␥20 Mexican Coral Reefs

W. David Liddell and John W. Tunnell Jr.

The Mexican reefs of the southern Gulf of Mexico occur Late 1800s in 2 distinct provinces, the Campeche Bank to the north Professor Angelo Heilprin led the first scientific expedi- and west of the Yucatan Peninsula and the Veracruz shelf tion to the coral reefs of the southern Gulf of Mexico in reefs near the coast in the Tuxpan–Veracruz area of the 1890. This expedition was sponsored by the Academy of southwestern Gulf of Mexico. Reefs differ greatly in size, Natural Sciences of Philadelphia (now Philadelphia Acad- geomorphology, and diversity, reflecting differences in emy of Sciences), which Heilprin directed. The expedi- the underlying geologic structures and ambient envi- tion’s purpose was to “investigate the natural history of ronments. The reefs of the Campeche Bank, generally, the Yucatan Peninsula and Mexico” (Heilprin 1890). The are better developed than those of the Tuxpan–Veracruz expedition documented the tropical nature of marine area, perhaps due to the deleterious effects of high clas- biota of the southern Gulf and described corals and coral tic sedimentation and cooler temperatures in the latter reefs (Heilprin 1890) and many other marine inverte- area. brates (Ives 1890, 1891; Baker 1891). Heilprin (1890) also described reefs and islands occurring off the city of Vera- History of Research cruz and noted the “vast quantity of coral” used in con- struction there. The reefs of the southeastern Gulf of Mexico (Fig. 20.1) have a long prehistory, dating back to the Mesozoic 1900s–1970s opening of the Gulf of Mexico (Liddell 2007). Modern historical accounts began with the English adventurer With the exception of Joubin’s 1912 map of coral reefs, William Dampier, who mentioned the prolific nesting little research was conducted on southern Gulf reefs until seabirds occurring on the islands of Alacrán and Trián- Smith’s (1954) work. Smith utilized Heilprin’s (1890) and gulos (Dampier 1699). A century would pass before oth- Joubin’s (1912) works and various unpublished sources ers again noted the vegetation, seabirds, sea turtles, and and nautical charts (refer to Tunnell 2007a, his Table 1.1) West Indian monk seal associated with these and other to create a coral reef distribution map. Smith also com- Campeche Bank islands (Smith 1838; Marion 1884; Ward piled a list of coral species from the southern Gulf. In 1887; Agassiz 1888). 1955, Moore (1958) briefly visited Blanquilla Reef off Tux- pan. 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 During the 1960s, interest and research on all south- offshore of the city of Veracruz. His work was published ern Gulf of Mexico coral reefs increased greatly (Tables in both English and Spanish, raising the level of interest in 20.1, 20.2). Because the reef localities were remote, field this unique (terrigenous) setting for coral reefs. Kornicker studies tended to be performed during extended expedi- et al. (1959) took part in a large, collaborative effort to tions to the reef systems. Huerta M. and Barrientos (1965) study Alacrán, the most northerly reef on the Campeche reported on the algae of Blanquilla and Isla de Lobos reefs, Bank (Fig. 20.1). Although the expedition largely focused near Tuxpan. Rigby and McIntire (1966) described the on geology, several biological studies also resulted from geology and ecology of Isla de Lobos Reef, while Cham- it, making Alacrán Reef one of the best-­known Gulf reefs berlain (1966) reported on gorgonians from Lobos that (Table 20.1). were studied during the same Brigham Young Univer- From 1959 to 1963, the Department of Oceanography sity expedition. Hidalgo (Hidalgo and Chávez 1967) and at Texas A&M University undertook a major, multiyear Chávez (Chávez et al. 1970; Chávez 1973; Bautista-­ Gil­ and study of the Campeche Bank or Yucatan Shelf in the south- Chávez 1977) conducted multiple expeditions to Isla de eastern Gulf. The project was funded by the National Sci- Lobos Reef with their students from the Instituto Politéc- ence Foundation, the American Petroleum Institute, the nico Nacional in Mexico City. Chávez (1966) reported on Office of Naval Research, Shell Development Company, the fishes of Triángulos and Cayo Arenas reefs. and Mobil Oil Company. The project was the longest and Students from the Universidad Nacional Autónoma de most productive study of southern Gulf reefs to that date Mexico conducted their “professional theses” on a vari- and contributed greatly to the knowledge of coral reefs ety of topics within the Veracruz Reef system. Villalobos and reef and shelf sediments of the region (Logan 1962, summarized much of this work in the proceedings of an 1969a, b; Logan et al. 1969). In addition, Cayo Arenas was international symposium on the Caribbean Sea and adja- studied in more detail as a side project (Busby 1966). cent 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 References Alacrán bathymetric mapping Liceaga-­Correa and Euan-­Avila 2002 communities benthic communities Solís 1990 reef communities Liddell and Ohlhorst 1988 geology Kornicker et al. 1959; Kornicker and Boyd 1962; Hoskin 1962, 1963, 1966, 1968; Bonet 1967; Macintyre et al. 1977 organisms algae Múzquiz 1961 birds Fosberg 1961, 1962; Folk 1967; Boswall 1978; Tunnell and Chapman 1988, 2001 fish Hildebrand et al. 1964; ­González-­Gandara et al. 1999; González-­ ­Gandara and Arias-González­ 2001a, 2001b; Brulé et al. 2003; González-­ ­Gandara et al. 2003 foraminiferans Davis 1964 mollusks Rice and Kornicker 1962; Aranda et al. 2003 stomatopods and decapods Martínez-­Guzmán and ­Hernández-­Aguilera 1993 vegetation Marion 1884; Millspaugh 1916; Fosberg 1961, 1962; Bonet and Rzedowski 1962; Folk 1967 reef growth model Bosscher and Schlager 1992 sediments Folk 1962, Folk and Robles 1964; Folk 1967; Folk and Cotera 1971; Novak 1992; Novak et al. 1992 spatial distribution Torruco et al. 1993 Cayo Arenas geology Busby 1966 Triangulos and Cayo Arenas organisms coral Carricart-­Ganivet and ­Beltrán-­Torres 1993 fish Chávez 1966 sponges Hernández 1997 environmental Ferre-­D’Amare 1995 Cayos Arcas organisms coral Farrell et al. 1983 environmental Ferre-­D’Amare 1995 meteorology Salas de Leon et al. 1992

(Villalobos-­Figueroa 1980). Other studies on southern late 1980s and 1990s, Universidad Veracruzana in Jalapa, Veracruz reefs from this time included algae (Huerta M. and Centro de Investigaciones y Estudios Avanzados 1960) and foraminiferans (Lidz and Lidz 1966). More- (CINVESTAV)–Unidad Mérida of the Instituto Politéc- lock and Koenig (1967), Edwards (1969), and Freeland nico Nacional, also contributed to research on Veracruz (1971) conducted geologic studies during the 1960s on reefs and Alacrán Reef, respectively. the southern Veracruz reefs. The reefs of northern Veracruz State are the least stud- ied of those in the southwestern Gulf region due to their 1970s–1990s remoteness. Of the 6 northernmost reefs, Isla de Lobos has received the most attention (Table 20.2). From 1970 to 1990, three groups were largely responsible Over 20 coral reefs exist in the extreme southwestern for research on the southern Gulf of Mexico coral reefs: Gulf, located offshore of the city of Veracruz and offshore (1) Secretaría de Marina, Dirección General de Ocean- of the fishing village of Antón Lizardo. Since these reefs ografía, (2) Texas A&M University-­ Corpus­ Christi, and are more accessible than the northern Veracruz reefs, they (3) Universidad Nacional Autónoma de México. In the 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 Subject References Tuxpan Reef system Blanquilla algae Moore 1958; Huerta M. and Barrientos 1965 Isla de Lobos algae Huerta M. and Barrientos 1965; Rigby and McIntire 1966; Hidalgo and Chávez 1967 crustaceans Ray 1974 crabs Allen 1982 fish Márquez-­Espinosa 1976 foraminiferans Bautista-­Gil and Chávez 1977 gorgonians Chamberlain 1966 mollusks Tunnell 1974, 1977 polychaetes Roberts 1981 reef communities Chávez et al. 1970; Chávez 1973 Veracruz Reef system Veracruz Area algae Huerta M. 1960 geology Emery 1963; Morelock and Koenig 1967; Freeland 1971; Hernandez 1987 foraminiferans Lidz and Lidz 1966 benthic communities Kühlmann 1975 La Blanquilla coral ecology Santiago 1977 ecology Villalobos 1971 fish Reséndez-­Medina 1971 hydrography Rodríguez and Fuentes 1965 sponges Green 1977 Verde algae Mateo-­Cid et al. 1996 coral taxonomy Beltrán-­Torres and ­Carricart-­Ganivet 1993 lead pollution Horta-­Puga and ­Ramírez-­Palacios 1996 stomatopods & decapods Morales-­García 1986 Sacrificios stomatopods & decapods Morales-­García 1987 Enmedio algae Huerta M. et al. 1977; Lehman and Tunnell 1992a, 1992b; Lehman 1993 ciliates Lubel 1984 corals Rannefeld 1972; Nelson 1991 crabs Rickner 1975, 1977; Allen 1982 echinoderms Henkel 1982 fish Choucair 1992; Riley and Holt 1993 IXTOC I oil impacts Tunnell and Dokken unpublished; Baca et al. unpublished mollusks Tunnell 1974, 1977 shrimps White 1982 sponges Stinnett 1989

haps the most studied of all reefs in this region is Enme- y Mar Caribe). This program first operated from a small dio Reef. Gutiérrez et al. (1993) compared the coral reefs station (Estación Oceanográfica) in the city of Veracruz, of the Veracruz region to those within the Sian Ka’an but it is now housed within a new institute building at Biosphere Reserve located along the Caribbean coast of Antón ­Lizardo. Quintana Roo, Mexico. The relatively great distance from shore continued to The Mexican Navy has contributed oceanographic limit the number of reef studies on the Campeche Bank. information, as well as information on reef biology, for Macintyre et al. (1977) published a report on the thick- the Veracruz reef system through its oceanographic pro- est recorded Holocene reef section at Alacrán Reef. Dur- gram (Instituto de Investigación Oceanográfica del Golfo ing the 1980s, coral diversity and zonation were charac- Mexican Coral Reefs ~ 345 terized for Cayos Arcas (Farrell et al. 1983) and Alacrán terrigenous sediment shed from the Sierra Madre Orien- (Liddell and Ohlhorst 1988) reefs. Additionally, several tál and Trans-­Mexican Neovolcanic Belt greatly influences authors reviewed all Mexican Atlantic coral reefs, includ- the nearshore reefs. Also, during the rainy season lowered ing environmental problems and human impacts (Tables salinities may occur in some areas (e.g., practical salin- 20.1, 20.2). ity of 18.3 at Veracruz compared to 36 for normal oceanic During the 1990s an increasing number of both organ- waters) (Ferre-­D’Amare 1985; Tunnell 1988). The ­Tuxpan– ismal and ecological studies continued on the Campeche Veracruz area is subject to low winter temperatures with Bank and Veracruz Reef system. Other studies focused on lows ranging from 8–10 °C (air) at Veracruz down to, reef sediments and a computer simulation model of reef infrequently, 0 °C (air) at Tampico (Ferre-D’Amare­ 1985). growth (Tables 20.1, 20.2). Reefs are absent to the north of the Tuxpan Reef system, most likely due to low winter water temperatures. Reefs are 2000s to the Present also absent to the south of the Veracruz Reef system, per- haps due to upwelling (Ferre-D’Amare­ 1985). A number of fish studies were conducted during the early In the southeastern Gulf, reefs are located far (130– 2000s, especially on Alacrán Reef (Tables 20.1, 20.2). 200 km) offshore on a wide carbonate shelf, the Campeche Other studies included Montastraea annularis growth Bank, primarily along the 55-­m depth contour (Logan rates, other invertebrates, and seabirds on Campeche 1969a, b). In contrast to the southwestern Gulf, southeast- Bank reef islands. In recent studies, geologic facies were ern Gulf reefs occur in a semiarid climate, surrounded by charted through time on Campeche Bank reefs (Tables oceanic water from the Caribbean and the Gulf of Mexico, 20.1, 20.2). and they are not affected by mainland drainage. The gen- Jordán-Dahlgren­ and Rodríguez-­ Martínez­ (2003) eral absence of reefs along the Gulf coastal areas of the summarized work on all Mexican coral reefs within the Yucatan Peninsula may be due to low winter tempera- Gulf of Mexico and Caribbean. ­Jordán-­Dahlgren (2004) tures (8–10 °C air temperatures; Ferre-D’Amare­ 1985), provided a sobering, though brief, environmental sta- the effects of which would be more pronounced along tus report on southern Gulf reefs. Reefs in the extreme the coastline than farther out on the shelf where the bank southwestern Gulf, located off the city of Veracruz reefs occur. Freshwater discharge is also possible through and the village of Antón Lizardo, have been negatively the Yucatan karst system into coastal areas. Fringing or affected the most, and those on the Campeche Bank have ­barrier-type­ reefs are conspicuously absent from the been affected the least. The reefs near Tuxpan in north- northern and western (Gulf) coastal areas of the Yucatan ern Veracruz State are intermediate in terms of negative Peninsula, although these reef types are extensively devel- effects (Tunnell 1992). oped along the eastern (Caribbean) side of the Yucatan Peninsula and southward near Belize. In contrast to adjacent, low-energy­ coastal areas such Reef Distribution and Structure as the Florida Keys and Belize where mangroves line the shoreline and seagrasses predominate as nearshore sub- There are 46 named coral reefs in the southern Gulf of marine vegetation, shoreline areas of the mainland consist Mexico: 31 in the southwestern Gulf off the state of Vera- of ­moderate-­energy sandy beaches or rocky shores (vol- cruz (Veracruz Shelf reefs) and 15 in the southeastern canic) in the southwestern Gulf and low-­energy sandy Gulf on the Campeche Bank (refer to Tunnell 2007b, his beaches or rocky shores (limestone) in the southeastern Table 2.1) (Fig. 20.1). In addition, Dahlgren (1993) listed Gulf. Also, unlike the scattered, ­shallow-­water patch reefs approximately 10 named and over 25 unnamed banks on associated with the former, the southern Gulf reefs are the Campeche Bank for which there is little or no scien- largely submerged, ­mountain-­like structures scattered tific information. across the continental shelf. Coral reefs in the southwestern Gulf are typically located at nearshore (<200 m) to mid-shelf­ (22 km) dis- Southwestern Gulf Reefs tances on a narrow, terrigenous shelf (Morelock and Koe- nig 1967; Ferre-D’Amare­ 1985). The Veracruz region The southwestern Gulf coral reefs are clustered in 2 sys- is characterized by subhumid to humid climates of high tems, the Tuxpan Reef system to the north (Fig. 20.2) and rainfall and substantial mainland drainage. Consequently, the Veracruz Reef system to the south (Fig. 20.3). Most of 346 ~ Liddell and Tunnell

Reef system have islands associated with them (La Blan- quilla, 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 light- house 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 Vera- cruz Reef system consists of 12 larger reefs occurring rela- tively farther offshore than those of the northern group. These include emergent platform-­ 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 Liz- ardo. 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 (San- tiaguillo, Cabezo, Isla de Enmedio, and Chopas). Chopas Figure 20.2. The Tuxpan Reef system (TRS) including 6 emer- Reef actually has 2 islands, Isla Salmedina and Isla Blanca; gent platform reefs, 3 off Cabo Rojo and 3 northeast of Tuxpan, both of these islands have only low natural vegetation. Veracruz, Mexico (reprinted from Tunnell 2007b, with permis- Isla de Enmedio has low, natural vegetation, as well as a sion of Texas A&M Press. dense canopy of transplanted shade trees. El Aguila (on Cabezo) and Santiaguillo islands have little vegetation these reefs are relatively small (only a few square kilome- and are composed almost totally of coral rubble with little ters), exhibit low relief, and are located near shore (Ferre-­ or no sand. Topatillo Reef used to have a coral rubble cay D’Amare 1985). There are 6 emergent platform reefs in consisting of Acropora cervicornis (staghorn coral) rubble, the Tuxpan Reef system (Fig. 20.2). Isla de Lobos Reef is but it no longer exists. After the death of A. cervicornis the only one in the Tuxpan Reef system that has an island and A. palmata (elkhorn coral) in the late 1970s and early and, thus, has served as a base camp for several expedi- 1980s (Tunnell 1992), this small island began to erode tions (e.g., Rigby and McIntire 1966; Chávez et al. 1970; because of the lack of living Acropora, which protected the Tunnell 1974). At 21°30′14″N, Blanquilla Reef is the most island from wave action. northerly emergent coral reef in the western Gulf of Mex- ico (Moore 1958). Southeastern Gulf Reefs In the Veracruz Reef system in the far southwestern Gulf of Mexico, 25 coral reefs occur in 2 subgroups, one In the southeastern Gulf of Mexico reefs occur in a located off the city of Veracruz and one off the village of broad arc running from the ­north-­central portion of the Antón Lizardo (Fig. 20.3). The first, or northern, group Campeche Bank (Alacrán Reef) to the southwest (Cayos consists of smaller reefs located more onshore than those Arcas). The reefs occur near the shelf edge, some 100– of the other grouping. This northern group of 13 reefs 200 km from the coastline of the Yucatan Peninsula (Fig. is located off the city of Veracruz and includes 8 emer- 20.1). The absence of reefs on northeastern portion of the gent platform reefs (Gallequilla, Anegada de Adentro, La shelf may be due to upwelling of cold (17–18 °C) water Blanquilla, La Gallega, Pájaros, Isla Verde, Tierra Nueva, (Logan 1969b; Glynn 1973; Merino 1997). and Isla de Sacrificios), 2 submerged bank or patch reefs Eleven named emergent reefs and 4 named submerged (Bajo Mersey and Lavandera), and 3 fringing reefs (Punta reefs are located along the outer continental shelf of the Gorda–Punta Majahua, Hornos, and Punta Mocambo). Campeche Bank. The Campeche Bank reefs have been Three of the reef platforms in the northern Veracruz 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, Vera- cruz, Mexico (reprinted from Tunnell 2007b, with permission of Texas A&M Press.

with the principal subdivision between submerged reef water depths up to 22 m. Alacrán Reef is the most studied banks (e.g., Bancos Ingleses, Banco Pera, Bajo Nuevo, Campeche Bank reef and has been the subject of numer- and Bajos Obispos) and emergent reef banks or “walls” ous geological and biological investigations (Table 20.1). (see Tunnell 2007b, his Table 2.1) (Fig. 20.1). The latter Liddell and Ohlhorst (1988), as part of a CINVESTAV are further subdivided into (1) solitary reef knolls (Nuevo and Armada de Mexico expedition, quantitatively char- and Triángulo Oeste), (2) linear reef walls (Triángulo acterized community structure and diversity of a 20-m­ Este-­Sur), (3) crescent reef walls (e.g., windward reef at windward reef site. They found that the bottom consisted Alacrán Reef and parts of Cayo Arenas and Cayos Arcas), of 76% total living cover comprised of macroalgae (45%), and (4) reef complexes with multiple emergent reef walls. corals (11%), non-coral­ cnidarians (chiefly gorgonians, Cayo Arenas, Triángulos, and Cayos Arcas are all consid- 5.4%), coralline algae (5.2%), and sponges (5.2%). Corals ered complexes having multiple emergent platforms and consisted of 12 species with a diversity of 1.86 (H′, natu- islands. Nuevo Reef has a single emergent platform and ral log). Torruco et al. (1993) characterized the structure ephemeral island. The majority of these reefs form arcs of benthic communities inhabiting the lagoon at Alacrán. that are convex to the northeast, reflecting the prevalent Alacrán Reef has 5 vegetated islands: Desterrada, ­northeast-­southwest wave progression (Logan 1969b). Desertora, Perez, Chica, and Pájaros. Arenas has one Alacrán Reef is the most northerly reef in the entire large, named vegetated island, Cayo Arenas, and 3 smaller southern Gulf of Mexico. It is also, by far, the largest reef in unvegetated and unnamed coral rubble cays. Triángu- the southern Gulf, occupying some 13 × 25 km. It has an los has at least 2 islands, Triángulo Oeste and Triángulo extensive ­shallow-­water lagoon with small patch reefs and Este. Finally, Cayos Arcas has 3 vegetated islands: Cayo 348 ~ Liddell and Tunnell del Centro, Cayo del Oeste, and Cayo del Este. Manned and perpendicular to the prevailing wave-­approach direc- lighthouses are found at Isla Perez, Cayo Arenas, Trián- tion. They are strongly influenced by storm waves and are gulo Oeste, and Cayo del Centro. largely barren of sandy sediment on their windward sides, In addition to the “true” reefs mentioned above, bio- where well-developed­ boulder ramparts occur. Again, the stromes lacking scleractinian frameworks also occur on reefs are thought to have been established on anteced- the Campeche Bank, particularly in the 18–60 m depth ent Pleistocene dunes. The authors concluded that much range (Logan 1969b). These hard banks form relatively of the terrigenous sand and gravel present on the shelf is thin veneers over previously existing topography. These late Pleistocene (Wisconsinan) in age. ­Present-­day terrig- veneers consist of 1–18–cm diameter, concentrically lay- enous sedimentation rates are very low, thus allowing the ered nodules of crustose coralline algae and foraminifera. reefs to develop. Even so, carbonate sediments are found These nodular crusts are most common on the northern to accumulate only in the immediate area of the coral shelf and are much less common on the western shelf, per- reefs and rarely exceed 50% calcium carbonate by weight haps due to increased terrigenous sedimentation there. (Morelock and Koenig 1967).

Southeastern Gulf Reefs Reef Sediment Composition Logan (1969a) sampled sediments from both reef and Numerous studies have examined the composition of open shelf settings on the Campeche Bank, noting that Holocene reef sediment from the southern Gulf of Mex- sediments from reefs and associated environments (e.g., ico, particularly Alacrán Reef. Studies of certain areas, lagoons) were dominated by coral, coralline and calcare- such as Cayos Arcas and Triángulos on the Campeche ous algae, and foraminifera. In contrast, mollusks, echi- Bank and the Tuxpan-­ Tampico­ portion of the western noids, and bryozoans were much more important sedi- Gulf coastal plain, are lacking. Although both geographic ment constituents on the open shelf. (e.g., increased terrigenous sediment in the southwestern Alacrán Reef is the most studied of the Campeche Gulf) and bathymetric variation in sediment composition Bank reefs. Kornicker and Boyd (1962) provided a gen- occur, generalizations may be made. eral overview of the sedimentology and biotic zonation of Alacrán Reef and lagoon. They noted that considerable Southwestern Gulf Reefs lateral variation occurred in the organisms present and the bottom character around the reef complex. Relatively little is known about the geology of the Tuxpan Folk and Robles (1964) examined the sedimentology reefs. The seafloor north and south of the Isla de Lobos of Isla Perez, the largest leeward island on Alacrán Reef. Reef at Tuxpan (Fig. 20.2) is dominated by reef-­derived The general composition of the sand-sized­ fraction was calcareous sand; whereas, the seafloor to the west of the 60% Halimeda, 25% coral, and 15% foraminifera and reef has both reef-derived­ and terrigenous material (Rigby other grains, whereas the coarser fractions were domi- and McIntire 1966) consisting of quartz, ferromagnesium nated by coral (e.g., coral stick ramparts generated by minerals, volcanic rock fragments, and obsidian, presum- storms). Folk (1967) and Folk and Cotera (1971) focused ably transported from the interior by the Río Tuxpan or on the origin, geomorphology, and size sorting occurring Río Panuco. The seafloor immediately east of the reef is on sand cays off the leeward side of Alacrán Reef, noting dominated by rocky reef debris. The linear trend of Isla de that these were areas of intense biotic sediment produc- Lobos, Medio, and Blanquilla reefs might reflect the con- tion. trol of relict Pleistocene or older topography (Rigby and A core taken from Isla Perez by Macintyre et al. (1977) McIntire 1966). documented 30 m of Holocene reef growth (principally by The Veracruz reefs are somewhat more studied from the coral Acropora cervicornis). This translates into accu- a geologic standpoint than those to the north. Emery mulation rates of 12 m / 1000 yr, considerably higher than (1963), Edwards (1969), and Freeland (1971) examined the rates previously described for Jamaica by Land (1974). the mixed terrigenous-­ carbonate­ sediments occurring The Holocene reef that was cored was situated on top of a off the city of Veracruz (Fig. 20.3). Morelock and Koenig ­similar-­appearing pre-Holocene­ reef that extended from (1967) studied the reefs to the south of Veracruz at Antón 33 m down to 50–60 m and the shelf platform (Bonet Lizardo. These reefs are located parallel to the shoreline 1967). Mexican Coral Reefs ~ 349

Hoskin (1963) conducted a detailed facies analysis of (e.g., Alacrán). The overall development of reefs in the Alacrán Reef. Seventeen environments were character- southern Gulf of Mexico may be limited by winter tem- ized, based on sediment composition and size, includ- peratures. In addition, reefs along the Tuxpan–Veracruz ing (from east to west) windward shelf, lower windward coast may be limited by the abundant supply of terrige- slope, upper windward slope, surge channels, boulder nous sediment being shed from the Sierra Madre Orien- rampart, windward reef flat, moat, Thalassia beds, cellu- tál and Trans-­Mexican Neovolcanic Belt and possibly by lar reef surface, cellular reef deep, pinnacle reef, lagoon reduced salinities in the rainy season. Unfortunately, most proper, patch reefs, shallow sand, leeward reef surface, of these reefs increasingly are being affected by economic lower leeward slope, and leeward shelf. Halimeda, coral, development (e.g., Farrell et al. 1983; Ferre-­D’Amare and fecal pellets were the dominant components of the 1985; Tunnell 1992; Chávez and Tunnell 1993) (Tables sand-­sized sediment fraction. Halimeda was most abun- 20.1, 20.2), particularly those in the southwestern Gulf off dant on the windward and leeward reefs; coral reached its Veracruz (city) and Antón Lizardo. 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 Acknowledgments aggregate grains varied little across these environments. Hoskin (1966) examined eodiagenesis (near-surface­ We dedicate this chapter to the memory of our good diagenesis) occurring on pinnacle reefs in the Alacrán friend and colleague, Mauricio Garduno Andrade, who Reef lagoon. Diagenetic processes observed included pel- participated on many of our expeditions to the reefs of letization of lagoon floor muds, macroboring of coral by Mexico and whose research contributed to turtle con- lithophagid bivalves, microboring of coral and mollusk servation biology. We thank the Texas A&M University fragments by endolithic algae, aragonite cementation in Press for permission to use figures 20.1–20.3. Portions of cavities, and recrystallization of coral grains. this article were originally published in a different form in Novak (1992) and Novak et al. (1992) provided a quan- Liddell (2007) and Tunnell (2007a, 2007b). titative description of Alacrán Reef sediment from fore-­ reef and back-­reef settings. For all samples, coral, the calcareous green alga, Halimeda, and coralline algae con- References stituted well over 70% of all grains by volume. The dis- tribution of grains was similar to the distribution of the Agassiz, A. 1888. Three Cruises of the United States Coast and living, ­sediment-­producing organisms, with coral grains Geodetic Survey Steamer “Blake,” in the Gulf of Mexico, in the being relatively more abundant in fore-reef­ samples and Caribbean Sea, and Along the Atlantic Coast of the United Halimeda grains in back-­reef samples. In contrast to the States, from 1877 to 1880, vol.1. Boston and New York: findings of Hoskin (1963), fecal pellets were relatively Houghton, Mifflin and Company. 314 pp. minor sedimentary constituents of their samples and Allen, R. L. 1982. The reptant decapods of Enmedio and Lobos coralline algae were more important (Novak et al. 1992). coral reefs, Southwestern Gulf of Mexico. M.S. thesis, The back-­reef samples studied by Novak et al. (1992) Corpus Christi State University, Corpus Christi, Tex. 65 pp. were from the shallower, eastern portion of the lagoon, Aranda, D. A., E. B. Cárdenas, I. M. Morales, A. Z. Zárate, and not the deeper, pellet-­ rich­ lagoon areas and pinnacle reefs T. Brulé. 2003. A review of the reproductive patterns of sampled by Hoskin (1963). Cluster analysis delineated gastropod mollusks from Mexico. Bulletin of Marine Science 3 Alacrán Reef lithofacies: back reef, shallow fore reef 73:629–41. (<10 m), and deep fore reef (>10m) (Novak et al. 1992). 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. Conclusions Bautista-­Gil, M. L., and E. A. Chávez. 1977. Contribución al conocimiento de los foraminíferos del arrecife de Lobos, The reefs of the southern Gulf of Mexico have a long his- Veracruz. Pp. 90–103 in F. Manrique (ed.), Memorias del V tory of study. They exhibit a diversity of forms, including Congreso Nacional de Oceanografía. Guaymas, Sonora, small, nearshore shelf reefs (e.g., Veracruz Shelf Reefs), México, 22–25 de Octubre 1974. fringing reefs (Veracruz), and large, open-­shelf atolls Beltrán-­Torres, A., and J. P. ­Carricart-­Ganivet. 1993. Skeletal 350 ~ Liddell and Tunnell

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