Coral boring Aka-species (Porifera: Phloeodictyidae) from Mexico with description of Aka cryptica sp. nov

Abstract

This paper focuses on Aka species boring coral of the East Pacific Ocean and the Caribbean Sea (Mexico). The new species Aka
cryptica sp. nov. is described from Mexican Pacific coral reefs, which constitutes the first time that a species of the genus Aka is reported from the East Pacific Ocean. The new species lives cryptically boring coral species of the genus Pocillopora. It is characterized by the small size of their papillae (from 1 to 4.5 mm high and from 0.8 to 1.8 mm in diameter), and oxeas (from 67 to 120 μm), and their exposed parts blend in well with background colours which tend to be overlooked during benthic marine surveys. In addition, Aka
coralliphaga and Aka
brevitubulata from Mexican Caribbean coral reefs are redescribed. The latter species is reported for the first time in Mexico.

Full text

J. Mar. Biol. Ass. U.K. (2007), 87, 1477–1484
Printed in the United Kingdom
Journal of the Marine Biological Association of the United Kingdom (2007)
doi: 10.1017/S0025315407055373
Coral boring Aka-species (Porifera: Phloeodictyidae) from Mexico
with description of Aka cryptica sp. nov.
This paper focuses on Aka species boring coral of the East Pacific Ocean and the Caribbean Sea (Mexico).
The new species Aka cryptica sp. nov. is described from Mexican Pacific coral reefs, which constitutes the f irst
time that a species of the genus Aka is reported from the East Pacific Ocean. The new species lives cryptically
boring coral species of the genus Pocillopora. It is characterized by the small size of their papillae (from 1 to 4.5
mm high and from 0.8 to 1.8 mm in diameter), and oxeas (from 67 to 120 µm), and their exposed parts blend
in well with background colours which tend to be overlooked during benthic marine surveys. In addition,
Aka coralliphaga and Aka brevitubulata from Mexican Caribbean coral reefs are redescribed. The latter species is
reported for the first time in Mexico.
INTRODUCTION
The genus Aka Laubenfels, 1936 comprises a small group
of excavating sponges living in calcareous substrata, with a
skeleton formed only by oxeas, and characterized by fistulose
tubes which are externally visible (Desqueyroux-Faúndez &
Valentine, 2002).
The genus was originally included in the family
Clionaidae (Hancock, 1849), but it was later moved to the
family Phloeodictyidae by Rützler & Stone (1986), who
recognized characters described for Siphonodictyon Bergquist,
1965 and consequently referred Aka as a senior synonym of
Siphonodictyon. Even though there is still some debate as to
the validity of this synonymy (see for example Desqueyroux-
Faúndez & Valentine, 2002), we considered it valid until
more information is available.
Aka-species bore calcitic substrata such as mollusc shells,
but they have also been described as an important agent
of destruction of coral reef ecosystems in the Indo-Pacific
(Bergquist, 1965; Thomas, 1968, 1973) and Caribbean areas
(Rützler, 1971; Pang, 1973). Cliona-species usually infest the
non-living basal parts of corals and only harm the living
colony if the attachment is weakened and the coral breaks
loose (Goreau & Hartman, 1963). In contrast, the genus
Aka attacks living corals and harms the polyps (Goreau &
Hartman, 1963; Rützler, 1971).
These species have been reported from different localities
in the Caribbean, West Africa, Mediterranean, Atlantic
coast and central Pacific (Johnson, 1899; Laubenfels, 1954;
Thomas, 1968; Fromont, 1993; Desqueyroux-Faúndez &
Valentine, 2002), but curiously, no present records of Aka
exist in the north-eastern Pacific Ocean, since the only
reference found is about a fossil Aka in Arizona (USA)
(Reitner & Keupp, 1991).
The eastern Pacific Ocean has been explored for boring
sponges for several years now and this has provided a general
vision of the diversity and distribution of this very important
sponge group in the area (Carballo et al., 2004; Carballo
& Cruz-Barraza, 2005; Bautista-Guerrero et al., 2006).
This paper focuses on Aka-species boring coral reefs species
from the East Pacific Ocean and the Caribbean (Mexico).
Three species were identified; Aka cryptica sp. nov., in the
Pacific Ocean and Aka coralliphaga (Rützler, 1971) and Aka
brevitubulata (Pang, 1973), in the Caribbean; the latter species
is reported for first time in Mexico. This is the first time that
a species of the genus Aka has been reported from the East
Pacific Ocean.
MATERIALS AND METHODS
The material was collected by SCUBA diving along the
Pacific and Caribbean coast of Mexico. Sampling locations
are presented in Figure 1.
The specimens were preserved with their coral substrate
in 10% formalin for 24 hours and later transferred to 75%
ethanol for storage. External morphology such as colour,
texture and size of the papillae, skeletal elements and
their arrangement were recorded for each species. Spicule
preparation for light microscopy (LM) involved dissolving
sponge tissue in 60% nitric acid, rinsing with water,
dehydrating in ethanol and mounting in Canada balsam.
Spicule measurements were obtained from a minimum of
25 spicules chosen at random for each specimen. Length,
width of oxeas were measured under light microscope. Coral
José Luis Carballo*‡, Leanne Hepburn†, Héctor H. Nava*, José Antonio Cruz-Barraza*
and Eric Bautista-Guerrero*
*Instituto de Ciencias del Mar y Limnología, (Estación Mazatlán), Universidad Nacional Autónoma de México,
Apartado postal 811, Mazatlán 82000, México. †Department of Biological Sciences, University of Essex,
Wivenhoe Park, Colchester, CO4 3SQ, UK. ‡Corresponding author, e-mail: jlcarballo@ola.icmyl.unam.mx

1478 J.L. Carballo et al. Coral boring Aka-species from Mexico
Journal of the Marine Biological Association of the United Kingdom (2007)
pieces were broken and examined in detail for excavating
patterns. Scanning electron micrographs (SEM) were made
on sodium hypochloride-cleaned limestone coral skeletons.
The specimens examined were deposited in the Museo
Nacional de Ciencias Naturales (MNCN; Madrid, Spain),
Natural History Museum (BMNH; London, UK), in the
Colección de Esponjas of the Instituto de Ciencias del Mar
y Limnología, Universidad Nacional Autónoma de México
(LEB-ICML-UNAM), (Mazatlán, México).
SYSTEMATICS
Order HAPLOSCLERIDA Topsent, 1928
Family PHLOEODICTYIDAE Carter, 1882
Genus Aka de Laubenfels, 1936
Diagnosis
Excavating sponges boring calcitic substrata. Spicules are
oxeas forming sinuous bands arranged in bundles, producing
fistulose tubes. Externally visible by their fistulose tubes.
Skeleton formed only of spicule-bundles in sinuous bands,
not in a network, lining walls of fistules. Spicules short,
slender, curved oxeas. Without microscleres (Desqueyroux-
Faúndez & Valentine, 2002).
Acca Johnson, 1899: 461 (preoccupied)
Aka de Laubenfels, 1936: 155; Thomas, 1968: 250; Rützler
& Stone, 1986: 663; Reitner & Keupp, 1991: 102.
Aka cryptica sp. nov.
(Figures 2–4)
Material examined
Holotype: [MNCN 1.01/363]. Bahía de San Agustín
(Huatulco, Oaxaca), 15°41'09"N 96°13'46"W; 6 m depth; 09
May 2005.
Paratypes: [BMNH 2006.6.30.1]. Bahía de La Entrega
(Huatulco, Oaxaca), 15°44'34"N 96°07'35"W; 6 m depth;
04 July 2005. Specimen excavating alive and dead Pocillopora
damicornis colonies. LEB-ICML-UNAM-1196; Bahía de San
Agustín (Huatulco, Oaxaca), 15°41'09"N 96°13'46"W; 7 m
depth; 09 May 2005. Specimen excavating alive and dead
Pocillopora damicornis colonies. LEB-ICML-UNAM-1233;
Bahía de La Entrega (Huatulco, Oaxaca), 15°44'34"N
96°07'35"W; 6 m depth; 04 July 2005. LEB-ICML-UNAM-
1325; Cerro Pelón (Isla Isabel), 21°51'21"N 105°52'42"W; 21
m depth; 28 October 2005. Specimen excavating alive Pavona
sp. colonies. LEB-ICML-UNAM-1328; Antiguo Corral del
Figure 1. Sampling localities (letters) and distribution of the Aka-species along the coast of Mexico (numbers in parentheses) (1) Aka
cryptica sp. nov.; (2) Aka brevitubulata; (3) Aka coralliphaga.

Journal of the Marine Biological Association of the United Kingdom (2007)
1479Coral boring Aka-species from Mexico J.L. Carballo et al.
Risco (Punta Mita, Nayarit), 20°46'20"N 105°32'49"W; 2 m
depth; 18 March 2006. Specimen excavating alive and dead
Pocillopora verrucosa colonies. LEB-ICML-UNAM-1340; Isla
Cacaluta (Huatulco, Oaxaca), 15°43'08"N 96°09'43"W; 3 m
depth; 04 July 2005.
Description
Boring sponge found within corals, visible externally only
by the presence of erect, hollow fistules from 1 to 4.5 mm high
and from 0.8 to 1.8 mm in diameter (Figure 2A,B). Fistule
walls are from 31.5 to 75.6 µm in thickness. Most specimens
have from 2 to 4 fistules, but the holotype has numerous
fistules (from 2 to 5 fistules cm-2) and covers an area of 10
cm in diameter. Fistules are from 2 to 7.5 mm apart and they
are regularly spread on the substratum surface. The tops of
the fistules are covered with a sieve-like reticulation bearing
many ostia (Figure 3C), or end in an oscule of 0.7 to 1 mm
in diameter. Ostial fistules are circular or oval in section,
and characteristically distally widened (Figure 2A). Oscular
fistules are commonly straight or slightly curved, sometimes
gradually tapering from the middle (Figure 2B). Fistules
are compressible, brittle, and easily broken, and fusion (two
fistules) has only been observed in one specimen. Fistules
are white and pale yellow at the base close to the substrate,
and they are hardly distinguishable at first sight because this
species lives cryptically, and their exposed parts often blend
in well with background colours. In alcohol fistules turn
dark yellow. Inside the coral the sponge tissue is beige, and
pale brown in alcohol. The tissue inside the coral consists of
a pulpy organic mass secreting mucus.
Spicules: the spicules are slender oxeas, mostly slightly
curved at the centre or straight, ending in symmetrical
points, sometimes with mucronate tips. Very slim oxeas that
are also common in some specimens probably represent
young spicules (Figure 2C). They measure 67.5–(101.3)–
122.5 µm long and 1.3–(3.9)–6.3 µm in width (Table 1).
Figure 3. SEM images of skeletal characteristics of Aka cryptica
sp. nov. (A) External view of the oscular fistule; (B) cross section of
the ostial fistule showing the internal structure; (C) top view of an
ostial fistule.
Figure 2. Aka cryptica sp. nov. (A) Detail of the ostial fistule; (B) detail
of the oscular fistule; (C) SEM images of oxeas and derived spicules;
(D, E) choanosomal structure of the basal mass inside the coral.
Aka cryptica specimens Oxeas
Holotype: Length×width shaft
MNCN 1.01/363 90–(106)–117×2.5–(4)–5
Paratypes
BMNH 2006.6.30.1 87.5–(101.5)–120×3–(4.5)–5.5
LEB-ICML-UNAM-1196 90–(100.5)–112.5×2–(3.7)–5
LEB-ICML-UNAM-1233 80–(101.8)–115×2–(4)–5
LEB-ICML-UNAM-1325 90–(98)–103×2.5–(2.7)–3.8
LEB-ICML-UNAM-1328 87.5–(103.5)–122.5×1.3–(3.5)–5
LEB-ICML-UNAM-1340 67.5–(94.3)–110×2.5–(3.9)–6.3
Table 1. Comparative data for the dimensions of the oxeas (in µm) of
Aka cryptica sp. nov. specimens. Values in parentheses are means.

1480 J.L. Carballo et al. Coral boring Aka-species from Mexico
Journal of the Marine Biological Association of the United Kingdom (2007)
Figure 4. Excavating characteristics of Aka cryptica sp. nov. (A) Excavating pattern in a cross section of a coral branch; (B) excavating pat-
tern in a longitudinal section of a coral branch, (the arrows show the diaphragms); (C) SEM image of patterns erosion; (D) SEM images of
erosion scars on the walls of excavated chambers.
Skeleton: spicules are strewn in confusion or grouped in
directionless loose tracts in the basal mass inside the coral
(Figure 2D,E). In the outer wall of the fistule the oxeas form
a confused ill-def ined tangential ectosomal skeleton (Figure
3A). Towards the basal part of the f istule, and below the
surface it is possible to distinguish sinuous bands of spicule
bundles (from 15 to 18 spicules) running up, connected
transversally by narrower bundles (from 5 to 8 spicules), and
forming meshes from 150 to 240 µm wide, but without a clear
network. It is possible to distinguish this structure up to the
lower third of the fistule length, but after that, the skeleton
becomes a somewhat isodictyal reticulation of multispicular
bundles (from 20 to 45 µm thick), forming meshes from 125
to 255 µm wide (Figure 3B).
Erosion pattern: network of reticulate galleries with
irregularly spherical or ovoid chambers densely distributed
in the coral (Figure 4A). They measure from 0.6 to 2 mm
long and from 0.5 to 1 mm wide. Wall between adjacent
chambers is from 0.2 to 0.8 mm in thickness, and the
interconnection between the chambers is by means of a
diaphragm from 0.2 to 0.4 mm in diameter. Chambers
can be fused, forming long chambers parallel to the coral
surface. This species also occupies the natural pores of the
coral boring only the walls that separate the septa of the
coral and producing long chambers parallel to the coral
surface (Figure 4B), from 3.7 to 11 mm long and from 0.7 to
1 mm wide. The walls of the chambers and tunnels, present
a pitted surface where chips have been removed (Figure

Journal of the Marine Biological Association of the United Kingdom (2007)
1481Coral boring Aka-species from Mexico J.L. Carballo et al.
4C,D). They are sub-spherical in form and measure 20–50
µm in diameter.
Etymology
The specific name refers to the cryptic habitat where the
new species lives.
Distribution and ecology
Mexican Pacific Ocean (Nayarit and Oaxaca) (Figure 1).
Common in shallow water, from 2 to 7 m depth. The species
mainly bores dead and live stems and branches of the species
Pocillopora damicornis and P. verrucosa.
Remarks
This new species can be compared with the Aka-species
from the Indo-Pacific region. Aka diagonoxea Thomas, 1968
described from the Gulf of Mannar (Thomas, 1968) has
fistules that branch dichotomously or polychotomously, and
they are longer (up to 50 mm long) and wider (up to 4 mm
in diameter) than those in the new species. Moreover, it has
double angulated oxeas, and it excavates chambers 2 cm or
more in diameter inside coral. Aka paratypica Fromont, 1993
is similar to A. diagonoxea in colour, shape and size of oxeas
(Fromont, 1993), and it also has fistules longer than the
new species (40–60 mm). Aka mucosa (Bergquist, 1965) has
Figure 5. Aka brevitubulata (Rützler, 1974). (A) Ocular fistule (ex-current); (B) ostial fistule (in-current); (C) oxeas; (D) tissue filled boreholes
showing mucus covered tissue and fistules (the arrows show the fistules); (E) internal tissue and canals; (F) SEM image of erosion scars
(sponge chips removed).

1482 J.L. Carballo et al. Coral boring Aka-species from Mexico
Journal of the Marine Biological Association of the United Kingdom (2007)
very long (up to 50 mm high) and wide (7 mm in diameter)
black fistules and two categories of oxeas, characteristics
that are clearly different from the new species. Finally, Aka
maldiviensis Calcinai et al., 2000 also has longer fistules (5–7
cm high). The species nearest to ours is Aka minuta Thomas,
1972 recorded from the Gulf of Mannar, Seychelles and
Mozambique (Thomas, 1972, 1973, 1979), which also bores
colonies of Pocillopora damicornis (Linnaeus). However, this
species has oxeas with a sharp angle at the centre, it does
not form f istules, and it excavates chambers of up to 5 mm
in diameter, characteristics that are very different in the new
species. The species Aka trachys de Laubenfels, 1954 from
the central Pacific does not seem to be a valid species of
Aka since it has spined oxeas (named acanthostyles by the
author) (de Laubenfels, 1954).
The new species is also different from the species in the
Caribbean in the size of the fistule and excavating patterns.
For example, A. xamaycaensis Pulitzer-Finali, 1986 has white
fistules up to 11 cm long, and oxeas slightly curved (110–
125×4.5–6 µm) (excavation pattern is not described). Aka
brevitubulata (Pang, 1973) has yellow and longer fistules (9
mm high) than the new species, and produces a single large
spherical chamber completely filled with sponge tissue (Pang,
1973 and present study) (oxea dimensions: 119–148×6.9–9.1
µm); A. cachacrouensis (Rützler, 1971) has larger oxeas (up to
203.9×6.9 µm) than A. cryptica sp. nov., and dark greyish
brown fistules (up to 2.5 cm high) and produces a single
large chamber of 500 cm3. Aka coralliphaga (Rützler, 1971)
also has longer yellow fistules (up to 4 cm high) and oxeas
(142.1–156.3×5.0–6.4 µm), and it excavates a single chamber
of 20–130 cm3. Aka siphona has yellow to brown fistules up
to 15 cm long, excavates large galleries up to 20 cm across,
and has oxeas from 120 to 200 µm long (Laubenfels, 1949;
Hofman & Kielman, 1992).
Aka brevitubulata (Pang, 1973)
Synonymy: Siphonodictyon brevitubulatum Pang, 1973
(Figure 5)
Material examined
Holotype: [YPM No. 8717]. Discovery Bay, Jamaica;
Montastraea annularis; 15 m depth; collected by R. Keeley; 16
October 1968.
Specimens: BMNH 2006.6.29.12, Puerto Morelos
(Quintana Roo), 20.5°24'16.5"N 86°49.5'26.67"W, 09
September 2002, excavating Acropora palmata, back-reef 2
m. BMNH 2006.6.29.13, BMNH 2006.6.29.14, BMNH
Figure 6. Aka coralliphaga (Rützler, 1971). (A) Internal chamber filled with mucus-secreting tissue. The arrow shows a longitudinal-section
through fistule; (B) excavation chamber completely filled with tissue still adhering to the erosion scars (also Aka brevitubulata and unidenti-
fied nestling species); (C) oxeas; (D) SEM image of erosion scars created by chemical etching of sponge chips inside excavation galleries.

Journal of the Marine Biological Association of the United Kingdom (2007)
1483Coral boring Aka-species from Mexico J.L. Carballo et al.
2006.6.29.15, BMNH 2006.6.29.16, Puerto Morelos (Q.
Roo), 20.5°24'16.5"N 86°49.5'26.67"W, 11 November
2002, Acropora palmata, reef-front 5 m. BMNH 2006.6.29.17,
BMNH 2006.6.29.18, BMNH 2006.6.29.19, Puerto Morelos
(Q. Roo), 20.5°24'16.5"N 86°49.5'26.67"W, 28 July 2003,
Acropora palmata, reef crest, 1m. BMNH 2006.6.29.20, BMNH
2006.6.29.21, Puerto Morelos (Q. Roo), 20.5°24'16.5"N
86°49.5'26.67"W, 14 September 2003, Montastraea annularis
back-reef, 3 m.
Description
The tissue has a glistening appearance due to mucus
secretion. Fistules (between 3 and 9 mm high) are relatively
few in number (1–2 cm apart) due to the nature of excavation
activity of this sponge, which usually produces a large single
excavation. Oscular and ostial papillae do not fuse and, nor
has this species been observed to overgrow the substrate
(Figure 5A,B). Oscules are 2–5 mm wide. In life, fistules are
yellow. The internal sponge tissue is a dull, dark yellow.
Spicules: the oxeas are abundant and smooth, although
some with rather abrupt points at either end. Oxeas are
curved around the mid-region, with the degree of curvation
varying between individual oxeas. Some are almost straight
and some have a sharper bent mid-region, giving a more
curved ‘boomerang’ type appearance (Figure 5C). However,
there is little variation in observed length/width or curvation
of oxeas. Oxea length(s): 109.5–142 µm (mean: 133.7 µm)
width: 3.7–5.5 µm (mean: 4.2 µm).
Skeleton: oxea are abundant but without orientation in
the choanosome. Spicule arrangement within fistules is
confused, although the basal part of the f istule shows a
tendancy for oxeas to be placed tangentially.
Erosion pattern: this species characteristically forms a large
single spherical/subspherical excavation completely filled
with tissue (Figure 5D). It constructs galleries without clearly
outlined chambers or ducts. Long narrow channels connect
the chamber with the substrate surface (Figure 5E). These
canals may be up to 15 mm reaching an excavation of up to
4–5 cm in massive substrates. Borehole diameter is less in
branching substrates.
Distribution and ecology
This is a relatively common species at Puerto Morelos,
Mexican Caribbean (Figure 1). It was previously described
by Pang (1973) in Jamaica, and Hofman & Kielman (1992)
in Santa Marta (Colombia).
Remarks
First record in Mexico.
Aka coralliphaga (Rützler 1971)
(Figure 6)
Synonymy: Siphonodictyon coralliphagum forma obruta
Rützler, 1971
Material examined
Holotype: [USNM 24098]. Barbados, West Indies; 25 m
depth; 5 July 1969.
Specimens: BMNH 2006.6.29.22, Puerto Morelos
(Q. Roo), 20.5°24'16.5"N 86°49.5'26.67"W, 02
September 2002, Siderastrea siderea, reef-front 5 m.
BMNH 2006.6.29.23, BMNH 2006.6.29.24, BMNH
2006.6.29.25, Puerto Morelos (Q. Roo), 20.5°24'16.5"N
86°49.5'26.67"W, 14 November 2002, Siderastrea
siderea, reef front 8 m. BMNH 2006.6.29.26, BMNH
2006.6.29.27, BMNH 2006.6.29.28, Puerto Morelos (Q.
Roo), 20.5°24'16.5"N 86°49.5'26.67"W, 05 August 2003,
Diploria strigosa, reef crest, 1.5 m. BMNH 2006.6.29.29,
BMNH 2006.6.29.30, BMNH 2006.6.29.31, Puerto
Morelos (Q. Roo), 20.5°24'16.5"N 86°49.5'26.67"W, 19
September 2003, Diploria strigosa, back reef, 3 m.
Description
Cream/tan/yellow mucus-secreting tissue fills chambers.
This species does not encrust substrate. Ostial and oscular
fistules are separate. The ostial fistules are conspicuous,
creamy coloured globular protrusions of 2–4 mm height
and similar diameter (Figure 6A). Ostia are distributed
over the entire surface of the f istules with fusion of ostial
tubes observed in some samples. Oscular structures are of
lower relief (1–2 mm), but comparable diameter; they are
open-ended cylindrical tubes which bear a single osculum.
Fistules are circular and irregularly spaced with at least one
per bored chamber.
Spicules: tissue is dense in this species with abundant oxeas
exhibiting no clear orientation. Spiculation consists solely of
oxeas which are bent in the mid-region (Figure 6C). There
may be slight variation in the extent of the curvation with
some being almost straight. Oxea length(s): 98.5–149.6 µm
(mean: 145.5 µm) width: 1.8–7.3 µm (mean: 4.7 µm).
Skeleton: spicules are strewn in confusion forming no clear
skeletal pattern.
Erosion pattern: canals and connecting tunnels may be long
(3–8 mm length/1.5–2.5 mm diameter). Endolithic chambers
are large, although diameter can be variable (average 13–
20 mm). They are roughly subspherical, with some being
slightly more elongate than others; longest axis parallel to
substrate surface (Figure 6B). Chambers extend deep from
the substrate surface where the substrate allows (e.g. 1.5–2
cm), and may be 2–3 cm apart. The chambers are covered
by erosion scars created by the sponge (Figure 6D).
Distribution and ecology
This is a common species at Puerto Morelos reef (Figure
1). It was found in dead rubble substrate of Acropora palmata,
Montastraea annularis complex and Diploria strigosa. Aka
coralliphaga is widely distributed throughout the Caribbean.
The species has been cited in Barbados, Puerto Rico,
Jamaica, Dominica (Rützler, 1971), Colombia (Hofman
& Kielman, 1992) and Mexico (Cozumel, Quintana Roo)
(Lehnert, 1993).
Remarks
Rützler (1971) described two new species of the genus
Aka (A. cachacrouensis and A. coralliphaga) from the Caribbean
Sea (both as Siphonodictyon). The present specimens are most
closely related to A. coralliphaga, however, oxeas are longer
and thinner than those found in the present species.

1484 J.L. Carballo et al. Coral boring Aka-species from Mexico
Journal of the Marine Biological Association of the United Kingdom (2007)
We are grateful to the following sources of funding CONACYT
SEP-2003-C02-42550, CONABIO FB666/S019/99, CONABIO
FB789/AA004/02 and CONABIO DJ007/26. We thank Israel
Gradilla Martínez (Centro de Ciencias de la Materia Condensada)
for the SEM photographs, Clara Ramírez Jáuregui (ICML-
Mazatlán) for help with the literature, Alejandra Torres Ariño
for her hospitality during the sampling in Oaxaca. We are also
grateful to the personnel of the National Park isla Isabel for the
availability and the permission conferred for the collection of the
samples in the Isabel Island, and especially to Cayetano Robles
Carrillo, Gonzalo Perez Lozano and Jorge Castrejón for their help
during the samplings.
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Submitted 22 July 2006. Accepted 2 May 2007.