Description of a new species ofHirsutonuphis (Annelida: Onuphidae) fromthe Mexican Pacific, with notes on the worlddistribution of the genus

alejandra estrella-ruiz, pablo herna’ ndez-alca’ ntara and vivianne soli’s-weiss

Laboratorio de Ecologıa y Biodiversidad de Invertebrados Marinos, Instituto de Ciencias del Mar y Limnologıa, UniversidadNacional Autonoma de Mexico. Circuito Exterior S/N, Cd. Universitaria, D. F. 04510, Mexico

A new species of Hirsutonuphis (Annelida: Onuphidae) is described from the continental shelf of the Gulf of California,Mexican Pacific. Hirsutonuphis paxtonae sp. nov. is easily distinguished from eight previously described species belongingto this genus, by the bi- and tri-dentate pseudocompound hooded falcigers on the first 5–6 chaetigers, the pectinatechaetae from chaetigers 6–11, the bidentate subacicular hooded hooks from chaetigers 18–21, and its colour pattern: adark transverse band in the dorsal part of the peristomium and on each chaetiger, which gradually fades from chaetigers12–27. Most species of Hirsutonuphis have been found in the Pacific Ocean, and more particularly in Australian waters,although this new onuphid is the third species of the genus described from the Mexican Pacific, in the Tropical EasternPacific. A taxonomic key is provided for all the species of the genus.

Keywords: Polychaeta, taxonomy, Eastern Pacific, geographical distribution

Submitted 1 October 2012; accepted 12 March 2013; first published online 3 May 2013

I N T R O D U C T I O N

The genus Hirsutonuphis was erected by Paxton (1986), in thesubfamily Onuphinae, as part of one of the most importantgeneric revisions of the taxonomy and relationships of thefamily Onuphidae. In addition, she established thatHirsutonuphis is the only genus in the Onuphidae where themaxillary carriers are of diagnostic importance. Paxton (1986)included in this genus the onuphids with modified parapodiawith long dorsal cirri, giving them a ‘hairy’ appearance. This,combined with the name Onuphis, the first genus described inthis family, provided the name for the newly erected genus.Initially, five species were included in the genus Hirsutonuphis:four from the Pacific Ocean (Onuphis zebra Berkeley &Berkeley, 1939 and Onuphis acapulcensis Rioja, 1944 from theMexican Pacific; Onuphis gygis Paxton, 1979 and Onuphis mar-iahirsuta Paxton, 1979 from Australia), and only one speciesfrom the Atlantic Ocean, in Belize: Onuphis geminataFauchald, 1980. Subsequently, Paxton (1996) described threenew species from Australian waters: Hirsutonuphis armillata,H. intermedia and H. macrocerata, thus increasing the presentlyrecognized species in the genus to eight.

The members of the family Onuphidae have been reportedfrom all the seas of the world and at all depths (Paxton, 1986);however, it seems that the species of Hirsutonuphis mainlyoccur in the Pacific Ocean. In fact, the two species recordedfrom the Eastern Tropical Pacific were collected in Mexican

waters: H. zebra is from Punta Gorda, Baja California, and H.acapulcensis is from Acapulco, Guerrero. Recently, in thecourse of a revision of biological material collected on the con-tinental shelf of the Gulf of California, we found specimensbelonging to the genus Hirsutonuphis, which had evident mor-phological differences, not only from the species recorded in theMexican Pacific, but also from all the species so far described.

Therefore, the aim of this study is to describe this newspecies of Hirsutonuphis from the Mexican Pacific, toprovide a taxonomic key to all species of the genus, and todiscuss the particularities of the known world distribution ofthe genus.

M A T E R I A L S A N D M E T H O D S

The polychaete specimens analysed here were collected on thecontinental shelf of the Gulf of California, on-board the RV ‘ElPuma’ from the Instituto de Ciencias del Mar y Limnologıa(ICMyL), Universidad Nacional Autonoma de Mexico(UNAM), during the oceanographic expedition ‘Cortes 2’ ofMarch 1985. The samples were collected with a Smith–McIntyre grab (0.1 m2), and the geographical position of thestations was determined by Global Positioning System(GPS). The polychaetes were fixed in 4% formaldehyde andpreserved in ethanol 70%. Scanning electron microscope(SEM) photographs of important diagnostic characters ofthe species were taken with JEOL JSM6360LV equipment, fol-lowing standard methodology.

Paxton (1986) standardized the nomenclature of the mor-phological characters used until then under different names.

Corresponding author:V. Solıs-WeissEmail: solisw@cmarl.unam.mx

1777

Journal of the Marine Biological Association of the United Kingdom, 2013, 93(7), 1777–1783. # Marine Biological Association of the United Kingdom, 2013doi:10.1017/S0025315413000441

To describe the new species, we used her terminology comple-mented, when appropriate, with her own additions of a laterstudy (Paxton, 1998) and those of Budaeva & Fauchald(2011). The numbers and measurements annotated in thedescription correspond to the holotype, but variations foundin the paratypes are also included; the total length of the speci-mens (without antennae), the length of the antennae andpalps, and the body width (without parapodia or chaetae) atchaetiger 10, were measured.

The holotype and paratypes are deposited in the NationalPolychaete Collection located at the Instituto de Ciencias delMar y Limnologıa (ICMyL), Universidad NacionalAutonoma de Mexico (UNAM) (CNP-ICML, UNAM;DFE.IN.061.0598), and at the Los Angeles County Museumof Natural History (LACM-AHF POLY), USA.

systematics

Phylum ANNELIDA Lamarck, 1809Family ONUPHIDAE Kinberg, 1865

Subfamily ONUPHINAE Kinberg, 1865Genus Hirsutonuphis Paxton, 1986

Hirsutonuphis paxtonae sp. nov.(Figures 1A–I, 2A–H)

Onuphis sp. 1 Gonzalez-Ortiz et al., 1997: 392.

type material

Holotype: CNP-ICML POH-52-002: Expedition Cortes 2;Station 3, Santa Marıa Bay, Gulf of California, 2582.4′N

Fig. 1. Hirsutonuphis paxtonae sp. nov. (holotype): (A) anterior end, showing pigmentation pattern, dorsal view (palpostyles details omitted); (B) parapodium 6,anterior view; (C) maxillae, ventral view; (D) median parapodium, anterior view; (E) limbate chaetae; (F) pseudocompound bidentate hooded falciger; (G)pseudocompound tridentate hooded falciger; (H) bidentate subacicular hook; (I) pectinate chaeta. Scale bars: A, 1 mm; B–D, 0.2 mm; E, H, 20 mm; F, G, I, 10 mm.

1778 alejandra estrella-ruiz et al.

108831.7′W; 32 m depth, on fine sands; coll. P.Hernandez-Alcantara, 10 March 1985.

Paratypes: CNP-ICML POP-52-002: two specimens, one ofthem coated with gold for SEM studies; same station as holo-type. LACM-AHF POLY 5319: two specimens; same station asholotype.

diagnosis

Body large, slender, with transversal dark band on dorsum ofperistomium and on each chaetiger, gradually fading fromchaetigers 12–27. Ceratophores of antennae and palps with7–11 rings and moderately long styles. Ventral cirri cirriformon anterior 6–7 chaetigers. Pseudocompound bi- and tri-dentate hooded falcigers on first 5–6 chaetigers. Pectinatechaetae from chaetigers 6–11, slightly oblique with tenteeth. Bidentate subacicular hooded hooks from chaetigers18–21. Branchiae from chaetiger 6 with up to 4–9 filaments.

description

Holotype incomplete with 52 chaetigers, 15 mm long, 1.6 mmwide (not including chaetae). All paratypes incomplete with59–73 chaetigers, 10–25 mm long, 1–2 mm wide. Bodylarge, slender, with transversal dark band on dorsum of peri-stomium and on each chaetiger, gradually disappearing fromchaetiger 18 (12–27 in paratypes).

Prostomium short, with anterior median incision; one pair ofeyes located between palps and lateral antennae. Dark brownpigmented area present on bases of frontal lips and distalparts of ceratophores of antennae and palps (Figure 1A).Frontal lips oval; upper lips oval with median section. Nuchalgrooves separated in midline, curved towards base of medianantenna. Palps 1 mm long (0.5–1 mm in paratypes), reachingback to chaetiger 2 (2–3 in paratypes), much shorter thanantennae. Peristomium without distinct anterior fold, similarin size to first chaetiger and appearing to extend along nuchalgrooves to median antenna. Peristomial cirri inserted in anteriorregion of peristomium, between median and lateral antennae,reaching back to first chaetiger (Figure 2A, B). Antennae withmoderately long styles about equal in length, measuring about4 mm long (3–4 mm in paratypes), reaching back to chaetiger14 (9–16 in paratypes). Lateral antennae inserted slightly infront of median antenna. Ceratophores of lateral antennaewith eight rings (7–11 in paratypes); ceratophores of medianantenna also with eight rings (7–8 in paratypes) (Figures 1A& 2A, B).

Anterior five pairs of parapodia modified (6 in a paratype),moderately elongated, directed anterodorsally (Figure 2A).Dorsal cirri digitiform, very long, with basal swelling: thoseof chaetigers 1 and 2 reaching back to chaetiger 5 (5–6 inparatypes), those of chaetiger 3 back to chaetiger 7 (7–8 inparatypes), gradually reduced in length; in median and pos-terior region of body, dorsal cirrus reduced so that its lengthis shorter than that of segment (Figure 2G). Postchaetallobes subulate, best developed on first five parapodia(Figures 1B & 2C, D), gradually reduced in length toward pos-terior part of the body. Ventral cirri cirriform on first sevenchaetigers (6–7 in paratypes) (Figure 1B), as long as the chae-tiger’s length; then replaced by glandular pads without tran-sition zone of glandular pads and reduced cirri. Interramalpapillae absent on all parapodia. Branchiae from chaetiger 6,with thick stem and pectinate filaments; first branchia withtwo filaments, increasing to maximum of six filaments (4–9in paratypes) (Figures 1B & 2A, C, G).

Parapodia sub-biramous with notopodia reduced to dorsalcirri (Figures 1B & 2A). Neuropodia with two bundles ofsimple limbate chaetae (Figures 1B & 2C, D), in the mediansegments, neuropodia have an upper bundle of thin andslender limbate chaetae and a lower bundle of thick andshort limbate chaetae which are replaced by hooded subacicu-lar hooks posteriorly. First five modified chaetigers (six in oneparatype) with bi- and tri-dentate pseudocompound hoodedfalcigers. Bidentate falcigers with a large slender upper fangand a secondary inferior tooth, shorter but wider than upperfang (Figures 1F & 2E), and tridentate falcigers with a largeslender upper fang, wide secondary tooth, and small thirdtooth at the base of second tooth (Figures 1G & 2F).Pectinate chaetae absent on anterior modified parapodia,occurring as 1–2 per parapodium from chaetiger 6 (7–11 inparatypes), slightly oblique with about 10 teeth of similarsize (Figures 1I & 2D). A pair of bidentate hooded subacicularhooks from chaetiger 21 (18–21 in paratypes) (Figures 1H &2H). Pygidium unknown, posterior segments missing.

Mandibles whitish, with slender shafts; maxillae with mod-erately long Mx III, small Mx V; Mx VI absent. Carriers heart-shaped: lateral triangular area between MX I and carriers, andanterior median region of carriers strongly sclerotized.Maxillary formula: Mx I ¼ 1 + 1; Mx II ¼ 7 + 7; Mx III ¼9 + 0; Mx IV ¼ 6 + 7; Mx V ¼ 1 + 1 (Figure 1C).

Fig. 2. Hirsutonuphis paxtonae sp. nov. (paratype) (A) anterior end,dorsolateral view; (B) anterior end, front view; (C) parapodia 6 and 7, lateralview; (D) parapodium 10, postchaetal lobe and chaetae, without cirri orbranchiae, posterior view; (E) pseudocompound bidentate hooded falcigers;(F) pseudocompound tridentate hooded falciger; (G) central body,dorsolateral view; (H) bidentate subacicular hooks. Scale bars: A, G, 0.5 mm;B, 0.2 mm; C, 0.1 mm; D, H, 20 mm; E, 10 mm; F, 5 mm.

new hirsutonuphis from the mexican pacific 1779

Table 1. Comparison of taxonomic characters in Hirsutonuphis species.

Diagnostic characters H. acapulcensis(Rioja, 1944)

H. armillataPaxton, 1996

H. geminata(Fauchald, 1980)

H. gygis(Paxton, 1979)

H. intermediaPaxton, 1996

Rings in ceratophores Median antenna 10–12.Lateral antennae 12–14

Median antenna 5–8. Lateralantennae 10–13

Median and lateral antennae9–10

Median and lateral antennae10–15

Median antenna 7–8. Lateralantennae 8–9

Branchiae from chaetiger 6 6 6 8–9 6Branchial filaments Maximum 12 Maximum 4–6 Maximum 8–9 Maximum 10–12 Maximum 6–9Pseudocompound hooded falcigers type Tridentate Tridentate Bi- and tridentate Bidentate Bi- and tridentateChaetigers with pseudocompound

hooded falcigers1 to 5 1 to 5 1 to 7 1 to 7 (8) 1 to 6

Ventral cirri cirriform First 5 chaetigers First 5–7 chaetigers First 9 chaetigers First 11 (9) chaetigers First 6–8 chaetigersDistribution of pectinate chaetae and

number of teethWith 15–20 teeth From about chaetiger 20, distally

oblique with 7–9 teethChaetigers 2–6, transverse

with 8 teeth. Absent inchaetigers 7 to 18.Posteriorly distally obliquewith 12 teeth

From chaetiger 1, distallyoblique with 10–16 teeth

From chaetiger 20, distallyoblique with 8–10 teeth

Subacicular hooded hooks from chaetiger 15–18 11–12 23 28–29 13–17Maxillary formula ? MxI ¼ 1 + 1

MxII ¼ 7–10 + 9–10MxIII ¼ 9–10 + 0MxIV ¼ 5–6 + 8–10MxV ¼ 1 + 1

MxI ¼ 1 + 1MxII ¼ 7 + 8MxIII ¼ 6 + 0MxVI ¼ 6 + 7MxV ¼ 1 + 1

MxI ¼ 1 + 1MxII ¼ 7 + 9MxIII ¼ 10 + 0MxIV ¼ 6 + 8MxV ¼ 1 + 1

MxI ¼ 1 + 1MxII ¼ 9 + 9MxIII ¼ 9 + 0MxVI not visibleMxV ¼ 1 + 1

Body colour pattern Peristomium and firstchaetiger with a browndark band. Two dark spotsat each side of following 4or 5 chaetigers.

Five morphs; basic mostcommon morph, brownpigment forming semi-circles around tips of anteriorparapodia

Most ceratophoral rings withnarrow bands; caratostyleswith speckled, irregularspots. Peristomium with aband across the dorsal side

Dark brown pigment on basesof frontal lips andprechaetal lobes of anteriorparapodia

Brown pigments as dorsallateral oval patches onanterior parapodia

Diagnostic characters H. macrocerataPaxton, 1996

H. mariahirsuta(Paxton, 1979)

H. zebra(Berkeley & Berkeley,1939)

H. paxtonae sp. nov.

Rings in ceratophores of antennae Median antenna 8–14. Lateralantennae 11–14

Median and lateral antennae 8–13

Median antenna 10. Lateralantennae 15

Median antenna 7–8. Lateralantennae 7–11

Branchiae from chaetiger 6 6 6 6Branchiae filaments Maximum 6–10 Maximum 9–12 Maximum 8 Maximum 4–9Pseudocompound hooded falcigers type Bi- and tridentate Bi- and tridentate Tridentate Bi- and tridentateChaetigers with pseudocompound

hooded falciger1 to 5 1 to 7 1 to 5 1 to 5 or 6

Ventral cirri cirriform First 6–7 chaetigers First 7–9 chaetigers First 10 chaetigers First 6 or 7 chaetigersDistribution of pectinate chaetae and

number of teethFrom chaetiger 1, distally

oblique with 7–9 teethFrom chaetiger 1, distally

oblique with 15–20 teethPosterior to fifth chaetiger,

slightly oblique with 12teeth

From chaetigers 6–11, slightlyoblique with 10 teeth

Subacicular hooded hooks from chaetiger 15–19 20–26 After chaetiger 35 12–21

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Hirsutonuphis paxtonae sp. nov. belongs to a group of specieswith bi- and tri-dentate pseudocompound falcigers and thuscan be clearly separated from the species with exclusivelytridentate pseudocompound falcigers (H. acapulcensis andH. zebra, described from the Mexican Pacific coasts, and H.armillata collected in Australia) or exclusively bidentate falci-gers (H. gygis recorded in Australia) ( Table 1).

Hirsutonuhis paxtonae sp. nov. can also be differentiatedfrom the other five Hirsutonophis species with bi- and tri-dentate pseudocompound hooded falcigers, by being theonly species lacking pectinate chaetae (10 teeth) in theanterior modified chaetigers, since they appear from chaeti-gers 6–11 (Table 1). In both H. macrocerata and H. mariahir-suta, from Australia, pectinate chaetae are present fromchaetiger 1 (7–9 teeth and 15–20 teeth, respectively), whilein H. intermedia also from Australia, they are found poster-iorly, from chaetiger 20 (8–10 teeth). Hirsutonuhis geminatafrom Belize, is a unique species in the genus, in having a dis-continuous distribution of the pectinate chaetae: they arepresent in chaetigers 2 to 6 (with 8 teeth), but absent fromchaetiger 7 through to about chaetiger 18 (Table 1).

According to the descriptions and remarks about theAustralian species made by Paxton (1979, 1996), the colourpattern can be used to distinguish the species of this genus.The colour pattern in H. paxtonae sp. nov., a dark brownband in the dorsal region of the peristomium and on eachchaetiger, which gradually fades from chaetigers 12–27,differs from other species with bi- and tri-dentate pseudocom-pound falcigers (Table 1), even from H. mariahirsuta, inwhich large variations in pigmentation are present: the peri-stomium and following five chaetigers without pigmentation,or with varying degrees of dark brown segmental bands, basi-cally on the central region of each chaetiger.

etymology

This species is named in honour of Dr Hannelore Paxton, oneof the most important Australian polychaetologists, whoseextensive and excellent studies on the family Onuphidaehave added considerably to our knowledge of the polychaetes.

type locality

Santa Marıa Bay (2582.4′N 108831.7′W), Gulf of California,eastern Pacific.

habitat

The specimens of H. paxtonae sp. nov. have only been col-lected from the type locality at 32 m depth, on fine sands(96%), salinity of 35.04 psu, temperature of 148C, 1.02 ml/lof dissolved oxygen and 5.7% of organic carbon content inthe sediment.

D I S C U S S I O N

Hirsutonuphis paxtonae sp. nov. from the Gulf of California,Mexican Pacific, is the ninth described species in the genus.Even though the polychaetes of the family Onuphidae havebeen widely recorded in all the seas of the world, so far thegenus Hirsutonuphis has been essentially restricted to thePacific Ocean, and the distribution of each of the ninespecies described (including H. paxtonae sp. nov.) isM

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new hirsutonuphis from the mexican pacific 1781

practically limited to their type locality (Table 2). Despite this,a distribution pattern can be inferred from the geographicalposition and depths under which those species were collected,i.e. the genus is better represented at depths shallower than40 m in the Pacific Ocean, since only H. macrocerata and H.intermedia have been recorded on the outer continentalshelf (75 and 95 m, respectively), and only H. geminata hasbeen described from the Atlantic Ocean (Belize, CaribbeanSea). Besides, with the exception of H. geminata, which wascollected in sea grasses and mangroves in Belize, the speciesof this genus have been basically found in sand substrates ofdifferent textures, but rarely in muds.

The restricted distribution of each of the species ofHirsutonuphis is more evident when looking at their recordsin the Pacific Ocean, where they have been found exclusivelyon Australian shores (south-western Pacific) (Paxton, 1979,1986) and in Mexico (north-eastern Pacific) (Hernandez-Alcantara et al., 2008). Obviously, the small number ofknown species and the scarce records so far available makeit necessary to do more in-depth research on the presence ofHirsutonuphis worldwide, to verify its distribution rangesand try to prove if this so far apparent disjunctive distributioncan be a geographical attribute distinctive of the genus.

K E Y T O D E S C R I B E D S P E C I E S O FH I R S U T O N U P H I S ( E M E N D E D F R O MP A X T O N , 1 9 8 6 , 1 9 9 6 )

1. Anterior dorsal cirri as thick as antennae; pseudocom-pound falcigers on anterior chaetigers bidentate . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. gygis (Paxton, 1979)

– Anterior dorsal cirri much thinner than antennae; pseudo-compound falcigers on anterior chaetigers tridentate or bi-and tridentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. (1) Conical interramal papillae on chaetigers 11–18; biden-tate subacicular hooks after chaetiger 35 . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . .H. zebra (Berkeley & Berkeley, 1939)

– Interramal papilla absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3. (2) All pseudocompound falcigers tridentate . . . . . . . . . . . . 4– Pseudocompound falcigers bi- and tridentate . . . . . . . . . . . 5

4. (3) Bidentate subacicular hooks from chaetigers 15–18;tips of anterior parapodia with no pigmentation; medianantenna with 10–12 rings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. acapulcensis (Rioja, 1944)– Bidentate subacicular hooks from chaetigers 11 to 12;

brown pigmentation on bases around tips of anterior para-podia; median antenna with 5–8 rings . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .H. armillata Paxton, 1996

5. (3) First bidentate subacicular hooks before chaetiger 19. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

– First bidentate subacicular hooks after chaetiger 20 . . . . . 7

6. (5) Colour pattern of brown lateral spots; pectinate chaetaefrom chaetiger 1; lateral ceratophores with 11–14 rings;very long antennae, median as long as laterals, to chaetigers7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. macrocerata Paxton, 1996

– Colour pattern of brown lateral oval patches; pectinatechaetae from about chaetiger 20; lateral ceratophoreswith 8–9 rings; moderately long antennae, median halfas long as laterals, to chaetigers 4–6 . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. intermedia Paxton, 1996

7. (5) Pseudocompound falcigers bi- and tridentate in first5 – 6 chaetigers; pectinate chaetae from chaetigers 6 – 11. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. paxtonae sp. nov.

– Pseudocompound falcigers bi- and tridentate in first sevenchaetigers; pectinate chaetae from chaetiger 1 or 2 . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

8. (7) Pectinate chaetae from chaetiger 1, distally oblique with15–20 teeth; branchiae with up to 9–12 filaments . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . H. mariahirsuta (Paxton, 1979)– Pectinate chaetae from chaetiger 2, distally transverse with

eight teeth, absent in chaetiger 7 through to about chaetiger

Table 2. World distribution, depth and substrate recorded for all species in the genus Hirsutonuphis.

Species Distribution Depth Habitat Source

H. acapulcensis ∗ Acapulco Guerrero, Mexican Pacific Intertidal Rioja, 1944H. armillata ∗ Halifax Bay, N of Townsville, Queensland,

to Jervis Bay, southern New South WalesIntertidal to 40 m Usually sands, rarely mud; Paxton, 1996

H. geminata ∗ Halfway between Colson Point and SaltCreek, Dangriga District, Belize

1 m Thalassia, and fine sands in amangrove area

Fauchald, 1980

H. gygis ∗ Fraser Island, Queensland Paxton, 1979Sydney Harbour, New South Wales Intertidal to 19 m Sand Paxton, 1996

H. intermedia ∗ Port Phillip Bay, Melbourne, Victoria, toBass Strait

4–95 m Usually in fine to mediumsands, rarely fine shells and

sands

Paxton, 1996

H. macrocerata ∗ Sydney area to Jervis Bay, New South Wales 20–70 m Sand Paxton, 1996H. mariahirsuta ∗ Northern New South Wales and

Queensland; Camden Haven, New SouthWales, to Yeppoon, Queensland

Intertidal Paxton, 1979

H. zebra ∗ Punta Gorda, Lower California, MexicanPacific

Berkeley & Berkeley, 1939

H. paxtonae sp. nov. Santa Marıa Bay, Gulf of California, MexicanPacific

32 m Fine sands Present study

∗, type locality.

1782 alejandra estrella-ruiz et al.

18, posteriorly each is distally oblique with 12 teeth; bran-chiae with up to 8–9 filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . .H. geminata (Fauchald, 1980)

A C K N O W L E D G E M E N T S

Thanks are due to Yolanda Hornelas for the electronic micro-scope photographs, to the staff of the Laboratorio deMicrocine of the Sciences Faculty, UNAM for their help inthe drawings processing and to Samantha Rupit for editingthe figures. The two anonymous referees are also thankedfor their comments.

R E F E R E N C E S

Berkeley E. and Berkeley C. (1939) On a collection of Polychaeta, chieflyfrom the West Coast of Mexico. Annals and Magazine of NaturalHistory, Series 11 3, 321–346.

Budaeva N. and Fauchald K. (2011) Phylogeny of the Diopatra genericcomplex with a revision of Paradiopatra Ehlers, 1887 (Polychaeta:Onuphidae). Zoological Journal of the Linnean Society 163, 319–436.

Fauchald K. (1980) Onuphidae (Polychaeta) from Belize, CentralAmerica, with notes on related taxa. Proceedings of the BiologicalSociety of Washington 93, 797–829.

Gonzalez-Ortiz L., Hernandez-Alcantara P. and Solıs-Weiss V. (1997)The Onuphidae of the Gulf of California and the Gulf ofTehuantepec, Mexican Pacific. Bulletin of Marine Science 60, 389–395.

Hernandez-Alcantara P., Tovar-Hernandez M.A. and Solıs-Weiss V.(2008) Polychaetes (Annelida: Polychaeta) described from theMexican Pacific: an historical review and an updated checklist. LatinAmerican Journal of Aquatic Research 36, 37–61.

Paxton H. (1979) Taxonomy and aspects of the life history of Australianbeachworms (Polychaeta: Onuphidae). Australian Journal of Marineand Freshwater Research 30, 265–94.

Paxton H. (1986) Generic revision and relationships of the familyOnuphidae (Annelida: Polychaeta). Records of the AustralianMuseum 38, 1–74.

Paxton H. (1996) Hirsutonuphis (Polychaeta: Onuphidae) from Australia,with a discussion of setal progression in juveniles. InvertebrateTaxonomy 10, 77–96.

Paxton H. (1998) The Diopatra chiliensis confusion—redescription of D.chiliensis (Polychaeta, Onuphidae) and implicated species. ZoologicaScripta 27, 31–48.

and

Rioja E. (1944) Estudios Anelidologicos. XI. Notas sobre algunas especiesde poliquetos de las costas Mexicanas del Pacifico. Anales del Institutode Biologıa, Universidad Nacional Autonoma de Mexico 15, 139–145.

Correspondence should be addressed to:V. Solıs-WeissLaboratorio de Ecologıa y Biodiversidad de InvertebradosMarinos, Instituto de Ciencias del Mar y Limnologıa,Universidad Nacional Autonoma de Mexico, Mexico D.F.04510, Mexicoemail: solisw@cmarl.unam.mx

new hirsutonuphis from the mexican pacific 1783