␥ 20 Mexican Coral Reefs Late 1800s

␥20
Mexican Coral Reefs
W. David Liddell and John W. Tunnell Jr.
The Mexican reefs of the southern Gulf of Mexico occur
in 2 distinct provinces, the Campeche Bank to the north
and west of the Yucatan Peninsula and the Veracruz shelf
reefs near the coast in the Tuxpan–Veracruz area of the
southwestern Gulf of Mexico. Reefs differ greatly in size,
geomorphology, and diversity, reflecting differences in
the underlying geologic structures and ambient environments. The reefs of the Campeche Bank, generally,
are better developed than those of the Tuxpan–Veracruz
area, perhaps due to the deleterious effects of high clastic sedimentation and cooler temperatures in the latter
area.
History of Research
The reefs of the southeastern Gulf of Mexico (Fig. 20.1)
have a long prehistory, dating back to the Mesozoic
opening of the Gulf of Mexico (Liddell 2007). Modern
historical accounts began with the English adventurer
William Dampier, who mentioned the prolific nesting
seabirds occurring on the islands of Alacrán and Triángulos (Dampier 1699). A century would pass before others again noted the vegetation, seabirds, sea turtles, and
West Indian monk seal associated with these and other
Campeche Bank islands (Smith 1838; Marion 1884; Ward
1887; Agassiz 1888).
Late 1800s
Professor Angelo Heilprin led the first scientific expedition to the coral reefs of the southern Gulf of Mexico in
1890. This expedition was sponsored by the Academy of
Natural Sciences of Philadelphia (now Philadelphia Academy of Sciences), which Heilprin directed. The expedition’s purpose was to “investigate the natural history of
the Yucatan Peninsula and Mexico” (Heilprin 1890). The
expedition documented the tropical nature of marine
biota of the southern Gulf and described corals and coral
reefs (Heilprin 1890) and many other marine invertebrates (Ives 1890, 1891; Baker 1891). Heilprin (1890) also
described reefs and islands occurring off the city of Veracruz and noted the “vast quantity of coral” used in construction there.
1900s–1970s
With the exception of Joubin’s 1912 map of coral reefs,
little research was conducted on southern Gulf reefs until
Smith’s (1954) work. Smith utilized Heilprin’s (1890) and
Joubin’s (1912) works and various unpublished sources
and nautical charts (refer to Tunnell 2007a, his Table 1.1)
to create a coral reef distribution map. Smith also compiled a list of coral species from the southern Gulf. In
1955, Moore (1958) briefly visited Blanquilla Reef off Tuxpan. He described 44 invertebrate species from this, the
most northerly emergent reef in the southwestern Gulf of
341
342 ~ Liddell and Tunnell
Mexico. In 1956, Emery (1963) sampled reef sediments
offshore of the city of Veracruz. His work was published
in both English and Spanish, raising the level of interest in
this unique (terrigenous) setting for coral reefs. Kornicker
et al. (1959) took part in a large, collaborative effort to
study Alacrán, the most northerly reef on the Campeche
Bank (Fig. 20.1). Although the expedition largely focused
on geology, several biological studies also resulted from
it, making Alacrán Reef one of the best-­known Gulf reefs
(Table 20.1).
From 1959 to 1963, the Department of Oceanography
at Texas A&M University undertook a major, multiyear
study of the Campeche Bank or Yucatan Shelf in the southeastern Gulf. The project was funded by the National Science Foundation, the American Petroleum Institute, the
Office of Naval Research, Shell Development Company,
and Mobil Oil Company. The project was the longest and
most productive study of southern Gulf reefs to that date
and contributed greatly to the knowledge of coral reefs
and reef and shelf sediments of the region (Logan 1962,
1969a, b; Logan et al. 1969). In addition, Cayo Arenas was
studied in more detail as a side project (Busby 1966).
During the 1960s, interest and research on all southern Gulf of Mexico coral reefs increased greatly (Tables
20.1, 20.2). Because the reef localities were remote, field
studies tended to be performed during extended expeditions to the reef systems. Huerta M. and Barrientos (1965)
reported on the algae of Blanquilla and Isla de Lobos reefs,
near Tuxpan. Rigby and McIntire (1966) described the
geology and ecology of Isla de Lobos Reef, while Chamberlain (1966) reported on gorgonians from Lobos that
were studied during the same Brigham Young University expedition. Hidalgo (Hidalgo and Chávez 1967) and
Chávez (Chávez et al. 1970; Chávez 1973; B
­ autista-­Gil and
Chávez 1977) conducted multiple expeditions to Isla de
Lobos Reef with their students from the Instituto Politécnico Nacional in Mexico City. Chávez (1966) reported on
the fishes of Triángulos and Cayo Arenas reefs.
Students from the Universidad Nacional Autónoma de
Mexico conducted their “professional theses” on a variety of topics within the Veracruz Reef system. Villalobos
summarized much of this work in the proceedings of an
international symposium on the Caribbean Sea and adjacent regions (Villalobos 1971) and in a later review article
Figure 20.1. Geographic distribution of coral reefs within the southern Gulf of Mexico. Tuxpan and Veracruz systems are in the
southwestern Gulf of Mexico and Campeche Bank Reefs are in the southeastern Gulf of Mexico (reprinted from Tunnell 2007b,
with permission of Texas A&M Press.
Mexican Coral Reefs ~ 343
Table 20.1. Principal literature by reef or island for certain coral reefs and islands on the Campeche Bank. Listed
counterclockwise from Alacrán in the north to Cayos Arcas in the south.
Subject by area
Alacrán
bathymetric mapping
communities
benthic communities
reef communities
geology
organisms
algae
birds
fish
foraminiferans
mollusks
stomatopods and decapods
vegetation
reef growth model
sediments
spatial distribution
References
Liceaga-­Correa and Euan-­Avila 2002
Solís 1990
Liddell and Ohlhorst 1988
Kornicker et al. 1959; Kornicker and Boyd 1962; Hoskin 1962, 1963, 1966, 1968; Bonet 1967;
Macintyre et al. 1977
Múzquiz 1961
Fosberg 1961, 1962; Folk 1967; Boswall 1978; Tunnell and Chapman 1988, 2001
Hildebrand et al. 1964; ­González-­Gandara et al. 1999; G
­ onzález-­Gandara and Arias-­González
2001a, 2001b; Brulé et al. 2003; G
­ onzález-­Gandara et al. 2003
Davis 1964
Rice and Kornicker 1962; Aranda et al. 2003
Martínez-­Guzmán and ­Hernández-­Aguilera 1993
Marion 1884; Millspaugh 1916; Fosberg 1961, 1962; Bonet and Rzedowski 1962; Folk 1967
Bosscher and Schlager 1992
Folk 1962, Folk and Robles 1964; Folk 1967; Folk and Cotera 1971; Novak 1992; Novak et al. 1992
Torruco et al. 1993
Cayo Arenas
geology
Busby 1966
Triangulos and Cayo Arenas
organisms
coral
fish
sponges
environmental
Carricart-­Ganivet and ­Beltrán-­Torres 1993
Chávez 1966
Hernández 1997
Ferre-­D’Amare 1995
Cayos Arcas
organisms
coral
environmental
meteorology
Farrell et al. 1983
Ferre-­D’Amare 1995
Salas de Leon et al. 1992
(Villalobos-­Figueroa 1980). Other studies on southern
Veracruz reefs from this time included algae (Huerta M.
1960) and foraminiferans (Lidz and Lidz 1966). Morelock and Koenig (1967), Edwards (1969), and Freeland
(1971) conducted geologic studies during the 1960s on
the southern Veracruz reefs.
1970s–1990s
From 1970 to 1990, three groups were largely responsible
for research on the southern Gulf of Mexico coral reefs:
(1) Secretaría de Marina, Dirección General de Oceanografía, (2) Texas A&M U
­ niversity-­Corpus Christi, and
(3) Universidad Nacional Autónoma de México. In the
late 1980s and 1990s, Universidad Veracruzana in Jalapa,
and Centro de Investigaciones y Estudios Avanzados
(CINVESTAV)–Unidad Mérida of the Instituto Politécnico Nacional, also contributed to research on Veracruz
reefs and Alacrán Reef, respectively.
The reefs of northern Veracruz State are the least studied of those in the southwestern Gulf region due to their
remoteness. Of the 6 northernmost reefs, Isla de Lobos
has received the most attention (Table 20.2).
Over 20 coral reefs exist in the extreme southwestern
Gulf, located offshore of the city of Veracruz and offshore
of the fishing village of Antón Lizardo. Since these reefs
are more accessible than the northern Veracruz reefs, they
have received considerably more study (Table 20.2). Per-
344 ~ Liddell and Tunnell
Table 20.2. Principal literature by reef or island for certain coral reefs and islands on the Tuxpan–Veracruz area.
Listed from Blanquilla Reef in the north to Enmedio Reef in the south.
Reef or island
Tuxpan Reef system
Blanquilla
Subject
References
algae
Moore 1958; Huerta M. and Barrientos 1965
algae
crustaceans
crabs
fish
foraminiferans
gorgonians
mollusks
polychaetes
reef communities
Huerta M. and Barrientos 1965; Rigby and McIntire 1966; Hidalgo and Chávez 1967
Ray 1974
Allen 1982
Márquez-­Espinosa 1976
Bautista-­Gil and Chávez 1977
Chamberlain 1966
Tunnell 1974, 1977
Roberts 1981
Chávez et al. 1970; Chávez 1973
algae
geology
foraminiferans
benthic communities
Huerta M. 1960
Emery 1963; Morelock and Koenig 1967; Freeland 1971; Hernandez 1987
Lidz and Lidz 1966
Kühlmann 1975
La Blanquilla
coral ecology
ecology
fish
hydrography
sponges
Santiago 1977
Villalobos 1971
Reséndez-­Medina 1971
Rodríguez and Fuentes 1965
Green 1977
Verde
algae
coral taxonomy
lead pollution
stomatopods & decapods
Mateo-­Cid et al. 1996
Beltrán-­Torres and ­Carricart-­Ganivet 1993
Horta-­Puga and ­Ramírez-­Palacios 1996
Morales-­García 1986
Sacrificios
stomatopods & decapods
Morales-­García 1987
Enmedio
algae
ciliates
corals
crabs
echinoderms
fish
IXTOC I oil impacts
mollusks
shrimps
sponges
Huerta M. et al. 1977; Lehman and Tunnell 1992a, 1992b; Lehman 1993
Lubel 1984
Rannefeld 1972; Nelson 1991
Rickner 1975, 1977; Allen 1982
Henkel 1982
Choucair 1992; Riley and Holt 1993
Tunnell and Dokken unpublished; Baca et al. unpublished
Tunnell 1974, 1977
White 1982
Stinnett 1989
Isla de Lobos
Veracruz Reef system
Veracruz Area
haps the most studied of all reefs in this region is Enmedio Reef. Gutiérrez et al. (1993) compared the coral reefs
of the Veracruz region to those within the Sian Ka’an
Biosphere Reserve located along the Caribbean coast of
Quintana Roo, Mexico.
The Mexican Navy has contributed oceanographic
information, as well as information on reef biology, for
the Veracruz reef system through its oceanographic program (Instituto de Investigación Oceanográfica del Golfo
y Mar Caribe). This program first operated from a small
station (Estación Oceanográfica) in the city of Veracruz,
but it is now housed within a new institute building at
Antón ­Lizardo.
The relatively great distance from shore continued to
limit the number of reef studies on the Campeche Bank.
Macintyre et al. (1977) published a report on the thickest recorded Holocene reef section at Alacrán Reef. During the 1980s, coral diversity and zonation were charac-
Mexican Coral Reefs ~ 345
terized for Cayos Arcas (Farrell et al. 1983) and Alacrán
(Liddell and Ohlhorst 1988) reefs. Additionally, several
authors reviewed all Mexican Atlantic coral reefs, including environmental problems and human impacts (Tables
20.1, 20.2).
During the 1990s an increasing number of both organismal and ecological studies continued on the Campeche
Bank and Veracruz Reef system. Other studies focused on
reef sediments and a computer simulation model of reef
growth (Tables 20.1, 20.2).
2000s to the Present
A number of fish studies were conducted during the early
2000s, especially on Alacrán Reef (Tables 20.1, 20.2).
Other studies included Montastraea annularis growth
rates, other invertebrates, and seabirds on Campeche
Bank reef islands. In recent studies, geologic facies were
charted through time on Campeche Bank reefs (Tables
20.1, 20.2).
Jordán-­Dahlgren and ­Rodríguez-­Martínez (2003)
summarized work on all Mexican coral reefs within the
Gulf of Mexico and Caribbean. ­Jordán-­Dahlgren (2004)
provided a sobering, though brief, environmental status report on southern Gulf reefs. Reefs in the extreme
southwestern Gulf, located off the city of Veracruz
and the village of Antón Lizardo, have been negatively
affected the most, and those on the Campeche Bank have
been affected the least. The reefs near Tuxpan in northern Veracruz State are intermediate in terms of negative
effects (Tunnell 1992).
Reef Distribution and Structure
There are 46 named coral reefs in the southern Gulf of
Mexico: 31 in the southwestern Gulf off the state of Veracruz (Veracruz Shelf reefs) and 15 in the southeastern
Gulf on the Campeche Bank (refer to Tunnell 2007b, his
Table 2.1) (Fig. 20.1). In addition, Dahlgren (1993) listed
approximately 10 named and over 25 unnamed banks on
the Campeche Bank for which there is little or no scientific information.
Coral reefs in the southwestern Gulf are typically
located at nearshore (<200 m) to mid-­shelf (22 km) distances on a narrow, terrigenous shelf (Morelock and Koenig 1967; Ferre-­D’Amare 1985). The Veracruz region
is characterized by subhumid to humid climates of high
rainfall and substantial mainland drainage. Consequently,
terrigenous sediment shed from the Sierra Madre Orientál and Trans-­Mexican Neovolcanic Belt greatly influences
the nearshore reefs. Also, during the rainy season lowered
salinities may occur in some areas (e.g., practical salinity of 18.3 at Veracruz compared to 36 for normal oceanic
waters) (Ferre-­D’Amare 1985; Tunnell 1988). The ­Tuxpan–
Veracruz area is subject to low winter temperatures with
lows ranging from 8–10 °C (air) at Veracruz down to,
infrequently, 0 °C (air) at Tampico (Ferre-­D’Amare 1985).
Reefs are absent to the north of the Tuxpan Reef system,
most likely due to low winter water temperatures. Reefs are
also absent to the south of the Veracruz Reef system, perhaps due to upwelling (Ferre-­D’Amare 1985).
In the southeastern Gulf, reefs are located far (130–
200 km) offshore on a wide carbonate shelf, the Campeche
Bank, primarily along the 55-­m depth contour (Logan
1969a, b). In contrast to the southwestern Gulf, southeastern Gulf reefs occur in a semiarid climate, surrounded by
oceanic water from the Caribbean and the Gulf of Mexico,
and they are not affected by mainland drainage. The general absence of reefs along the Gulf coastal areas of the
Yucatan Peninsula may be due to low winter temperatures (8–10 °C air temperatures; Ferre-­D’Amare 1985),
the effects of which would be more pronounced along
the coastline than farther out on the shelf where the bank
reefs occur. Freshwater discharge is also possible through
the Yucatan karst system into coastal areas. Fringing or
­barrier-­type reefs are conspicuously absent from the
northern and western (Gulf) coastal areas of the Yucatan
Peninsula, although these reef types are extensively developed along the eastern (Caribbean) side of the Yucatan
Peninsula and southward near Belize.
In contrast to adjacent, low-­energy coastal areas such
as the Florida Keys and Belize where mangroves line the
shoreline and seagrasses predominate as nearshore submarine vegetation, shoreline areas of the mainland consist
of m
­ oderate-­energy sandy beaches or rocky shores (volcanic) in the southwestern Gulf and low-­energy sandy
beaches or rocky shores (limestone) in the southeastern
Gulf. Also, unlike the scattered, ­shallow-­water patch reefs
associated with the former, the southern Gulf reefs are
largely submerged, ­mountain-­like structures scattered
across the continental shelf.
Southwestern Gulf Reefs
The southwestern Gulf coral reefs are clustered in 2 systems, the Tuxpan Reef system to the north (Fig. 20.2) and
the Veracruz Reef system to the south (Fig. 20.3). Most of
346 ~ Liddell and Tunnell
Figure 20.2. The Tuxpan Reef system (TRS) including 6 emergent platform reefs, 3 off Cabo Rojo and 3 northeast of Tuxpan,
Veracruz, Mexico (reprinted from Tunnell 2007b, with permission of Texas A&M Press.
these reefs are relatively small (only a few square kilometers), exhibit low relief, and are located near shore (Ferre-­
D’Amare 1985). There are 6 emergent platform reefs in
the Tuxpan Reef system (Fig. 20.2). Isla de Lobos Reef is
the only one in the Tuxpan Reef system that has an island
and, thus, has served as a base camp for several expeditions (e.g., Rigby and McIntire 1966; Chávez et al. 1970;
Tunnell 1974). At 21°30′14″N, Blanquilla Reef is the most
northerly emergent coral reef in the western Gulf of Mexico (Moore 1958).
In the Veracruz Reef system in the far southwestern
Gulf of Mexico, 25 coral reefs occur in 2 subgroups, one
located off the city of Veracruz and one off the village of
Antón Lizardo (Fig. 20.3). The first, or northern, group
consists of smaller reefs located more onshore than those
of the other grouping. This northern group of 13 reefs
is located off the city of Veracruz and includes 8 emergent platform reefs (Gallequilla, Anegada de Adentro, La
Blanquilla, La Gallega, Pájaros, Isla Verde, Tierra Nueva,
and Isla de Sacrificios), 2 submerged bank or patch reefs
(Bajo Mersey and Lavandera), and 3 fringing reefs (Punta
Gorda–Punta Majahua, Hornos, and Punta Mocambo).
Three of the reef platforms in the northern Veracruz
Reef system have islands associated with them (La Blanquilla, Isla Verde, and Isla de Sacrificios). The island of La
Blanquilla is a sandy cay, always changing shape, whereas
Isla Verde and Isla Sacrificios are densely vegetated and
fairly stable. Low and mostly natural vegetation is found
on Isla Verde. Isla Sacrificios has a large, important lighthouse as well as a public visitor’s area, and includes many
exotic transplants among the natural vegetation.
The second, or southern, group of reefs in the Veracruz Reef system consists of 12 larger reefs occurring relatively farther offshore than those of the northern group.
These include emergent p
­ latform-­type reefs (Anegada
de Afuera, Topatillo, Santiaguillo, Anegadilla, Polo, Isla
de Enmedio, Aviso, Blanca, Chopas, El Rizo, Cabezo,
and El Giote) located off the fishing village of Antón Lizardo. This group of reefs contains the 3 largest reefs in
the southwestern Gulf (Afuera, Chopas, and Cabezo). El
Giote Reef is the smallest and nearest to shore and has a
navigational light stand on it.
Four of the reefs off Antón Lizardo have islands (Santiaguillo, Cabezo, Isla de Enmedio, and Chopas). Chopas
Reef actually has 2 islands, Isla Salmedina and Isla Blanca;
both of these islands have only low natural vegetation.
Isla de Enmedio has low, natural vegetation, as well as a
dense canopy of transplanted shade trees. El Aguila (on
Cabezo) and Santiaguillo islands have little vegetation
and are composed almost totally of coral rubble with little
or no sand. Topatillo Reef used to have a coral rubble cay
consisting of Acropora cervicornis (staghorn coral) rubble,
but it no longer exists. After the death of A. cervicornis
and A. palmata (elkhorn coral) in the late 1970s and early
1980s (Tunnell 1992), this small island began to erode
because of the lack of living Acropora, which protected the
island from wave action.
Southeastern Gulf Reefs
In the southeastern Gulf of Mexico reefs occur in a
broad arc running from the ­north-­central portion of the
Campeche Bank (Alacrán Reef) to the southwest (Cayos
Arcas). The reefs occur near the shelf edge, some 100–
200 km from the coastline of the Yucatan Peninsula (Fig.
20.1). The absence of reefs on northeastern portion of the
shelf may be due to upwelling of cold (17–18 °C) water
(Logan 1969b; Glynn 1973; Merino 1997).
Eleven named emergent reefs and 4 named submerged
reefs are located along the outer continental shelf of the
Campeche Bank. The Campeche Bank reefs have been
subdivided (Logan 1969b) into geomorphic categories
Mexican Coral Reefs ~ 347
Figure 20.3. The Veracruz Reef system, including reefs offshore from the city of Veracruz and fishing village of Anton Lizardo, Veracruz, Mexico (reprinted from Tunnell 2007b, with permission of Texas A&M Press.
with the principal subdivision between submerged reef
banks (e.g., Bancos Ingleses, Banco Pera, Bajo Nuevo,
and Bajos Obispos) and emergent reef banks or “walls”
(see Tunnell 2007b, his Table 2.1) (Fig. 20.1). The latter
are further subdivided into (1) solitary reef knolls (Nuevo
and Triángulo Oeste), (2) linear reef walls (Triángulo
Este-­Sur), (3) crescent reef walls (e.g., windward reef at
Alacrán Reef and parts of Cayo Arenas and Cayos Arcas),
and (4) reef complexes with multiple emergent reef walls.
Cayo Arenas, Triángulos, and Cayos Arcas are all considered complexes having multiple emergent platforms and
islands. Nuevo Reef has a single emergent platform and
ephemeral island. The majority of these reefs form arcs
that are convex to the northeast, reflecting the prevalent
­northeast-­southwest wave progression (Logan 1969b).
Alacrán Reef is the most northerly reef in the entire
southern Gulf of Mexico. It is also, by far, the largest reef in
the southern Gulf, occupying some 13 × 25 km. It has an
extensive ­shallow-­water lagoon with small patch reefs and
water depths up to 22 m. Alacrán Reef is the most studied
Campeche Bank reef and has been the subject of numerous geological and biological investigations (Table 20.1).
Liddell and Ohlhorst (1988), as part of a CINVESTAV
and Armada de Mexico expedition, quantitatively characterized community structure and diversity of a 20-­m
windward reef site. They found that the bottom consisted
of 76% total living cover comprised of macroalgae (45%),
corals (11%), non-­coral cnidarians (chiefly gorgonians,
5.4%), coralline algae (5.2%), and sponges (5.2%). Corals
consisted of 12 species with a diversity of 1.86 (H′, natural log). Torruco et al. (1993) characterized the structure
of benthic communities inhabiting the lagoon at Alacrán.
Alacrán Reef has 5 vegetated islands: Desterrada,
Desertora, Perez, Chica, and Pájaros. Arenas has one
large, named vegetated island, Cayo Arenas, and 3 smaller
unvegetated and unnamed coral rubble cays. Triángulos has at least 2 islands, Triángulo Oeste and Triángulo
Este. Finally, Cayos Arcas has 3 vegetated islands: Cayo
348 ~ Liddell and Tunnell
del Centro, Cayo del Oeste, and Cayo del Este. Manned
lighthouses are found at Isla Perez, Cayo Arenas, Triángulo Oeste, and Cayo del Centro.
In addition to the “true” reefs mentioned above, biostromes lacking scleractinian frameworks also occur on
the Campeche Bank, particularly in the 18–60 m depth
range (Logan 1969b). These hard banks form relatively
thin veneers over previously existing topography. These
veneers consist of 1–18–cm diameter, concentrically layered nodules of crustose coralline algae and foraminifera.
These nodular crusts are most common on the northern
shelf and are much less common on the western shelf, perhaps due to increased terrigenous sedimentation there.
Reef Sediment Composition
Numerous studies have examined the composition of
Holocene reef sediment from the southern Gulf of Mexico, particularly Alacrán Reef. Studies of certain areas,
such as Cayos Arcas and Triángulos on the Campeche
Bank and the T
­ uxpan-­Tampico portion of the western
Gulf coastal plain, are lacking. Although both geographic
(e.g., increased terrigenous sediment in the southwestern
Gulf) and bathymetric variation in sediment composition
occur, generalizations may be made.
Southwestern Gulf Reefs
Relatively little is known about the geology of the Tuxpan
reefs. The seafloor north and south of the Isla de Lobos
Reef at Tuxpan (Fig. 20.2) is dominated by reef-­derived
calcareous sand; whereas, the seafloor to the west of the
reef has both reef-­derived and terrigenous material (Rigby
and McIntire 1966) consisting of quartz, ferromagnesium
minerals, volcanic rock fragments, and obsidian, presumably transported from the interior by the Río Tuxpan or
Río Panuco. The seafloor immediately east of the reef is
dominated by rocky reef debris. The linear trend of Isla de
Lobos, Medio, and Blanquilla reefs might reflect the control of relict Pleistocene or older topography (Rigby and
McIntire 1966).
The Veracruz reefs are somewhat more studied from
a geologic standpoint than those to the north. Emery
(1963), Edwards (1969), and Freeland (1971) examined
the mixed t­errigenous-­carbonate sediments occurring
off the city of Veracruz (Fig. 20.3). Morelock and Koenig
(1967) studied the reefs to the south of Veracruz at Antón
Lizardo. These reefs are located parallel to the shoreline
and perpendicular to the prevailing wave-­approach direction. They are strongly influenced by storm waves and are
largely barren of sandy sediment on their windward sides,
where well-­developed boulder ramparts occur. Again, the
reefs are thought to have been established on antecedent Pleistocene dunes. The authors concluded that much
of the terrigenous sand and gravel present on the shelf is
late Pleistocene (Wisconsinan) in age. ­Present-­day terrigenous sedimentation rates are very low, thus allowing the
reefs to develop. Even so, carbonate sediments are found
to accumulate only in the immediate area of the coral
reefs and rarely exceed 50% calcium carbonate by weight
(Morelock and Koenig 1967).
Southeastern Gulf Reefs
Logan (1969a) sampled sediments from both reef and
open shelf settings on the Campeche Bank, noting that
sediments from reefs and associated environments (e.g.,
lagoons) were dominated by coral, coralline and calcareous algae, and foraminifera. In contrast, mollusks, echinoids, and bryozoans were much more important sediment constituents on the open shelf.
Alacrán Reef is the most studied of the Campeche
Bank reefs. Kornicker and Boyd (1962) provided a general overview of the sedimentology and biotic zonation
of Alacrán Reef and lagoon. They noted that considerable
lateral variation occurred in the organisms present and
the bottom character around the reef complex.
Folk and Robles (1964) examined the sedimentology
of Isla Perez, the largest leeward island on Alacrán Reef.
The general composition of the sand-­sized fraction was
60% Halimeda, 25% coral, and 15% foraminifera and
other grains, whereas the coarser fractions were dominated by coral (e.g., coral stick ramparts generated by
storms). Folk (1967) and Folk and Cotera (1971) focused
on the origin, geomorphology, and size sorting occurring
on sand cays off the leeward side of Alacrán Reef, noting
that these were areas of intense biotic sediment production.
A core taken from Isla Perez by Macintyre et al. (1977)
documented 30 m of Holocene reef growth (principally by
the coral Acropora cervicornis). This translates into accumulation rates of 12 m / 1000 yr, considerably higher than
the rates previously described for Jamaica by Land (1974).
The Holocene reef that was cored was situated on top of a
­similar-­appearing pre-­Holocene reef that extended from
33 m down to 50–60 m and the shelf platform (Bonet
1967).
Mexican Coral Reefs ~ 349
Hoskin (1963) conducted a detailed facies analysis of
Alacrán Reef. Seventeen environments were characterized, based on sediment composition and size, including (from east to west) windward shelf, lower windward
slope, upper windward slope, surge channels, boulder
rampart, windward reef flat, moat, Thalassia beds, cellular reef surface, cellular reef deep, pinnacle reef, lagoon
proper, patch reefs, shallow sand, leeward reef surface,
lower leeward slope, and leeward shelf. Halimeda, coral,
and fecal pellets were the dominant components of the
sand-­sized sediment fraction. Halimeda was most abundant on the windward and leeward reefs; coral reached its
peak abundance on the tops of lagoon pinnacle reefs; and
fecal pellets were most abundant in deeper lagoonal areas.
In contrast, foraminifera, coralline algae, mollusks, and
aggregate grains varied little across these environments.
Hoskin (1966) examined eodiagenesis (near-­surface
diagenesis) occurring on pinnacle reefs in the Alacrán
Reef lagoon. Diagenetic processes observed included pelletization of lagoon floor muds, macroboring of coral by
lithophagid bivalves, microboring of coral and mollusk
fragments by endolithic algae, aragonite cementation in
cavities, and recrystallization of coral grains.
Novak (1992) and Novak et al. (1992) provided a quantitative description of Alacrán Reef sediment from fore-­
reef and back-­reef settings. For all samples, coral, the
calcareous green alga, Halimeda, and coralline algae constituted well over 70% of all grains by volume. The distribution of grains was similar to the distribution of the
living, s­ ediment-­producing organisms, with coral grains
being relatively more abundant in fore-­reef samples and
Halimeda grains in back-­reef samples. In contrast to the
findings of Hoskin (1963), fecal pellets were relatively
minor sedimentary constituents of their samples and
coralline algae were more important (Novak et al. 1992).
The back-­reef samples studied by Novak et al. (1992)
were from the shallower, eastern portion of the lagoon,
not the deeper, p
­ ellet-­rich lagoon areas and pinnacle reefs
sampled by Hoskin (1963). Cluster analysis delineated
3 Alacrán Reef lithofacies: back reef, shallow fore reef
(<10 m), and deep fore reef (>10m) (Novak et al. 1992).
Conclusions
The reefs of the southern Gulf of Mexico have a long history of study. They exhibit a diversity of forms, including
small, nearshore shelf reefs (e.g., Veracruz Shelf Reefs),
fringing reefs (Veracruz), and large, open-­shelf atolls
(e.g., Alacrán). The overall development of reefs in the
southern Gulf of Mexico may be limited by winter temperatures. In addition, reefs along the Tuxpan–Veracruz
coast may be limited by the abundant supply of terrigenous sediment being shed from the Sierra Madre Orientál and Trans-­Mexican Neovolcanic Belt and possibly by
reduced salinities in the rainy season. Unfortunately, most
of these reefs increasingly are being affected by economic
development (e.g., Farrell et al. 1983; Ferre-­D’Amare
1985; Tunnell 1992; Chávez and Tunnell 1993) (Tables
20.1, 20.2), particularly those in the southwestern Gulf off
Veracruz (city) and Antón Lizardo.
Acknowledgments
We dedicate this chapter to the memory of our good
friend and colleague, Mauricio Garduno Andrade, who
participated on many of our expeditions to the reefs of
Mexico and whose research contributed to turtle conservation biology. We thank the Texas A&M University
Press for permission to use figures 20.1–20.3. Portions of
this article were originally published in a different form in
Liddell (2007) and Tunnell (2007a, 2007b).
References
Agassiz, A. 1888. Three Cruises of the United States Coast and
Geodetic Survey Steamer “Blake,” in the Gulf of Mexico, in the
Caribbean Sea, and Along the Atlantic Coast of the United
States, from 1877 to 1880, vol.1. Boston and New York:
Houghton, Mifflin and Company. 314 pp.
Allen, R. L. 1982. The reptant decapods of Enmedio and Lobos
coral reefs, Southwestern Gulf of Mexico. M.S. thesis,
Corpus Christi State University, Corpus Christi, Tex. 65 pp.
Aranda, D. A., E. B. Cárdenas, I. M. Morales, A. Z. Zárate, and
T. Brulé. 2003. A review of the reproductive patterns of
gastropod mollusks from Mexico. Bulletin of Marine Science
73:629–41.
Baker, F. C. 1891. Notes on a collection of shells from southern
Mexico. Proceedings of the Academy of Natural Sciences of
Philadelphia 43:45–55.
Bautista-­Gil, M. L., and E. A. Chávez. 1977. Contribución al
conocimiento de los foraminíferos del arrecife de Lobos,
Veracruz. Pp. 90–103 in F. Manrique (ed.), Memorias del V
Congreso Nacional de Oceanografía. Guaymas, Sonora,
México, 22–25 de Octubre 1974.
Beltrán-­Torres, A., and J. P. C
­ arricart-­Ganivet. 1993. Skeletal
350 ~ Liddell and Tunnell
morphologic variation in Montastrea cavernosa (Cnidaria:
Scleractinia) at Isla Verde Coral Reef, Veracruz, México.
Revista de Biología Tropical 41(3):559–62.
Bonet, F. 1967. Biogeología subsuperficial del Arrecife
Alacranes, Yucatán, Boletín del Instituto de Geología
Universidad Nacional Autónoma de México 80:1–192.
Bonet, F., and J. Rzedowski. 1962. La vegetación de las islas del
Arrecife Alacranes, Yucatán (México). Anales de la Escuela
Nacional de Ciencias Biológicas (México) 6(1–4):15–59.
Bosscher, H., and W. Schlager. 1992. Computer simulation of
reef growth. Sedimentology 39:503–12.
Boswall, J. 1978. The birds of Alacran Reef, Gulf of Mexico.
Bulletin of the British Ornithologists’ Club 98(3):99–109.
Brulé, T., X. Renán, T. Colás-­Marrufo, Y. Hauyon, and A. N.
Tuz-­Sulub. 2003. Reproduction in the protogynous black
grouper (Mycteroperca bonaci (Poey)) from the southern
Gulf of Mexico. Fishery Bulletin 101:463–75.
Busby, R. F. 1966. Sediments and reef corals of Cayo Arenas,
Campeche Bank, Yucatan, Mexico. Technical Report
TR-­187. Washington, D.C.: U.S. Naval Oceanographic
Office. 58 pp.
Carricart-­Ganivet, J. P., and A. U. ­Beltrán-­Torres. 1993.
Zooxanthellae and chlorophyll a responses in the scleractinian coral Montastrea cavernosa at T
­ riangulos-­W. Reef,
Campeche Bank, Mexico. Revista de Biología Tropica
41(3):491–94.
Chamberlain, C. K. 1966. Some Octocorallina of Isla de Lobos,
Veracruz, Mexico. Brigham Young University Geology
Studies 13:47–54.
Chávez, E. A. 1973. Observaciones generales sobre las
comunidades del arrecife de Lobos, Veracruz. Anales de la
Escuela Nacional de Ciencias Biológicas (México) 20:13–21.
Chávez, E. A., E. Hidalgo, and M. L. Sevilla. 1970. Datos acerca
de las comunidades bentónicas del arrecife de Lobos,
Veracruz. Revista de la Sociedad Mexicana de Historia
Natural 31:211–80.
Chavez, E. A., and J. W. Tunnell Jr. 1993. Needs for management and conservation of the southern Gulf of México. Pp.
2040–53 in O. T. Magoon, W. S. Wilson, H. Converse, and
L. T. Tobin (eds.), Coastal Zone ’93: Proceedings of the 8th
Symposium on Coastal and Ocean Management, vol. 2. New
York: American Society of Civil Engineers.
Chávez, H. 1966. Peces colectados en el Arrecife Triángulos
Oeste y en Cayo Arenas, Sonda de Campeche, México. Acta
Zoológica Mexicana 8(1):1–12.
Choucair, P. C. 1992. A quantitative survey of the ichthyofauna
of Arrecife de Enmedio, Veracruz, Mexico. M.S. thesis,
Corpus Christi State University, Corpus Christi, Tex. 59 pp.
Dahlgren, E. J. 1993. El ecosistema arrecifal coralino del
Atlántico mexicano. Revista de la Sociedad Mexicana de
Historia Natural, Mexico City 44:157–75.
Dampier, W. 1699. Voyages and Descriptions, Volume 2, Part 2:
Two Voyages to Campeachy, 2nd ed. (microfiche). London
James: Knapton.
Davis, R. A. Jr. 1964. Foraminiferal assemblages of Alacran
Reef, Campeche Bank, Mexico. Journal of Paleontology
38(2):417–21.
Edwards, G. S. 1969. Distribution of shelf sediments, offshore
from Anton Lizardo and the Port of Veracruz, Veracruz,
Mexico. M.S. thesis, Texas A&M University, College Station,
Tex. 74 pp.
Emery, K. O. 1963. Coral reefs off Veracruz, Mexico. Geofísica
Internacional 3:11–17.
Farrell, T. M., C. F. D’Elia, L. Lubbers, and L. J. Pastor. 1983.
Hermatypic coral diversity and reef zonation at Cayo Arcas,
Campeche, Gulf of Mexico. Atoll Research Bulletin no. 270.
7 pp.
Ferre-­D’Amare, A. R. 1985. Coral reefs of the Mexican Atlantic:
a review. Pp. Proceedings of the Fifth International Coral Reef
Congress 6:349–54.
———. 1995. Prospección ecológica de los arrecifes coralinos
de los Cayos Arcas y Triángulos, Campeche, México:
impacto ambiental de una década de actividades de la
industria petrolera. Reporte al Gobierno Constitucional del
Estado de Campeche. 24 pp.
Folk, R. L. 1962. Sorting in some carbonate beaches of Mexico.
New York Academy of Sciences 25:222–44.
———. 1967. Sand cays of Alacran Reef, Yucatan, Mexico:
Morphology. Journal of Geology 75:412–437.
Folk, R. L., and A. S. Cotera. 1971. Carbonate sand cays of
Alacran Reef, Yucatan, Mexico: Sediments. Atoll Research
Bulletin no.137. 35 pp.
Folk, R. L., and R. Robles. 1964. Carbonate sands of Isla Perez,
Alacran Reef Complex, Yucatan. Journal of Geology
72(3):255–92.
Fosberg, F. R. 1961. Atoll news and comments. Atoll Research
Bulletin 84:6–9.
———. 1962. A brief study of the cays of Arrecife Alacran, a
Mexican atoll. Atoll Research Bulletin no. 93. 25 pp.
Freeland, G. L. 1971. Carbonate sediments in a terrigenous
province: the reefs of Veracruz, Mexico. Ph.D. diss., Rice
University, Houston, Tex. 253 pp.
Glynn, P. W. 1973. Aspects of the ecology of coral reefs in the
western Atlantic region. Pp. 271–324 in O. A. Jones and
R. Endean (eds.), Biology and Geology of Coral Reefs. New
York: Academic Press.
González-­Gándara, C., and J. E. Arias-­González. 2001a. Lista
actualizada de los peces del arrecife Alacranes, Yucatán,
Mexican Coral Reefs ~ 351
México. Anales del Instituto de Biología, Serie Zoología,
Universidad Nacional Autonóma de México 72(2):245–58.
González-­Gándara, C., and J. E. Arias-­González. 2001b.
Nuevos registros de peces en el Arrecife Alacrán, Yucatán,
México. Revista de Biología Tropical 49:770–71.
González-­Gándara, C., N. ­Membrillo-­Venegas, E. Nuñez-­Lara,
and J. E. Arias-­González. 1999. The relationship between
fish and reefscapes in the Alacranes Reef, Yucatan, Mexico:
a preliminary trophic functioning analysis. Vie et Milieu
49(4):275–86.
González-­Gándara, C., E. Pérez-­Díaz, L. ­Santos-­Rodriguez,
and J. E. Arias-­González. 2003. L
­ ength-­weight relationships
of coral reef fishes from the Alacranes Reef, Yucatan,
Mexico. Naga 26(1):14–16.
Green, G. 1977. Sinopsis taxonómica de trece especies de
esponjas del arrecife La Blanquilla, Veracruz, México.
Anales del Centro de Ciencias del Mar y Limnología,
Universidad Nacional Autónoma de México, Mexico City
4(1):79–98.
Gutiérrez, D., C. ­García-­Sáez, M. Lara, and C. Padilla. 1993.
Comparación de arrecifes coralinos: Veracruz y Quintana
Roo. Pp. 787–806 in S. I. ­Salazar-­Vallejo and N. E. González
(eds.), Biodiversidad Marina y Costera de México. México,
D.F.: Comisión Nacional para la Biodiversidad y Centro de
Investigaciones de Quintana Roo (CIQRO).
Heilprin, A. 1890. The corals and coral reefs of the western
waters of the Gulf of Mexico. Proceedings of the Academy of
Natural Sciences of Philadelphia 42:303–16.
Henkel, D. H. 1982. Echinoderms of Enmedio Reef, southwestern Gulf of Mexico. M.S. thesis, Corpus Christi State
University, Corpus Christi, Tex. 78 pp.
Hernández, L. R. 1997. Estructura de la comunidad faunística
asociada a la esponja Iricinia strobilina (Lamark, 1816)
Porifera: Demospongiae: Thoredtidae, del arrecife
Triángulo Oeste, Banco de Campeche, México. Tesis
profesional, Universidad Veracruzana, Xalapa, Veracruz.
29 pp.
Hernandez, R. C. 1987. Primer Informa del Estudio de Caracterizacion de los Sedimentos en el Sistema Arrecifal Veracruzano. México, D.F.: Secretaría de Marina, Dirección General
de Oceanografía. 21 pp.
Hidalgo, E., and E. A. Chávez. 1967. Prospección ecológica en
el arrecife de Lobos,Veracruz. Contribución del Instituto de
Investigaciones ­Biológico-­Pesqueras al III Congreso Nacional
de Oceanografía. Campeche, Campeche. 21 pp.
Hildebrand, H. H., H. Chávez, and H. Compton. 1964. Aporte
al conocimiento de los peces del Arrecife Alacranes,
Yucatán, México. Ciencia, México 23(3):107–34.
Horta-­Puga, G., and R. ­Ramírez-­Palacios. 1996. Niveles de
plomo en el esqueleto del coral arrecifal Montastrea
annularis. Pp. 823–830 in O. ­Rivero-­Serrano (ed.), La
Situación Ambiental en México. México, D.F.: Universidad
Nacional Autónoma de México.
Hoskin, C. M. 1962. Recent carbonate sedimentation on
Alacran Reef, Mexico. Ph.D. diss., University of Texas,
Austin, Tex. 254 pp.
———. 1963. Recent Carbonate Sedimentation on Alacran Reef,
Yucatan, Mexico. Publication No.1089. Washington, D.C.:
National Academy of Sciences–National Research Council.
160 pp.
———. 1966. Coral pinnacle sedimentation, Alacran Reef
Lagoon, Mexico. Journal of Sedimentary Petrology
36(4):1055–74.
———. 1968. Magnesium and strontium in mud fraction of
recent carbonate sediment, Alacran Reef, Mexico.
Bulletin of American Association of Petroleum Geologist
52:2170–177.
Huerta M., L. 1960. Lista preliminar de las algas marinas del
litoral del estado de Veracruz. Anales del Instituto de
Biología, Serie Botánica 25:39–45.
Huerta M., L., and A. G. Barrientos. 1965. Algas marinas de la
Barra de Tuxpan y de los arrecifes Blanquilla y Lobos.
Anales de la Escuela Nacional de Ciencias Biológicas, México
13(1–4):5–21.
Huerta M., L., M. L. Chávez, and M. E. Sánchez. 1977. Algas
marinas de la Isla de Enmedio, Veracruz. Pp. 314–25 in F. A.
Manrique (ed.), Memorias de V Congreso Nacional de
Oceanografía, Guaymas, Sonora, México, 22–25 de Octubre
1974.
Ives, J. E. 1890. Echinoderms from the northern coast of
Yucatan and the harbor of Veracruz. Proceedings of the
Academy of Natural Sciences of Philadelphia 42: 317–40.
———. 1891. Crustacea from the northern coast of the Yucatan,
the harbor of Veracruz, the west coast of Florida and the
Bermuda Islands. Proceedings of the Academy of Natural
Sciences of Philadelphia 1891:176–207.
Jordán-­Dahlgren, E. 2004. Los arrecifes coralinos del Golfo de
México: caracterización y diagnóstico. Pp. 555–71 in
M. Caso, I. Pisanty, and E. Ezcurra (eds.), Diagnóstico
Ambiental del Golfo de México. México, D.F.: Instituto
Nacional de Ecología.
Jordán-­Dahlgren, E., and R. E. ­Rodríguez-­Martínez. 2003. The
Atlantic coral reefs of Mexico. Pp. 131–158 in J. Cortés (ed.),
Latin American Coral Reefs. Amsterdam: Elsevier Science.
Joubin, M. L. 1912. Carte des bancs et récifs de Coraux
(Madrépores). Annales de L’Institut Océanographique 4 no.
2. 6 pp.
Kornicker, L. S., and D. W. Boyd. 1962. S­ hallow-­water geology
352 ~ Liddell and Tunnell
and environments of Alacran Reef Complex, Campeche
Bank, Mexico. Bulletin of the American Association of
Petroleum Geologists 46:640–73.
Kornicker L. S., R. Bonet, R. Cann, and C. M. Hoskin. 1959.
Alacran Reef, Campeche Bank, Mexico. Publications of the
Marine Science Institute, University of Texas 6:1–22.
Kühlmann, D. H. H. 1975. Charakterisierung der korallenriffe
von Veracruz, Mexico. Musseum für naturkunde on der
Humboldt Universitat zu Berlin. Internationale Revue der
gesamten Hydrobiologie 60(4):495–521.
Land, L. S. 1974. Growth rate of a West Indian (Jamaican) reef.
Proceedings of the Second International Coral Reef Symposium 2:409–12.
Lehman, R. L. 1993. Field and laboratory investigations of the
macroalgae of Enmedio coral reef, with specific taxonomic
reference to the genus Caulerpa. Ph.D. diss., Texas A&M
University, College Station, Tex. 161 pp.
Lehman, R. L., and J. W. Tunnell Jr. 1992a. A topographic
description of Enmedio Reef, Veracruz, Mexico. Paper
presented at: Gulf Estuarine Research Society Annual
Meeting; 2–4 April 1992; Port Aransas, Texas.
Lehman, R. L., and J. W. Tunnell Jr. 1992b. Species composition
and ecology of the macroalgae of Enmedio Reef, Veracruz,
Mexico. Texas Journal of Science 44:445–57.
Liceaga-­Correa, M. A., and J. I. Euan-­Avila. 2002. Assessment
of coral reef bathymetric mapping using visible Landsat
Thematic Mapper data. International Journal of Remote
Sensing 23(1):3–14.
Liddell, W. D. 2007. Chapter 3, Origin and geology. Pp. 23–33
in J.W. Tunnell Jr., E. A. Chávez, and K. Withers (eds.), Coral
Reefs of the Southern Gulf of Mexico. College Station, Tex:
Texas A&M University Press.
Liddell, W. D., and S. L. Ohlhorst. 1988. Comparison of western
Atlantic coral reef communities. Proceedings of the Sixth
International Coral Reef Symposium 3: 281–86.
Lidz, L., and B. Lidz. 1966. Foraminiferal biofaces of Veracruz
reefs. Bulletin of the American Association of Petroleum
Geologists 50(7):1514–517.
Logan, B. W. 1962. Submarine topography of the Yucatan
Platform. Pp. 101–104 in G. E. Murray and A. E. Weidie Jr.
(eds.), Guide Book Field Trip to Peninsula of Yucatan. New
Orleans, La.: New Orleans Geological Society.
———. 1969a. Part 1. Late Quaternary carbonate sediments of
Yucatan Shelf, Mexico. Pp. 1–128 in B. W. Logan, J. L.
Harding, W. M. Ahr, J. D. Williams, and R. S. Snead (eds.),
Carbonate Sediments and Reefs, Yucatan Shelf, Mexico.
Memoir 11. Tulsa, Okla.: American Association of
Petroleum Geologists.
———. 1969b. Part 2. Coral reefs and banks, Yucatan shelf,
Mexico (Yucatan Reef Unit). Pp. 129–198 in B. W. Logan,
J. L. Harding, W. M. Ahr, J. D. Williams, and R.S. Snead
(eds.), Carbonate sediments and reefs, Yucatan shelf, Mexico.
Memoir 11. Tulsa, Okla.: American Association of
Petroleum Geologists
Logan, B. W., J. L. Harding, W. M. Ahr, J. D. Williams, and R. S.
Snead (eds.). 1969. Carbonate Sediments and Reefs, Yucatan
Shelf, Mexico. Memoir 11. Tulsa, Okla.: American Association of Petroleum Geologists
Lubel, M. A. A. 1984. Algunos ciliados intersticiales de Isla
Enmedio, Veracruz, México. Anales Instituto de Biología,
Universidad Nacional Autónoma de México, Serie Zoología
55(1):1–59.
Macintyre, I. G., R. B. Burke, and R. Stuckenrath. 1977.
Thickest recorded Holocene reef section Isla Perez core
hold, Alacran Reef, Mexico. Geology 5:749–54.
Marion, A. F. 1884. Excursion aux isles Alacrans. Bulletin of
Societe Academique de Brest, (France, Serie 2) 9:5–21.
Martínez-­Guzmán, L. A, and J. L. ­Hernández-­Aguilera. 1993.
Crustáceos estomatópodos y decápodos del Arrecife
Alacrán, Yucatán. Pp. 609–629 in S. I. ­Salazar-­Vallejo and
N. E. Gonzalez (eds.), Biodiversidad Marina y Costera de
México. México, D.F.: Comisión Nacional para la Biodiversidad y Centro de Investigaciones de Quintana Roo
(CIQRO).
Márquez-­Espinoza, A. 1976. Peces de importancia económica
del Arrecife de Lobos, Veracruz, México. Pp. 133–40 in J. L.
­Castro-­Aguirre (ed.), Memorias de la Reunión Sobre
Recursos de la Pesca Costera, México.
Mateo-­Cid, L. E., A. C. ­Mendoza-­Gonzalez, and C. G. Garcia.
1996. Algas marinas de Isla Verde, Veracruz, Mexico. Acta
Botanica Mexicana 36:59–75.
Merino, M. 1997. Upwelling on the Yucatan Shelf: hydrographic
evidence. Journal of Marine Systems 13:101–21.
Millspaugh, C. F. 1916. Vegetation of Alacran Reef. Field
Museum of Natural History, Publication, Botanical Series
2(2):421–31.
Moore, D. R. 1958. Notes on Blanquilla Reef, the most
northerly coral formation in the Western Gulf of Mexico.
Publications of the Institute of Marine Science, University of
Texas 5:151–55.
Morales-­García, A. 1986. Estudio de los Crustáceos Estomatópodos y Decápodos de Isla Verde, Veracruz, México. México,
D.F.: Secretaría de Marina, Dirección General de Oceanografía. 78 pp.
———. 1987. Estudio de los Crustáceos Estomatópodos y
Decápodos de Isla Sacrificios, Veracruz, México. Report no.
87. Secretaría de Marina, Estación Veracruz, México.
Morelock, J., and K. J. Koenig. 1967. Terrigenous sedimentation
Mexican Coral Reefs ~ 353
in a shallow water coral reef environment. Journal of
Sedimentary Petrology 37:1001–1005.
Múzquiz, L. H. 1961. Flora marina de los alrededores de la isla
Pérez, Arrecife Alacranes, sonda de Campeche, México.
Anales de la Escuela Nacional de Ciencias Biológicas
(México) 10(14):11–22.
Nelson, T. J. 1991. A quantitative comparison of the community
structure of two forereefs in the southwestern Gulf of
Mexico. M.S. thesis, Corpus Christi State University, Corpus
Christi, Tex. 55 pp.
Novak, M. J. 1992. Sedimentology and Community Structure
of Reefs of the Yucatan Peninsula, Mexico. M.S. thesis, Utah
State University, Logan, Utah. 96 pp.
Novak, M. J., W. D. Liddell, and D. Torruco. 1992. Sedimentology and community structure of reefs of the Yucatan
Peninsula. Proceedings of the Seventh International Coral
Reef Symposium 1:265–72.
Rannefeld, J. W. 1972. The stony corals of Enmedio Reef off
Veracruz, Mexico. M.S. thesis, Texas A&M University,
College Station, Tex. 104 pp.
Ray, J. P. 1974. A study of the coral reef crustaceans (Decapoda
and Stomatopoda) of two Gulf of Mexico reef systems: West
Flower Garden, Texas and Isla de Lobos, Veracruz, Mexico.
Ph.D. diss., Texas A&M University, College Station, Tex.
323 pp.
Reséndez-­Medina, A. 1971. Peces colectados en el Arrecife La
Blanquilla, Veracruz, México. Anales del Instituto de
Biología (Universidad Nacional Autónoma de México), 42.
Serie Ciencias del Mar y Limnología 1:7–30.
Rice, W. H., and L. S. Kornicker. 1962. Mollusks of Alacran reef,
Campeche Bank, Mexico. Publications of the Institute of
Marine Science, University of Texas 8:366–403.
Rickner, J. A. 1975. Notes on members of the Family Porcellanidae (Crustacea: Anomura) collected on the east coast of
Mexico. Proceedings of the Biological Society of Washington
88(16):159–66.
———. 1977. Notes on a collection of crabs (Crustacea:
Brachyura) from the east coast of Mexico. Proceedings of the
Biological Society of Washington 90(4):831–38.
Rigby, J. K., and W. G. McIntire. 1966. The Isla de Lobos and
associated reefs, Veracruz, Mexico. Brigham Young
University Geology Studies 13:3–46.
Riley, C. M., and G. J. Holt. 1993. Gut contents of larval fishes
from light trap and plankton net collections at Enmedio
Reef near Veracruz, Mexico. Revista de Biologia Tropical
41:53–57.
Roberts, K. A. 1981. Polychaetes of Lobos Reef, Veracruz,
Mexico. M.S. thesis, Corpus Christi State University, Corpus
Christi, Tex. 40 pp.
Rodríguez, F. V., and V. A. Fuentes. 1965. Resultados preliminares sobre la distribución de plancton y datos hidrográficos
del arrecife La Blanquilla, Veracruz, Veracruz. Sección de
Hidrobiología del Instituto de Biología, Universidad Nacional
Autónoma de México 36:53–59.
Salas de Leon, D. A., M. A. M. Gomez, and J. A. Ramirez. 1992.
Periodos característicos en las oscilaciones de parámetros
meteorológicos en Cayo Arcas, México. Atmósfera 5:193–205.
Santiago, V. 1977. Algunos estudios sobre las madréporas del
Arrecife “La Blanquilla” Veracruz, México. Tesis profesional, Universidad Autónoma Metropolitana. 103 pp.
Smith, T. 1838. Description of Alacran and Cay Arenas, in the
Gulf of Mexico. Nautical Magazine 7:804–805.
Smith, W. F. G. 1954. Gulf of Mexico Madreporaria. Pp.
291–295 in P. S. Galtsoff (ed.), Gulf of Mexico, Its Origin,
Waters and Marine Life. Fishery Bulletin 89, Fishery
Bulletin of the U.S. Fish and Wildlife Service, Volume 55.
Washington, D.C.: U.S. Government Printing Office.
Solís, W. R. S. R. 1990. Estructura de la comunidad bentonica
de la parte sur del Arrecife Alacranes, Yucatán, México. M.S.
tesis, Recursos del Mar del Centro de Investigación y
Estudios Avanzados del IPN, Unidad Mérida. 39 pp.
Stinnett, T. L. 1989. Species composition and ecological
zonation of the sponge fauna of Enmedio Reef, Veracruz,
Mexico. M.S. thesis, Corpus Christi State University, Corpus
Christi, Tex. 23 pp.
Torruco, D., M. A. González, and W. D. Liddell. 1993. Integracion ecologica de grupos funcionales en la laguna arrecifal
de Alacranes, Yucatan, Mexico. Brenesia 39–40:37–49.
Tunnell, J. W. 1974. Ecological and geographical distribution of
Mollusca of Lobos and Enmedio coral reefs, southwestern
Gulf of Mexico. Ph.D. diss., Texas A&M University, College
Station, Tex. 158 pp.
———. 1977. Coral reef mollusks of the southwestern Gulf of
Mexico. Bulletin of the American Malacological Union 43:91.
———. 1988. Regional comparison of southwestern Gulf of
Mexico to Caribbean Sea coral reefs. Proceedings of the Sixth
International Coral Reef Symposium 3: 303–308.
———. 1992. Natural versus human impacts to Southern Gulf
of Mexico coral reef resources. Proceedings of the Seventh
International Coral Reef Symposium 1: 300–306.
———. 2007a. Chapter 1, Research history. Pp. 5–13 in J. W.
Tunnell Jr., E. A. Chávez, and K. Withers (eds.), Coral Reefs
of the Southern Gulf of Mexico. College Station, Tex.: Texas
A&M University Press.
———. 2007b. Chapter 2, Reef distribution. Pp. 14–22 in J. W.
Tunnell Jr., E. A. Chávez, and K. Withers (eds.), Coral Reefs
of the Southern Gulf of Mexico. College Station, Tex.: Texas
A&M University Press.
354 ~ Liddell and Tunnell
Tunnell, J. W., and B. R. Chapman. 1988. First record of
Red-­footed Boobies nesting in the Gulf of Mexico.
American Birds 42(3):380–81.
Tunnell, J. W., and B. R. Chapman. 2001. Seabirds of the
Campeche Bank islands, Southeastern Gulf of México. Atoll
Research Bulletin no.482. 50 pp.
Villalobos, A. 1971. Estudios ecológicos en un arrecife coralino
en Veracruz, México. pp. 531–545 in Symposium on
Investigations and Resources of the Caribbean Sea and
Adjacent Regions. Willemstad, Curaçao.
Villalobos-­Figueroa, A. 1980. Informe del programa de
arrecifes coralinos. Anales del Instituto de Biología. México,
D.F.: Universidad Nacional Autónoma de México. 26 pp.
Ward, H. L. 1887. Notes on the life-­history of Monachus
topicalis, the West Indian seal. American Naturalist
21:257–64.
White, R. B. 1982. A study of the natantid decapod crustaceans
collected from Enmedio Reef, Veracruz, Mexico, M.S.
thesis, Texas A&I University, Kingsville, Tex. 114 pp.