Galland, Grantly R

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Comments on Microhabitat Specialization and a Depth Range Extension for a Chaenopsid Tube Blenny in the Gulf of California, Mexico

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Bulletin of the Southern California Academy of Sciences
Volume 110 | Issue 2 Article 3
October 2012
Comments on Microhabitat Specialization and a
Depth Range Extension for a Chaenopsid Tube
Blenny in the Gulf of California, Mexico
Grantly R. Galland
Marine Biology Research Division, Scripps Institution of Oceanography, University of California San Diego,
ggalland@ucsd.edu
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Recommended Citation
Galland, Grantly R. (2011) "Comments on Microhabitat Specialization and a Depth Range Extension for a Chaenopsid Tube Blenny
in the Gulf of California, Mexico," Bulletin of the Southern California Academy of Sciences: Vol. 110: Iss. 2.
Available at: https://scholar.oxy.edu/scas/vol110/iss2/3
Comments on Microhabitat Specialization and a Depth Range Extension
for a Chaenopsid Tube Blenny in the Gulf of California, Mexico
Cover Page Footnote
I thank P.A. Hastings, H.J. Walker, J.S. Stephens, and two anonymous reviewers for reading and commenting
on early manuscripts describing this work. B.E. Erisman and O. Aburto-Oropeza organized field expeditions
that allowed me to conduct these observations and served as dive buddies, along with J. Lund. C. Sanchez and
P. Hastings helped in procuring the necessary permits for collecting fishes in Mexico. Funding for expeditions
was provided by Instituto Nacional de Ecología (INE), Fondo Mexicano para la Conservación de la
Naturaleza, a Pew Marine Conservation Fellowship awarded to E. Ezcurra, and anonymous donors.
Additional funding to support this work was provided by the Center for Marine Biodiversity and
Conservation.
This article is available in Bulletin of the Southern California Academy of Sciences: https://scholar.oxy.edu/scas/vol110/iss2/3
Comments on Microhabitat Specialization and a Depth Range
Extension for a Chaenopsid Tube Blenny in the Gulf of
California, Mexico
Grantly R. Galland
Marine Biology Research Division, Scripps Institution of Oceanography, University of
California San Diego, ggalland@ucsd.edu
Acanthemblemaria balanorum Brock (Clubhead Blenny; Fig. 1) is a chaenopsid tube
blenny endemic to the Tropical Eastern Pacific. Adults of this species, as all members of
the Chaenopsidae, inhabit vacated invertebrate tubes or tests (Stephens 1963; Lindquist
1985). In the case of A. balanorum, the shelter of choice is the vacated test of Megabalanus
Hoek barnacles, a genus characterized in the Tropical Eastern Pacific by a complex of
species (Henry and McClaughlin 1986) that typically live in the upper 10 m on shallow
rocky reefs (Brusca and Hendrickx 2008). In the Gulf of California (GOC), Mexico, A.
balanorum overlaps in distribution with two congeners, A. crockeri Beebe and Tee-Van
and A. hastingsi Lin and Galland, and these species are known to exhibit depth
partitioning, with A. balanorum inhabiting relatively shallower depths, A. crockeri
inhabiting relatively deeper depths, and A. hastingsi overlapping near the edges of the
depth ranges of the other two species at intermediate depths (Lindquist 1985). A detailed
study of the relationships among these three congeners in the southern GOC (Lindquist
1985) reported that A. balanorum inhabits shelters (5barnacles) down to approximately
7 m depth. Guides to the fishes of the region (e.g., Allen and Robertson 1994; Humann
and DeLoach 2004) report a similar depth range.
In November 2010, I observed and collected several individuals of A. balanorum at a
depth of 21 m at the base of a pinnacle off the south end of Maria Cleofas, the
southernmost point in the Islas Marias archipelago, southern GOC. These individuals,
like all individuals of this species that I have observed, inhabited vacant barnacles
(Megabalanus). This observation represents a significant depth range extension for this
normally shallow subtidal fish (and may also represent an extension for the barnacle;
Brusca and Hendrickx 2008). This ability of a microhabitat specialist to colonize
abnormal macrohabitats (in this case much deeper than normal waters) when its
microhabitat (5barnacles) is available supports a hypothesis that these specialists are
resource (5shelter) limited. Similar shelter limitation has already been experimentally
demonstrated in the GOC congener, A. crockeri, which increases in average density with
shelter addition (Hastings and Galland 2010).
I observed additional evidence that A. balanorum is a shelter-limited microhabitat
specialist in July 2009 at Las Animas, a small island and a series of small pinnacles in the
central GOC. That site proved to be ideal A. balanorum habitat, with several large
boulders completely covered by broad, very dense Megabalanus fields down to 5 m depth.
Within these barnacle fields, I observed large numbers of A. balanorum, more densely
distributed than any other chaenopsid population reported to date (e.g. in Lindquist
1985; Clarke 1996; Thomson and Gilligan 2002; P.A. Hastings, pers. comm.). During an
opportunistic survey of the area, I placed a 0.25 m2 quadrat on five randomly selected
areas on the top of a large, flat boulder at 5 m depth in order to survey chaenopsids and
Bull. Southern California Acad. Sci.
110(2), 2011, pp. 52–55
E Southern California Academy of Sciences, 2011
52
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Galland: Microhabitat Specialization and Depth Range Extension in Tube Blenny
Published by OxyScholar, 2011
ascertain densities. Numbers of A. balanorum ranged (mean +/2 SD) from 8 – 28 (20.4
+/2 8.6) individuals per 0.25 m2. These high densities may reflect the very high barnacle
densities at this site, though it is difficult to quantify available shelters, as it is not
immediately obvious what a chaenopsid considers to be a sufficient shelter.
To date, no experiments have been designed to determine what a second limiting factor
is for these or similar shelter-dwelling microhabitat specialists, but it is quite possibly an
issue of territory size. Males of Acanthemblemaria and several other chaeonpsid genera
actively court females, which, upon choosing a suitable male, enter the male’s shelter and
deposit eggs (Hastings 1986; Hastings 1988; Hastings and Peterson 2011). The males
guard the eggs and may simultaneously protect clutches from multiple females (Hastings
1986; Hastings 1988; Hastings and Peterson 2011). Under these conditions, males
compete for female choice and for the most desirable shelters (Hastings 1988; Hastings
1992). The high densities of A. balanorum observed at Las Animas could lead to agonistic
interactions that secondarily limit the population size (or density) of this species.
In order to begin quantifying the distance between occupied barnacle shelters at the
densely populated Las Animas site, I randomly chose an individual A. balanorum and
measured the distance to its nearest neighbor. I then measured the distance to that
individual’s nearest neighbor and repeated the process until I reached ten total
individuals. On average (SD), these ten individuals were only 4.4 cm (1.6) from their
nearest neighbor. Upon collecting the specimens, I obtained sex and standard length (SL)
for each individual and determined that they were, on average (SD), 28.9 mm SL (3.8)
and were all sexually mature adults. In fact, to the best of my knowledge, juvenile habitat
preference in this species is unknown. The male to female sex ratio was 7:3, indicating the
ability of several adult males to live in close proximity to one another and to mature
females, which were somewhat evenly dispersed among the males, within this set of ten
individuals (numbers one, three, and eight out of ten; 5, 2.5, and 5 cm from their nearest
neighbors, respectively). The presence of some shelters inhabited by less competitive
Fig. 1. Acanthemblemaria balanorum in shelter at Las Animas in the central Gulf of California,
Mexico. Photo taken in July 2009 at 5 m depth during nearest neighbor surveys. Photo by J. Lund.
MICROHABITAT SPECIALIZATION AND DEPTH RANGE EXTENSION IN TUBE BLENNY 53
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Bulletin of the Southern California Academy of Sciences, Vol. 110 [2011], Iss. 2, Art. 3
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chaenopsid species, Coralliozetus angelicus (Bo¨hlke and Meade) and Protemblemaria
bicirrus (Hildebrand), increases the overall chaenopsid density, and the availability of
some uninhabited shelters perhaps implies that the community at that site is approaching
some maximum. It is difficult, however, to determine whether or not empty barnacles
represent a choice to avoid high densities, a preference for other more highly desirable
shelters, or some additional factor.
While the preliminary evidence presented here supports a hypothesis that these
microhabitat specialists are first limited by the presence of their preferred shelter and
then by some other factor (possibly territoriality) that prevents higher densities, even
with greater shelter availability, it is necessary to design and implement experiments to
test this and similar hypotheses empirically. Species like A. balanorum that rely on
biologically derived microhabitats may be able to utilize a wider range of macrohabitats
as their ‘‘hosts’’ move to new areas. In the case of the depth range extension at Maria
Cleofas, the ability of Megabalanus barnacles to survive down to at least 21 m at this
site allows a species that specializes on its empty tests to do the same, and this ability is
not unique to A. balanorum. A blenniid (Hypsoblennius brevipinnis Gu¨nther) and another
chaenopsid (A. macrospilus Brock) were observed and collected at the Maria Cleofas
site, inhabiting barnacle tests. Hypsoblennius brevipinnis is also a microhabitat specialist,
its preferred shelter is also Megabalanus, and it is typically confined to the upper 3–4 m
of the subtidal zone (Allen and Robertson 1994; Humann and DeLoach 2004; though it
may be known from as deep as 10 m; De la Cruz Agu¨ero et al. 1997). Like in A.
balanorum, this observation represents a significant depth range extension for this
specialist and an interesting finding for the Blenniidae, which are characterized by few
species reaching depths greater than approximately 25 m (Springer and Smith-Vaniz
1970). Acanthemblemaria macrospilus, sister to the more northern A. hastingsi discussed
above (see Lin and Galland 2010), shares its ability to utilize multiple microhabitat
shelters (Lindquist 1985) and typically lives to depths of 15–18 m (Allen and Robertson
1994). Observation of individuals at a depth of 21 m may represent a minor change in
depth range, but its flexibility in shelter choice probably allows it to occupy a greater
diversity of habitats than A. balanorum. Further research into the intra- and inter-
specific relationships within this group of species that utilize (and potentially compete
for) similar microhabitats will reveal more of the processes at play in determining the
makeup of this assemblage and may provide further insight into the relative advantages
and disadvantages of microhabitat specialization.
All specimens of all species discussed here were collected using quinaldine and are
archived at the Scripps Institution of Oceanography Marine Vertebrate Collection (Las
Animas collection number 5 SIO 09-261; Maria Cleofas collection number 5 SIO 10-
132).
Acknowledgments
I thank P.A. Hastings, H.J. Walker, J.S. Stephens, and two anonymous reviewers for
reading and commenting on early manuscripts describing this work. B.E. Erisman and O.
Aburto-Oropeza organized field expeditions that allowed me to conduct these
observations and served as dive buddies, along with J. Lund. C. Sanchez and P.
Hastings helped in procuring the necessary permits for collecting fishes in Mexico.
Funding for expeditions was provided by Instituto Nacional de Ecologı´a (INE), Fondo
Mexicano para la Conservacio´n de la Naturaleza, a Pew Marine Conservation
54 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
3
Galland: Microhabitat Specialization and Depth Range Extension in Tube Blenny
Published by OxyScholar, 2011
Fellowship awarded to E. Ezcurra, and anonymous donors. Additional funding to
support this work was provided by the Center for Marine Biodiversity and Conservation.
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