R. HOUART & M.E.

HENDRICKX NOVAPEX 21(1): 17–33, 10 mars 2020

Three new species of Muricidae (Ocenebrinae, Pagodulinae) from the Gulf

of California, Mexico and update of the living muricids from the area
Roland HOUART1
Research Associate
Institut royal des Sciences naturelles de Belgique
and
Muséum national d'Histoire naturelle, Paris, France, UMR7205 IsyEB
[email protected]

Michel E. HENDRICKX
Laboratorio de Invertebrados Bentónicos, Unidad Académica Mazatlán,
Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México,
P.O. Box 811, Mazatlán, Sinaloa 82000, Mexico
[email protected]
1
Corresponding author

KEYWORDS. Gastropoda, Muricidae, Zacatrophon, Nodulotrophon, Gulf of California,

distribution, new taxa.

ABSTRACT. Three new species of Muricidae are described from the Gulf of California. Two
species, Zacatrophon coani n. sp. and Z. scotti n. sp. are assigned to the subfamily Ocenebrinae
and one, Nodulotrophon pumae n. sp., is assigned to Pagodulinae. The genus Nodulotrophon is
transferred from Trophoninae to Pagodulinae based on radula characters. The total of extant
species of Muricidae in the Gulf is updated to 98 and all are listed here.

RESUME. Trois nouvelles espèces de Muricidae sont décrites du golfe de Californie. Deux
espèces, Zacatrophon coani n. sp. et Z. scotti n. sp., sont assignées à la sous-famille des
Ocenebrinae et une, Nodulotrophon pumae n. sp., est assignée aux Pagodulinae. Le genre
Nodulotrophon est transféré des Trophoninae aux Pagodulinae sur base des caractères de la radula.
Le total des espèces de Muricidae vivant dans le golfe est actualisé à 98 et toutes sont listées ici.

INTRODUCTION Hendrickx & Ayón-Parente 2014, Tovar-Hernández et

al. 2016, Gómez & Díaz 2017, Salgado-Barragán et al.
The Gulf of California, an enclosed sea located on 2017, Berschauer et al. 2018).
the west coast of Mexico, is considered as one of the Since the early 19th century, many expeditions
richest natural ecosystem in the world (Brusca et al. have explored the Gulf of California diverse habitats,
2005). About 1200 km long, it extends from its mouth with special emphasis in shallow waters (Brusca
(at about 22°N) to the Colorado River estuary 1980), but a sizable number of expeditions have also
(Ledesma-Vázquez & Carreño 2010). It includes a focussed, although rather occasionally, on deeper
wide variety of habitats, from the shallow sandy and waters (Schwartzlose & Hendrickson 1983, Hendrickx
rocky beaches, coastal lagoons and estuaries, to the 2012a). Intensive offshore sampling of pelagic and
great depths found in the series of basins and trenches benthic fauna below the continental shelf depth and to
that increase in depths towards its mouth, to a 2500 m in the Gulf of California is, however, very
maximum of about 3000 m at the entrance to the Gulf recent (1989-2011) (Hendrickx 2012a, 2012b).
(Alvarez-Borrego 2010). The affinity of the Gulf of As in other parts of western Mexico, the Gulf of
California fauna is markedly tropical (Hendrickx California is characterized by the presence of a wide,
1992, Brusca & Hendrickx 2010), with a relatively intensive Oxygen Minimum Zone (OMZ) (Hendrickx
high percentage of endemic species: 15.8% overall & Serrano 2010, Papiol & Hendrickx 2016). In the
and about 26% in the case of the Gastropoda core of this OMZ, anoxia or severe hypoxia conditions
(Hendrickx & Brusca 2007). Although many attempts prevail (< 0.2 ml O2/l), thus reducing the number of
have been made in order to divide the Gulf of species able to inhabit certain depth ranges (Gage &
California into faunistic units, distribution patterns of Tyler 1992, Papiol & Hendrickx 2016). The
benthic invertebrates is far from being settled due to progressive fading of the OMZ towards the northern
the frequent discovery of new species and reports on Gulf of California, where it completely disappears in
new distributions (e.g., Massin & Hendrickx 2011, latitudes higher that 28°30'N (Hendrickx & Serrano

17
R. HOUART & M.E. HENDRICKX New species of Muricidae from the Gulf of California

2014), has two major consequences: 1) even in deep Methods

water (i.e. > 200 m) level of oxygen is no longer a
major limiting factor for pelagic and benthic fauna; 2) The characters used to describe shell morphology
the presence of the wide and intensive OMZ south of address the general aspect of the shell, its shape, size
28°30'N represents a physiological barrier that isolate and colour, the shape of the spire including the
the deep fauna of the northern Gulf from the deep number and features of the teleoconch whorls, details
fauna occurring in the central and southern Gulf. of the suture and of the subsutural ramp, details of
During exploratory surveys in the northern Gulf of axial and spiral sculpture, the aperture, the siphonal
California, deep-water invertebrates were collected to canal and, when available, the operculum. The
a maximum depth of 1346 m. Among the material, 12 descriptions are based on the type material.
specimens belong to undescribed species of Regarding the length/width ratio, the width is always
Muricidae. This material is studied in the present measured including the spines.
contribution and three new species are described. The bathymetric ranges given are the inner values of
the recorded depths: the deepest minimum and the
Material and Methods shallowest maximum of each recorded depth range
(Bouchet et al. 2008). The terminology used to
Material describe the spiral cords follows Merle (2001, 2005)
(Fig. 2A-B).
The material reported in this study was obtained while
sampling with the R/V “El Puma”, Universidad
Nacional Autónoma de México, in the northern Gulf
of California, roughly between 28°10’ and 29°10’N
(Fig. 1). A total of 30 stations were visited, with
depths ranging from 148 to 1346 m (Hendrickx,
2012b). Specimens of Muricidae and other benthic
gastropods were collected with a 2.35 m wide by 0.95
m high, standard benthic sledge equipped with an
outer collecting net of ca 5.5 cm (2 1/4") stretch mesh
and an inner net of ca 2.0 cm (3/4") stretch mesh.
Trawling lasted 30 minutes at an average speed of
1.75 knots [approx. 3.25 km/h]. Sampling depths were
estimated with a digital Simrad echo sounder.
Epibenthic temperature and oxygen concentration
were measured ca 10 m above bottom level with a
Seabird CTD-O2 probe. Oxygen concentrations were
also double-checked with the Winkler method using
water samples collected in closing bottles near the
bottom.
The specimens examined, including the holotype and Figure 1. Distribution of the new described species
paratypes, are deposited in the Regional Collection of (●) Zacatrophon coani n. sp.; (■) Zacatrophon scotti
Marine Invertebrates at the Mazatlán Marine Station, n. sp.; (▲) Nodulotrophon pumae n. sp.
UNAM, in Mazatlán, Mexico (ICML-EMU), in the
Institut royal des Sciences naturelles de Belgique Abbreviations
(IRSNB), in Brussels, Belgium, in the Muséum
national d'Histoire naturelle (MNHN) in Paris, France, Repositories
with their respective catalogue number and in the IRSNB: Institut royal des Sciences naturelles de
research collection of the first author. A compilation Belgique, Bruxelles, Belgium;
of species of Muricidae known for the Gulf of ICML-EMU: Instituto de Ciencias del Mar y
California was performed based on Hendrickx et al.
Limnología, UNAM, Mazatlán, Sinaloa 82000,
(2005), on an invertebrates data base available online
(Brusca & Hendrickx 2008 and onward) and on Mexico;
recently published literature. A result of this LACM: Los Angeles County Museum of Natural
compilation is included in this contribution History, California, U.S.A.;
(Appendix). MNHN: Muséum national d'Histoire naturelle, Paris,
The Gulf of California division in three areas France;
proposed by Hendrickx et al. (2005) was used in order NMNZ: Museum of New Zealand Te Papa
to compare species richness among the southern, Tongarewa, Wellington, New Zealand;
central, northern Gulf, and with the Southwestern RH: research collection of Roland Houart;
portion of west Baja California shelf which is SBMNH: Santa Barbara Museum of Natural History,
considered by Brusca & Hendrickx (2010) an California, U.S.A.
extension of the Cortés Zoogeographic Province.
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R. HOUART & M.E. HENDRICKX NOVAPEX 21(1): 17–33, 10 mars 2020

Terminology used to describe the spiral cords 71.7 × 35.6 mm, Baja California, Mexico, Cedros
morphology (Fig. 2) Island, 80 m, date unknown (RH).

P, primary cord; P1, shoulder cord; P2-P6, primary Type locality. Northern Gulf of California,
cords of the convex part of the teleoconch whorl, 28°15'38"N‒111°58'33"W, 208‒212 m.
starting adapically; s6, secondary cord of the convex
part of the teleoconch whorl, below P6. Distribution. Northern Gulf of California and Cedros
Island, northern Baja California, 80‒208 m depth.

Description. Shell large for the genus, up to 86.4 mm

in length at maturity (holotype), length/width ratio
1.8‒2.2. Slender, elongate, narrow, heavy. Subsutural
ramp broad, tabulate, lightly sloping, weakly convex.
Light tan, tan or light cream, occasionally with a
broad, weakly darker or light brown spiral band just
above suture and midway of last teleoconch whorl.
Aperture glossy white or light brown within. Spire
high, up to 7+ narrow, strongly shouldered, spinose
whorls. Earlier teleoconch whorls eroded. Suture
deeply impressed. Protoconch unknown, eroded in all
Figure 2 (spiral sculpture) examined specimens. Axial sculpture of teleoconch
A-B. Nodulotrophon pumae n. sp. Holotype (IMCL- whorls consisting of ribs and low, broad, flattened
EMU-12599), 29.0 × 14.4 mm. lamellae, more strongly developed at shoulder,
producing short or moderately long, broad, flat,
Others guttered, spinelike projections on two or three last
abapical teleoconch whorls; adapical whorls with
more or less rounded ribs. First teleoconch whorl
ISyEB: Institut de Systématique, Évolution, Biodiversité;
eroded in all examined specimens; second whorl with
OMZ: Oxygen Minimum Zone;
9 ribs in holotype; third whorl with 7‒9 rounded ribs;
SEM: Scanning Electron Microscope;
fourth whorl with 9 ribs or lamellate ribs; fifth with
dd: empty shell;
9‒11 flatter, lamellate ribs; sixth with 9‒12 lamellate
lv: collected alive;
ribs; last whorl with 7‒10 low, flat, strongly dorsally
st.: sampling station.
reflected axial, lamellate ribs, generally ending as
long, adapically curved, flat, guttered, open, spine-like
SYSTEMATICS projections at shoulder. Spiral sculpture consisting of
several low or almost indistinguishable, narrow, close-
Family MURICIDAE Rafinesque, 1815 set spiral threads, extending also on lamellae and on
Subfamily OCENEBRINAE Cossmann, 1903 siphonal canal, less apparent on canal. Subsutural
Genus Zacatrophon Hertlein & Strong, 1951 ramp smooth, except axial lamellae and growth striae.
Zacatrophon Hertlein & Strong, 1951: 86. Aperture large, broad, roundly ovate. Columellar lip
Type species by original designation: Trophon broad, entirely smooth, rim completely adherent.
(Boreotrophon) beebei Hertlein & Strong, 1948, Gulf Outer lip erect, smooth, with smooth surface within.
of California (Fig. 6C‒F). Siphonal canal moderately long, 22‒28% of total shell
length, broad, straight or weakly dorsally curved,
Zacatrophon coani n. sp. broadly open, with low axial lamellae over whole
Figs 3A‒J, 4A‒F length. Operculum dark brown with lateral nucleus on
lower right.
Austrotrophon cerrosensis ― Houart, 2010: fig. 16
(only) (not Trophon cerrosensis Dall, 1891). Ecology. (TALUD XIV, st. 1). Zacatrophon coani n.
sp. was collected in 208–212 m depth, on sandy
Material examined. Holotype, 86.4 × 43.0 mm, bottom (sand, 93.3%; silt, 5.7%; clay, 0.9%) with
TALUD XIV, st. 1, northern Gulf of California, 0.56% of organic matter content. Epibenthic
28°15'38"N‒111°58'33"W, 208-212 m, lv, April 07, parameters measured near bottom were: 12.2°C; 34.36
2011 (IMCL-EMU-11261). ups; 0.61 ml O2/l.
Paratypes, 6 specimens from same locality and date,
lv; 35.1 × 17.0 mm (ICML-EMU-11262); 53.3 × 25.9 Remarks. Two species of Zacatrophon are currently
mm (ICML-EMU-12596); 62.5 × 28.0 mm (ICML- known from the Gulf of California: Z. beebei (Hertlein
EMU-12597); 63.3 × 31.1 mm (ICML-EMU-12598); & Strong, 1948) and Z. skoglundae Houart, 2010. At
67.2 × 37.7 mm (IRSNB I.G. 34092/MT.3825); 73.9 × first sight Zacatrophon coani n. sp. resembles Z.
40.6 mm (MNHN-IM-2014-7980); one specimen, skoglundae (Fig. 4G‒L), but Z. coani n. sp. differs in

19
R. HOUART & M.E. HENDRICKX New species of Muricidae from the Gulf of California

Figure 3 (scale bars 10 mm)

A-J. Zacatrophon coani n. sp. A-D. Holotype (IMCL-EMU-11261), 86.4 × 43.0 mm; E-G. Paratype (IRSNB
I.G. 34092/MT.3825), 67.2 × 37.7 mm; H-J. Paratype (MNHN-IM-2014-7980), 73.9 × 40.6 mm.

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R. HOUART & M.E. HENDRICKX NOVAPEX 21(1): 17–33, 10 mars 2020

having broader and lower teleoconch whorls, also Subsutural ramp moderately broad, tabulate, lightly
obviously less loosely coiled whorls and each whorl sloping, weakly convex. Shell entirely light tan.
less abapically constricted. The suture in Z. coani n. Aperture white within. Spire high, up to 6 narrow,
sp. is also less deeply excavated, less impressed and strongly shouldered, spinose whorls. Suture deeply
clearly on a more horizontal plane, while it is strongly impressed. Protoconch unknown, eroded in all
abapically tilted from the first to the last whorl in Z. examined specimens. Axial sculpture of teleoconch
skoglundae. whorls consisting of very low, broad, flattened
Zacatrophon coani n. sp. also differs from Z. beebei lamellae, adherent to shell on abapical part,
(Fig. 6C–F) in having a comparatively much larger inconspicuous up to tip of siphonal canal, but strongly
shell, less loosely coiled and less abapically developed at shoulder, then producing long, broad,
constricted whorls, and more obvious, broad axial flattened, guttered, spine-like projections, abaxially
lamellae ending as broader spinelike projections at bent on penultimate and last whorls; shoulder spines
shoulder, while Z. beebei has obviously narrower on antepenultimate whorl less abaxially bent; other
spines and axial lamellae adherent to the shell. adapical whorls with lamellae or rounded ribs. First
Zacatrophon coani n. sp. also has a broader last teleoconch whorl badly eroded; second whorl partly
teleoconch whorl and a broad, short, siphonal canal as eroded and with some ribs; third whorl with rounded
opposed to an abapically constricted last whorl and a ribs, starting lamellae; fourth with 7 flat, lamellate
narrower, comparatively longer siphonal canal in Z. ribs; fifth with 7 or 8 flat lamellate ribs; last whorl
beebei, reaching 33‒35% of the total shell length. with 9 or 10 low, flat, dorsally reflected, axial,
Zacatrophon coani n. sp. was erroneously reported lamellate ribs, ending as long, adapically curved, flat,
and illustrated as Austrotrophon cerrosensis (Dall, guttered, open, abaxially bent, spinelike projections at
1891) by Houart (2010: fig. 16). Austrotrophon shoulder. Spiral sculpture indistinguishable.
cerrosensis also occurs in the Gulf of California, but Subsutural ramp smooth, except axial lamellae
Z. coani n. sp. differs in having a relatively larger strongly adherent to shell and growth striae. Aperture
shell with a broader and shorter siphonal canal, a moderately large, broadly ovate. Columellar lip
narrower, less sloping subsutural ramp and in having narrow, smooth, rim entirely adherent to shell. Outer
comparatively narrower and more convex teleoconch apertural lip thin, smooth, with smooth surface within.
whorls, opposed to angular and straight in A. Siphonal canal long, 34‒36% of total shell length,
cerrosensis. narrow to moderately broad, straight, broadly open,
with growth striae and strongly adherent, almost
Etymology. This new species of Zacatrophon is invisible, low axial lamellae over whole length.
named after our colleague Dr. Eugene ("Gene") V. Operculum dark brown with lateral nucleus on lower
Coan, in recognition for his outstanding contribution right (Fig. 5E).
to our knowledge of marine mollusks.
Ecology. Zacatrophon scotti n. sp. was collected in
Zacatrophon scotti n. sp. 390–414 m depth, on sandy (sand, 90.4%; silt, 8.5%;
Figs 5A‒M, 6A‒B clay, 1.2%) and silty-sand (sand, 43.5%; silt, 48.9%;
clay, 7.7%) bottoms with 0.94% and 4.86% of organic
Material examined. Holotype, 51.1 × 24.4 mm, matter content, respectively. Epibenthic parameters
TALUD XIV, st. 22, northern Gulf of California, measured near bottom were: 11.39‒11.40°C; 34.90
29°05'27"N‒112°46'44"W, 380-390 m, lv, April 9, ups; 1.92‒1.94 ml O2/l.
2011 (ICML-EMU-11263).
Paratypes, 42.7 × 22.0 mm, TALUD XIV, st. 20, north Remarks. Zacatrophon scotti n. sp. is a strange
of the northern Gulf of California, 28°46'29"N‒ looking species which can probably reach a larger
112°45'40"W, benthic sledge, 414-415 m, lv, April 9, size. The holotype has 5 intact teleoconch whorls but
2011 (IMCL-EMU-12600); 34.7 × 19.5 mm (ICML- is badly eroded adapically.
EMU-12595) and 35.3 × 15.16 mm (RH), TALUD The shell, with a comparatively lower and broader
XIV, st. 22, north of the Gulf of California, spire, less disjunct and less abapically constricted
29°05'27"N‒112°46'44"W, 380-390 m, lv, April 9, teleoconch whorls, a narrower subsutural ramp, long,
2011. broad, flattened, less adapically but strongly abaxially
bent shoulder spines and a relatively long, moderately
Type locality. Northern Gulf of California, narrow, straight siphonal canal is distinguishable from
29°05'27"N‒112°46'44"W, 380‒390 m. Zacatrophon skoglundae (Fig. 4G‒L).
Zacatrophon scotti n. sp. also differs from Z. coani n.
Distribution. Northern Gulf of California, 390‒414 m sp. (Figs 3A‒J, 4A‒F) in having a lighter and fragile
depth. shell with a much lower spire, close set axial lamellae
with strongly abaxially bent spines, as opposed to
Description. Shell medium sized for the genus, up to more largely spaced lamellae with adapically curved
51.1 mm in length (holotype), length/width ratio spines at shoulder in Z. scotti n. sp. and in having a
2.1‒2.3. Slender, elongate, narrow, delicate. much narrower and longer siphonal canal.

21
R. HOUART & M.E. HENDRICKX New species of Muricidae from the Gulf of California

Figure 4 (scale bars 10 mm)

A-F. Zacatrophon coani n. sp. A. Paratype (ICML-EMU-12598), 63.3 × 31.1 mm; B. Paratype (ICML-EMU-
12597), 62.5 × 28.0; C. Paratype (ICML-EMU-12596), 53.3 × 25.9 mm; D. Paratype (ICML-EMU-11262), 35.1
× 17.0 mm; E-F. Paratype (RH), 71.7 × 35.6 mm.
G-L. Zacatrophon skoglundae Houart, 2010, Baja California Sur, Mexico, San Juanico, by shrimpers. G-I.
Holotype (SBMNH 423912), 72 mm (damaged); J-L. (RH), 52.8 × 24.9 mm. Scale bars 10 mm.
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R. HOUART & M.E. HENDRICKX NOVAPEX 21(1): 17–33, 10 mars 2020

Figure 5 (scale bars 10 mm)

A-M. Zacatrophon scotti n. sp. A-E. Holotype (ICML-EMU-11263), 51.1 × 24.4 mm (Fig. E not to scale); F-I.
Paratype (IMCL-EMU-12600), 42.7 × 22.0 mm (tip of siphonal canal broken); J-M. Paratype (RH), 35.3 × 15.6
mm.

23
R. HOUART & M.E. HENDRICKX New species of Muricidae from the Gulf of California

Zacatrophon scotti n. sp. differs from Z. beebei (Fig. Distribution. Only known from the type locality.
6C–F) in many aspects, namely in having a shell with
a lower spire, less disjunct teleoconch whorls, a Description. Shell small for the genus, 29.0 mm in
narrower subsutural ramp, much flatter and close-set length, length/width ratio 1.9. Slender, last teleoconch
axial lamellae with longer, flatter, abaxially bent whorl broadly ovate with long, narrow siphonal canal.
spines at shoulder as opposed to narrower and shorter Strongly spinose at shoulder, lightly built. Subsutural
spines in Z. beebei, and in having a smaller, more ramp moderately broad, weakly sloping, lightly
ovate aperture. convex. Shell entirely white. Spire low. Teleoconch
consisting of 4+, broad, convex, weakly angular,
Etymology. This new species of Zacatrophon is strongly shouldered, spinose whorls. Suture
named after our colleague Dr. Paul Valentisch-Scott, impressed. Protoconch and first teleoconch whorls
in recognition for his outstanding contribution to our eroded. Axial sculpture of teleoconch whorls
knowledge of marine mollusks of the East Pacific, consisting of broad, lamellae, ending as long, strongly
particularly Bivalvia, and for the significant impact he adapically bent, flat, guttered spines at shoulder,
has had on training and helping young malacologists forming coronate teleoconch whorls. Last whorl with
in Mexico. 10 lamellae, penultimate whorl with 11,
antepenultimate with 16 or 17 low, narrow lamellae.
Subfamily PAGODULINAE Barco, Schiaparelli, Lamellae increasing in strength and height abapically.
Houart & Oliverio, 2012 Other axial sculpture of low, weak growth lamellae.
Genus Nodulotrophon Habe & Ito, 1965 Spiral sculpture of low, narrow cords, with visible P1,
Nodulotrophon Habe & Ito, 1965: 32. P2 and P3 from first intact whorl to penultimate (Fig.
Type species by original designation: Trophon dalli 2B). Last whorl with P1, P2, P3, P4, P5, P6, s6 spiral
Kobelt, 1878 (= Trophon coronatum H. & A. Adams, cords (Fig. 2A). Shoulder P1 cord crossed by low,
1864), New Zealand (erroneous) (Japan, Hokkaido; broad, axial lamellae, more obvious on penultimate
Okhotsk Sea to Eastern Bering Sea; Chukchi Sea, and last whorls. P2 cord obvious on antepenultimate
Arctic Ocean; British Columbia, Vancouver) (Fig. 6N- whorl, becoming shallow, almost indistinct on
P). penultimate and last whorls, merging into the lower
part of the shoulder spines, P3 near suture of
Remarks. The inclusion of Boreotrophon and succeeding whorl, ending as narrow, rounded,
Abyssotrophon in Pagodulinae was based exclusively moderately high, squamous cord on last whorl,
on radula morphology of the type species (Houart immediately followed by a similar P4 cord. P5, P6 and
2001; Barco et al. 2012; Houart et al. 2019). The s6 cords indistinguishable on last whorl, only visible
radula of Pagodulinae is muricine with the rachidian as low knobs at edge of aperture. Aperture broad,
tooth bearing a major triangular central cusp, with on roundly ovate. Columellar lip narrow, entirely
each side a small intermediate denticle and a lateral adherent. Outer lip partly broken, smooth within.
cusp slightly smaller than the central one. The Siphonal canal long, 46% of total shell length,
rachidian base is subrectangular, broad, and anteriorly moderately narrow, weakly recurved dorsally, broadly
concave and has no marginal cusps (Fig. 7A). In open, smooth except low, axial growth striae.
Trophoninae the radula is ocenebrine-like with the Ecology. Nodulotrophon pumae n. sp. was collected
rachidian teeth bearing 5 major cusps and 2 small in 1150–1165 m depth, on mixed silty bottom (sand,
denticles, one central cusp, with on each side a small 14.8%; silt, 71.3%; clay, 13.9%) with 7.03% of
denticle, a small lateral cusp and a marginal cusp as in organic matter content. Epibenthic parameters
Trophon Montfort, 1810 (Fig. 7C) and in measured near bottom were: 11.25°C; 34.87 ups; 1.38
Scabrotrophon McLean, 1996 (Fig. 7D). ml O2/l.
The inclusion of Nodulotrophon in Pagodulinae is also
based exclusively on radula morphology of the type Remarks. The choice of the genus and subfamily for
species (Sysoev 1992: 167, Egorov 1993: 4, and this this species is speculative and only based on the shell
paper, Fig. 7B). characters, as nothing is known about the animal,
radula or operculum. Four species are currently
Nodulotrophon pumae n. sp. assigned to Nodulotrophon: N. coronatum, from
Figs 2A‒B, 6G‒J Hokkaido, Japan to the Bering Sea, Alaska to British
Columbia; N. raymondi (Moody, 1916) from Queen
Material examined. Holotype, 29.0 × 14.4 mm, Charlotte Sound, British Columbia [LACM 69-71.11]
TALUD XIV, st. 26, northern Gulf of California, (52° N), to San Clemente Island, California; N.
29°02'11"N‒113°17'12"W, 1150–1165 m, dd, April scolopax (Watson, 1882) from the Kerguelen Islands;
10, 2011 (IMCL-EMU-12599). and N. septus (Watson, 1882) from the Kerguelen and
Crozet Islands. The resemblance of N. pumae n. sp.
Type locality. Northern Gulf of California, with the four other known species of Nodulotrophon is
29°02'11"N‒113°17'12"W, 1150‒1165 m. striking, having a broad, coronate last teleoconch
whorl, a broad aperture, axial lamellae extending as

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R. HOUART & M.E. HENDRICKX NOVAPEX 21(1): 17–33, 10 mars 2020

Figure 6 (scale bars 10 mm)

A-B. Zacatrophon scotti n. sp. Paratype (ICML-EMU-12595), 34.7 × 19.5 mm; C-F. Zacatrophon beebei
(Hertlein & Strong, 1948). Cabo San Lucas, Baja California, Mexico (RH), 43.4 × 20.2 mm; G-J.
Nodulotrophon pumae n. sp. Holotype (IMCL-EMU-12599), 29.0 × 14.4 mm; K-M. Nodulotrophon scolopax
(Watson, 1882). E. Kerguelen Islands, 620-650 m (RH), 25.0 × 12.9 mm; N-P. Nodulotrophon coronatus (H. &
A. Adams, 1864). Okhotsk Sea, S E Sakhalin Islands (RH), 41.2 × 20.2 mm.

25
R. HOUART & M.E. HENDRICKX New species of Muricidae from the Gulf of California

flattened spines at shoulder periphery, a long, narrow, Nodulotrophon, had previously been reported for the
broadly open and almost straight siphonal canal, a Gulf.
narrow spiral shoulder cord and numerous axial A comparative richness of deep-water mollusks (200
growth striae. However, N. pumae n. sp. differs from to 4200 m) occurring in both the Gulf of California
the other species of the genus in having more obvious (East of the Baja California Peninsula) and in the
axial lamellae ending as extremely long, broad spines California Current area (West of the peninsula) was
at shoulder periphery, a quite broader siphonal canal, performed by Zamorano & Hendrickx (2011). Of the
less tapered abapically, and in having obvious P3 and 282 species and 201 genera recorded for both sides of
P4 spiral cords. Nodulotrophon scolopax (Fig. 6K‒M) the peninsula, 202 species and 148 genera were
from the Kerguelen Islands also has spiral cords, but recorded within the Gulf vs. only 118 (104 genera) in
these are more distant from each other and obviously the California Current area, but only 39 species (51
narrower, while the axial lamellae are also less genera) were common to both. The discovery of three
numerous and less obvious with less upward bent and new species within the Gulf would therefore increase
shorter, flat spines at shoulder. the disparity between the two areas. However, new
recent discoveries in deep water along the west coast
Etymology. This new species of Nodulotrophon is of the Baja California Peninsula (Hendrickx et al.
named after the R/V "El Puma" of the Universidad 2016, Suárez-Mozo & Geiger 2017, Suárez-Mozo et
Nacional Autónoma de México, which was the al. 2018, Hendrickx & Suárez-Mozo 2018) clearly
research vessel used during the TALUD XIV compensate for this and might have decreased the
operations in the northern Gulf of California. disparity.
The extant Muricidae are composed of 12 subfamilies
Discussion comprising 205 genera and subgenera and 1820 to
The northern Gulf of California has been extensively 1827 species (Houart 2018, updated). Of these, 10
sampled in the past, but scientists have focused mostly subfamilies and 49 genera or subgenera are present in
on the intertidal zone and on shallow portions of the the Gulf of California (Hendrickx et al. 2005, Brusca
continental shelf where most fishing activities take & Hendrickx 2008 and onward, present contribution).
place (Herrera-Valdivia et al. 2016). The deeper areas In what has remained as the classical reference for the
located near or north of Tiburón Island (ca tropical eastern Pacific mollusks, Keen (1971) listed
29°N‒112°23'W), one of the two large islands 47 species of Muricidae with a geographic range
separating the upper Gulf from the central and limited to the Gulf of California or extending either
southern Gulf, have been rarely explored. Based on south or north of the Gulf. A subsequent work by
the recent exploratory cruise TALUD XIV, during Radwin & D'Attilio (1976) and several additional
which the material reported herein was collected, papers, collated and referenced by Skoglund (2002),
several new species and one new genus have been added some 15 species to this total. Herbert (2005)
discovered in these areas. This includes large and added a new species of Eupleura, E. vokesorum
abundant species of decapod crustaceans (Hendrickx Herbert, 2005 (type locality, Guaymas), while Kaiser
2012c, Hendrickx & Ayón-Parente 2012) and (2005) reported the presence of Tripterotyphis lowei
amphipods (Hendrickx & Ayón-Parente 2014). New lowei (Pilsbry, 1931) at Isla Partida, thus bringing the
data have also been obtained on composition of total of muricids in the Gulf to 64.
meiofauna (Alvarez-Castillo et al. 2018) and of Hendrickx et al. (2005) listed 83 species of Muricidae
decapod crustaceans fauna for the same areas and 9 species of Coralliophilidae, now assigned to
(Hendrickx 2011, 2012b). It is therefore not surprising Muricidae as a subfamily. One species, Plicopurpura
that new taxa within other major groups of marine patula pansa (Gould, 1853) is considered synonym of
species, like the Mollusca, have now been discovered P. columellaris (Lamarck, 1816) by Wellington &
in the northern Gulf of California. Kuris (1983). The total in Hendrickx et al. (2005) is
An extensive study of the deep-water mollusks fauna therefore adjusted to 91, including the 9
of the central Gulf of California by Zamorano et al. coralliophilines. Eupleura limata Dall, 1890, added by
(2006) revealed the presence of 24 species, occurring Herbert (2005), was not included in this list and three
between 731 and 2250 m, including new geographic additional species were more recently described for
and depth records (Zamorano et al. 2007). A the Gulf of California by Wiedrick (2009), Houart
complementary study was also made on two sympatric (2010) and Berschauer et al. (2018), bringing the
species of deep-water Vesicomyidae in the same area updated total of muricids in the Gulf to 95, increased
(Hendrickx & Valentisch-Scott 2018). In a review of here by the three new species, to reach a total of 98
the deep-water (> 200 m depth) mollusks fauna of the species (Tables 1‒10).
Gulf of Californa, Zamorano & Hendrickx (2009) Information in Brusca & Hendrickx (2008 and
reported 614 records corresponding to 225 species. onward) used in this contribution is constantly updated
Overall, richness was lower in the northern Gulf (at 30 and some recent data considered herein might not yet
and 31°N) than in the central and southern Gulf. Their be available in this web page. The list of species
list included 102 Gastropoda but no species belonging proposed here has been elaborated to the best of our
to the genera reported herein, i.e. Zacatrophon and knowledge based on all sources available to date.

26
R. HOUART & M.E. HENDRICKX NOVAPEX 21(1): 17–33, 10 mars 2020

Figure 7. Radulae (scale bars: A–B: 50 µm; C–D. 100 µm).

A. Pagodula echinata (Kiener, 1840), off Capri, Italy (NMNZ) SEM B. Marshall (Pagodulinae);
B. Nodulotrophon coronatus (H. & A. Adams, 1864), Beaufort Sea, Alaska (RH), SEM P. Bouchet
(Pagodulinae); C. Trophon geversianus (Pallas, 1774), Patagonia (RH), SEM P. Bouchet (Trophoninae);
D. Scabrotrophon maltzani (Kobelt & Kuster, 1878), Tutka Bay, Cook Inlet, Alaska (RH), SEM P. Bouchet
(Trophoninae).

The OMZ occurring off western Mexico, including reported herein would have been collected and
most part of the Gulf of California, certainly reported much earlier. Maybe they were but did not
represents a strong limiting factor for species received proper attention or were confused with more
dispersion and settlement (Gage & Tyler 1992, commonly found species. Further exploration within
Hendrickx & Serrano 2010, 2014), thus favoring the depth range visited during this study (i.e. 146 to
organisms tolerant to oxygen-deficient (< 0.2‒0.5 ml 1346 m) and deeper is therefore needed in order to
O2/l) habitats. Among mollusks, crustaceans and increase our knowledge of the deep-water mollusks
echinoderms, many deep-water species occurring in fauna of the Gulf of California.
the area have proved to be able to adapt to such The list of species of Muricidae in the Appendix
stressing environment (Hendrickx et al. 2011, Papiol (Tables 1‒10) includes 98 species. Subfamilies
& Hendrickx 2016, Suárez-Mozo et al. 2018). Among include from three (Pagodulinae) to 22 (Muricopsinae)
the three new species reported herein, only one was species. A total of seven species were not considered
collected in stressing conditions: Z. coani sp. nov in by Hendrickx et al. (2005): one Muricopsinae, five
0.61 ml O2/l. The other two were found in habitats Ocenebrinae (but this includes two of the three new
with dissolved oxygen concentrations of 1.38‒1.94 ml species described herein), and one Pagodulinae (the
O2/l, well above generally recognized mild hypoxic new species of Nodulotrophon) (Tables 5‒7).
conditions (i.e., < 0.5 ml O2/l) (Helly & Levin 2004). When examining the repartition of the 96 species of
As shown in this contribution, the Gulf of California Muricidae (2 species have no precise distribution data)
mollusk fauna inhabiting in water deeper than the among the three sections of the Gulf of California
continental shelf is far from being adequately known. (north, central and south), the central Gulf is clearly
One would have expected that large species like those richer (69 species) than the two other sections (40

27
R. HOUART & M.E. HENDRICKX New species of Muricidae from the Gulf of California

species in the south and 56 in the north). This APPENDIX

coincides with what was concluded by Hendrickx et
al. (2007) who reported a higher richness for the entire The complete list and distribution of Muricidae known
Mollusca group in the central Gulf (1479 species) to occur in the Gulf of California is provided below
compared to the northern (990 species) and southern (Tables 1-10). Distribution of each species is reported
(1376 species) sections of the Gulf. The affinity of the for the Gulf of California and the southern tip of the
Gulf of California Muricidae with the Southern tip of Baja California Peninsula as follows: NGC, Northern
the Baja California Peninsula is very low, and only 11 Gulf of California; CGC, Central Gulf of California;
of the species listed for the Gulf also occur in that SGC, Southern Gulf of California; SWB,
area. Southwestern Baja California shelf. Sources are as
follows: (1) Hendrickx et al. 2005; (2) Herbert 2005;
ACKNOWLEDGEMENTS (3) Wiedrick 2009; (4) Houart 2010; (5) Berschauer et
We thank the crew members, scientists and students al. 2018. New records are indicated in "bold face".
who took an active part in the TALUD XIV cruise. Two subfamilies of Muricidae are absent in the Gulf
Shiptime aboard the R/V "El Puma" was provided by of California: Haustrinae Tan, 2003 and Trophoninae
the Coordinación de la Investigación Científica, Cossmann, 1903. The geographical distribution of
UNAM. Thanks to Bruce A. Marshall, Museum of Haustrinae, which comprise two genera, Haustrum
New Zealand Te Papa Tongarewa, Wellington, New Perry, 1811 and Bedeva Iredale, 1924, is limited to
Zealand for the image of the radula of Pagodula New Zealand and South Australia. However, due to
echinata and to Philippe Bouchet, Muséum national maritime transport, Bedeva is now found in many
d'Histoire naturelle, Paris, France for the preparation parts of the world as an invasive species, as far as the
and SEM work of the other radulae in the eighties. We Canary Islands (Houart & Abreu, 1994), but not in the
thank José Salgado-Barragán for calling our attention Gulf of California. Species of Trophoninae are widely
on the material of the TALUD XIV cruise reported distributed but none has so far been recorded in the
herein and for taking and editing photographs of Central eastern Pacific and in the Gulf of California.
Figures 3H-J; 4A-D; 5A-I and 6A-B. We also thank The classification used here follows largely
Mercedes Cordero for preparing the map of Figure 1 MolluscaBase (2018) with a few exceptions on
and Christiane Delongueville, Roland Scaillet and generic level.
Claude Vilvens for their comments on the manuscript.

APPENDIX. Distribution of Muricidae in the Gulf of California

TABLE 1. Aspellinae Keen, 1971 (8 species)

SPECIES NGC CGC SGC SWB Ref.

Aspella pollux Radwin & D'Attilio, 1976 x 1
Aspella pyramidalis (Broderip, 1833) x x 1
Attiliosa nodulosa (A. Adams, 1855) x x 1
Dermomurex (Dermomurex) indentatus (Carpenter, 1857) x x 1
Dermomurex (Dermomurex) myrakeenae (Emerson & D'Attilio, x x 1
1970)
Dermomurex (Dermomurex) obeliscus (A. Adams, 1853) x 1
Dermomurex (Dermomurex) cunninghamae (Berry, 1964) x x 1
Dermomurex (Gracilimurex) bakeri (Hertlein & Strong, 1951) x 1

TABLE 2. Coralliophilinae Chenu, 1859 (9 species)

SPECIES NGC CGC SGC SWB Ref.

Babelomurex hindsi (Carpenter, 1857) x x x 1
Coralliophila aspera (A. Adams, 1855) No data 1
Coralliophila californica (A. Adams, 1855) No data 1
Coralliophila costata (Blainville, 1832) x x x 1
Coralliophila macleani Shasky, 1970 x x x 1
Coralliophila monodonta (Blainville, 1832) = Quoyula x x 1
madreporarum
Coralliophila nux (Reeve, 1846) x x 1
Coralliophila orcuttiana Dall, 1919 x x 1
Coralliophila parva (E.A. Smith, 1877) x 1

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R. HOUART & M.E. HENDRICKX NOVAPEX 21(1): 17–33, 10 mars 2020

TABLE 3. Ergalataxinae Kuroda, Habe & Oyama, 1971 (3 species)

SPECIES NGC CGC SGC SWB Ref.

Claremontiella adiakritos Houart, Zuccon & Puillandre, 20191 x x 1
Pascula rufonotata (Carpenter, 1864) x x 1
Trachypollia lugubris (C.B. Adams, 1852) x x x 1
1
As Pascula ferruginosa in Hendrickx et al. (2005)

TABLE 4. Muricinae Rafinesque, 1815 (17 species)

SPECIES NGC CGC SGC SWB Ref.

Calotrophon turritus (Dall, 1919) x 1
Hexaplex (Trunculariopsis) brassica (Lamarck, 1822) x x 1
Hexaplex (Trunculariopsis) princeps (Broderip, 1833) x x x 1
Muricanthus ambiguus (Reeve, 1845) x x 1
Muricanthus nigritus (Philippi, 1845) x x x 1
Muricanthus radix (Gmelin, 1791) x 1
Phyllocoma scalariformis (Broderip, 1833) x x 1
Phyllonotus erythrostomus (Swainson, 1831) x x x 1
Phyllonotus peratus Keen, 1960 x x 1
Phyllonotus regius (Swainson, 1821) x x 1
Purpurellus macleani (Emerson & D'Attilio, 1969) x x 1
Purpurellus pinniger (Broderip, 1833) x x 1
Vokesimurex elenensis (Dall, 1909) x x 1
Vokesimurex lividus (Carpenter, 1857) x x 1
Vokesimurex recurvirostris (Broderip, 1833) x x 1
Vokesimurex ruthae (Vokes, 1988) x 1
Vokesimurex tricoronis (Berry, 1960) x x 1

TABLE 5. Muricopsinae Radwin & D'Attilio, 1971 (22 species)

SPECIES NGC CGC SGC SWB Ref.

Acanthotrophon carduus (Broderip, 1833) x x 1
Acanthotrophon sorenseni (Hertlein & Strong, 1951) x x x 1
Bizetiella carmen (Lowe, 1935) x x 1
Bizetiella shaskyi Radwin & D'Attilio, 1972 x 1
Favartia diomedaea (Dall, 1908) x 1
Favartia erosa (Broderip, 1833) x 1
Favartia humilis (Broderip, 1833) x x x 1
Favartia keenae (Vokes, 1970) x 1
Favartia lappa (Broderip, 1833) x x x 1
Favartia laurae (Vokes, 1970) x x 1
Favartia perita (Hinds, 1844) x x x 1
Favartia vittata (Broderip, 1833) x 1
Murexsul armatus (A. Adams, 1854) x 1
Murexsul mildredae (Poorman, 1980) x x 1
Murexsul skoglundae (Myers, Hertz & D'Attilio, 1993) x 1
Murexsul tulensis (Radwin & D'Attilio, 1976) x 1
Muricopsis mcleani Wiedrick, 2009 x 3
Muricopsis pauxilla (A. Adams, 1854) x x 1
Muricopsis zeteki Hertlein & Strong, 1951 x x 1
Pazinotus advenus Poorman, 1980 x 1
Pygmaepterys poormani (Radwin & D'Attilio, 1976) x 1
Vitularia salebrosa (King, 1832) x x 1

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R. HOUART & M.E. HENDRICKX New species of Muricidae from the Gulf of California

TABLE 6. Ocenebrinae Cossmann, 1903 (17 species)

SPECIES NGC CGC SGC SWB Ref.

Austrotrophon cerrosensis (Dall, 1891) x x 1
Calcitrapessa leeana (Dall, 1890) x 1
Ceratostoma nuttalli (Conrad, 1837)2 x 1
Eupleura limata Dall, 1890 x 2
Eupleura muriciformis (Broderip, 1833) x x 1
Eupleura nitida (Broderip, 1833) x 1
Eupleura triquetra (Reeve, 1844) x x 1
Forreria corteziana Berschauer, Petuch & Clark, 2018 x 5
Mexacanthina angelica (Oldroyd, 1918) 1
Mexacanthina lugubris (Sowerby I, 1822) x x x 1
Pteropurpura centrifuga (Hinds, 1844) x x x 1
Pteropurpura erinaceoides (Valenciennes, 1832) x x 1
Pteropurpura macroptera (Deshayes, 1839) x 1
Zacatrophon beebei (Hertlein & Strong, 1948) x x 1
Zacatrophon coani n. sp. x
Zacatrophon scotti n. sp. x
Zacatrophon skoglundae Houart, 2010 x 4
2
as Ceratostoma unicorne in Hendrickx et al. (2005) (= Murex unicornis Reeve, 1849), a synonym of C.
nuttalli.

TABLE 7. Pagodulinae Barco, Schiaparelli, Houart & Oliverio, 2012 (3 species)

SPECIES NGC CGC SGC SWB Ref.

Abyssotrophon lorenzoensis (Durham, 1942) x 1
Nodulotrophon pumae n. sp. x
Trophonopsis diazi (Durham, 1942) x 1

TABLE 8. Rapaninae Gray, 1853 (9 species)

SPECIES NGC CGC SGC SWB Ref.

Acanthais brevidentata (Wood, 1828) x 1
Acanthais triangularis (Blainville, 1832) x x 1
Neorapana muricata (Broderip, 1832) x x 1
Neorapana tuberculata (Sowerby I, 1835) x x x 1
Plicopurpura columellaris (Lamarck, 1816)3 x x x x 1
Stramonita biserialis (Blainville, 1832) x x x 1
Thaisella kiosquiformis (Duclos, 1832) x x x x 1
Tribulus planospira (Lamarck, 1822) x 1
Vasula speciosa (Valenciennes, 1832) x x x 1
3
Purpura pansa Gould, 1853 is a junior synonym.

TABLE 9. Tripterotyphinae D'Attilio & Hertz, 1988 (5 species)

SPECIES NGC CGC SGC SWB Ref.

Cinclidotyphis myrae DuShane, 1969 x 1
Pterotyphis fimbriatus (A. Adams, 1854) x 1
Tripterotyphis arcana (DuShane, 1969) x 1
Tripterotyphis fayae (Keen & Campbell, 1964) x 1
Tripterotyphis lowei lowei (Pilsbry, 1931) x x x 1

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R. HOUART & M.E. HENDRICKX NOVAPEX 21(1): 17–33, 10 mars 2020

TABLE 10. Typhinae Cossmann, 1903 (5 species)

SPECIES NGC CGC SGC SWB Ref.

Haustellotyphis cumingii (Broderip, 1833) x 1
Typhina latipennis (Dall, 1919) x 1
Typhisala clarki (Keen & Campbell, 1964) x x x 1
Typhisala grandis (A. Adams, 1855) x 1
Typhisopsis coronatus (Broderip, 1833) x x x 1

REFERENCES Brusca, R.C. & Hendrickx, M.E. 2010. Invertebrate

Biodiversity and Conservation in the Gulf of
Alvarez-Borrego, S. 2010. Physical, Chemical, and California. Chapter 4, pp. 72–95. In: Brusca, R.C.
Biological Oceanography of the Gulf of California. (ed.) The Gulf of California. Biodiversity and
Chapter 2, pp. 24–48. In: Brusca, R.C. (ed.). The Conservation. Arizona-Sonora Desert Museum
Gulf of California. Biodiversity and Conservation. Studies in Natural History. ASDM Press and
Arizona-Sonora Desert Museum Studies in Natural University of Arizona Press, 354 pp.
History. ASDM Press and University of Arizona Egorov, R.V. 1993. Trophoninae (Muricidae) of
Press, 354 pp. Russian and adjacent waters. Ruthenica, suppl. 1:
Alvarez-Castillo, L., Hermoso-Salazar, M., Estradas- 1–48.
Romero, A., Rivas, G. & Prol-Ledesma, R.M. Gage, J.D. & Tyler, P.A. 1992. Deep-Sea Biology: A
2018. Composition and spatial distribution of the Natural History of Organisms at the Deep-Sea
meiofauna in the Wagner and Consag basins, Gulf Floor. Cambridge University Press, 504 pp.
of California, Mexico. Cahiers de Biologie Gómez, S. & Díaz, K. 2017. On some new species of
Marine, 59(3): 245‒256. Ancorabolidae Sars, 1909 from the Gulf of
Barco, A., Schiaparelli, S., Houart, R. & Oliverio M. California: the genera Ceratonotus Sars, 1909, and
2012. Cenozoic evolution of Muricidae (Mollusca, Dendropsyllus Conroy-Dalton, 2003 (Crustacea,
Neogastropoda) in the Southern Ocean, with the Copepoda, Harpacticoida). ZooKeys, 657: 43–65.
description of a new subfamily. Zoologica Scripta. Habe, T. & Ito, K. 1965. New genera and species of
The Norwegian Academy of Science and Letters, shells chiefly collected from the North Pacific.
41(6): 596–616. Venus, 24(1): 16–45.
Berschauer, D.P., Petuch, E.J. & Clark, R.N. 2018. A Helly, J.J. & Levin, L.A. 2004. Global distribution of
new species of Forreria (Gastropoda: Muricidae: naturally occurring marine hypoxia on continental
Ocenebrinae) from the Sea of Cortez, Mexico. The margins. Deep Sea Research Part I: Oceanographic
Festivus, 50(3): 139–146. Research Papers, 51(9): 1159‒1168.
Bouchet, P., Héros, V., Lozouet, P. & Maestrati, P. Hendrickx, M.E. 1992. Distribution and zoogeographic
2008. A quarter-century of deep-sea malacological affinities of decapod crustaceans of the Gulf of
exploration in the South and West Pacific: Where California, Mexico. Proceedings of the San Diego
do we stand? How far to go? In: Héros, V., Cowie, Society of Natural History, 20: 1–11.
R. H. & Bouchet, P. (eds), Tropical Deep-Sea Hendrickx, M.E. 2011. Distribution and abundance of
Benthos 25. Mémoires du Muséum national Neocrangon resima (Rathbun, 1902) (Crustacea:
d’Histoire naturelle 196: 9–40. Decapoda: Caridea: Crangonidae) collected during
Brusca, R.C. 1980. Common intertidal invertebrates the TALUD XIV cruise in the Gulf of California,
of the Gulf of California. 2d edition. The Mexico. Nauplius, 19 (2): 129‒134.
University of Arizona Press. Tucson, AZ, USA, Hendrickx, M.E. 2012a. Operaciones oceanográficas
513 pp. en aguas profundas: los retos del pasado, del
Brusca, R.C., Findley, L.T., Hastings, P.A., presente y del proyecto TALUD en el Pacífico
Hendrickx, M.E., Torre Cosio, J. & van der mexicano (1989-2009), pp. 23-104. In: P.
Heiden, A.M. 2005. Macrofaunal diversity in the Zamorano, M.E. Hendrickx & M. Caso (eds.)
Gulf of California, pp. 179–203. In: Cartron, J.-L. Biodiversidad y comunidades del talud continental
E., G. Caballos & R.S. Felger (eds.) Biodiversity, del Pacífico mexicano. Secretaría del Medio
Ecosystems and Conservation in Northern Mexico. Ambiente y Recursos Naturales (SEMARNAT),
Oxford University Press, 514 pp. Instituto Nacional de Ecología (INE), 468 pp.
Brusca, R.C. & Hendrickx, M.E. 2008 and onward. http://www2.ine.gob.mx/publicaciones/libros/690/
The Gulf of California Invertebrate Database: The operaciones.pdf
Invertebrate Portion of the Macrofauna Golfo Hendrickx, M.E. 2012b. Pandalid shrimp (Crustacea:
Database. Decapoda: Caridea: Pandalidae) collected during
http://www.desertmuseum.org/center/seaofcortez/d the TALUD XIV cruise in the Gulf of California,
atabase.php (accessed October 2019). Mexico, and rediscovery of Plesionika

31
R. HOUART & M.E. HENDRICKX New species of Muricidae from the Gulf of California

carinirostris Hendrickx, 1989. Cahiers de Biologie Biología Marina y Oceanografía, 53(2): 251‒260.
Marine, 53: 495–504. https://doi.org/10.22370/rbmo.2018.53.2.1297.
Hendrickx, M.E. 2012c. Distribution and abundance Hendrickx, M.E., Valentich-Scott, P. & Suárez Mozo,
of the pelagic processid, Processa pippinae N.Y. 2016. Deep-water bivalve mollusks collected
Wicksten & Méndez, 1985 (Crustacea, Decapoda, during the TALUD XV cruise off the west coast of
Caridea, Processidae), collected during the the southern Baja California Peninsula, Mexico.
TALUD XIV cruise in the Gulf of California, Biodiversity Data Journal, 4: 1‒22. doi:
Mexico, and description of a new genus. 10.3897/BDJ.4.e8661
Crustaceana, 85 (4‒5): 463‒472. Hendrickx, M.E. & Valentich-Scott, P. 2018. Notes on
doi:10.1163/156854012X623773 two sympatric species of deep-water Vesicomyidae
Hendrickx, M.E. & Ayón-Parente, M. 2012. First (Mollusca, Bivalvia) from the central Gulf of
record of Prionocrangon Wood Mason & Alcock, California, western Mexico. Pan-American
1981 (Crustacea: Decapoda: Caridea: Crangonidae) Journal of Aquatic Sciences, 13(3): 216‒225.
in the East Pacific and description of a new deep- Herbert, G. 2005. Systematic revision of the genus
water species from western Mexico. Zootaxa, Eupleura H. and A. Adams, 1853 (Gastropoda:
3205: 63‒68. Muricidae) in the Neogene to Recent of Tropical
Hendrickx, M.E. & Ayón-Parente, M. 2014. Two new America. The Veliger, 47(4): 294–331.
species of deep-water Caprella (Peracarida: Herrera-Valdivia, E., López-Martínez, J., Castillo
Amphipoda: Caprellidae) from the Pacific coast of Vargasmachuca, S. & García-Juárez, A. R. 2016.
Mexico collected during the TALUD XIV cruise, Taxonomic and functional diversity of the bycatch
with a checklist of species of Caprellidea recorded fishes community of trawl fishing from Northern
for the East Pacific. Crustaceana, 87(1): 41–63. Gulf of California, Mexico. Revista de biologia
doi: 10.1163/15685403-00003277. Tropical, 64(2): 587‒602.
Hendrickx, M.E., Brusca, R. C. & Findley, L.T. (eds.) Hertlein, L.G. & Strong, A.M. 1951. Eastern Pacific
2005. A Distributional Checklist of the expeditions of the New York Zoological Society.
Macrofauna of the Gulf of California, Mexico. Mollusks from the west coast of Mexico and
Part I. Invertebrates [Listado y Distribución de la Central America. Pt. 10, vol. 36(2): 67–120.
Macrofauna del Golfo de California, México, Houart, R. 2001. A review of the Recent
Parte I. Invertebrados]. Arizona-Sonora Desert Mediterranean and Northeastern Atlantic species
Museum and Conservation International, 429 pp. of Muricidae. Evolver, 227 pp.
Hendrickx, M.E. & Brusca, R.C. 2007. Distribución Houart, R. 2010. A remarkable new species of
de invertebrados marinos endémicos en el golfo de Zacatrophon Hertlein & Strong, 1951
California, México, pp. 1–4. In: Memorias del XII (Gastropoda: Muricidae: Ocenebrinae) from the
Congresso Latino-Americano de Ciêncas do Mar. Gulf of California. Novapex, 11(1): 21–27.
XII COLACMAR, Florianópolis, 15 a 19 de abril Houart, R. 2018. Historique et classification des
de 2007. (Resumen 10097 y Texto en CD). espèces actuelles de Muricidae (Neogastropoda,
Hendrickx, M.E., Brusca, R.C., Cordero, M. & Muricoidea). Novapex, 19(2): 37–66.
Germán Ramírez R. 2007. Marine and brackish- Houart, R. & Abreu, A.D. 1994. The Muricidae
water molluscan biodiversity in the Gulf of (Gastropoda) from Madeira with the description of
California, Mexico. Scientia Marina, 71 (4): a new species of Ocenebra (Ocinebrina)
637‒647. (Muricidae: Ocenebrinae). Apex 9(4): 119-130.
Hendrickx, M.E., Mah, C. & Zárate Montes, C.M. Houart, R., Vermeij, G. & Wiedrick, S. 2019. New
2011. Deep-water Asteroidea (Echinodermata) taxa and new synonymy in Muricidae
collected during the TALUD cruises in the Gulf of (Neogastropoda: Pagodulinae, Trophoninae,
California, Mexico. Revista Mexicana de Ocenebrinae) from the Northeast Pacific.
Biodiversidad, 82: 798‒824. Zoosymposia, 13: 184–241.
Hendrickx, M.E. & Serrano, D. 2010. Impacto de la Kaiser, K.L. 2005. Notes on Pterotyphis lowei lowei
zona de mínimo de oxígeno sobre los corredores (Pilsbry, 1931) with spawn mass (Gastropoda:
pesqueros en el Pacífico mexicano. Interciencia, Muricidae). The Festivus, 37(6): 63–64.
35(1): 12–18. Keen, A.M. 1971. Sea Shells of Tropical West
Hendrickx, M.E. & Serrano, D. 2014. Effects of the America, 2d ed. Stanford University Press,
oxygen minimum zone on squat lobsters Stanford, California, i-xiv + 1064 pp.
distribution in the Gulf of California, Mexico. Ledesma-Vázquez, J. & Carreño, A.L. 2010. Origin,
Central European Journal of Biology, 9(1): 92– Age, and Geological Evolution of the Gulf of
103. doi: 10.2478/s11535-013-0165-6 California. Chapter 1, pp. 7-23. In: Brusca, R.C.
Hendrickx, M.E. & Suárez-Mozo, N.Y. 2018. (ed.) The Gulf of California. Biodiversity and
Distribution and abundance of Bathybembix Conservation. Arizona-Sonora Desert Museum
bairdii (Dall, 1889) (Mollusca: Gastropoda: Studies in Natural History. ASDM Press and
Calliotropidae) off the west coast of the Baja University of Arizona Press, 354 pp.
California Peninsula and SW Mexico. Revista de

32
R. HOUART & M.E. HENDRICKX NOVAPEX 21(1): 17–33, 10 mars 2020

Massin, C. & Hendrickx, M.E. 2011. Deep-water Suárez Mozo, N.Y., Valentich-Scott, P. & Hendrickx,
Holothuroidea (Echinodermata) collected during M.E. 2018. Deep-water bivalves from the Oxygen
the TALUD cruises off the Pacific coast of Minimum Zone area off the western Peninsula of
Mexico, with the description of two new species. Baja California, Mexico. Molluscan Research,
Revista Mexicana de Biodiversidad, 82: 413–443. 39(2): 99‒109.
Merle, D. 2001. The spiral cords and the internal https://doi.org/10.1080/13235818.2018.1508550.
denticles of the outer lip in the Muricidae: Sysoev, A.V. 1992. A new hadal species of
terminology and methodological comments. Boreotrophon (Gastropoda, Muricidae) from
Novapex, 2(3): 69–91. Northwestern Pacific. Ruthenica, 2(2): 166–167.
Merle, D. 2005. The spiral cords of the Muricidae Tovar-Hernández, M.A., Villalobos-Guerrero, T.F.,
(Gastropoda, Neogastropoda): importance of Kupriyanova, E.K. & Sun, Y. 2016. A new fouling
ontogenetic and topological correspondences for Hydroides (Annelida, Sabellida, Serpulidae) from
delineating structural homologies. Lethaia, 38: southern Gulf of California. Journal of the Marine
367–379. Biological Association of the United Kingdom,
MolluscaBase 2018. Muricidae Rafinesque, 1815. 96(3): 693–705.
http://www.molluscabase.org/aphia.php?p=taxdeta Wellington, G.M. & Kuris, A.M. 1983. Growth and
ils&id=148 Accessed on 2019-10-30. shell variation in the tropical eastern Pacific
Papiol, V. & Hendrickx, M.E. 2016. Community intertidal genus Purpura: ecological and
structure and ecology of deep-water decapod evolutionary implications. Biological Bulletin,
crustaceans below the Oxygen Minimum Zone in 164: 518–535.
the SE Gulf of California, Mexico. Marine and Wiedrick, S.G. 2009. Description of a new Muricopsis
Freshwater Research, 67(12): 1862–1879. species (Gastropoda: Muricidae: Muricopsinae)
doi.org/10.1071/MF15040 from the Panamic Province. Novapex, 10(2): 47–
Radwin, G. & D'Attilio, A. 1976. Murex shells of the 52.
world. An illustrated guide to the Muricidae. Zamorano, P. & Hendrickx, M.E. 2009. Análisis
Stanford University Press, Stanford, 284 pp. latitudinal y batimétrico de la comunidad de
Salgado-Barragán, J., Ayón-Parente, M. & Zamora- moluscos de mar profundo en el golfo de
Tavares, P. 2017. New records and description of California, México. Brenesia, 71–72: 41‒54.
two new species of caridean shrimps from Bahía Zamorano, P. & Hendrickx, M.E. 2011. State of
Santa María-La Reforma lagoon, Gulf of knowledge about the community of mollusks on
California, Mexico (Crustacea, Caridea, Alpheidae both sides of the Baja California Peninsula,
and Processidae). ZooKeys, (671): 131‒153. Mexico: A comparative analysis. Cahiers de
Schwartzlose, R. A. & Hendrickson, J. R. 1983. Biologie Marine, 52: 13‒22.
Bibliografía del Golfo de California: Ciencias Zamorano, P., Hendrickx, M.E. & Toledano
Marinas (hasta el final de 1981). Anales del Granados, A. 2006. Distribution and ecology of
Instituto de Ciencias del Mar y Limnología, deep water mollusks from the continental slope,
UNAM, Mexico. Publ. Esp. 7: 1–212. southeastern Gulf of California, Mexico. Marine
Skoglund, C. 2002. Panamic province molluscan Biology, 150 (5): 883‒892.
literature additions and changes from 1971 through Zamorano, P., Hendrickx, M.E. & Toledano
2001. The Festivus, 33, suppl., i-xi: 1–286. Granados, A. 2007. New geographic and depth
Suárez-Mozo, N.Y. & Geiger, D.L. 2017. A new records for deep-water mollusks in the Gulf of
species of Fissurellidae (Gastropoda: California, Mexico. Nuevos registros geográficos y
Vetigastropoda) from the deep-sea off the eastern batimétricos para moluscos de mar profundo en el
Pacific coast of Mexico. Nautilus, 131(4): Golfo de California, México. Revista Mexicana de
240‒245. Biodiversidad, 78: 311‒318.

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