Mar Biodiv (2017) 47:167–177
DOI 10.1007/s12526-016-0459-4
ORIGINAL PAPER
Ophiuroidea (Echinodermata) from the Central Mexican Pacific:
an updated checklist including new distribution records
R. Granja-Fernández 1 & A. P. Rodríguez-Troncoso 2 & M. D. Herrero-Pérezrul 3 &
R. C. Sotelo-Casas 2 & J. R. Flores-Ortega 4 & E. Godínez-Domínguez 5 & P. Salazar-Silva 6 &
L. C. Alarcón-Ortega 2 & A. Cazares-Salazar 7 & A. L. Cupul-Magaña 2
Received: 16 December 2015 / Revised: 21 January 2016 / Accepted: 4 February 2016 / Published online: 27 February 2016
# Senckenberg Gesellschaft für Naturforschung and Springer-Verlag Berlin Heidelberg 2016
Abstract The Central Mexican Pacific is an oceanographic
transitional region with complex habitats and important conservation areas, but knowledge of its Ophiuroidea fauna is
limited. A total of 61 localities on a variety of substrata were
sampled between 2008 and 2014 using different methodology
techniques. Twenty-four species were collected and members
of the families Ophiocomidae, Ophiotrichidae and
Ophiactidae were the most widespread. The new records of
28 species have relevance in terms of filling distribution gaps
along the Mexican Pacific or extending their geographical
distribution ranges. This considerable number can be attributed to a higher number of prospected localities and the diversification of collecting methods. An updated checklist from
the study area is provided, including previous literature records and those found during this work, summarising 57 species. Islas Marías was the area with the highest number of
species and with the highest value of taxonomic distinctness,
which can be explained since these islands are considered as
stepping stones for the dispersal of species in the Mexican
Pacific. Despite the important diversity and composition of
ophiuroids in our study area, a sample-based rarefaction curve
suggests that at least 104 species can inhabit the area, so we
recommend conducting more research in the region.
Keywords Brittle stars . Checklist . New distribution
records . Substrata . Taxonomic distinctness
Introduction
The Ophiuroidea is the largest group among the
Echinodermata, with 2064 described species worldwide
(Stöhr et al. 2012). The Mexican Pacific possesses 125 species
of ophiuroids, where the less diverse and explored are the
areas of Jalisco (21), Nayarit (20), Michoacán (nine), Colima
(eight) and Isla Isabel (nine), all of them located at the Central
Mexican Pacific (CMP) (Granja-Fernández et al. 2015a). This
region has a high marine biodiversity potential, as it is
Communicated by: P. Martinez Arbizu
* R. Granja-Fernández
beckygranja@gmail.com
1
Doctorado en Ciencias Biológicas y de la Salud, Universidad
Autónoma Metropolitana, San Rafael Atlixco 186, Col. Vicentina,
09340 Ciudad de México, D.F., México, México
2
Laboratorio de Ecología Marina del Centro de Investigaciones
Costeras, Centro Universitario de la Costa, Universidad de
Guadalajara, Av. Universidad 203, 48280 Puerto Vallarta, Jalisco,
México
3
Centro Interdisciplinario de Ciencias Marinas, Instituto Politécnico
Nacional, Ave. Instituto Politécnico Nacional s/n Col. Playa Palo de
Santa Rita, 23096 La Paz, BCS, México
4
Unidad Académica Escuela Nacional de Ingeniería Pesquera,
Universidad Autónoma de Nayarit, Bahía de Matanchén Km 12,
63740 San Blas, Nayarit, México
5
Departamento de Estudios para el Desarrollo Sustentable de Zonas
Costeras, Universidad de Guadalajara, V. Gómez Farías 82,
48980 San Patricio-Melaque, Jalisco, México
6
Laboratorio de Zoología, Instituto Tecnológico de Bahía de
Banderas, Crucero a Punta de Mita SN, 63763 La Cruz de
Huanacaxtle, Nayarit, México
7
Departamento de Acuicultura, Laboratorio de Microalgas, Centro de
Investigación Científica y de Educación Superior de Ensenada
(CICESE), Carretera Ensenada-Tijuana 3918, Zona Playitas,
22860 Ensenada, Baja California, México
168
considered an oceanographic transitional area that combines
fauna from the North and the South regions of the Mexican
Pacific. Moreover, the CMP is complex due to the presence of
extensive bays, coastal lagoons, estuaries, mangroves and coral reefs (Arriaga-Cabrera et al. 1998); furthermore, the CMP is
important in conservation terms because it includes National
Parks, Biosphere Reserves, Sanctuaries and Marine Priority
Regions (Arriaga-Cabrera et al. 1998; Comisión Nacional de
Areas Naturales Protegidas, CONANP 2005, 2007a, b).
The study of the ophiuroids in the CMP region began when
Lütken and Mortensen (1899) recorded for the first time, from
Islas Marías and at depths up to 1230 m, the species Amphiura
serpentina, Astrodia excavata, Dougaloplus notacanthus,
Ophiacantha costata, Ophiacantha hirta, Ophiacantha
moniliformis, Ophiernus adspersus annectens, Ophiomitra
granifera, Ophiomitra partita, Ophiura (Ophiura) scutellata
and Ophiomusium variabile. Later, Clark (1940) contributed
with the study of the ophiuroids of the region, adding the first
records for Jalisco (Diopederma daniana, Ophioderma
variegata, Ophiolepis variegata) and Colima (Ophiolepis
variegata, Ophiothrix (Ophiothrix) spiculata), all of them distributed in shallow waters. In the same year, Ziesenhenne
(1940) described the new species Amphichondrius laevis,
Amphiodia sculptilis, Ophiolepis plateia and
Ophiophragmus lonchophorus, with their type locality in
Tenacatita Bay, Jalisco. Caso (1951) recorded for the first time
at the CMP (Puerto Vallarta, Jalisco) the presence of Ophiactis
savignyi, Ophiocoma aethiops, Ophiocoma alexandri,
Ophiolepis pacifica, Ophioderma teres and Ophionereis
annulata. Nowadays, the contributions on ophiuroids have
been mostly focused on ecology and checklists from specific
geographical areas. In this regard, Fuentes-Farías et al. (2005)
provided the first records of ophiuroids (D. daniana, O.
aethiops, O. alexandri, O. annulata, O. savignyi, Ophiactis
simplex, Ophiothela mirabilis, O. (O.) spiculata and
Ophiothrix (Ophiothrix) rudis) from Michoacán, all of them
being the only species recorded to date for that state. Other
important additions to the knowledge of ophiuroids in terms
of taxonomy and new distribution records in the CMP were
those by Luke (1982), Caso (1986) and Granja-Fernández et
al. (2014, 2015a).
It is important to note that most of the studies on ophiuroids
in the CMP had been mostly conducted in specific areas with a
status of protection (Nayarit and Jalisco), such as Isla Isabel
National Park, Islas Marías Biosphere Reserve (CONANP
2005, 2007a, b; Ríos-Jara et al. 2008a, b; Granja-Fernández
et al. 2014, 2015a), Tenacatita Bay and Chamela Bay, which is
considered as the first marine sanctuary in the Mexican
System of Natural Protected Areas (Ziesenhenne 1940;
López-Uriarte et al. 2009; Ríos-Jara et al. 2013; GranjaFernández et al. 2014). Despite the above, there are areas such
as Colima, Michoacán, the coast of Nayarit and Islas Marietas
National Park that have been remained almost unstudied. It is
Mar Biodiv (2017) 47:167–177
important to highlight that most of the recent works that mention the presence of ophiuroids in the CMP are not exclusive
of this taxon, since they provide general checklists of echinoderms and/or marine invertebrates, and their sampling effort is
not conducted to collect especially ophiuroids (López-Uriarte
et al. 2009; Ríos-Jara et al. 2008a, b, 2013). Due to the scarce
numbers of studies focused specifically on ophiuroids, to the
sampling effort conducted in just specific areas and because
the CMP has biotic and abiotic mixed conditions with important protected areas, the goal of this work is to fill the gap of
the knowledge of the Ophiuroidea in the CMP, providing an
updated checklist, as well as important biological-ecological
and geographical features of the species.
Materials and methods
Study area
The surveyed area comprises the continental shelf off
the coast of Nayarit, Jalisco, Colima and Michoacán
and their islands (Islas Marías, Isla Isabel, Islas
Marietas), in the central portion of the Mexican Pacific
(between 17°N and 22°N). The area is considered a
conservation hotspot as it includes National Parks (Isla
Isabel and Islas Marietas), Biosphere Reserves (Islas
Marías), Sanctuaries (islands from Chamela Bay) and
Marine Priority Regions (Bahía de Banderas,
Mismaloya-Punta Soledad, Chamela-El Palmito, Punta
Graham-El Carrizal, Cuyutlán-Chupadero, MaruataColola) (Arriaga-Cabrera et al. 1998; CONANP 2005,
2007a, b).
The water masses of the CMP are exposed to the
transisthmic wind jets on the southwestern coast of
México, where a pool of warm water is generated
(Kessler 2006). The water circulation in the CMP has
an annual variability characterised by the convergence
of three current systems: (1) the cold California Current
flowing to the Southeast, (2) the warm Coastal Costa
Rica Current flowing northward and the warm water
from the Gulf of California (Badán 1997), and (3) a
semi-annual variability determined by average northwesterly flow during summer and a northeasterly flow during
winter (De la Lanza 1991). The area has a narrow continental shelf with a width of only 7–10 km; the bottom
of the continental shelf is irregular and increases abruptly
towards the Middle American Trench that extends along
the continent. The maxim depth in the CMP is up to
3500 m. The CMP is influenced by cyclones, tropical
storms and ENSO (El Niño-Southern Oscillation) events,
which produce abnormal increases and decreases in the
mean temperature along the region (Filonov et al. 2000).
Mar Biodiv (2017) 47:167–177
Sampling
A total of 61 localities from the states of Nayarit (12 on the
coast of the state, five in Islas Marietas, three in Islas Marías,
two in Isla Isabel), Jalisco (24), Colima (10) and Michoacán
(five) were sampled between 2008 and 2014 (Table 1, Fig. 1).
This work takes its results from a larger multi-disciplinary and
inter-institutional programme assessing Ophiuroidea along
the Mexican Pacific. The collecting methods were different,
but all of them were next to coral and/or rocky reefs
environments.
Ophiuroids are usually associated with other organisms;
therefore, to avoid damaging healthy coral branches when
collecting specimens, we used only dead natural fragmented
branches of the coral Pocillopora spp. (~5 cm long) as recruitment substrata. Clean fragments were attached with plastic
cables to two semi-spherical concrete structures with six steel
rods, each placed on a sandy bottom at 5 m depth, next to the
healthy Pocillopora coral colonies. After two months, the
fragments were collected and packed into plastic bags, and
all the collected organisms were separated and taxonomically
identified. This sampling was conducted only in Islas
Marietas.
During 2009–2014, in the South of Jalisco (7–27 m depth),
organisms associated with sand and rock were collected using
the following sampling method. Each month, with the help of
a suction pump, 15 cm of sand and surface of rocks were
pumped in an area of 50 × 50 cm (three replicates per station).
The pump consisted of a PVC tube (180 cm long and 7.5 cm
diameter) attached to a stainless steel diffusor (30 cm long)
and connected to an air compressor. In the extreme of the tube,
a bag with a mesh size of 0.5 mm was attached. The samples
were sieved on the field with a mesh of 0.5 mm and packed
into plastic bags for further identification. In 2014, in Nuevo
Vallarta, Nayarit, samples on sand were obtained with the help
of a Van Veen dredger of 2500 cm3 (3–15 m depth).
Finally, collections were carried out in a variety of
available substrata (algae, gorgonians, hydrozoan, live
and dead stony corals, rhodoliths, rock, sand and
sponges) in all sampling sites and at different depths (0
to 11 m). The sampling was carried out by hand collection with scuba diving. All the collected ophiuroids were
anaesthetised using magnesium chloride buffered on seawater in order to prevent autotomy; afterwards, all the
specimens were fixed and preserved in 70 % ethanol.
The specimens were identified after Lütken (1856),
Verrill (1867), Nielsen (1932), Ziesenhenne (1937,
1940), Clark (1939), Hendler et al. (1995), Hendler
(2011) and Granja-Fernández et al. (2014). The specimens were deposited in the Colección de Equinodermos
de la Universidad Autónoma Metropolitana-Iztapalapa,
Colección de Invertebrados del Instituto Tecnológico de
Bahía de Banderas, Colección de Invertebrados del
169
Laboratorio de Ecología Marina del Departamento de
Estudios para el Desarrollo Sustentable de Zonas
Costeras, Universidad de Guadalajara, and in the
Colección Nacional de Equinodermos BDra. María
Elena Caso Muñoz^, Instituto de Ciencias del Mar y
Limnología, Universidad Nacional Autónoma de México.
Analysis data
With all the collected specimens, a matrix of presence/
absence per locality and a database with information specifying locality, substrata, depth and qualitative abundance
of the species were elaborated. Due to the different
collecting methods used during this study, a qualitative
estimate of the abundance consisted merely of ranking
the taxa within samples into a series of abundance classes,
which, in this case, were employed as: very abundant
(>1500 organisms), abundant (700–401 organisms), common (400–100 organisms) and rare (<50 organisms). An
updated checklist including the collected species during
this work was elaborated and completed with literature
records (Lütken and Mortensen 1899; Clark 1940, 1970;
Ziesenhenne 1940; Caso 1951, 1986; Luke 1982;
CONANP 2005, 2007a, b; Fuentes-Farías et al. 2005;
Maluf and Brusca 2005; Honey-Escandón et al. 2008;
Ríos-Jara et al. 2008a, b, 2013; López-Uriarte et al.
2009; Hendler 2011; Granja-Fernández et al. 2013, 2014,
2015a) in order to provide a complete checklist of the
ophiuroids inhabiting the CMP. The valid names and the
systematics are according to Granja-Fernández et al.
(2015a).
In order to determine if the observed species richness
of ophiuroids from the CMP is complete, a sample-based
rarefaction curve was performed using the non-parametric
species richness estimator Chao2. For statistical purposes,
the localities were considered as replicates inside each area
(coast of Nayarit, Colima, Jalisco, Michoacán, Islas
Marías, Isla Isabel, Islas Marietas). The expected number
of the species as well as its confidence interval (95 %)
was determined with a randomised order of samples with
replacement for 1000 runs (Colwell et al. 2004). The analysis was performed with the software EstimateS 9.1.
The average taxonomic distinctness (Δ+) and its variation (Λ+) were estimated in order to have a better understanding of the Ophiuroidea diversity along the CMP.
This index evaluates the taxonomic distance between each
pair of individuals, defined by a Linnean classification
tree (species, genus, family, orders, class) and can be used
with a presence/absence matrix without taking into consideration the sampling methodology (Clarke and
Warwick 2001). The analysis was performed with the
PRIMER-E 7 software.
170
Table 1 Geographic coordinates
of prospected areas along the
Central Mexican Pacific
Mar Biodiv (2017) 47:167–177
No.
Area
Locality
1
ISA
Las Monas
21°51′06.00″ - 105°52′47.00″
2
3
4
ISA
ISA
MAR
Bahía Rabijuncos
Las Pozas
Caleta, María Cleofas
21°50′35.49″ - 105°52′50.94″
21°50′29.00″ - 105°53′03.00″
21°19′12.70″ - 106°13′21.90″
5
MAR
Cleofas, María Cleofas
21°18′10.00″ - 106°13′30.00″
6
7
MAR
NAY
Carrizal, María Cleofas
Careyeros
21°17′32.70″ - 106°13′52.60″
20°46′10.00″ - 105°30′48.00″
8
NAY
El Anclote
20°46′15.19″ - 105°31′03.02″
9
10
NAY
NAY
Punta de Mita
La Lancha
20°46′02.60″ - 105°31′25.90″
20°45′34.70″ - 105°29′19.20″
11
NAY
El Tizate
20°45′31.30″ - 105°21′56.90″
12
NAY
NAY
Playa Paladium
Punta Pelícanos
20°45′05.46″ - 105°27′17.22″
20°45′02.32″ - 105°22′23.94″
NAY
NAY
NAY
Playa del CRIP
La Manzanilla
Bajo de las Viudas
20°44′42.30″ - 105°22′58.96″
20°44′34.52″ - 105°23′13.75″
20°43′59.90″ - 105°23′58.70″
17
18
19
20
21
22
23
24
25
26
27
28
MAT
MAT
MAT
MAT
MAT
NAY
JAL
JAL
JAL
JAL
JAL
JAL
Playa del Amor, Isla Redonda
Plataforma Pavonas, Isla Redonda
Túnel Amarradero, Isla Redonda
Área de Restauración, Isla Larga
Cueva del Muerto, Isla Larga
Nuevo Vallarta
Muelle de los Muertos
Los Arcos
Colomitos
Punta Caletas
Pelícanos
Isla Pajarera
20°42′14.10″ - 105°33′50.10″
20°42′05.20″ - 105°33′54.00″
20°41′58.10″ - 105°33′57.10″
20°41′55.38″ - 105°34′51.77″
20°41′50.60″ - 105°34′58.10″
20°41′24.31″ - 105°17′51.12″
20°36′02.22″ - 105°14′21.45″
20°32′44.83″ - 105°17′34.79″
20°30′44.07″ - 105°19′33.81″
20°30′23.46″ - 105°23′34.79″
19°33′27.00″ - 105°06′28.00″
19°33′23.87″ - 105°06′42.84″
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
JAL
JAL
JAL
JAL
JAL
JAL
JAL
JAL
JAL
JAL
JAL
JAL
JAL
JAL
JAL
JAL
JAL
JAL
La Palma
Isla Cocinas
Tenacatita poza de marea
Tenacatita zona arrecifal
Tenacatita intermareal
La Calechosa
La Virgencita
Centro de Caleta de Cuastecomates, zona somera
Laboratorio
Centro de Caleta de Cuastecomates, zona profunda
Zona de arena cercano a la zona de arrecife
Cuastecomate
Zona de arrecife cercano a la zona de playa
Cuastecomatito
Frente al Hotel Náutico-El Dorado
Frente a la Laguna del Tule
Las Monjas
Morro Chino, Laguna de Barra de Navidad
19°33′02.00″ - 105°06′23.00″
19°32′46.00″ - 105°06′28.00″
19°16′53.58″ - 104°52′21.66″
19°16′49.62″ - 104°52′21.24″
19°16′45.99″ - 104°52′21.83″
19°14′18.29″ - 104°44′18.01″
19°14′09.00″ - 104°44′57.00″
19°14′05.21″ - 104°44′06.59″
19°13′52.50″ - 104°45′21.34″
19°13′46.90″ - 104°44′27.28″
19°13′46.54″ - 104°44′07.80″
19°13′45.33″ - 104°43′59.18″
19°13′44.06″ - 104°44′07.23″
19°13′12.90″ - 104°43′28.02″
19°13′09.75″ - 104°42′25.73″
19°13′01.89″ - 104°41′55.81″
19°12′54.54″ - 104°42′55.68″
19°11′41.91″ - 104°41′35.68″
47
48
49
COL
COL
COL
La Audiencia 2
La Boquita, zona somera
La Boquita, poza de marea
19°06′17.99″ - 104°21′06.63″
19°06′17.56″ - 104°23′53.58″
19°06′12.57″ - 104°23′48.87″
13
14
15
16
Coordinates: latitude
north - longitude west
Mar Biodiv (2017) 47:167–177
Table 1 (continued)
171
No.
Area
Locality
Coordinates: latitude
north - longitude west
50
51
52
COL
COL
COL
La Boquita
La Audiencia 1
Playa Veleros
19°06′10.08″ - 104°23′37.92″
19°06′09.87″ - 104°21′01.15″
19°06′02.58″ - 104°21′03.76″
53
COL
L′Recif
19°06′01.00″ - 104°24′24.00″
54
COL
Punto B
19°05′55.00″ - 104°23′24.00″
55
56
57
COL
COL
MICH
Carrizales 2
Carrizales 1
Morro de Enmedio
19°05′48.30″ - 104°26′07.24″
19°05′47.05″ - 104°26′20.10″
18°21′11.00″ - 103°31′02.00″
58
MICH
Isla Pájaros
18°21′09.00″ - 103°31′09.00″
59
60
MICH
MICH
Faro de Bucerías
Morro Chino
18°21′08.00″ - 103°30′58.00″
18°11′56.00″ - 103°08′10.00″
61
MICH
Caleta de Campos
18°04′11.00″ - 102°44′37.00″
ISA Isla Isabel; MAR Islas Marías; NAY coast of Nayarit; MAT Islas Marietas; JAL Jalisco; COL Colima; MICH
Michoacán
Fig. 1 Study area representing
the 61 localities along the Central
Mexican Pacific. a Isla Isabel; b
Islas Marías; c Islas Marietas; d
coast of Nayarit and Jalisco; e, f
South coast of Jalisco; g Colima;
h Michoacán. The numbers
correspond to those listed in
Table 1
172
Mar Biodiv (2017) 47:167–177
Results
A total of 24 species of Ophiuroidea from 61 localities from
the CMP were collected (Table 2). The species were classified
in seven families (Amphiuridae, Ophiotrichidae, Ophiactidae,
Ophionereididae, Ophiocomidae, Ophiodermatidae and
Ophiolepididae) and 16 genera (Amphiodia, Amphipholis,
M i c ro p h i o p h o l i s , O p h i o c n i d a , O p h i o p h r a g m u s ,
Ophiostigma, Ophiothela, Ophiothrix, Hemipholis,
Ophiactis, Ophionereis, Ophiocoma, Ophiocomella,
Ophioderma, Ophiuroconis and Ophiolepis). The families
with the highest number of species were Amphiuridae
(eight) and Ophiodermatidae (four); the family
Ophionereididae was represented by one species (Table 2).
At the species level, the ophiuroids Ophiocoma alexandri,
Ophiocoma aethiops, Ophiothrix (Ophiothrix) rudis,
Ophiothrix (Ophiothrix) spiculata, Ophiothela mirabilis,
Ophiactis savignyi, Ophiactis simplex and Ophionereis
annulata were the most widespread in the CMP and were
collected in a higher type of substrata (i.e. algae, live and dead
stony coral, gorgonian, rock, sponge) (Table 2). On the other
hand, Amphiodia sculptilis, Ophiophragmus lonchophorus,
Ophiophragmus marginatus, Hemipholis gracilis,
Ophioderma sp. and Ophiuroconis bispinosa were collected
in just one locality. It is important to note that the following
species were found inhabiting only one substrate, sand: A.
sculptilis, Microphiopholis platydisca, O. lonchophorus, O.
marginatus, H. gracilis, O. bispinosa, Ophiolepis crassa,
Table 2 Species of ophiuroids collected during this study, including geographical and ecological features. The numbers of the localities correspond to
those in Table 1
Species
Localities
Amphiodia sculptilis
38
sd
27
R
Amphipholis squamata
8, 12, 13, 20, 21, 31, 33, 36, 48, 55
al, cr, dc, rk, sd
0–18
R
Microphiopholis platydisca
22, 55
sd
4
R
Ophiocnida hispida
34, 37, 42, 43, 46, 52
dc, rk, sd
3–15
R
Ophiophragmus lonchophorus
22
sd
9–10
R
Ophiophragmus marginatus
22
sd
3–6
C
Ophiophragmus papillatus
12, 19, 41
rk
6–7
R
Ophiostigma tenue
19, 39, 41
rk, sd
7–8
R
Ophiothela mirabilis
16, 17, 18, 19, 20, 21, 23, 24, 25, 26, 28, 34, 36, 37, 40, 42, 47,
50, 52, 55, 56, 58
5, 6, 8, 9, 10, 11, 12, 14, 15, 16, 34, 41, 61
gr, sp
0–18
VA
al, cr, gr, rk
1.5–11
A
1, 2, 14, 16, 17, 18, 19, 20, 21, 22, 23, 24, 25, 26, 27, 28, 29,
34, 35, 37, 39, 40, 41, 42, 45, 46, 47, 48, 49, 50, 51, 52, 55,
56, 57, 58, 59, 60
22
al, cr, dc, gr, rk, sp
0–15
A
sd
3–15
R
al, cr, dc, gr, rk, rd, sp
0–9
A
Ophiocomella schmitti
1, 2, 7, 9, 11, 12, 13, 14, 15, 16, 18, 19, 20, 21, 24, 25, 27, 28, 29,
30, 33, 34, 35, 37, 41, 42, 48, 50, 52, 55, 56, 58, 59
2, 3, 11, 14, 16, 17, 18, 19, 20, 21, 22, 23, 24, 25, 26, 27, 28, 29,
30, 31, 34, 35, 36, 37, 39, 40, 41, 42, 46, 50, 52, 56, 58, 59, 60
5, 6, 7, 8, 9, 10, 11, 13, 14, 15, 19, 20, 27, 28, 31, 33, 37, 40, 42,
45, 46, 48, 49, 50, 52, 54, 55, 56, 57
4, 5, 6, 7, 8, 9, 11, 12, 13, 14, 15, 18, 19, 20, 21, 25, 27, 28, 30,
31, 33, 34, 35, 37, 40, 41, 42, 45, 46, 47, 48, 49, 50, 51, 52, 53,
54, 55, 56, 57, 58, 59
1, 2, 5, 6, 7, 8, 9, 12, 13, 14, 15, 18, 19, 20, 21, 26, 27, 28, 29, 30, 31,
32, 33, 34, 35, 36, 37, 39, 40, 41, 42, 45, 46, 47, 48, 49, 50, 51, 52,
53, 54, 55, 56, 57, 58, 59
7, 9, 11
Ophioderma panamensis
8, 9, 14, 15, 19, 20, 21, 31, 33, 37, 40, 42, 45, 47, 48, 49, 51, 52
Ophioderma teres
47, 51, 56
Ophiothrix (Ophiothrix) rudis
Ophiothrix (Ophiothrix)
spiculata
Hemipholis gracilis
Ophiactis savignyi
Ophiactis simplex
Ophionereis annulata
Ophiocoma aethiops
Ophiocoma alexandri
Substrata
Depth
(m)
Qualitative
abundance
al, cr, dc, gr, hz, rd, rk, sd, sp 0–11
VA
sd, rk
0–15
C
al, cr, dc, hz, rk
0–15
C
cr, dc, gr, hz, rd, rk, sd
0–15
A
al, cr, sp
R
rk, sd
0.10–
0.30
0–10
C
rk
0–2
R
Ophioderma sp.
5
rk
9.6
R
Ophiuroconis bispinosa
36
sd
18
R
Ophiolepis crassa
18, 19, 20, 21
sd
5–10
R
Ophiolepis pacifica
5, 19, 21
sd
5.5–10.8
R
Substrata = al algae; cr stony coral; dc dead stony coral; gr gorgonian; hz hydrozoan; rd rhodolithe; sd sand; rk rock; sp sponge
Qualitative abundance = A abundant; C common; R rare; VA very abundant
Mar Biodiv (2017) 47:167–177
Ophiolepis pacifica; rock: Ophiophragmus papillatus,
Ophioderma teres and Ophioderma sp. According to the qualitative abundances obtained during this study, two species can
be considering as very abundant, four as abundant, four as
common and 14 as rare (Table 2).
Nayarit and Jalisco were the areas with the highest number
of species (16), followed by Colima (12) and Michoacán
(eight). Regarding the islands, 13 species were collected at
Islas Marietas, six at Islas Marías and four at Isla Isabel.
Islas Marietas localities represented the highest number of
species of the entire region: Túnel Amarradero (12), Área de
Restauración (10) and Cueva del Muerto (10) (Table 2).
According to previous records of ophiuroids from the CMP,
we found a total of 28 new distribution records: ten from Islas
Marietas (Amphipholis squamata, O. panamensis, O. crassa,
O. papillatus, O. tenue, O. mirabilis, O. annulata, O. savignyi,
O. simplex and O. pacifica), one from Islas Marías
(Ophioderma sp.), six from the coast of Nayarit (A. squamata,
H. gracilis, Ophiocomella schmitti, O. panamensis, O.
loncophorus and O. papillatus), five from Jalisco (A.
squamata, O. papillatus, Ophiostigma tenue, Ophiothrix
(Ophiothrix) rudis and Ophiuroconis bispinosa) and six from
Colima (A. squamata, Microphiopholis platydisca,
Ophiocnida hispida, Ophioderma panamensis, O. teres and
O. mirabilis).
The number of collected species during this work and the
records from the literature sum a total of 57 from the CMP
(Table 3). For the entire study area, according to the estimator
Chao2 and its confidence intervals, the expected number of
species was higher (up to 104) than the number of observed
species (57), indicating that the checklist of ophiuroids from
this area is not yet complete (Fig. 2). The taxonomic distinctness index (Δ+) indicated that all the areas comprising the
CMP are inside the 95 % confidence limit; in this regard, the
coast of Nayarit showed a value near to the lower limit and the
Islas Marías displayed a value above the upper limit of confidence (Fig. 3). The results indicated that Islas Marías had the
highest value of taxonomic distinctness, as well as the highest
number of species of all the areas in the CMP, which suggests
that these islands are represented by different taxonomical
groups (genera). If we compare the areas of the CMP with
that of the Southern Mexican Pacific (Guerrero and Oaxaca)
in terms of taxonomic distinctness and the number of species
of ophiuroids, the analysis showed that both areas displayed
similar patterns (Fig. 3).
Discussion
This study includes new species distribution records for Islas
Marietas (ten), the coast of Nayarit (six), Colima (six), Jalisco
(five) and Islas Marías (one), and many of them have
173
Table 3 Updated checklist of valid species of Ophiuroidea from the
Central Mexican Pacific region according to literature records and the
collected species during this work
Phylum Echinodermata Brugiére, 1791
Class Ophiuroidea Gray, 1840
Order Euryalida Lamarck, 1816
Family Asteroschematidae Verrill, 1899
Asteroschema sublaeve Lütken and Mortensen, 1899 MAR
Family Asteronychidae Verrill, 1899
Astrodia excavata (Lütken and Mortensen, 1899) MAR
Family Gorgonocephalidae Ljungman, 1867
Astrocaneum spinosum (Lyman, 1875) MAR
Order Ophiurida Müller and Troschel, 1840
Family Ophiacanthidae Ljungman, 1867
Ophiacantha costata Lütken and Mortensen, 1899 MAR
Ophiacantha hirta Lütken and Mortensen, 1899 MAR
Ophiacantha moniliformis Lütken and Mortensen, 1899 MAR
Ophiomitra granifera Lütken and Mortensen, 1899 MAR
Ophiomitra partita Lütken and Mortensen, 1899 MAR
Family Ophiuridae Müller and Troschel, 1840
Gymnophiura mollis Lütken and Mortensen, 1899 MAR
Ophiernus adspersus annectens Lütken and Mortensen, 1899 MAR
Ophiura (Ophiura) scutellata (Lütken and Mortensen, 1899) MAR
Ophiura (Ophiuroglypha) irrorata irrorata (Lyman, 1878) MAR
Family Amphiuridae Ljungman, 1867
Amphichondrius laevis Ziesenhenne, 1940 JAL
Amphiodia assimilis (Lütken and Mortensen, 1899) MAR
Amphiodia occidentalis (Lyman, 1860) NAY
Amphiodia platyspina Nielsen, 1932 NAY
Amphiodia sculptilis Ziesenhenne, 1940 JAL
Amphiodia violacea (Lütken, 1856) NAY
Amphiodia (Amphispina) digitata Nielsen, 1932 NAY
Amphiodia (Amphispina) urtica (Lyman, 1860) NAY, MAR
Amphipholis elevata Nielsen, 1932 MAR
Amphipholis pugetana (Lyman, 1860) NAY
Amphipholis squamata (Delle Chiaje, 1828) MAT, NAY, JAL, COL
Amphiura serpentina Lütken and Mortensen, 1899 MAR
Dougaloplus notacanthus (Lütken and Mortensen, 1899) MAR
Microphiopholis platydisca (Nielsen, 1932) NAY, COL
Microphiopholis puntarenae (Lütken, 1856) MAR
Ophiocnida hispida (Le Conte, 1851) MAR, JAL, COL
Ophiophragmus lonchophorus Ziesenhenne, 1940 NAY, JAL
Ophiophragmus marginatus (Lütken, 1856) NAY, JAL
Ophiophragmus papillatus Ziesenhenne, 1940 MAT, NAY, JAL
Ophiostigma tenue Lütken, 1856 ISA, MAT, JAL
Family Ophiotrichidae Ljungman, 1867
Ophiothela mirabilis Verrill, 1867 ISA, MAT, NAY, JAL, COL, MICH
Ophiothrix galapagensis Lütken and Mortensen, 1899 MAR
Ophiothrix (Ophiothrix) rudis Lyman, 1874 MAR, MAT, NAY, JAL,
MICH
Ophiothrix (Ophiothrix) spiculata Le Conte, 1851 ISA, MAR, MAT,
NAY, JAL, COL, MICH
174
Mar Biodiv (2017) 47:167–177
Table 3 (continued)
Family Ophiactidae Matsumoto, 1915
Hemipholis gracilis Verrill, 1867 NAY, JAL
Ophiactis savignyi (Müller and Troschel, 1842) ISA, MAR, MAT,
NAY, JAL, COL, MICH
Ophiactis simplex (Le Conte, 1851) ISA, MAR, MAT, NAY, JAL,
COL, MICH
Family Ophionereididae Ljungman, 1867
Ophionereis albomaculata E.A. Smith, 1877 ISA, MAR
Ophionereis annulata (Le Conte, 1851) MAR, MAT, NAY, JAL, COL,
MICH
Family Ophiocomidae Ljungman, 1867
Ophiocoma aethiops Lütken, 1859 ISA, MAR, MAT, NAY, JAL,
COL, MICH
Ophiocoma alexandri Lyman, 1860 ISA, MAR, MAT, NAY, JAL,
COL, MICH
Ophiocomella schmitti A.H. Clark, 1939 NAY, COL
Ophiocomella sexradia (Duncan, 1887) MAR
Family Ophiodermatidae Ljungman, 1867
Diopederma daniana (Verrill, 1867) JAL, MICH
Ophioderma panamensis Lütken, 1859 ISA, MAR, MAT, NAY, JAL,
COL
Ophioderma teres (Lyman, 1860) MAR, NAY, JAL, COL
Ophioderma sp. MAR
Ophioderma variegata Lütken, 1856 MAR, NAY, JAL
Ophiuroconis bispinosa Ziesenhenne, 1937 JAL
Family Ophiolepididae Ljungman, 1867
Ophiolepis crassa Nielsen, 1932 MAR, MAT
Ophiolepis pacifica Lütken, 1856 MAR, MAT, NAY, JAL
Ophiolepis plateia Ziesenhenne, 1940 JAL
Ophiolepis variegata Lütken, 1856 MAR, NAY, COL, JAL
Ophiomusium lymani Wyville-Thomson, 1873 MAR
Ophiomusium variabile Lütken and Mortensen, 1899 MAR, JAL
ISA Isla Isabel; MAR Islas Marías; MAT Islas Marietas; NAY coast of
Nayarit; JAL Jalisco; COL Colima; MICH Michoacán
relevance in terms of filling their distribution gap in México or
extending their general geographical distribution range.
Some ophiuroids have been previously reported for the CMP
in other studies (Granja-Fernández et al. 2015a) and, of
Fig. 2 Sample-based rarefaction curve of the Ophiuroidea from the
Central Mexican Pacific region according to the number of localities.
Dotted lines confidence interval (±95 %); solid line expected average
richness; dashed line observed richness
Fig. 3 Taxonomic distinctness analysis of the Ophiuroidea from the
Central Mexican Pacific region. a Average (delta: Δ+). b Variation
(lambda: Λ+). Solid line 95 % confidence limit; dotted line Δ+ average
value
these, the presence of ten species was confirmed for the CMP
during this work in the same as well as new areas: H. gracilis,
O. savignyi, O. simplex, O. hispida, O. panamensis, O. teres,
O. pacifica, O. annulata, O. mirabilis and O. (O.) rudis. These
species are conspicuous not only in the CMP but also in the
rest of the Mexican Pacific, with a wide distribution along the
Eastern Pacific (Solís-Marín et al. 2013; Granja-Fernández et
al. 2015a). Ophionereis annulata has a symbiotic relationship
with the polynoid polychaete Malmgreniella cf. variegata in
two localities from Jalisco and one from Colima (GranjaFernández et al. 2013). In this work, this association was
found in Carrizales (Colima) and Túnel Amarradero, Isla
Redonda, Islas Marietas (Nayarit), being the latest the northernmost record observed for this association in the Eastern
Pacific.
The new report of A. squamata, M. platydisca and O. tenue
in the study area is relevant, as these species had not been
previously reported in the CMP, notwithstanding they had
their distributions in both the northernmost (Gulf of
California, Baja California and Baja California Sur) and
southernmost areas (Southern Mexican Pacific region) of the
Mexican Pacific (Honey-Escandón et al. 2008; GranjaFernández et al. 2015a, b). There are three species having their
distribution in most of the countries comprising the Tropical
Eastern Pacific (Solís-Marín et al. 2013). Nevertheless, in
Mar Biodiv (2017) 47:167–177
México, they have been reported exclusively in the northern
(Ophiuroconis bispinosa and Ophiolepis crassa) or the central
parts (Ophiocomella schmitii) of the country (GranjaFernández et al. 2015a). Despite the sampling effort along
the Mexican Pacific since 2007, the only areas in which O.
crassa and O. bispinosa were found were Islas Marietas and
Jalisco (personal observation), which represent the southernmost records of the species in México. On the other hand, O.
schmitii, a six-armed species of the family Ophiocomidae, was
previously reported from Colima and Islas Revillagigedo
(Granja-Fernández et al. 2015a), but during this work, it was
collected in the coast of Nayarit, representing the northernmost record of the species in the Mexican Pacific.
Considering that the latest ophiuroids have been recorded in
Central and South America (Solís-Marín et al. 2013), there is a
high probability of finding them in the Southern Mexican
Pacific region.
The most remarkable new records correspond to
O. lonchophorus, O. papillatus and Ophioderma sp.
Ophiophragmus lonchophorus is only known from
Tenacatita (Jalisco) and had not been reported since its description by Ziesenhenne (1940) in other areas from the
Eastern Pacific. During this work, this species was collected
on the coast of Nayarit; therefore, these findings represent a
range extension of almost 165 km north of the type locality.
On the other hand, the only report of O. papillatus in the
Eastern Pacific is from the states of Oaxaca and Guerrero in
Mexico (Granja-Fernández et al. 2014), but in this work, we
collected it in Jalisco, Islas Marietas and other localities on the
coast of Nayarit, which represents also a range extension of
740 km north. Finally, Ophioderma sp. is a new species in the
process of description and had been recorded from Guerrero
and Oaxaca; therefore, its new record in Islas Marías during
this work represents a range extension of 670 km north (see
Granja-Fernández et al. 2015a for more information).
Although the CMP is characterised as a highly biodiverse
region, the considerable number of new records of ophiuroids
can be attributed to a more intensive–extensive sampling; in
other words, a higher number of prospected localities and the
diversification of the collecting methods allowed recording
species that were not collected in other studies (i.e. Ríos-Jara
et al. 2008a, b, 2013; López-Uriarte et al. 2009). The different
sampling techniques allowed us to observe that collecting
specimens by hand is the best method because it obtained
75 % of the total of the collected species (A. squamata,
O. aethiops, O. alexandri, O. annulata, O. crassa, O. hispida,
O. mirabilis, O. pacifica, O. panamensis, O. papillatus,
O. teres, O. savignyi, O. schmitti, O. simplex, Ophioderma
sp., O. tenue, O. (O.) rudis, O. (O.) spiculata). Furthermore,
the advantage of the latest collecting method is the ability of
finding organisms in a greater variety of substrata (live
and dead stony corals, gorgonians, hydrozoan, rock, sand,
algae, rhodoliths and sponges), but, generally, the most
175
conspicuous and/or bigger species (disc diameter > 10 mm)
are the only ones collected. Other sampling techniques such
as suction pump and dredge in specific substrata (sand and
rock) allowed collecting some species that were not found
by hand collection, that are buried in sand and that are of a
smaller size (disc diameter < 5 mm). Using a suction pump, a
total of 54 % of the species were collected (A. sculptilis, A.
squamata, O. aethiops, O. alexandri, O. annulata, O.
hispida, O. lonchophorus, O. bispinosa, O. savignyi, O.
simplex, O. tenue, O. (O.) rudis, O. (O.) spiculata).
Furthermore, two species, A. sculptilis and O. bispinosa,
were collected using only this method and in a specific
substrata (sand). The use of a dredge was also important,
even though only 17 % of the total collected species were
obtained by this method (H. gracilis, M. platydisca, O.
lonchophorus, O. marginatus), and all of them (except O.
lonchophorus) were collected just by dredging. It is important to highlight that dredging allowed obtaining a major
quantity of organisms and of a larger size (disc
diameter < 5 mm) in comparison to the suction pump method. Finally, the species associated with the coral Pocillopora
spp. were A. squamata, O. savignyi, O. simplex and O. (O.)
spiculata, being this common since Granja-Fernández et al.
(2014) reported the association of these species (except A.
squamata) with stony corals in the Mexican Pacific. It is of
relevance to note that all of the collected specimens in this
substratum were juvenile, which corroborates with the fact
that stony corals are an important key habitat in the Mexican
Pacific, as they provide nursering, concealment, breeding
space and feeding for juvenile organisms (Spalding et al.
2001; Granja-Fernández et al. 2014).
According to the results obtained during this work, the
species with the highest qualitative abundance in the CMP
were O. mirabilis, O. (O.) spiculata, O. (O.) rudis, O.
savingyi, O. simplex and O. alexandri, all of them being the
most conspicuous too, distributed in almost all the sampled
localities (>50 %). The high abundance as well as the wide
distribution of these species is very common along the Eastern
Tropical Pacific (Ríos-Jara et al. 2008a; Alvarado et al. 2010;
Granja-Fernández et al. 2014), since Ophiocomidae,
Ophiotrichidae and Ophiactidae are the most specious families in tropical shallow waters (Hendler et al. 1995). A total of
14 species were considered as rare in the CMP; of these, eight
inhabited exclusively on sand. Despite the suction pump and
dredging representing effective sampling methods in terms of
finding new distribution records, the low number of performed replicates can denote the lowest abundance of the rare
collected species. A major sampling effort in this substratum
by this methodology has to be considered in further studies in
order to obtain a most real relative abundance of the species.
In the most recent updated checklist of ophiuroids from the
Mexican Pacific, Granja-Fernández et al. (2015a) recorded 20
species from the coast of Nayarit, 36 from Islas Marías, 21
176
from Jalisco, nine from Michoacán, nine from Isla Isabel and
eight from Colima, and despite the authors not including Islas
Marietas in their checklist, the CONANP (2007a) documented
four species from these islands. If the number of species per
area reported in Granja-Fernández et al. (2015a) and
CONANP (2007a) is summarised with the present results,
the numbers are as follows: 38 from Islas Marías, 26 from
the coast of Nayarit, 26 from Jalisco, 14 from Colima and
14 from Islas Marietas. We did not find new records of species
in Michoacán and Isla Isabel, so the number of species in these
sites remains as nine, respectively.
The CMP region comprises a total of 57 species of ophiuroids and Islas Marías is the area with the highest richness, and
Michoacán and Isla Isabel with the lowest. The highest richness recorded at Islas Marías also represents the highest value
of taxonomic distinctness of the entire CMP, which can be
explained because 18 species inhabit only in these islands,
all of them from deeper waters (>500 m). The above is not
unusual since Islas Marías is considered a stepping stone for
the dispersal route of species and individuals to and from the
Islas Revillagigedo, the Gulf of California and the Central and
Southern Mexican Pacific (López-Pérez et al. 2015). In fact,
all the collections in deep waters in the CMP had been performed only at Islas Marías (Lütken and Mortensen 1899),
and since most of the deeper species inhabiting these islands
have been recorded in Central and South America as well
(Solís-Marín et al. 2013), it is highly probable to find the same
species during a deep water exploration along the Mexican
Pacific. In the case of the low species richness observed at
Michoacán, sampling has not been effective, mostly due to
the difficulty in accessing the area. This could explain the
lowest number of species and, probably, the numbers might
differ if the sampling increased. Moreover, the low number of
species for Isla Isabel was related to the limited number of
sampling localities and to the fact that only a few types of
substrata were explored (i.e. Ríos-Jara et al. 2008a, b;
Granja-Fernández et al. 2014).
On the other hand, if the CMP (57 species) is compared
with the Southern Mexican Pacific (46 species) (GranjaFernández et al. 2015b) in terms of Ophiuroidea composition,
the former possesses more species despite a smaller coastal
line (~613 km and ~1,125 km, respectively). The higher number of species in the CMP can be attributed to species from
deep waters that have been collected only in Islas Marías.
Although they have differences in area extension, both regions
are similar in species composition as they share 43 % of the
species, 67 % of the genera and 77 % of the families. It is
important to emphasise that the analysis suggests that the
CMP and the Southern Mexican Pacific are similar in taxonomic distinctness, regardless of the total number of species
attributed to their similarity on oceanographic and substrata
conditions (Arriaga-Cabrera et al. 1998; Kessler 2006). It
could be suggested that it is highly probable that the following
Mar Biodiv (2017) 47:167–177
ophiuroids distributed in the southernmost part of the Mexican
Pacific can inhabit the CMP too: Amphichondrius granulatus,
Amphiodia psara, Amphiophiura superba, Amphiura
arcystata, Asteronyx longiffisus, Ophiomisidium leurum,
Ophiomusium glabrum, Ophiomyxa panamensis,
Ophionereis perplexa and Stegophiura ponderosa. In addition, there is a significant number of species of ophiuroids that
can be potentially collected in the CMP, since they are distributed from the northernmost part of the Mexican Pacific to
South America (Solís-Marín et al. 2013; Granja-Fernández
et al. 2015a).
The results evidence that the CMP region is very important in terms of Ophiuroidea diversity and composition in the Mexican Pacific, but, despite that, this study
recorded only 57 species and 28 new distribution records; according to our models, at least 104 species can
inhabit the area. We detected some points that we consider relevant in order to obtain a most complete checklist of the ophiuroids from the CMP in future works: (1)
the current study is a multi-disciplinary and interinstitutional work, which allowed combining different
collecting methods, resulting in the collection of species
that had never been reported in previous studies. For this
reason, we recommend to increase sampling by different
methods in as many types of substrata as possible in
order to collect a higher number of species; (2) despite
the fact that we carried out the collections in 61 localities
(most of them unexplored areas), there are still many
other areas and habitats (i.e. mangroves, small islands)
with a high potential for finding ophiuroids; (3) most of
the studies conducted in the CMP come from shallow
waters, so we suggest to conduct exploration in deep
waters. This study contributes to the knowledge of
Ophiuroidea fauna within protected and unprotected
areas of the CMP, and might provide useful information
for conservation and management purposes and enhances
the relevance of the region. Despite the improvement, the
CMP must be studied thoughtfully, since it plays a significant biogeographic role contributing to the dispersal
and permanence of species, and, moreover, possesses
ophiuroids that are only found in this region considering
the entire Eastern Tropical Pacific.
Acknowledgements This work is part of the PhD thesis of RGF at
UAM supported by a Consejo Nacional de Ciencia y Tecnología scholarship (no. 336853), and was funded by Comisión Nacional para el
Conocimiento y Uso de la Biodiversidad (JF047 to MDHP), Programa
para el Desarrollo Profesional Docente (220265 to APRT) and Dirección
General de Educación Superior Tecnológica (2353-P to PSS) projects.
We thank Andrés López-Pérez, Alejandro Hernández-Morales,
Emerson Martínez-Zavala, Diego Rangel-Solís, Adolfo TortoleroLangarica and Karla del Castillo for helping in the collection and identification of the specimens. We thank the reviewers and editor for
commenting and improving the manuscript.
Mar Biodiv (2017) 47:167–177
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