater monitoring studies. Dif- ficulty distinguishing N. signifera from juvenile N. ferruginea Hartman, 1940 led to the discovery that counts of distal paired papillae from monitoring samples dif- fered from the original and subsequent descriptions of N.ferruginea. This discovery prompted a review of the proboscidial features of type material for six Nephtys species reported from California: N. caecoides Hartman, 1938; N. californiensis Hart- man, 1938; N. cornuta Berkeley and Berkeley, 1945; N.ferruginea Hartman, 1940; N. parva Clark and Jones, 1955; and N. signifera Hilbig, 1992. The proboscis of Nephtys is eversible, cylindrical, and has three regions (Fig. IA). The proximal region is smooth or covered with small wart-like processes. The proximal region was smooth for all species studied. The subdistal region has longitudinal rows of papillae. The papillae within a row are progressively shorter anterior to posterior and adjacent rows may converge posteriorly. The number of subdistal papillae in a row is consistent within a narrow range for each species (Table 1). In addition, the subdistal region of the proboscis may have a long unpaired middorsal papilla just anterior to and between the rows of papillae (Fig. IA). The distal region of the proboscis has arcs of paired papillae along each side. The use of the term paired to describe the distal papillae is preferred over the term bifid. Both the inner and outer papillae originate separately on the distal rim of the proboscis (paired), rather than originating from a common area (bifid). The number of paired distal papillae and number of rows of subdistal papillae are

350 LOVELL: REVIEW OF NEPHTYS FROM EASTERN PACIFIC 351

Figure 1. A. Lateral view, proboscis of Nephtys caecoides Hartman, 1938, showing the distal, subdistal, and proximal regions. Goleta, station B-4, replicate 2 (MBC). pdp = paired distal papillae, mup = median unpaired papilla, sp = subdistal papillae. B. Closeup of paired distal papillae of Neph- tys californiensis Hartman, 1938. Holotype (USNM 20320). Bars I mm. 352 BULLETIN OF MARINE SCIENCE. VOL. 60. NO.2. 1997

Table 1. Comparison of selected taxonomic characters between four eastern Pacific specie~ of Nephtys (Cuvier, 1817) based on examination of type material

Subdistal papillae Unpaired middorsal Species Distal paired papillae (rowsfno./row) papillae Intcrrumal cirri begin N. caecoides Hartman, 20 22/3-6 yes setiger 4 1938 N. californiensis Hartman, 20+ 22/5-8 no setiger 3 1938 unpaired middorsal N. cornuta Berkeley and 18 20/4-8 yes setiger :'i Berkeley, 1945 N. ferruginea Hartman, 20 22/3-6 yes seliger 3 1940 invariant for each species (Table 1), The outer papilla of each pair is longer than the inner papilla (Fig. IB). The middorsal and midventral areas are bare (Fig. 4B) or may have a short unpaired papilla (Fig 3B).

MATERIALS AND METHODS Type material was obtained on loan from the Allan Hancock polychaete collection of the Los Angeles County Museum of Natural History (LACM-AHF), the Santa Barbara Museum of Natural History (SBMNH), and the National Museum of Natural History, Smithsonian Institution (USNM). Additional material was provided by the County Sanitation Districts of Orange County (CSDOC), MBC Ap- plied Environmental Services (MBC), MEC Analytical Systems (MEC), and the private collection of the author (LLL). A midventral incision was made to expose each non-everted proboscis, and an additional incision was made along the proboscis to expose proboscidial features (Fig. 2A). Specimens were examined with dissecting and compound microscopes. Methyl green stain was used to highlight brain structures (see Clark, 1955, 1957, 1958a, 1958b; and Hilbig, 1992). Specimens were placed in a saturated solution of methyl green and 70% ethanol for 5-30 min, and then allowed to destain in 70% ethanol for 1-15 min. The anterior nervous system may be seen through the dorsal body wall in small specimens, but ventral dissection may be necessary to expose the anterior nervous system in large specimens (Fig. 6A). Photographs were taken on Nikon SMZ-lO and Nikon Optiphot microscopes equipped with a Nikon FX-35A photo head and Nikon HFX-II exposure meter using Fujichrome Sensia 100 film. Photos were annotated with Photoshop.

SYSTEMATICS Family Nephtyidae Grube, 1850 Genus Nephtys Cuvier, 1817

Type Species.-Nephthys [sicl hombergii Savigny, 1818, designated by Hartman (1950)

Nephtys caecoides Hartman, 1938

Nephtys caecoides Hartman, 1938: 148-149, fig. 63; 1950: 101-102; 1968: 577, figs. 1-3: Banse and Hobson, 1974: 75, fig. 19 e-f; Hilbig, 1994 (in part): 344-346, fig. 13.6 A-E, G, H. Nephthys [sic] caecoides Hartman, 1940: 240. Nephtys californiensis Hartman, 1950 (in part): 103. Material Examined.-CALIFORNIA: Marin Co., Tomales Bay, Williams, col., holotype (USNM 20319); Marin Co., Tomales Bay, Jul. 1935,8 paratypes (LACM-AHF 0794); Orange Co., 33°35.15'N, l.OVELL: REVIEW OF NI:PIITYS FROM EASTERN PACIFIC 353

Figure 2. A. Ventral view, dissected specimen of Nephfys caecoides Hartman, 1938, showing proboscidial features. Paratype (LACM-AHF 0794). pdp = paired distal papillae, mup = median unpaired papilla. sp = subdistal papillae. B. Frontal view, proboscis of Nephfys caecoides Hartman, 1938, showing paired distal papillae. Paratype (LACM-AHF 0794). pdp = paired distal papillae. Bars I mm. 354 BULLETIN OF MARINE SCIENCE, VOL. 60. NO.2. 1997

118°02.53'W,Aug. 1985,Sta. 13, Rep. 4, 59 m., 1 specimen (CSDOC-P475), Rep. 3, 59 m" 1 specimen (CSDOC-P476),Oct. 1991,Sta. 13, Rep. 4, 60 m., I specimen (LLL), Jan. 1995,Sta. 13, Rep. 2, 60 m" 2 specimens (LLL), 33°35.59'N, 118°00.05'W,luI. 1994, Sta, 11,30 m., 1 specimen (LLL), 33°35.50'N, 117°58.15'W,luI. 1994, Sta. IS, 30 m., 3 specimens (LLL), 33°34.29'N, 118°00.12'W, luI. 1994,Sta. ZB, Rep. 2, 56 m., 3 specimens (LLL); Santa Barbara Co., GOLETA, 4 Oct. 1994, Sta. B2, Rep. 1, 90 ft., 1 specimen (MBC), Sta. B4, Rep. 2, 1 specimen (MBC), Sta. B6, Rep, 3, 2 specimens (MBC); San Luis Obispo Co., just north of Morro Bay, 2\ May 1993,Sta. B1, Rep. 1, 2 specimens (MBC), Rep. 4, 1 specimen (MBC), Sta. B2, Rep. 2, 2 specimens(MBC); San Diego Co., Fisher, col., 33°\5.17'N, 117°28.09'W,17 m., 2 specimens (LLL). Corrections and Additions to the Description.-Proboscis with 20 paired distal papillae (Fig. 2B), middorsal and midventral distal areas without papilla. Subdis- tal area with 22 rows of papilla, 3-6 per row, unpaired middorsal papillae present (Fig. lA). Proximal area smooth. Interramal cirri begin on setiger four, recurved. Remarks.-Hartman (1938) provided no count for the distal papillae, and reported 22 rows of subdistal papillae, but no count per row. Hartman (1950) stated that some specimens of N. californiensis have interramal cirri beginning on setiger 4 rather than 3. Specimen lots noted by Hartman as having interramal ci.rri beginning from seti.ger 4 were reexamined. Lots 1882-49 and 1886-49 proved to be N. caecoides, lot 1620-48 could not be found (pers. com.-L. Harris). Later Hartman (1968) added the presence of a subdistal middorsal papillae to the description. Hilbig (1994) reported 22 paired distal papillae, 22 rows of subdistal papillae, with 3-5 papillae per row, and the presence of subdistal long, unpaired middorsal and shorter unpaired midventral papilla. All of the specimens examined have 20 rather than 22 paired distal papillae, and none have an unpaired midventral subdistal papilla. In the same paper Hilbig placed Nephtys parva Clark and Jones (1955) in synonymy with N. caecoides. The holotype of N. parva was examined and found to have bifid ventral antennae on the prostomium and interramal cirri beginning on setiger 5, which are not recurved. This suite of characters places N. parva in synonymy with N. cornuta Clark and Jones (1955) (see below) rather than N. caecoides. Clark and Jones (1955) discussed the possibility that N. parva is a juvenile N. caecoides. The material examined from MBC originalIy identi- fied as N. parva are juvenile N. caecoides. Specimens matching the original description of N. parva have not been found by me. Distribution.-British Columbia to western Mexico, intertidal to 110 m, in mud.

Nephtys californiensis Hartman, 1938

Nephtys californiensis Hartman, 1938: 150-151, fig. 64; 1950 (in part): 103; \968: 579, fig. 1-3; Banse and Hobson, 1974: 75, fig. 19 g-h; Hilbig, 1994: 346-348, fig. 13.7. Nephthys [sic] californiensis Hartman, 1940:240. .Material Exarnined.-CALIFORNIA: Marin Co., Dillon Beach, Intertidal, Hartman, col., Jul. 1935, holotype (USNM 20320), luI. 1934, 6 paratypes (part, specimens with proboscis everted) (LACM- AHF 0642); VenturaCo., Mandalay Beach, Intertidal, Scanland and Parr, col., Nov. 1972, 1 specimen (MEC P1309). Corrections and Additions to the Description.-Proboscis with 20 paired distal papillae (Fig. 3A), short, unpaired middorsal distal papilla present (Fig. 3B), midventral unpaired distal papilla absent (Fig. 3B). Proboscis with 22 rows of subdistal papillae, 5-8 per row; unpaired middorsal papilla absent. Proximal area smooth. Interramal cirri begin on setiger 3, recurved. Remarks.-Hartman (1938) reported no count for paired distal papillae and 22 rows of subdistal papillae with 6-8 per row. She also stated "proboscis. . . usu- ally without a median papilla (rarely one of the paired papillae seems to occupy a LOVELL: REVIEW OF NEPIfTYS FROM EASTERN PACIFIC 355

Figure 3. A. Frontal view, proboscis of Nephtys califomiensis Hartman, 1938, showing paired distal papillae. Paratype (USNM 20320). pdp = paired distal papillae. B. Dorsal view, proboscis of Nephtys californiensis Hartman, 1938, showing paired distal papillae, single middorsal distal papilla, and rows of subdistal papillae. Paratype (USNM 20320). pdp = paired distal papillae, smdp = single middorsal distal papilla, sp = subdistal papillae. Bars I mm. 356 BULLETIN OF MARINE SCIENCE, VOL 60. NO.2, 1997 median position)". Examination of type material verified the presence of a short, middorsal distal papilla. Later (1950) she examined additional material and expanded the species description to include specimens with interramal cirri beginning on setiger 3 or 4. Several lots of that additional material were reexamined and proved to be N. caecoides (see above) or N. ferruginea (see below). Distribution.-California to Mexico, Gulf of Mexico, Japan, China, Korea; intertidal to 330 m; sand or silty sand.

Nephtys cornuta Berkeley and Berkeley, 1945

Nephthys [sic] cornuta Berkeley and Berkeley, 1945: 328-330, figs. 2-4. Nephtys cornuta Hartman, ]950: 106-]07; Hilbig, ]994 (in part ?): 348-35], fig. 13.8 A,B,E-J. Nephtys cornutafranciscana Clark and Jones, 1955: 146, fig, 2; Jones, 1961; Hartman, 1968: 581, fig. 1,2; Banse, 1972: 216-217. Nephlys parva Clark and Jones, 1955: 146, fig. IA-F, ? Jones, 1961. NEW SYNONYMY. Nephtys signifera Hilbig, 1992 (in part), NEW SYNONYMY. Material Examined.-WASHINGTON: Friday Harbor, Fishing Bay, Sta. 39-2, soft mud, Weese, col., holotype (USNM 32873); Eagle Harbor, 47°37.26'N, 122°30.10'W, 13.3 m., Sta, 30, Rep. 1.,4 specimens (LLL). CALIFORNIA: San Francisco Bay, Contra Costa Co., PI. Richmond. holotypc (Nephtys cornutafranciscana) (USNM 26466); Jones, Col.. 14 Sep. 1953, holotype (N. pan'a) (USNM 26464); Santa Maria Basin, Sta. 13,35° 14.54'N, 120059.77'W, 197 m., 1 paratype (Nephtys signifera) (SBMNH 35617), Sta. 30, 34°54.]9'N, ]20047.07'W, 98 m" 1 paratype (Nephfys signifera) (SBMNH 35618); Orange Co., 33°35.l5'N, 118°02.53'W, Sta. ]3, Rep. 3, 59 m., Jan. ]994,2 specimens (LLL). Corrections and Additions to the Description.-Proboscis with 18 paired distal papillae, middorsal and midventral distal areas without papillae. Subdistal area with 20 rows of papillae, 4-8 per row; unpaired middorsal papilla present (Fig. 4A). Proximal area smooth. Interramal cirri begin on setiger 5, not recurved. Remarks.-Berkeley and Berkeley (1945) and Clark and Jones (1955) did not report on the characters of the proboscis. Banse (1972) reported 18 paired distal papillae, 22 rows of subdistal papillae with 4-5 in a row, and an unpaired middorsal subdistal papilla present. Hilbig (1994) reported 14 paired distal papillae, 16 rows of subdistal papillae with about 5 in a row, and an unpaired middorsal papillae. The holotype of N. parva was examined and proved to be N. camllta (see N. caecoides above). Paratype material of N. parva consists of indetermin- able dried posterior fragments (pers. com.-L. Harris). Single specimens of N. cornuta were found in SBMNH paratype lots (35617 and 35618) of N. signifera. Hilbig (1994) expanded the definition of N. comuta to include specimens with paired ventral antenna of subequal length on the prostomium and interramal cirri beginning on setigers 5 or 6, and proposes Aglaophamlls neotellllS Noyes, 1980; and N. neotena Ohwada, 1985 as new synonymies. No variation in length of the bifid ventral antennae or the start of the interramal cirri was observed in the material examined or has been reported in specimens of N. comllta from British Co- lumbia to southern California (L. harris-pers. com., H. Jones-pers. com., G. Ruff-pers. com., and T. Phillips-pers. com.). The acceptance of these synonymies is questioned pending further investigation. The discontinued use of the trinomial names N. cornuta cornuta and N. cornuta franciscana suggested by Ellis and Ronaldson (1988) is followed. Distribution.-British Columbia to southern California, subtidal to 1600 m, silt or mud.

Nephtys ferruginea Hartman, 1940

Nephfhys [sic] caecoidesferruginea Hartman, 1940: 241, pI. 42, figs. 110-114, pI. 43, fig. 115. Nephlys californiensis Hartman, 1950 (in part): 103. LOVELL: REVIEW OF Nl,"I'lITYS T'ROM EASTERN PACIFIC 357 A

Figure 4. A. Lateral view, prohoseis or Neplllys ("omura Berkeley and Berkeley, 1945, showing paired distal papillae, median unpaired papilla, and subdistal papillae. Orange Co., station 13, replicate 3 (LLL). pdp ~ paired distal papillae, mup = median unpaired papilla, sp = subdistal papillae. Bar .1 mill. B. Frontal view, prohoscis or Neplltys fC'rmgillea Hariman, 1940, showing paired distal papillae. Orange Co., (, m. north or pipe (LLL). pdp = paired distal papillae. Bars A .1 111m, B I mm. 358 BULLETINOF MARINESCIENCE.VOL.60. NO.2. 1997

Nephtys ferruginea Berkeley and Berkeley, 1945: 327-328; Hartman. 1950: 102-103; 1968: 583, figs. 1-4; Banse and Hobson, 1974: 75, fig. 19. k-1; Hilbig, 1994: 351-352, fig. 13.9 A-H. Nephtys signifera Hilbig, 1992 (in part): 719-721, fig. 5A (?), C, D; Hilbig, 1994 (in part): 356- 358, fig. 13.12A (?), C, D. NEW SYNONYMY.

Material Examined.-PERU: Independencia Bay, 14°16'S, 76°IQ'W, VELERO Sta. 834-38, 10 Feb. 1938, 21 fms., mud, holotype (LACM-AHF 0798), 19 paratypes (LACM-AHF (799). CALIFORNIA: Anacapa Is., VELEROSta. 876-38, 1 specimen; southern California Bight, BLM Sta. 82517, 1 specimen; Santa Maria Basin, Sta. R-2, 35°05.50'N, 120049.16'W, 161 m., holotype (N. sigllifl'l'a) (USNM 148706), Sta. PJ-I, 34°55.79'N, 120°49.91 'w, 145 m., I paratype (N. sigllifera) (LACM-AHF 1(15), Sta. PJ-6, 34°54.71'N, 120049.91'W, 148 m., 1 paratype (N. signifera) (LACM-AHF 1616), Sta. 13, 35°14.54'N, 120059.77'W, 197 m., 3 paratypes (N. signifera) (SBMNH 35617), Sta. 30, 34°54.19'N, 120047.07'W, 98 m., 6 paratypes (N. signifera) (SBMNH 35618); Ventum Co., 34°12.29'N, I 19°30.43'W, 91 m., Sta. 661, 2 specimens, (CSDOC); Orange Co., 33°34.42'N, I 18°01.33'W, 59 m., Sta. 5, Rep. 4, Jan. 1993,3 specimens (LLL), 33°35.59'N, 117°55.57'W, 98 m., Sta. C3, Jul. 1993, 1 specimen (LLL), 33°34.29'N, 117°59.42'W, 56 m., Sta. 4, Rep. 2, Oct. 1993, ] specimen, (LLL), 33°34.20'N, 117°59.26'W, 59 m., Sta. 9, Rep. 3, Oct. 1993, 3 specimens (LLL), 6 m. North of pipe, 30 m., Jul. 1994, 1 specimen (LLL). Corrections and Additions to the Description.-Proboscis with 20 paired distal papillae, middorsal and midventral distal papilla absent. Subdistal area with 22 rows of papillae, 3-6 per row (Fig. 5A). Proximal area smooth. lnterrama] cirri begin on setiger 3, not recurved. Postacicular setae smooth to spinulose; upper- most and lowermost fascicles of both rami with a mix of smooth to spinulose setae, middle fascicles with spinulose setae only (Fig. 5B). Remarks.-Hartman's original description (1940) of N. caecoides ferruginea reported no presence of distal paired papillae, 22 rows of subdistal papillae without counts per row, presence of a subdistal unpaired middorsal papilla, proximal region smooth, and recurved interramal cirri starting on setiger 3. Berke]ey and Berkeley (1945) elevated N.ferruginea to specific rank without providing any additional information. Lots 1597-47 and 1759-49 of the material reported as N. californiensis by Hartman (1950) were reexamined and found to be specimens of N. ferruginea (pers. com.-L. Harris). The types of N. signifera were reexamined and the holotype and paratype lot (LACM-AHF) 16] 6 were found to be juvenile N.ferruginea. Para- type 10tLACM-AHF 1615 was found to be an Aglaophamus species (involute interramal cirri) too small to be determined to species. Paratype lots SBMNH 35617 and SBMNH 35618 were examined and found to contain one N. cornuta (see N. cO/'lluta above) each, with the other specimens in the lots being juvenile N. ferruginea. Hilbig (1992) used three characters to separate N. signifera from N.ferruginea: paired distal papillae counts, dorsal methyl green staining pattern, and distribution of post-acicular setal types. Corrected paired papillae counts for N. ferruginea are the same as those described for N. signifera. The dorsal methyl green staining area in Nephtys is composed of the anterior nervous system as described by Clark (1955, 1957, 1958a, 1958b). Clark (1955, 1958b) illustrates the anterior nervous system for N. ferruginea and other Eastern Pacific Nephtys species (Fig. 7). The dorsal staining pattern is visible through the body wall of small specimens, but not visible through the thicker body wall of large specimens. The stained area can be seen in a ventral dissection with the proboscis moved aside (Figs. 6A). All type material of N. sign(fera from Hilbig (1992) herein referred to N. ferruginea were small specimens. Reexamination of the type material of N. ferruginea and N. signifera reveal that the distribution of post-acicular setal types is the same for both species. Examination of all material indicates that smooth post- acicular setae are more abundant in juveniles and the median and posterior setigers of adults. This distributional pattern may be due to the effect of setal wear caused by sediment abrasion or setal replacement with age. Distrihution.-British Columbia to Peru; 16-450 m; in mud, sand, silt, and clay. LOVELL: REVIEW OF NEPIITYS FROM EASTERN PACIFIC 359

Figure 5. A. Dorsolateral view, proboscis of NepJllys ferruginea Hartman, 1940, showing paired distal papillae, median unpaired papilla, subdistal papillae. Orange Co., 6 m. north of pipe (LLL). pdp = paired distal papillae, mup = median unpaired papilla, sp = subdistal papillae. B. Posterior view, 10th notopodiallobe of Nepl1lys ferrllginea Hartman, 1940, showing smooth and spinulose setae. Ho- lotype (LACM-AHF 0798). Superior to inferior = left to right. Bar A 1 mm, B .1 mm. 360 BULLETIN OF MARINE SCIENCE, VOL. 60, NO.2. 1997

Figure 6. A. Dorsal and ventral (dissected) views of methyl green stained anterior nervous system of Nephtys ferruginea Hartman, 1940. Ventura Co., Station 661 (CSDOC). mgsa = methyl green staining area. B. Dorsal view of methyl green stained anterior nervous system of Nephtys ferruginea Hart- man, 1940. mgsa = methyl green staining area. Bars I mm. LOVELL: REVIEW OF NEPH7YS FROM EASTERN PACIFIC 361

N. californiensis N. cornuta N. ferruginea N. parva

Clark, 1958

Clark, 1955

Figure 7. Illustrations from Clark, 1955 and 1958, showing the anterior nervous systems of four species of Nephtys from the Eastern Pacific. The lower row of illustrations shows differentiation of staining intensity and includes far anterior staining areas. Bars as noted.

DISCUSSION The proboscidial characters of the six species of Nephtys presented have often been confused or misreported in the literature. The facts that specimens rarely fully evert their probosci and researchers rarely expose these structures by dissection, have been a hindrance to realizing and correcting this problem. Partially everted probosci have led to confusion in determining correct papillae counts for distal and subdistal regions. Authors have not reported, or have incorrectly reported, the distal paired papillae counts as equal to the number of subdistal rows. The number of paired distal papillae is two fewer than the number of subdistal rows in each of the species of Nephtys examined. The presence of a short, unpaired middorsal distal papilla in N. californiensis Hartman, 1938 has often been overlooked. Corrected proboscidial characters and start of the interramal cirri for N. caecoides Hartman, 1938; N. californiensis Hartman, 1938; N. cornuta Berkeley and Berkeley, 1945; and N. ferruginea Hartman, 1940 are presented in Table 1. Specimens matching the description of N. parva Clark and Jones, 1955 have not been seen by the author. Should those specimens be found, that material should be redescribed, with a new name, and new type material designated since the name N. parva is unavailable. Preliminary findings (pers. obs.) show that similar problems with reported proboscidial characters exist for other species of Nephtys and some species of Ag- laophamus from the Pacific. It is probable that this situation exists with other members of these two genera. The systematic importance of paired distal papillae counts will not be fully understood until a review of remaining type material from within the family is performed.

ACKNOWLEDGMENTS

I thank the curators and collections managers at the National Museum of Natural History, Smithso- nian Institution; the Los Angeles County Museum of Natural History, and the Santa Barbara Museum 362 BULLETIN OF MARINE SCIENCE. VOL. 60. NO.2. 1997 of Natural History for loans of type material. Additional material was kindly provided by the County Sanitation Districts of Orange County, MBC Applied Environmental Scienccs, and MEC Analytical Systems. R. Velarde at the City of San Diego Marine Biology Laboratory provided access to the pho- 10microscopy equipment. Special thanks are due Dr. K. Fauchald for his advice, and Ms. L. Harris for her examination of specimens and helpful comments during the preparation of this paper. This paper benefitted from reviews by Dr. K. Fitzhugh, Ms. L. Harris, and Mr. R. E. Ruff. Partial funding was provided by Southern California Association of Marine Invertebrate Taxonomists Publication Grant 96-1. This paper is Contribution No. 10 of the Southern California Association of Marinc Invcrtebrate Taxonomists.

LITERATURE CITED

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DATE ACCEPTED: September 23, ]996.

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