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Annotated checklist for the Gorgonocephalidae family
(Echinodermata: Ophiuroidea) from the National Echinoderm Collection
“Dra. Ma. E. Caso M.” of the Instituto de Ciencias
del Mar y Limnología, UNAM
Frida Jacqueline Ramírez-Moreno
1
Francisco Alonso Solís-Marín*
2
Alfredo Laguarda-Figueras
2
1. Facultad de Estudios Superiores Iztacala, Universidad Nacional Autónoma de Mexico. Mexico City, Mexico;
fridajacqueline95@gmail.com
2. Colección Nacional de Equinodermos “Dra. Ma. E. Caso Muñoz”, Laboratorio de Sistemática y Ecología de
Equinodermos. Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México. Ciudad de
México, México; fasolis@cmarl.unam.mx (*Correspondence), laguarda@cmarl.unam.mx
Received 02-II-2020. Corrected 12-IX-2020. Accepted 10-XII-2020.
ABSTRACT
Introduction: The order Euryalida is represented by fragile ophiuroids with long and slender arms that can
be ramified. Species within the family Gorgonocephalidae are characterized by the presence of tiny hooklets
on the dorsal aspect of their arms. There are very few published studies referring to species of the fam-
ily Gorgonocephalidae inhabiting Mexican waters. Objective: To review the taxonomic status of the species
belonging to this taxonomic family, identifying their diagnostic taxonomic characteristics to create an illustrated
guide of species kept at the CNE, UNAM. Methods: Specialized bibliography of the Gorgonocephalidae family
was gathered. Subsequently, all the specimens from the CNE were photographed using Scanning Electron and
Multifocal Microscopy photography, to show the structures of taxonomic relevance. Results: A total of 193
specimens corresponding to eight genera and nine species were reviewed. The genus Astrocaneum presented
the highest species richness. Conclusions: For the species identified in this study and for the sake of accurate
identification, morphological characters such as the shape of hooklets and arm plates were highlighted.
Keywords: Basket stars, taxonomy, biodiversity, deep-sea, Mexico.
Ramírez-Moreno, F.J., Solís-Marín, F.A., & Laguarda-
Figueras, A. (2021). Annotated checklist for the
Gorgonocephalidae family (Echinodermata:
Ophiuroidea) from the National Echinoderm
Collection “Dra. Ma. E. Caso M.” of the Instituto de
Ciencias del Mar y Limnologia, UNAM. Revista de
Biología Tropical, 69(S1), 287-303. DOI 10.15517/
rbt.v69iSuppl.1.46360
Current classifications of the Ophiuroidea
recognize two orders, the Euryalida (basket or
snake stars) with 5 families, and the Ophiu-
rida (brittle-stars) with 15 families (O´Hara,
Hugall, Thuy, Stöhr and Martynov, 2018).
Even though the order Euryalida is widely dis-
tributed the specimens are not easy to capture
since many live in the deep sea and display
cryptic behaviors (Hyman, 1955). Most of
the species are epizoic and are found among
gorgonians, antipatharians, and sponges. They
are occasionally associated with hard sub-
strates (Gondim, Dias & Manso, 2012). The
members of this order possess specialized
morphological attributes of the class Ophiuroi-
dea, including the presence of vertebrae with
DOI 10.15517/rbt.v69iSuppl.1.46360
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hourglass-shaped joints (of the streptospondyl-
ous type) that allow them to move their arms
vertically (Hendler and Miller, 1984).
The Gorgonocephalidae is conformed by
34 genera and is the largest of the three families
integrating the order Euryalida (Stöhr, 2015).
The species in this family have five intricately
branched arms, with rings that have few hook-
shaped spines, giving them a banded appear-
ance (Piepenburg, 2000). The disc and the arms
in this genus are covered by small granules,
except on the interradios (Baker, 1980). There
are studies addressing specimens of this family,
like O’Hara, Hugall, Thuy, Stöhr & Martynov
(2018) who reported a new classification with
the morphological diagnosis of the Ophiuroidea
taxa. However, the studies referring to species
of the family Gorgonocephalidae inhabiting
Mexican waters are scarce. An example is the
work published by Solís-Marín and collabora-
tors (2014) in which they only present a list of
echinoderm species found in Mexican waters
with no further analysis or description.
This study will bring together important
and specific information of some gorgonoce-
phalid species, the species under consideration
are distributed along the Pacific and Atlantic
coasts of Mexico, which are home to important
reef systems. The taxonomic information on
this family has been scarce in comparison with
other ophiuroid groups.
MATERIALS AND METHODS
Using the specialized library incorporated
in the Laboratory of Echinoderm Systematics
and Ecology (LESE, UNAM), some literature
specialized in the family Gorgonocephalidae
was gathered. Specimens described herein have
been cataloged and deposited at the Echino-
derm National Collection “Dra. Ma. Elena
Caso M.”. Materials were collected in the
Pacific and Atlantic Oceans between 1964 and
2007 through multiple expeditions (Table 1).
These specimens were preserved dry and in 70
% alcohol. The materials were examined using
an OLYMPUS SZX7 stereoscopic microscope
and pictures were taken using a Leica stereo-
scopic microscope at the Biology Institute
(BI), UNAM. Some specimens were dissected
to analyze and photograph their internal struc-
tures, using Scanning Electron Microscopy
(SEM) JEOL JSM-6360 LV at ICML, UNAM.
The fragments were immersed in commercial
house bleach, and then a 5% NaCIO solution
to remove muscle tissue. Finally, the mounting
was performed in an aluminum sample holder
and carbon labels for SEM. Finally, the photo-
graphs of the taxonomically important struc-
tures used for species identification were edited
using Adobe Photoshop CC and were edited to
add scales and group all the photographs.
The taxonomic terms used to describe the
euryalid brittle stars in this work follow Stöhr,
O’Hara, and Thuy (2012) and Okanishi (2016).
The taxonomic arrangement was made accord-
ing to O’Hara et al. (2018).
Abbreviations used in this work: Colec-
ción Nacional de Equinodermos, CNE. Univer-
sidad Nacional Autónoma de México, UNAM.
RESULTS
A total of 193 specimens were examined,
belonging to eight genera and nine species, with
a bathymetric distribution from 73 to 201 m.
The genus with the highest species richness in
the CNE, UNAM was Astrocaneum (two spe-
cies) and the ones with less specific richness
were Asteroporpa, Astrodictyum, Astracme,
Astrocyclus, Astrogomphus, Astrophyton, and
Gorgonocephalus (with one species each).
Taxonomic synopsis
Class Ophiuroidea Gray, 1840
Order Euryalida Lamarck, 1816
Family Gorgonocephalidae Ljungman, 1867
Genus Asteroporpa Örstedt & Lütken, 1856 in Lütken, 1856
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Asteroporpa (Asteroporpa) annulata (Örstedt & Lütken, 1856 in Lütken, 1856)
Genus Astracme Döderlein, 1927
Astracme mucronata (Lyman, 1869)
Genus Astrocaneum Döderlein, 1911
Astrocaneum herrerai (Clark, 1919)
Astrocaneum spinosum (Lyman, 1875)
Genus Astrocyclus Döderlein, 1911
Astrocyclus caecilia (Lütken, 1856)
Genus Astrodictyum Döderlein, 1927
Astrodictyum panamense (Verrill, 1867)
Genus Astrogomphus Lyman, 1869
Astrogomphus vallatus (Lyman, 1869)
Genus Astrophyton Fleming, 1828
Astrophyton muricatum (Lamarck, 1816)
Genus Gorgonocephalus Leach, 1815
Gorgonocephalus chilensis (Philippi, 1858)
TABLE 1
Species of the family Gorgonocephalidae reviewed at the Colección Nacional de Equinodermos,
Instituto de Ciencias del Mar y Limnología, UNAM
No. of
specimens
Locality Depth Preservation condition
Asteroporpa (Asteroporpa) annulata (Örstedt & Lütken 1856, in Lütken, 1856)
23 North of Olbox, Quintana Roo. 69.2 - 730 m Alcohol
1 North of Cabo Catoche, Quintana Roo. 57.5 m Dry
Astracme mucronata (Lyman, 1869)
5 North Atlantic Ocean, Bahamas, Yucatan Mexico. 305 - 400 m Alcohol
3 North of Progreso, Yucatan, Mexico. 336 m Dry
Astrocaneum herrerai (Clark, 1919)
2 Franco de Contoy, Quintana Roo. 38 m Dry
Astrocaneum spinosum (Lyman, 1875)
1 Bajo Marisal, Gulf of California. 40 m Alcohol
1 Las Ánimas Island, Gulf of California. 60 m Dry
Astrocyclus caecilia (Lütken, 1856)
2 Gulf of Mexico. 87 - 116 m Alcohol
Astrodictyum cf. panamense (Verrill, 1867)
5 The Gulf of California. 30 - 60 m Alcohol
12 Baja California and Oaxaca. 30 - 40 m Dry
Astrogomphus vallatus (Lyman, 1869)
1 Off Río Lagartos, Campeche’s Bank. 215 m Alcohol
Astrophyton muricatum (Lamarck, 1816)
16 Contoy Island, Quintana Roo. 33 m Alcohol
20 Contoy Island, Quintana Roo. 15 - 57 m Dry
Gorgonocephalus chilensis (Philippi, 1858)
1 Puerto Santa Cruz, Bahía Grande, South Atlantic Ocean. 50 m Alcohol
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Order Euryalida Lamarck, 1816
Diagnosis (O’Hara et al., 2018): arm spine
articulations with muscle and nerve openings
well separated. Arm spines ventral only. Lateral
arm plates devoid of ornamentation. Vertebrae
with hourglass-shaped articulations.
Family Gorgonocephalidae Ljungman, 1867
Diagnosis (O’Hara et al., 2018): dorsal
part of the disc and arms with scales and plates,
which bear thorny granules or spines. In the
arms, they present Girdles of small hyaline
hooks. Arms branched or simple. All vertebrae
without an oral bridge. Multiple teeth in irregu-
lar columns, dental plate whole. Arm spine
articulation with slit-shaped muscle opening
proximally bordered by a vertical ridge, small
nerve opening at a distance.
Genus Asteroporpa Örsted & Lütken 1856,
in Lütken, 1856
Diagnosis (Örstedt & Lütken, 1856 in
Lütken, 1856): disc with 10 short radial shields
and five arms, extremely long and tightly
articulated arms that are very narrow at their
ventral side. Both radial shields are elevated
from the arm region and covered by skin with
warts and articulated hooks. Along the borders
of the mouth, they exhibit sharp cone-shaped
papillae. They have short hooklets underneath
the arm rings.
Asteroporpa (Asteroporpa) annulata
(Örstedt and Lütken, 1856 in Lütken, 1856)
Fig. 1A, 1B, 1C, 1D, Fig. 6A, 6B.
Asteroporpa (Asteroporpa) annulata
Örstedt & Lütken, 1856 in Lütken, 1856: 17,
Figs. 4a-d, 5a-b; Lyman, 1883: 125; Verrill,
1899 (b): 320; Döderlein, 1911: 24, Fig. 2a-f;
Döderlein 1927: 89; Gondim et al., 2012: 142-
145, Fig. 1.
Diagnosis (Gondim et al., 2012): aboral
part of the disk covered by a thick skin, the
radial shields and their surrounding are tumid,
bar-like, extending more than half of the disk
diameter, and covered by five rows of rings
with sharp granules. The jaws and oral parts
are covered by a thick skin and show small,
flattened dermal ossicles. There are five simple
arms; the dorsal and lateral surfaces of the arms
are annulated, with inflated, white, and trans-
verse rows of hooklet-bearing plates.
Scanning Electron Microscopy: Asteropor-
pa (Asteroporpa) annulata can be distinguished
by its type of spines, which no other organism
possesses. These spines have a rectangular base
with five to seven spikes, the hooklets have
an open and marked base, with clear edges;
there are six very noticeable perforations found
along the hooklet, at the secondary tooth base
there is a groove, and the main hooklet is long
(Fig. 6A, 6B).
Genus Astracme Döderlein, 1927
Diagnosis (Döderlein, 1927): disc and
arms with thick spines, one madreporite; the
first section of the arms exhibits many rami-
fications. They exhibit small tentacle pores
before the first bifurcation; from one to three
arm spines (small-sized), beginning before or
immediately after the first arm bifurcation; all
the final bifurcation are thin, with hooks and
well-developed hook rows.
Astracme mucronata (Lyman, 1869)
Fig. 2A, 2B, 2C, 2D, Fig. 6D, 6H.
Astrophyton mucronatum Lyman, 1869:
348. Gorgonocephalus mucronatus Lyman,
1882: 265; Verrill, 1899: 85. Astrospartus
mucronatus Döderlein, 1911: 73. Astracme
mucronata Döderlein, 1927: 31.
Diagnosis (Manso, Gondim, & Ventura,
2014): disc covered by flattened plate-shaped
dermal ossicles of varied sizes and shapes,
and some granules. Large radial shields, bar-
like, covered flattened by plate-shaped dermal
ossicles and some granules and spines; there
are irregularly distributed spine-shaped dermal
ossicles on the radial shield, the interradius
is covered by flattened, plate-shaped dermal
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ossicles and some spine-shaped ossicles (simi-
lar to the radial shields). The jaw and oral part
are covered with granules larger than those on
the dorsal surface.
Scanning Electron Microscopy: Astracme
mucronata has spines at the radial shields,
which have a flat point and a narrow base (Fig.
6C). It has three different types of hooklets;
there are marked differences at the base of
each type of hooklet and also in the shape of
it, having one and two secondary teeth (Fig.
6F, 6G, 6H).
Genus Astrocaneum Döderlein, 1911
Diagnosis (Döderlein, 1911): disc and
arms with protuberances or thicker spines. Five
madreporites. The main internal and external
arm branching is lightly extended, the dis-
tal ramifications are thin, thread-shaped and
the lateral ramifications are cone-shaped. The
spine-shaped papillae are found along the arms,
are tiny, and are observable from the third or
fourth bifurcation.
Astrocaneum herrerai (Clark, 1919)
Fig. 2E, 2F, 2G, 2H, Fig. 6I, 6J.
Astrocynodus herrerai A.H. Clark, 1919:
638, Fig. 96. Astrocaneum herrerai.- Döderlein,
1927: 56; Durán-González, Laguarda-Figueras,
Solís-Marín, Buitrón-Sánchez, Ahearn, & Tor-
res-Vega, 2005: 5; Pomory, 2007: 3.
Diagnosis (Clark, 1919): the oral side
of the disc is covered by small granules, at
the external half of the radial shields present
crossed bands that are divided into sections and
can be found diagonally across the shield. At
the interradial oral side of the disc, it is covered
with very small granules, among which there
are bigger and brighter granules, grouped in a
way that they form a mesh-like pattern.
Astrocaneum spinosum (Lyman, 1875)
Fig. 3A, 3B, 3C, 3D, Fig. 6K, 6L, 6M.
Astrophyton spinosum Lyman, 1875: 29.
Astrocaneum spinosum.- Döderlein, 1911: 92;
Döderlein, 1927: 55; Béarez & Bujard, 2013:
2, Figs. 2-3.
Diagnosis (Lyman, 1875): at the aboral
side of the disc they have a small row of spines
that are scattered in groups of two or three, at
the internal angle of each of the oral interradial
spaces there is a madreporite; there are no signs
of arm spines close to the base of the arm, but
they appear at the fifth ramification, where
there are two to three arm spines per each pore;
the tentacle pores are narrowly contracted.
Scanning Electron Microscopy: The genus
Astrocaneum posses oblong spines, however,
between A. herrerai and A. spinosum there are
differences at the tip of the hooklet; A. spino-
sum has a flat tip and is located at the center
of the hooklet (Fig. 6L). The hooklets are the
main characteristic to differentiate all species,
which is why between these species (A. herre-
rai and A. spinosum) there is a morphological
difference at the base of the hooklet, been lon-
ger with a bulky protruding base in A. herrerai
(Fig. 6J).
Genus Astrocyclus Döderlein, 1911
Diagnosis (Döderlein, 1911): the radial
shields and arms show prominent transver-
sal bulges and slightly ramified arms. Five
madreporites at the interior edge of the oral
interradius, the tentacle pores are small and
appear before or immediately after the first
bifurcation. Each hook of the belt posses one
or two secondary teeth; the arm spines are
hook-shaped, they present one to two second-
ary teeth.
Astrocyclus caecilia (Lütken, 1856)
Fig. 3E, 3F, 3G, 3H, Fig. 6N, 6O.
Asterophyton caecilia Lütken, 1856: 18;
Lyman, 1882: 258.
Astrocyclus caecilia.- Tommasi, 1970: 13;
Gondim et al., 2015: 2, Fig. 1.
Diagnosis (Tommasi, 1970): slightly ele-
vated radial shields, without spines or tubercu-
les, only transversal lines; at the center of the
disc there are big, protruding, and elongated
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Fig. 1. Asteroporpa (Asteroporpa) annulata (ICML-UNAM 3.120.1). A. Ventral aspect of the central region of the disc
showing the rings over the radial shields. B. Dorsal aspect of the basal portion of the arm showing the ring. C. Mouth. D.
Ventral aspect of the basal portion of the arm. (a). Arms spines (b). Bursal slit (c). Granules.
granules. Five madreporites, each one located
in an internal angle of the interradial spaces.
The arms at the oral side exhibit a small and
truncated arm spine, at the tip of the arms, the
arm spine transforming into hook-shaped.
Genus Astrodictyum Döderlein, 1927
Diagnosis (Döderlein, 1927): the disc and
arms exhibit bulges or thick spines; the main
ramification of arms is slightly extended, the
distal ramifications are thin and feather-shaped.
There are five madreporites, the arm spines are
uniformly found along the arms, they are small
and begin at a certain distance from the disc,
not before the third or fourth bifurcation.
Astrodictyum cf. panamense (Verrill, 1867)
Fig. 4A, 4B, 4C, 4D, Fig. 7A, 7B, 7C.
Astrophyton panamense Verrill, 1867: 251.
Astrocaneum panamense.- Döderlein, 1911: 95.
Diagnosis (Verrill, 1867): the oral and
aboral surfaces are covered by rough granules,
at the radial shields and the dorsal side of the
arms the granules are round and uniform in
size, but on the ventral side they are flattened
and larger. The aboral surface is slightly con-
vex and has no presence of spines or tubercu-
les. From two to three tentacle pores that are
noted for being short, which begin at the third
or fourth bifurcation; they exhibit small hooks,
visible only at the terminal ramifications.
Genus Astrogomphus Lyman, 1869
Diagnosis (Lyman, 1869): radial shields
narrow and short, the peripheral part presents
numerous spines. Simple arms, the skin is
covered by a mosaic of small granules, the seg-
ments of the arms are distinguished for having
ridges, each one formed by a belt of granules,
some of which are very small. The arm spines
have the shape of a thorny stump, arranged in
groups just above the tentacle pores.
Astrogomphus vallatus Lyman, 1869
Fig. 4E, 4H, Fig. 7D, 7E, 7F, 7G.
Astrogomphus vallatus Lyman, 1869: 350;
Durán-González et al., 2005: 59.
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Fig. 2. Astracme mucronata (ICML-UNAM, 12308). A. detail of the interradi zone with conical spines. B. Distal region of
the radial shields showing a small notch at their base and first fork of the arm. C. Mouth. D. Ventral view of the arm showing
the tentacular pores. Astrocaneum herrerai (ICML-UNAM 3.118.15), E. Detail of the spines arranged in-band over radial
shields. F. Radial shields showing a small notch at their base, arm with bands of spines similar to those of the disc. G. Detail
of the tentacular pores at the first bifurcation. H. Arm spines after the third fork.
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Fig. 3. Astrocaneum spinosum (ICML-UNAM 9821), A. Series of six spines on the arms. B. Dorsal view of the base of
the disc and the arms. C. Mouth. D. Detail of the madreporite and tentacular pores. Astrocyclus caecilia (ICML-UNAM
3.184.1). E. Detail view of the radial shields. F. Dorsal view of the proximal portion of the arm. G. Mouth. H. Distal
ramifications.
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Fig. 4. Astrodictyum cf. panamense (ICML-UNAM 9822). A. Radial shield with a transverse notch. B. Dorsal view of
the branching base of the arms. C. Tentacular pores. D. Ventral view of the arms. Astrogomphus vallatus (ICML-UNAM
3.165.0). E. Aboral views of the disc. F. Arm base and disk periphery. G. Mouth. H. Arm spines from basal arm segments.
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Fig. 5. Astrophyton muricatum (ICML-UNAM 3.165.0). A. detail of the radial shields. B. Dorsal view of the basal arm
segments. C. Mouth. D. Dorsal and ventral view of the distal portion of the arm. Gorgonocephalus chilensis (ICML-UNAM
3.239.0). E. Partial view of the radial shields and the central region of the disc. F. Basal portion of the arm. G. Mouth. H.
Arm spines that extends into the oral disc.
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Fig. 6. SEM photographs of arm spines, hooks, and hooklet-bearing plates of different species of Gorgonocephalidae housed
at ICML. Asteroporpa (Asteroporpa) annulata (ICML-UNAM 30120.1). A. Hook on distal arm. B. Arm spine of the middle
portion of the arm. Astracme mucronata (ICML-UNAM 12308). C. Arm spine. D-E. Arm spine of the middle portion of the
arm. F. Hook located in the middle part of an arm. G-H. Hook on the distal portion of an arm. Astrocaneum herrerai (ICML-
UNAM 3.118.15). I. Arm spines. J. Simple hooks distributed along the arms. Astrocaneum spinosum (ICML-UNAM 9821).
K. External structures: hooklet-bearing plate. L. Arm spine. M. Hook of the arm. Astrocyclus caecilia (ICML-UNAM
3.184.1). N. Hooklet-bearing plate. O. Hook of the arms. Abbreviations: ST. Secondary tooth. sa, Arm spine articulation.
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Diagnosis (Lyman, 1869): the entire aboral
side is filled with short and thick spines, arranged
in concentric rows, the radial shields have a
rough and thorny appearance. Every ramifica-
tion is covered by a belt of four rows of gran-
ules; the oral side of the arms is flat, they have a
smooth skin that is uniformly covered by small
flat granules, that look like a rugged mosaic.
The interradial oral spaces are covered by small,
round, and flat granules of different sizes.
Scanning Electron Microscopy: Astrogom-
phus vallatus presents an elongated slightly
curved hooklet, with four secondary teeth, with
a narrow well-defined base (Fig. 7G).
Fig. 7. SEM photographs of external structures of different species of Gorgonocephalidae housed at ICML. Astrodictyum
cf. panamense (ICML-UNAM 9822). A. Hooklet-bearing plate. B. Distal dorsal plates found. C. Hook of the arm.
Astrogomphus vallatus (ICML-UNAM 3.165.0). D. hooklet-bearing plate. E-F. Arm spines. G. hook of the middle portion
of the arm. Astrophyton muricatum (ICML-UNAM 3.165.0). H. Arm spines. I. Hook of the arm. Gorgonocephalus chilensis
(ICML-UNAM 3.239.0). J. Ventral plates. K. Hooklet-bearing plate. L-M. Arm spines. N. Arm spine. O. Arm spine from
the base of the arm. Abbreviations: ST. secondary tooth. sa. spine articulation.
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Fig. 8. SEM photographs of the hooks and the arm spine of Gorgonocephalus chilensis (ICML-UNAM 3.239.0) A. Hook
from basal portion of the arm. B. Hook from the middle portion of the arm. C. arm spine from the middle portion of arm.
Abbreviations: ST. Secondary tooth.
Genus Astrophyton Fleming, 1828
Diagnosis (Fleming, 1828): ventral disc
covered only by skin, five arms divided dico-
tomically. At the periphery, the genital slit is
long, ranging from the peripheral end of the
oral shield to the lateral edge of the disc.
Astrophyton muricatum (Lamarck, 1816)
Fig. 5A, 5B, 5C, 5D, Fig. 7H, 7I.
Euryale muricatum Lamarck 1816: 538.
Astrophyton arborescens, Müller & Troschel,
1842: 124. Astrophyton muricatum.- Müller &
Troschel, 1842: 122; Döderlein, 1911: 7, 18,
53, 108, Fig. 1; Döderlein, 1927: 24, 25.
Diagnosis (Müller & Troschel, 1842): the
radial shields stand out at the aboral side, they
exhibit a thin granulation with conic spines that
have a sliced tip (Fig. 5A); the arm granula-
tion is thin and dense, covering the entire arm;
the first segment of the arms do not have arm
spine, only the thinner ramifications have two
or three short arm spines. The oral side of the
disc is characterized by having a thick arm
base with few small and fine granules at the
oral interradius, the arms can be exhibit either
tubercules or spines.
Scanning Electron Microscopy: Astrophy-
ton muricatum: the spines and hooklets are of
two types and easily distinguished; one spine
has a rectangular base and a short hooklet (Fig.
7H), meanwhile the other is curved with an
ovoid base (Fig. 7I).
Genus Gorgonocephalus Leach, 1815
Diagnosis (Leach, 1815): it presents a
convex body with five thick and dichotomic
arms, the distal ramifications are thin and more
bifurcated. The species have bursal slits at each
side of the base of the arm’s inferior side.
Gorgonocephalus chilensis (Philippi, 1858)
Fig. 5E, 5F, 5G, 5H, Fig. 7J, 7K, 7L, 7M,
7N, 7O, Fig. 8A, 8B, 8C.
Astrophyton chilense Philippi, 1858: 268.
Astrophyton pourtalesi Lyman, 1875: 28. Gor-
gonocephalus chilensis var. Novaezelandiae
Mortensen 1924: 91. Gorgonocephalus chilen-
sis.- Döderlein 1911: 30, 105, Fig. 5; Döder-
lein, 1927: 30; Monteiro & Tommasi, 1983:
56, figs. 1-6.
Diagnosis (Philippi, 1858): radial shields
protruding; the arms spread five times, and the
final ramifications are rosary-shaped, usually
every bifurcation is divided again into another
segment, but the ramifications are not identical.
It has 12 main ramifications, the arms at the
base get divided three more times.
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Scanning Electron Microscopy: Gorgono-
cephalus chilensis has three types of spines and
four types of hooklets, three of these hooklets
have a secondary tooth, but the shape of the
base of them is different (Fig. 7O, 8A, 8B), the
fourth hooklet exhibits two secondary teeth and
a narrow base (Fig. 8C), the perforations of the
hooklets change in number and position (Fig.
8A, 8B, 8C).
DISCUSSION
There have been published taxonomic lists
which include the gorgonocephalids of the Gulf
of Mexico and Gulf of California (Hernández-
Herrejón, 2007; Laguarda-Figueras, Hernán-
dez-Herrejón, Solís-Marín, & Durán-González,
2009) however, they have focused only on
external structures. Gorgonocephalids reported
in the present study were analyzed using main-
ly internal taxonomic characteristics; this paper
includes two genera and species for the Gulf of
California and seven genera and species for the
Mexican Caribbean and the Gulf of Mexico.
Asteroporpa (Asteroporpa) annulata is the
type species of this genus, it was described by
Örsted & Lütken (1856) in Lütken (1856), this
species was collected in the western islands of
the Caribbean (West Indies) (Örsted & Lütken
in Lütken, 1856), however, the description
based on the specimens collected from these
areas was very poor. They mentioned taxonom-
ic characters of disc highlighting the presence
of irregular tubercles and rows full of hooklet-
bearing plates in the arms. A. (Asteroporpa)
annulata is distinguished from these species:
A. (Asteroporpa) australiensis Clark, 1909, A.
(A.) pulchra Clark, 1915, A. (A.) affinis Lütken,
1859, for having simple arms, a convex disc
and thin arms, thin radial shields with lines
of granules, presence of protruding bands sur-
rounding the arms, oral part of the disc very
granulated and hooklets present throughout the
oral part of the arms.
According to Lyman (1869) and Manso,
Gondim, & Ventura (2014), Astracme mucro-
nata is defined by the following morphological
characters: a thick and conspicuous disc, and
radial shields with numerous spines found in
the aboral interradia. Döderlein (1927) mention
that A. mucronata can be distinguished from A.
mediterraneus by the latter spines being pres-
ent only on the radial shields. We examined
eight specimens collected in Yucatan, Mexico,
these specimens showed a spine (similar to the
disc) on the dorsal side of the arms, present up
until the third arm segment, one per segment;
according to Manso et al. (2014), these speci-
mens correspond to adult individuals.
Astrocaneum herrerai differs from A. spi-
nosum by having a granule-covered disc, with
conical spines arranged in crossed bands at
the radial shields; it is also possible to observe
ringed arms, each ring posses a hooklet bearing
plate with a short and conical spine. Mean-
while, A. spinosum possesses a smooth skin
covering the disc (except for the groups of
spines located between the radial shields), and
the arms possess a group of spines arranged in
rows (Lyman, 1875; Clark, 1919).
According to Baker, Okanishi & Pawson
(2018), Astrocyclus somaliensis differs from
A. caecilian in having the disk partially cov-
ered with more or less continuous, transverse,
thin white lines made up of closely packed
granules, surrounding the radial shields and
partially extending into the inter-shield and
interradial areas; this specie was collected in
the north-eastern coast of Somalia (Indian
Ocean); A. caecilia has been recorded from
the Caribbean Sea, Gulf of Mexico, and from
northeastern Brazil by Gondim, Días & Christ-
offersen (2015).
All the Astrocaneum cf. panamense speci-
mens examined in the present study, exhibited
calcareous plates covered with granules and
spines in crossed bands on top of their radial
shields; the interradial areas have scattered
packs of two to three spines; the arms are
ringed and at their base, there are fragmented
calcareous plates covered with spines. All
the previously stated is not mentioned in the
original description by Verrill (1867). It is very
important to review all these specimens in the
near future; it may be a new species of the
genus Astrocaneum.
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One of the most relevant contributions of
this study is to propose the synonymization of
Astrogomphus rudis Verrill, 1899 under Astro-
gomphus vallatus Lyman, 1869. It is important
to mention that this species was reported by
Laguarda-Figueras et al. (2009) as Astrochele
lymani. However, the material mentioned by
the later authors was reviewed, and their taxo-
nomic description does not agree with Verrill
(1878), they rather fit with the genus Astrogom-
phus Lyman, 1869 which is represented by two
species A. rudis Verrill, 1899 and A. vallatus
Lyman, 1869. At the moment of comparing
the type material of both species we noticed
that the morphological variation was minimal,
Verrill (1899) differentiated the species using
variations in the disk ornamentation, but both
species share the same disk structures, and both
species possess the same ventral ornamenta-
tion, in the oral side, close to the jaws, there
are small “arm spines” distributed in group of
five which also cover the entire ventral side of
the arms.
This is the first study that describes Astrac-
me mucronata, Astrocaneum herrerai, Astro-
caneum spinosum, Astrogomphus vallatus and
Gorgonocephalus chilensis external structures
(hooks, ventral plates and spines) with provid-
ing SEM photographs. Asteroporpa (Astero-
moana) muricatopatella and Asteroporpa
(Asteromoana) koyoae differ from A. (Astero-
porpa) annulata in that the first species has arm
hooks with two lateral teeth, while the second
species has three lateral teeth of the hooklets
located on the basal portion of the ventral arm.
When describing A. somaliensis, Baker et
al. (2018) mention the presence of three types
of spines and two simple hooks of varying
sizes; in the present study it was found that
A. caecilia possesses only simple hooks dis-
tributed along the arm, these structures have
an arched base with three well-defined pores,
these hooks (A. caecilia) are smaller than the
ones in A. somaliensis.
Ethical statement: authors declare that
they all agree with this publication and made
significant contributions; that there is no
conflict of interest of any kind; and that we
followed all pertinent ethical and legal proce-
dures and requirements. All financial sources
are fully and clearly stated in the acknowledge-
ments section. A signed document has been
filed in the journal archives.
ACKNOWLEDGEMENTS
We would like to thank Berenit Mendoza
(Instituto de Biología [IB], UNAM) and Laura
Gómez (ICML, UNAM) for their support tak-
ing the SEM photographs; Susana Guzmán
(Laboratorio de Microscopía y Fotografía de
la Biodiversidad, IB, UNAM) for her guidance
provided taking the multifocal photographies;
Adam J. Baldinger (Museum of Comparative
Zoology, Harvard, USA), for providing pho-
tographs of type material; and Alicia Durán-
González (ICML, UNAM) for her technical
support. Our appreciation goes to Carlos A.
Conejeros Vargas for preparing the figures that
illustrates the present work. To Itzel Rosales,
Francisco Márquez (Posgrado en Ciencias del
Mar y Limnología, UNAM), and Tania Pineda-
Enríquez (University of Florida, U.S.A.) for
their valuable constructive criticism during the
early stages of this work, and to Antonio Tinoco
for his reviews, time spent and support shown
throughout the development of this study.
RESUMEN
Lista anotada de la familia Gorgonocephalidae
(Echinodermata: Ophiuroidea) de la Colección
Nacional de Equinodermos “Dra. Ma. E. Caso M.”
del Instituto de Ciencias del Mar y Limnología, UNAM
Introducción: El orden Euryalida está formado por
ofiuros frágiles con brazos delgados, largos y a veces
ramificados. Las especies de la familia Gorgonocephalidae
se caracterizan por la presencia de diminutos ganchos en
el lado dorsal de sus brazos. Los estudios que refieren
especies de la familia Gorgonocephalidae que habitan
en aguas mexicanas son escasos. Objetivo: Realizar la
revisión taxonómica de dicha familia, reconociendo los
caracteres taxonómicos diagnósticos para generar una guía
ilustrada de cada una de las especies que se encuentran en
la CNE del ICML, UNAM. Métodos: Se recopiló toda la
bibliografía especializada de la familia, posteriormente con
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ayuda de fotografías de MEB y multifocal se fotografiaron
los especímenes de la CNE para destacar las estructuras
de importancia taxonómica. Resultados: se revisaron un
total de 193 ejemplares correspondientes a ocho géneros,
nueve especies, siendo el género Astrocaneum el de mayor
riqueza específica. Conclusiones: Para las especies iden-
tificadas en este estudio, para dar una identificación más
precisa, se destacaron los caracteres morfológicos forma de
los ganchos y forma de las placas de los brazos.
Palabras clave: Estrellas canasta, taxonomía, biodiversi-
dad, mar profundo, México.
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