Revista Mexicana de Biodiversidad 85: 463-471, 2014
DOI: 10.7550/rmb.34177
Revista Mexicana de Biodiversidad 85: 463-471, 2014
DOI: 10.7550/rmb.34177
463
Review of the geographic distribution of Hoffmannola hansi (Gastropoda:
Pulmonata) in the Mexican Pacific
Revisión del ámbito de distribución geográfica de Hoffmannola hansi (Gastropoda:
Pulmonata) en el Pacífico mexicano
Omar Hernando Avila-Poveda1 , Quetzalli Yasú Abadia-Chanona2, Raúl Herrera-Fragoso3 and Benoît Dayrat4
1
División de Estudios de Posgrado, Universidad del Mar, 70902 Puerto Ángel, Oaxaca, Mexico.
Licenciatura en Biología Marina, Universidad del Mar, 70902 Puerto Ángel, Oaxaca, Mexico. Present address: Centro Interdisciplinario de
Ciencias Marinas, Instituto Politécnico Nacional, 23096 La Paz, Baja California Sur, Mexico.
3
Universidad La Salle-Nezahualcoyotl. Av. Bordo de Xochiaca, 57300 Nezahualcóyotl, Estado de México, Mexico.
4
Department of Biology, Pennsylvania State University. University Park, 16802 PA, USA.
oavila@colombia.com
2
Abstract. Hoffmannola hansi (Mexican intertidal leather slug) is traditionally reported as an endemic species to the
Gulf of California, Mexico. However, its presence in the southern Mexican Pacific has been mentioned in regional
checklists and reports. Here we provide new records of H. hansi populations from at least 3 locations from Oaxaca,
Mexico. The anatomical characteristics useful for H. hansi identification are described for both, living and preserved
specimens. Specimen’s reports from the Gulf of California to Oaxaca, Mexico, are mentioned, yielding a revised
distribution throughout the Mexican Pacific. A map with the wider geographic distribution of H. hansi is also updated.
Therefore, this species is not “endemic” to the Gulf of California, but is distributed throughout the Mexican Pacific.
However its distribution is discontinuous with the possible existence of 2 cryptic species geographically separated by
the Sinaloan gap: 1 in the North (Gulf of California) and 1 in the South. Additional work is needed to explore in more
detail the complete geographical range of H. hansi populations on the entire Mexican Pacific Coast.
Key words: Mollusca, Onchidiidae, non-endemic, Oaxaca, Gulf of California, Tropical Eastern Pacific (TEP),
geographical range, cryptic species.
Resumen. Hoffmannola hansi (babosa de cuero intermareal mexicana) es tradicionalmente registrada como una
especie endémica del golfo de California, México. Sin embargo, su presencia en el sur del Pacífico mexicano ha sido
mencionada en listas de control y en informes regionales. Ofrecemos nuevos registros de poblaciones de H. hansi
de al menos 3 localidades de Oaxaca, México. Las características anatómicas que ayudan a identificar a H. Hansi se
describen para ejemplares vivos y conservados. Se mencionan los ejemplares desde el golfo de California hasta Oaxaca,
México, proporcionando una distribución ampliada a lo largo del Pacífico mexicano. Un mapa con la distribución
geográfica de H. Hansi también se actualiza. Por lo tanto, esta especie no es endémica del golfo de California, sino que
está bien distribuida en el Pacífico mexicano. Sin embargo, su distribución es discontinua, con la posible existencia de
2 especies crípticas geográficamente separadas por la brecha sinaloense: una en el Norte (golfo de California) y otra
en el Sur. Sigue siendo necesario realizar trabajo adicional para explorar en mayor detalle la distribución geográfica
completa de las poblaciones de H. hansi en toda la costa del Pacífico mexicano.
Palabras clave: Mollusca, Onchidiidae, no endémico, Oaxaca, golfo de California, Pacífico Oriental Tropical (POT),
ámbito geográfico, especies crípticas.
Introduction
Currently, the Onchidiidae (Mollusca: Gastropoda:
Pulmonata) is classified within the pulmonate gastropods
(Holznagel et al., 2010; Wu et al., 2010; Dayrat et al.,
2011a). However, its phylogenetic position has been
Recibido: 27 octubre 2012; aceptado: 01 julio 2013
heavily debated during the last 8 decades. Authors
have considered them as opisthobranchs (Fretter, 1943;
Boettger, 1954; Marcus, 1965; Marcus and Marcus, 1967;
1970), pulmonates (Bretnall, 1919; Baker, 1938; Baker,
1955; Ghiselin, 1965; Solem, 1978; Britton, 1984; Barker,
2001; Grande et al., 2004; Dayrat, 2009; Dayrat et al.,
2011a,b), euthyneurans (Marcus and Burch, 1965; Selmi et
al., 1988; Winnepenninckx et al., 1998; Dayrat and Tillier,
464
Avila-Poveda et al.- Geographic distribution of Hoffmannola hansi (Gastropoda)
2000; 2002; 2003), or even as a separate Order Onchidiida
(Starobogatov, 1976) and Silicodermatae (Labbé, 1934).
Regardless, Onchidiidae remains a poorly-known taxon
in many regards, especially in species diversity and
distribution.
The most interesting aspect of the geographic range for
Onchidiidae is that almost all the genera are exclusively
found in tropical and subtropical areas from the Indo-West
Pacific (Bretnall, 1919; Stringer, 1969; Wu et al., 2010)
and the Mediterranean Sea (Barletta and Ghisotti, 1978),
excepting 4 nominal Onchidella species and 2 nominal
Hoffmannola species (i.e., H. hansi and H. lesliei) that
are restricted to the tropical Eastern Pacific (Dayrat,
2009; Dayrat et al., 2011b). Onchidella has an extensive
geographical range in all the tropical Eastern Pacific; while
Hoffmannola hansi is restricted to the Northern Gulf of
California and Hoffmannola lesliei to the Galapagos (e.g.,
Marcus and Marcus, 1967, 1970; Keen, 1971; Hendrickx
et al., 2005; Dayrat, 2009; Dayrat et al., 2011a).
Hoffmannola hansi Marcus and Marcus, 1967 (for
nomenclatural information, see, Dayrat et al., 2011b) has
been known by its description as “endemic” commonly
found in the rocky, intertidal communities of the Gulf of
California, Mexico, across shores from Baja California
Norte, Sonora, and Sinaloa (Marcus and Marcus, 1967,
1970; Keen, 1971; Hendrickx et al., 2005; Zamora-Silva
and Naranjo-García, 2008; Dayrat, 2009; ASDM, 2011;
Dayrat et al., 2011a). Nevertheless, various publications,
reports, and regional checklists (none of which citing
any deposited material) have mentioned H. hansi in
other Mexican states and localities outside the Gulf of
California and further south into the Mexican Pacific,
such as, Jalisco (Esqueda-González, 1995; Esqueda et al.,
2000), Jalisco, Colima and Michoacán (Holguín-Quiñones
and González-Pedraza, 1994), Michoacán (VillaseñorGómez, 2005), Guerrero (Flores-Garza et al., 2005; 2007;
Flores-Rodríguez et al., 2001, 2003, 2007, 2012; FloresRodríguez, 2004; Valdés-González et al., 2004) and Oaxaca
(Rodríguez-Palacios et al., 1988; León-Herrera, 2000;
2001; Ramírez-González, 2008). Although they did not
cite any actual specimens, those publications suggest that
H. hansi may not be endemic to the Gulf of California.
The present study provides a review of the geographic
distribution of H. hansi from available original literature
(e.g., reports, grey literature, and regional checklists),
several online databases of museum collections and
specimens collected by our group Oaxaca, Mexico.
Material and methods
Specimens of different sizes of H. hansi were found
while sampling for Chiton (Chiton) articulatus at Oaxaca
(Avila-Poveda, 2013; Avila-Poveda and Abadia-Chanona,
2013). They were living next to 2 other species of mollusks
(Chiton (Chiton) articulatus and/or Plicopurpura pansa)
within crevices at high tide, or by themselves on the surface
of rocks during low tide (Fig. 1). Several specimens of H.
hansi were collected (n= 18, 14 ≤ TL ≤ 48 mm, total
length, Fig. 2), in March, April, August, and October 2011
from 3 localities in Oaxaca, Mexico: 1) Puerto Angel
(15°39’ N, 96°29’ W); 2) El Faro, Puerto Angel (15°39’
N, 96°30’ W), and 3) Conejos Bay, Huatulco (15°46’ N,
96°04’ W).
Three specimens (45 ± 3 mm TL, Fig. 3) were relaxed
and allowed to extend for 1 h, with gradual additions of
tap water to the seawater, until the salinity reached half the
salinity of the seawater of the sampling site (i.e., 50:50 in
volume: Avila-Poveda, 2013) and accelerating the relaxation
process by adding few milliliters of 10% ethanol (after
Lincoln and Sheals, 1979). Later, specimens were fixed
in 10% neutral formalin-saline solution in seawater for 2
weeks and subsequently preserved in 70% ethanol (AvilaPoveda and Baqueiro-Cárdenas, 2009). Other specimens
were preserved in 95% ethanol for future molecular work
(Dayrat lab). The external and internal morphologies were
studied under a dissecting microscope.
Specimens were (or shortly will be) deposited in
museums in the region: 1) Laboratorio de Sistemática de
Invertebrados Marinos (LABSIM), Universidad del Mar,
Puerto Angel, Oaxaca, Mexico; 2) California Academy of
Sciences (CAS), San Francisco, California, USA; 3) Santa
Barbara Museum of Natural History (SBMNH), Santa
Barbara County, California, USA; 4) Scripps Institution
of Oceanography (SIO), San Diego, California, USA;
5) Arizona-Sonora Desert Museum (ASDM), Tucson,
Arizona, USA; 6) Smithsonian National Museum of
Natural History (NMNH), NW Washington, D.C., USA.
A checklist and a map of the geographic distribution of
H. hansi were developed, based on records found for this
species in scientific databases and web sites (e.g., BHL,
BioStor, JSTOR, among many others), available original
literature (e.g., reports and checklists), online databases
of museum collections (e.g., ASDM, CAS, LABSIM,
NMNH, SBMNH, SIO) and with the samples collected
by our group Oaxaca, Mexico.
Results
The most important anatomical characters observed
that help to identify the species are (for additional
information, see: Dayrat et al., 2011b): 1) the total length
of live specimens ranges from 14 mm to 48 mm (Fig.
3A) while preserved material is on average 25% shorter
(e.g., the largest live specimen of 48 mm long measures
Revista Mexicana de Biodiversidad 85: 463-471, 2014
DOI: 10.7550/rmb.34177
465
Figure 1. Hoffmannola hansi. Live animals (~50 specimens in picture) observed in the rocky intertidal
of El Faro, Puerto Angel, Oaxaca, Mexico (15°39’N-96°30’W), USNM 1155112. The arrow indicates a
2 pesos Mexican coin (23 mm diameter).
only 33 mm once preserved; see, Fig. 3B); exceptionally,
we observed specimens in the field that were longer than
48 mm, but in areas with difficult access; 2) the total
width of the hyponotum (left and right side, H) relative
to the width of the pedal sole (S) is around the third or
fourth part (i.e., H <<< S) with H-S-H of 5-16/22-5 mm
for live organisms, while these are 4-13/18-4 mm for the
same preserved organisms; 3) there are generally about
14 marginal glands on the left and about 15 on the right;
4) the visceral cavity is divided in 2 by a septum (which
actually is a characteristic of Hoffmannola).
During the sampling months (i.e., March, April,
August, and October), early life stages of various species
of intertidal mollusks such as Chiton (Chiton) albolineatus,
Chiton (Chiton) articulatus, Echinolittorina aspera,
Echinolittorina modesta, Plicopurpura pansa, Nerita
scabricosta, and Hoffmannola hansi, among many others
mollusks, were observed in large stocks that resembled a
nursery.
A geographic distribution of H. hansi specimens was
found available broadly over the Mexican pacific, and not
just restricted to the Gulf of California (Table 1; Fig. 4).
Discussion
Figure 2. Preserved specimens of different sizes of Hoffmannola
hansi. The viscous fluid secreted by the marginal gland
(repugnatorial gland) is observed in the periphery of each
specimen.
The checklist and map presented here show many more
records of Hoffmannola hansi towards the Mexican south
Pacific (records No. 13 to 36, total 24: Table 1; Fig. 4) than
northern locations, where it has been termed as “endemic”
466
Avila-Poveda et al.- Geographic distribution of Hoffmannola hansi (Gastropoda)
Figure 3. Hoffmannola hansi. Live animals (A); and fixed and preserved individuals used for dissection (B). Collected at El Faro,
Puerto Angel, Oaxaca, Mexico (15°39’N-96°30’W). Same number of asterisks indicates the same specimen.
Figure 4. Geographical distribution of Hoffmannola hansi along the Tropical Eastern
Pacific. Records (R) according to Table 1. A star (⋆) marks 2 questionable localities,
outside the Tropical Eastern Pacific (catalog numbers USNM 771808 and USNM 710013;
NMNH 2011).
to the Gulf of California ((records No. 3 to 12, total 10:
Table 1; Fig. 4). Thus, H. hansi has been previously
recorded in Jalisco (Esqueda et al., 2000), Jalisco, Colima
and Michoacán (Holguín-Quiñones and González-Pedraza,
1994), Michoacán (Villaseñor-Gómez, 2005), Guerrero
(Flores-Garza et al., 2005; 2007; Flores-Rodríguez et al.,
2001, 2003, 2007, 2012; Flores-Rodríguez, 2004; ValdésGonzález et al., 2004) and Oaxaca (Rodríguez-Palacios et
al., 1988; León-Herrera, 2000; 2001; Ramírez-González,
2008).
Based on information presented here, the localities of
Puerto Angel, El Faro, and Conejos Bay of Huatulco, all
in the state of Oaxaca, Mexico, should be incorporated into
the species geographical distribution, as well as the states
of Jalisco, Colima, Michoacán, and Guerrero, in Mexico,
where H. hansi (Mexican intertidal leather slug) has been
mentioned from the rocky intertidal.
In any case, H. hansi is not endemic to the Gulf of
California, but is distributed throughout the Mexican
Pacific, although its distribution is quite discontinuous.
R
Catalog number
Coordinates
Geographical area and locality
Year of collection
Sources
1
2
USNM 771808
USNM 710013
45°N-125°W
41°N-71°W
USA, Washington, Fort Canby State Park ***
USA, New Hampshire, Pittsfield ***
1970
1974
NMNH 2011
NMNH 2011
3
USNM 678419
29°N-113°W
1967*
NMNH 2011, Marcus and Marcus 1967
4
----
29°N-113°W
----
ASDM 2011, Hendrickx et al. 2005
5
----
29°N-113°W
----
Keen 1971
6
----
29°N-113°W
Mexico, Baja California, Gulf of California,
San Agustín
Mexico, Baja California, Gulf of California,
Angel de la Guarda Island
Mexico, Baja California, Gulf of California,
Angel de la Guarda Island
Mexico, Baja California, Gulf of California,
San Agustín
1966
Marcus and Marcus 1970
7
8
9
---USNM-753651
USNM 753653
28°N-111°W
27°N-110°W
27°N-110°W
Mexico, Sonora, Gulf of California, Kino Bay
Mexico, Sonora, Gulf of California, El Sahuaral
Mexico, Sonora, Gulf of California , El
Sahuaral
---1966
1966
Keen 1971
NMNH 2011
NMNH 2011
10
CAS-81809
23°N-106°W
1953
CAS 2011
11
CNMO 1783 **
23°N-106°W
1987
Zamora-Silva and Naranjo-García 2008
----
23°N-106°W
----
ASDM 2011, Hendrickx et al. 2005
13
CMLE †
19°13′N- 104°42′W
Mexico, Jalisco, Melaque
1987-1989
14
----
19°14′N- 104°46′W
Mexico, Jalisco, Cuastecomate Bay, La
Calechosa
1993-1994
Holguín-Quiñones and González-Pedraza
1994
Esqueda-González 1995, Esqueda et al. 2000
15
CMLE †
19°06′N-104°20′W
Mexico, Colima, Las Hadas, Club de Yates
Beach
1987-1989
Holguín-Quiñones and González-Pedraza
1994
16
17
---CMLE †
18°N-103°W
18°20′N-103°30′W
Mexico, Michoacán
Mexico, Michoacán, Faro de Bucerías town
---1987-1989
18
CMLE †
18°16′N-103°20′W
Mexico, Michoacán, Maruata town
1987-1989
Villaseñor-Gómez 2005
Holguín-Quiñones and González-Pedraza
1994
Holguín-Quiñones and González-Pedraza
1994
19
20
21
22
CMLIECS §,b
----------
17°47'N-101°44'W
17°32'N-101°26'W
17°15'N-101°01'W
16°49'N-99°54'W
2000/2001/2005
2000/2001/2005
2000/2001/2005
2000/2001/2005
a, b, d, e, g
a, d, e, g
a, d, e, g
a, e, g, h
23
----
16°42'N-99°54'W
Mexico, Guerrero, Troncones Beach
Mexico, Guerrero, Barra de Potosí Beach
Mexico, Guerrero, Tlacoyunque Beach
Mexico, Guerrero, La Roqueta-"Palmitas"
Island
Mexico, Guerrero, La Roqueta-"Zoológico"
Island
2000/2001/2005
a, c, e, g, h
467
12
Mexico, Sinaloa, Gulf of California, Gaviotas
Beachs
Mexico, Sinaloa, Gulf of California, Cerro
Crestón Beachs
Mexico, Sinaloa, Gulf of California, Mazatlán
Revista Mexicana de Biodiversidad 85: 463-471, 2014
DOI: 10.7550/rmb.34177
Table 1. Records of Hoffmannola hansi along the Tropical Eastern Pacific. Coordinates were taken from each original source. Dashed lines indicate unavailable data.
Data was organized according to geographical coordinates and area according to country, state, region, and locality. R, is the progressive record for each locality. The gray
shadow helping to distinguish each group by state
24
25
26
----------
16°33'N-98°46'W
16°32'N-98°54'W
16°19'N-98°34'W
Mexico, Guerrero, Las Peñitas Beach
Mexico, Guerrero, Ventura Beach
Mexico, Guerrero, Punta Maldonado Beach
2000/2001/2005
2000/2001/2005
2000/2001/2005
a, e, g
a, e, f, g
a, e, g
27
28
29
30
-------------
15°46'N-96°05'W
15°46'N-96°04'W
15°45'N-96°07'W
15°45'N-96°07'W
Mexico, Oaxaca, Tangolunda Bay
Mexico, Oaxaca, Conejo Beach
Mexico, Oaxaca, Violín Beach
Mexico, Oaxaca, Arroyo Beach
---2011
-------
31
32
33
34
35
36
---------USNM 1155112
UMAR-Gast-001^
----
15°44'N-96°07'W
15°43'N-96°10'W
15°41'N-96°26'W
15°39'N-96°30'W
15°39'N-96°29'W
15°39'N-96°29'W
Mexico, Oaxaca, India Beach
Mexico, Oaxaca, Cacaluta Island
Mexico, Oaxaca, Tijera Bay
Mexico, Oaxaca, Puerto Ángel, El Faro
Mexico, Oaxaca, Puerto Ángel town
Mexico, Oaxaca, Puerto Ángel
---2005
---2011
2011
----
León-Herrera 2001, 2000
This study
León-Herrera 2001, 2000
Ramírez-González 2008, León-Herrera 2001,
2000
León-Herrera 2001, 2000
Ramírez-González 2008
León-Herrera 2001, 2000
This study
This study
Rodríguez-Palacios et al. 1988, León-Herrera
2001, 2000
Flores-Rodríguez et al., 2012
Flores-Rodríguez et al., 2007
c
Flores-Rodríguez et al., 2003
d
Flores-Rodríguez et al., 2001
e
Flores-Rodríguez 2004,
f
Flores-Garza et al. 2007,
g
Flores-Garza et al. 2005,
h
Valdés-González et al. 2004,
* Collected date only mentioned by Marcus and Marcus (1967)
** Colección Nacional de Moluscos (CNMO), Instituto de Biología, Universidad Nacional Autónoma de México (UNAM).
*** Questionable localities, outside of the Tropical Eastern Pacific Region, possible errors. Feedback was sent to NMNH for its review.
§
Colección de Moluscos del Laboratorio de Investigación en Ecología Costera y Sustentabilidad (CMLIECS), Unidad Académica de Ecología Marina, Universidad
Autónoma de Guerrero, Mexico.
†
Colección de Moluscos del Laboratorio de Ecología (CMLE), Escuela Nacional de Ciencias Biológicas, Mexico.
^
Colección Científica de Invertebrados, Laboratorio de Sistemática de Invertebrados Marinos (LABSIM), Universidad del Mar, campus Puerto Ángel, Oaxaca, Mexico.
b
Avila-Poveda et al.- Geographic distribution of Hoffmannola hansi (Gastropoda)
a
468
Table 1. Continues
Revista Mexicana de Biodiversidad 85: 463-471, 2014
DOI: 10.7550/rmb.34177
However, H. hansi requires rocky shores to live and
large parts of the coast between the Gulf of California
and southern Mexico are sandy beaches, especially the
Sinaloan gap (Hastings, 2000). However, additional work
is still needed to explore in more detail the complete
geographical range of H. hansi populations on the entire
Mexican Pacific Coast.
Currently, as only morphological data are available and
as populations from the Gulf of California and those from
Oaxaca are not distinguishable, it is more parsimonious
to regard them all as part of the same species, H. hansi.
However, it cannot be excluded that molecular data could
reveal the existence of 2 cryptic but separate species, one
in the North (Gulf of California) and one in the South,
both separated geographically by the long Sinaloan gap.
Various types of patterns of distribution have been found
by authors for rocky, intertidal species from the tropical
Eastern Pacific (e.g., Hastings, 2000; Wares, 2001; Craig
et al., 2006; Pitombo and Burton, 2007; Wares et al., 2009).
Both onchidiid species, H. hansi and Onchidella binneyi
(also distributed all along the Mexican Pacific), seem ideal
candidates for such population studies (Dayrat et al. 2011b).
On the other hand, some errors were unveiled in the
literature as well as online databases. Two museum records
for H. hansi (NMNH 2011: USNM 771808 and USNM
710013) are likely to be incorrect, or at least questionable,
since they are outside of the tropical Eastern Pacific and
even in a different region of the Western Atlantic (Fig. 4).
Feedback has been sent to NMNH Customer Service for
its review (Avila-Poveda, pers. comm.).
Acknowledgments
This work was personally assured by Avila-Poveda OH
with SNI-Conacyt stimulus (record No. 44175, from January
to July 2012). Avila-Poveda OH gives special thanks for
the family grant that covered his stay in the location during
the course of this study. Thanks to Saúl Jaime SerranoGuzmán for transporting the specimens to Benoît Dayrat
at the School of Natural Sciences, University of California
at Merced, California, USA. Thanks to Francisco BenitezVillalobos for transporting and depositing some specimens
in the NMNH (USNM 1155112). All rare references were
downloaded via the Biodiversity Heritage Library (http://
www.biodiversitylibrary.org/), Bioline International
(http://www.bioline.org.br/), BioStor (http://biostor.org/),
and Internet Archive (http://www.archive.org/).
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