Facies (2009) 55:2735 DOI 10.

1007/s10347-008-0162-4

O R I G I N A L A R T I CL E

Modern benthic foraminifera on Banco Chinchorro, Quintana Roo, Mexico

Eberhard Gischler Andrea Mder

Received: 12 June 2008 / Accepted: 22 September 2008 / Published online: 7 October 2008 Springer-Verlag 2008

Abstract Benthic foraminiferal assemblages may be used as environmental indicators on Banco Chinchorro, an isolated carbonate platform oV the eastern coast of the Yucatan Peninsula. Seventy-seven species from 44 genera were identiWed in 14 surface sediment samples, which were collected along an EW windward-to-leeward transect across the platform. A total of 15,493 foraminiferal tests (max. 1,200 tests per sample) were investigated and served as the basis of this study. As many taxa range throughout several platform zones, assemblages are better environmental indicators as compared to individual species. Four foraminiferal assemblages were identiWed using statistical methods including (1) a Homotrema assemblage, which occurs at the windward platform margin, (2) an Archaias-Homotrema assemblage which is found on the leeward margin and on platform interior coral patch reefs, (3) a Quinqueloculininae-Archaias-Rosalina assemblage of the western platform, and (4) an Archaias-Quinqueloculininae assemblage characteristic of the eastern platform interior. Environmental factors which inXuence variation in foraminiferal distribution and diversity on Banco Chinchorro include exposure to waves and currents and substrate (plant and algal growth). Sediment transport does not play a major role in Banco Chinchorro based on the observation that there are only limited amounts of taxa found outside their typical habitats,

and, that mean grain-size and sorting of foraminiferal tests do not exhibit clear patterns. Keywords Holocene Reef Mexico Chinchorro Foraminifera

Introduction Benthic foraminifera have been used as facies indicators in both modern and ancient carbonate depositional environments (e.g., Hallock and Glenn 1986; Murray 1991; Beavington-Penney and Racey 2004). Studies of modern benthic foraminifera in reefal and shallow-water carbonate platform environments in the Caribbean realm are numerous. Bandy (1964) studied foraminiferal distribution in the Gulf of Batabano, Cuba. Cebulski (1969) and Wantland (1975) detailed the distribution of benthic foraminifera on the Belize shelf. Brooks (1973) investigated foraminifera along the southern shore of Puerto Rico, and Rose and Lidz (1977) and Lidz and Rose (1989) in south Florida. Brasier (1975) described living foraminifera from the reefs around Barbuda. The distribution of living benthic foraminifera around the island of Tobago was investigated by Radford (1976a, b), and the importance of ecological factors such as salinity, food supply, oxygen, pH, sediment type, substrate stability, and water depth was evaluated. Hallock (1979) discussed how test shapes of individual benthic foraminiferal taxa may vary depending on water depth. Martin and Liddell (1988, 1989) investigated foraminifera in reefs on the northern Jamaica island slope and related distribution patterns to bathymetry and reef morphology. They distinguished between typical fore reef and back reef taxa. Culver (1990) studied benthic foraminifera in lagoons and mangrove areas of Puerto Rico. Exposure, substrate, and

Electronic supplementary material The online version of this article (doi:10.1007/s10347-008-0162-4) contains supplementary material, which is available to authorized users. E. Gischler (&) A. Mder Institut fr Geowissenschaften, Johann Wolfgang Goethe-Universitt, Altenhferallee 1, 60438 Frankfurt am Main, Germany e-mail: gischler@em.uni-frankfurt.de

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vegetation were identiWed as important controlling factors of distribution. Murray (1991) discussed general aspects of modern benthic foraminiferal ecology, typical assemblages, and ranges of environmental parameters such as water depth, salinity, temperature, and substrate. Lee and Anderson (1991) and Lee and Hallock (2000) presented results on biological studies on modern foraminifera including their ecology. Despite these numerous studies, the number of studies of benthic foraminifera on isolated carbonate platforms of the Caribbean is limited. Streeter (1963) investigated the distribution of foraminifera on Great Bahama Bank. Davis (1964) gave a brief report of benthic foraminifera distribution on Alacran Reef, a small atoll-like platform on Campeche Bank. Likewise, investigations on the use of foraminifera as tracers of sediment movement in Atlantic reef and platform systems are rather rare. Illing (1950) and Moore (1957) discussed foraminiferal transport based on observations regarding test occurring outside their common habitats in fore reef versus lagoonal environments in the Bahamas and south Florida, respectively. Recently, Li et al. (1997, 1998) interpreted patterns of textural parameters of foraminiferal tests such as grain-size and sorting as proxies of sediment movement among the reefs of Grand Cayman Island. Foraminiferal distribution and redeposition on the isolated platforms of Belize to the south of the study area were studied by Gischler et al. (2003). Here, we present the results of a project, in which we investigated the distribution of benthic foraminifera and their use as tracers of sediment movement in Banco Chinchorro, a remote isolated carbonate platform oV the eastern coast of the Yucatan Peninsula. Chinchorro is well suited for such a study because it includes a variety of diVerent reef and platform environments. A comprehensive volume on the fauna of Banco Chinchorro was published not too long ago (Suarez-Morales and Camarena-Luhrs 2003) including studies on corals, nematodes, arthropods, tunicates, chaetognaths, Wsh, gastropods, and birds, however, foraminifera were not investigated.

Fig. 1 Map of Banco Chinchorro showing environmental zones and locations of surface sediment samples analyzed during this study. Small inset map in lower left corner shows location of Chinchorro in the wider Caribbean realm (modiWed from Gischler and Lomando 1999)

Setting Banco Chinchorro is an isolated carbonate platform, which covers an area of 550 km2 (Fig. 1). The platform is surrounded by deep water. Three islands on the platform make up 1% of the total platform area. Banco Chinchorro is 46 km long and 19 km wide at the longest and widest points, respectively. Lagoon depths increase from 2 m in the shallow north to 12 m in the deeper south (Jordan and Martin 1987; Gischler and Lomando 1999). The seagrass Thalassia testudinum and codiacean algae Halimeda and Penicillus are common. In the northern and central parts of

the platform, three NNE-trending, elongated depressions up to 7 m deep can be found, which are separated by shallow lagoon (12 m deep) areas. This conWguration is presumably an expression of the underlying tectonic grain (Gischler and Lomando 1999). Surface-breaking reef margins are well developed and give way to a fore reef slope and wall on the seaward side. On the northern platform margin, a wide sand apron is developed. Patch reefs of coral are common in all parts of the platform. In the southern part, patch reefs are coalescing and form NNE and NW striking trends. Gischler and Lomando (1999) distinguished three types of surface sediments including coralgal grainstones on marginal and lagoonal reefs, mollusk-foram-Halimeda packstones and grainstones in the eastern part of the platform, and mixed peloidal-skeletal wackestones, packstones, and grainstones in the western interior lagoon area. No subsur-

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face studies on the geological development of Banco Chinchorro have been made as yet. Climate of the study area is subtropical with water temperatures ranging from 26.729.2C (De Jesus-Navarrete 2003). Salinity in the platform interior is elevated as compared to the open sea and ranges from 36 to 37.5 (De Jesus-Navarrete 2003). Annual precipitation reaches 1,400 mm on the adjacent Yucatan mainland. The rainy season lasts from June to October. Winds blow from the E and NE for most of the year. During winter and fall, cold fronts with winds from the NW and W may occur. Banco Chinchorro lies in the hurricane belt. Hurricane season is from July to October.

Methods Foraminifera in 14 surface sediment samples from the study of Gischler and Lomando (1999) were analyzed during the present study. Sediment samples were collected along an EW traverse across Banco Chinchorro by free diving in December 1996. Average abundance of foraminifera in the size-classes >0.125 mm in the 14 samples is 11% as determined by point counting of thin sections (Gischler and Lomando 1999). After washing in a dilute solution of Clorox, the samples were sieved into the sizeclasses >2, 21, 10.5, 0.50.25, 0.250.125, and <0.125 mm. We attempted to pick 300 foraminiferal tests from each of the four fractions 21, 10.5, 0.50.25, and 0.250.125 mm (
10, 01, 12, 23), i.e., a total of 1,200 tests from each sediment sample (sieve method or standard method of Martin and Liddell 1988, 1989). In a

number of samples, however, the size classes yielded less than 300 foraminifera per size class (see Electronic supplementary material). This was particularly the case for the larger size fractions. In the smaller size-classes, more than 300 tests were frequently present, necessitating the use of a sample splitter in order to obtain smaller, statistically similar sub-samples. According to Patterson and Fishbein (1989) the statistical error depends on the abundance of a taxon and ranges from approximately 16% for 300 individuals and from about 0.42% for 1,200 specimens. In the present study, a total of 15,493 foraminiferal tests were picked with small brushes and glued using gelatine into Plummer-type slides with 60 numbered Welds (Table 1). Individuals were identiWed using the standard reference of Loeblich and Tappan (1988a, 1988b) and other literature illustrating shallow-water Caribbean foraminifera (e.g., Streeter 1963; Brooks 1973; Sen Gupta and Schafer 1973; Rose and Lidz 1977; Lidz and Rose 1989) and foraminifera from the Belize shelf (Cebulski 1969; Wantland 1975). We did not diVerentiate between living and dead individuals as the samples were not stained after collection. Consequently, the foraminiferal distribution patterns represent the potential preservable rock record of foraminifera, not zonations of living taxa. The conspicuous red encrusting Homotrema rubrum was also included into the analysis as each Homotrema fragment would contribute to the potential rock record. Qmode cluster analyses were used to measure similarity between samples and foraminiferal taxa associations. Mean grain-size and sorting of tests of the common species Archaias angulatus were calculated using the formulas of Folk and Ward (1957).

Table 1 Depth, locations, and texture including abundance and diversity of foraminifera in surface sediment samples of Banco Chinchorro

Sample

Water depth (m) 10.0 2.2 3.5 3.5 4.0 2.5 2.5 7.5 6.0 3.8 5.0 4.0 3.5 2.0

Environment

Forams picked 1,017 1,112 1,002 996 997 1,200 1,200 1,044 949 1,200 1,176 1,200 1,200 1,200 1,107

Abundance (%) 1.6 6.3 1.7 5.9 0 53 19.5 2.9 4.6 31.0 32.0 6.3 5.8 32.0 4.3 11.0

No. of taxa 39 43 55 49 26 48 53 49 51 56 51 56 54 29 49

Mud (%) 4.7 1.8 7.2 8.0 4 4.9 4.2 33.3 54.8 4.4 6.8 12.6 20.1 0

15 16 19 20 21 22 23 24 25 26 27 28 29 30 Average

LM LM SL DL DL SL SL DL DL SL SL SL DL WM

Abundance based on point counting in thin sections (Gischler and Lomando 1999). Mud is grain-size fraction <125 m LM Leeward margin, WM Windward margin, SL Shallow lagoon, DL Deep lagoon

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Results We identiWed 77 species from 44 genera, which are almost exclusively benthic foraminifera. A few planktonic species were found as well (Globigerinoides trilobus, Orbulina universa). The most common taxa include several families of the miliolids, rotaliids, and textulariids (Electronic supplementary material). Average species richness is 49 taxa per sample, and mean abundance based on thin-section counts amounts to 11%. Species richness and foraminiferal abundance appear to be slightly higher on the windward platform as compared to the leeward platform side. Species richness is clearly lower at marginal stations as compared to platform interior sample stations (Table 1). There are no signiWcant trends or correlations detectable though when comparing occurrence, species richness, foraminiferal abundance, water depth, or sediment texture. The distribution of taxa exhibits certain patterns (Fig. 2). Archaias angulatus, species of Peneroplis, and the Quinqueloculininae (Quinqueloculina, Triloculina, Pyrgo) are abundant throughout the sample stations except for station 30 on the windward reef. Their abundances are also somewhat lower at the patch reef station 23. At reefal stations 30, 23, and 15, Homotrema rubrum is very abundant. Amphistegina gibbosa and Asterigerina carinata are common at platform mar-

gins and on the eastern interior platform. Species of Textularia and Planorbulina reach highest abundances in the platform interior, especially at the patch reef location. Discorbis is most abundant at the patch reef location and towards platform margins. Valvulina oviedoiana and Miliolinella species are also common in the platform interior and reach high abundances at stations 25 and 26. Articulina species occur in the platform interior but are lacking at the windward and leeward sample stations. Amphisorus hemprichi and Sorites marginalis are found throughout the sample transect and reach highest abundance at stations 22 and 29. Species of Rosalina also occur at all stations and are most common on the western platform side. Species of Elphidium and Cribroelphidium reach highest abundances in the deeper platform stations in the platform centre. A Q-mode cluster analysis of the data set allows for a separation of four dendrogram entities (Fig. 3). The unweighted pair-group average algorithm with Euclidean distances produced the most meaningful results. The dendrogram entities, which deWne foraminiferal assemblages, are quite stable as seen in the fact that an analysis with all taxa identiWed produced the same results as an analysis with only the 14 most abundant taxa, which reach percentages of >5% (Fig. 4). Foraminiferal assemblages include (1) a Homotrema assemblage, which characterizes the

Fig. 2 Distribution of most common benthic foraminifera along sample transect across Banco Chinchorro. Note that sample distances are not to scale; for location of samples see Fig. 1

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Fig. 3 Tree diagram of a Q-mode cluster analysis deWning foraminiferal assemblages. Algorithm used: unweighted pair-group average; distant coeYcient: Euclidean. n = 80

windward platform margin, (2) an Archaias-Homotrema assemblage that is found on the leeward margin and at the platform interior patch reef, (3) the QuinqueloculininaeArchaias-Rosalina assemblage of the western platform, and (4) the Archaias-Quinqueloculininae assemblage characteristic of the eastern platform interior (Fig. 5). For some of the most common foraminifers such as Archaias angulatus, Homotrema rubrum, Peneroplis, and the Quinqueloculininae we calculated sorting and mean grain-size along the sample transect (Fig. 6). Mean grainsizes and sorting in most taxa do not show any clear patterns. Only in Archaias angulatus, sorting of tests exhibits a general trend of better sorting towards the leeward margin of the platform.

Discussion The distribution of benthic foraminifera exhibits patterns that are comparable with Wndings of studies elsewhere in the Caribbean-western Atlantic realm. Archaias angulatus is the most common species in Banco Chinchorro, as it is in several other Caribbean reef and platform environments. It lives in symbiosis with the green alga Chlamydomonas hedleyi and usually attached to the seagrass T. testudinum (Martin 1986; Hallock et al. 1986; Lidz and Rose 1989). Archaias is also known to be quite tolerant with regard to water temperature (1433C) and salinity (2939) in south Florida (Hallock and Peebles 1993). It can usually be found in shallow and low-energy depositional environments (Brasier 1975; Martin and Liddell 1988, 1989; Hal-

lock et al. 1986; Hallock and Peebles 1993; Li and Jones 1997; Lee and Anderson 1991; Gischler et al. 2003). Because of their robustness, Archaias tests have a high preservation potential (Martin 1986; Martin and Wright 1988). Species of Peneroplis have comparable ecological demands. They also live in symbiosis with a green alga and live on seagrass or coarser sediment (Lidz and Rose 1989; Hallock and Peebles 1993). Peneroplis is common in shallow wind- and current-protected environments (Culver 1990), however, higher energy habitats (stations 28, 29) are colonized as well. Bandy (1964) also observed Peneroplis in higher energy environments in the Gulf of Batabano. The abundant Quinqueloculina and Triloculina also prefer lower energy platform environments and live preferentially on seagrass (Lidz and Rose 1989). In the nearby Belize platforms, Triloculina and Quinqueloculina were also very abundant in platform interiors (Gischler et al. 2003). Likewise, Rosalina is common in the Chinchorro platform interior. In south Florida and the Bahamas, Rosalina species are most common in patch reef areas (Rose and Lidz 1977). Species of Rosalina and Planorbulina are adapted to somewhat higher energy because they are able to attach themselves to seagrass with an organic glue (Langer 1993). Even so, Planorbulina is preferentially found in the more protected western platform interior of Chinchorro. Discorbis, which is common at patch reef and marginal areas of Chinchorro also prefers higher energy settings in Caribbean locations such as the Gulf of Batabano, Cuba, Grand Cayman, and TurneVe Islands, Belize (Bandy 1964; Li et al. 1997; Gischler et al. 2003). Like in the shallow Belize platform of Lighthouse Reef, Valvulina oviedoiana is

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Fig. 4 Pie diagrams of taxonomic composition of four foraminiferal assemblages

Fig. 5 Bathymetric cross-section over Banco Chinchorro including sample stations and spatial occurrence of foraminiferal assemblages

common in the platform interior of Chinchorro. In south Florida, it occurs both in back reef areas and within Florida Bay (Rose and Lidz 1977; Hallock and Peebles 1993). The elevated abundances of Amphisorus hemprichii and S. marginalis at sample stations 22 and 29 is somewhat enigmatic.

These species have dinoXagellate symbionts and are reported to live in lower energy settings on plants, algae, or mollusk shells (Lidz and Rose 1989; Langer and Hottinger 2000). Like in Chinchorro, species of Elphidium and Cribroelphidium, which are infaunal and lack photosymbionts,

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Fig. 6 Mean grain-size and sorting of Archaias angulatus along the sample transect. Y-axis indicates Phi values

are common in quiet and deeper water settings of other Caribbean reefs and platforms (Bandy 1964; Cebulski 1969; Wantland 1975; Gischler et al. 2003). Homotrema rubrum is an indicator of very high energy and shallow water settings both in Chinchorro and in other platform such as in Belize (Gischler and Ginsburg 1996; Gischler et al. 2003), or the Bahamas (Streeter 1963). Homotrema preferentially dwells in cryptic habitats and is able to attach itself to the hard substrate by an organic glue. Species of Amphistegina gibbosa and Asterigerina carinata are characteristic of fore reef slope environments (Brasier 1975; Martin and Liddell 1988; Li et al. 1998; Gischler et al. 2003). In Chinchorro, these taxa are most common at platform margins. Most of the tests in these areas and in the platform interior are potentially transported. Because most taxa occur over several platform zones, foraminiferal assemblages are more useful as environmental indicators. The Archaias-Quinqueloculininae and the Quinqueloculininae-Archaias-Rosalina assemblages of the Chinchorro platform interior are comparable to Archaiasand Quinqueloculina-dominated associations in the wider Caribbean and the Gulf of Mexico (Murray 1991, p. 96). Archaias associations are deWned by a rather narrow salinity range of 3437 and a high temperature range of 9 38C. Water depths are <12 m and the substrate is seagrass and carbonate sands and muds. Quinqueloculina associations have wide salinity (1047) and temperature ranges (1538C) and also occur in depths <12 m and seagrass and carbonate sand and mud substrates (Murray 1991). Gischler et al. (2003) described very similar Archaias-Quinqueloculina-Triloculina and Quinqueloculina-Triloculina assemblages from the platform interiors of Glovers, Lighthouse, and TurneVe (Belize). In Belize, the Archaias-dominated assemblage is more common in lower energy settings, whereas the Quinqueloculina-dominated assemblage appears to characterize areas of patch reefs and carbonate shoals. In Chinchorro, the environmental/bathymetrical relation between the two assemblages is not as clear. The

Archaias-dominated assemblage is found on the eastern platform side, whereas the Quinqueloculininae-dominated assemblages characterize the western platform interior. The Homotrema assemblage of Banco Chinchorro is also found occurring at the windward margins of the Belize platforms to the south (Gischler et al. 2003). Environmental factors that inXuence the variability of foraminiferal occurrence, abundance, and diversity on Banco Chinchorro largely include exposure to waves and currents and plant and algal growth on the bottom. The relatively clear spatial separation of the four assemblages supports the contention that parameters such as, e.g., submarine relief, substrate character (texture and plant/algal growth), and depositional energy are of importance. High degrees of exposure to waves and currents including sediment redeposition as well as the lower density of seagrass and algal growth presumably cause the lower diversities at platform margins as compared to the interior. Water depth does not appear to be of great importance because the two platform interior assemblages include shallower and deeper lagoon areas. Also, no statistically signiWcant correlations exist between bathymetry and foraminiferal abundance or diversity. As we did not systematically measure water temperature and salinity, it is diYcult to estimate the inXuence of these parameters on foraminiferal occurrence. Water temperature and salinity are probably both slightly higher and prone to larger Xuctuation in the platform interior as compared to the margins. Even so, despite potential elevated temperature and salinity, foraminiferal diversity is higher in the Chinchorro platform interior as compared to the margins. The fact that the sorting of tests of Archaias angulatus improves from windward to leeward and the occurrence of marginal taxa such as Homotrema rubrum, Asterigerina carinata, and Amphistegina gibbosa in the eastern platform interior support the contention that sediment is transported to some degree from east to west. This is presumably a consequence of the prevailing easterly and northeasterly winds and currents. According to Li et al. (1998), normal conditions are not suYcient in order to transport sediment from the fore reef to the back reef and lagoonal areas of Grand Cayman. Storms and cyclones are needed for this kind of sediment redeposition, according to these authors. The lack on any clear textural patterns in other taxa and the relatively clear patterns in foraminiferal distribution and occurrence of assemblages suggest, however, that sediment transport does not play a major role in Banco Chinchorro. Similar results were obtained in the Belize platforms Glovers, Lighthouse, and TurneVe to the south of the study area (Gischler et al. 2003). Occurrence and distribution of benthic foraminiferal assemblages apparently have high potential as environmental indicators, which in turn is of great importance for studies in the fossil record.

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Facies (2009) 55:2735 atoll-like coral reef system in the Mexican Caribbean Sea. Bull Mar Sci 73:514 Folk RL, Ward W (1957) Brazos river bar: a study in the signiWcance of grain size parameters. J Sediment Petrol 27:326 Gischler E, Ginsburg RN (1996) Cavity dwellers (coelobites) under coral rubble in southern Belize barrier and atoll reefs. Bull Mar Sci 58:570589 Gischler E, Lomando AJ (1999) Recent sedimentary facies of isolated carbonate platforms, Belize, Yucatan system, Central America. J Sediment Res 69:747763 Gischler E, Hauser I, Heinrich K, Scheitel U (2003) Characterization of depositional environments in isolated carbonate platforms based on benthic foraminifera, Belize, Central America. Palaios 18:236255. doi:10.1669/0883-1351 Hallock P (1979) Trends in test shape with depth in large symbiontbearing foraminifera. J Foraminiferal Res 9:6169 Hallock P, Glenn EC (1986) Larger foraminifera: a tool for paleoenvironmental analysis of Cenozoic carbonate depositional facies. Palaios 1:5564. doi:10.2307/3514459 Hallock P, Peebles MW (1993) Foraminifera with chlorophyte endosymbionts: habitats of six species in the Florida Keys. Mar Micropal 20:292299 Hallock P, Cottey TL, Forward LB, Halas J (1986) Population biology and sediment production of Archaias angulatus (Foraminiferida) in Largo Sound, Florida. J Foraminiferal Res 16:18 Illing MA (1950) Mechanical distribution of recent foraminifera in Bahamas sediments. Ann Mag Nat Hist Ser 2(3):757761 Jordan E, Martin E (1987) Chinchorro: morphology and composition of a Caribbean atoll. Atoll Res Bull 310:133 Langer MR (1993) Epiphytic foraminifera. Mar Micropal 20:235265. doi:10.1016/0377-8398(93)90035-V Langer MR, Hottinger L (2000) Biography of selected larger foraminifera. In: Lee, JJ, Hallock P (eds) Advances in the biology of foraminifera micropaleontology 46(Suppl 1):105126 Lee JJ, Anderson OR (eds) (1991) Biology of Foraminifera. Academic Press, London, 368 pp Lee JJ, Hallock P (eds) (2000) Advances in the biology of Foraminifera. Micropaleontology 46(Suppl 1):1198 Li C, Jones B (1997) Comparison of foraminiferal assemblages in sediments on the windward and leeward shelves of Grand Cayman, British West Indies. Palaios 12:1226. doi:10.2307/3515291 Li C, Jones B, Blanchon P (1997) Lagoon-shelf sediment exchange by storms: evidence from foraminiferal assemblages, east coast of Grand Cayman, British West Indies. J Sediment Res 67:1725 Li C, Jones B, KalbXeisch WB (1998) Carbonate sediment transport pathways based on foraminifera: case study from Frank Sound, Grand Cayman, British West Indies. Sedimentology 45:109120. doi:10.1046/j.1365-3091.1998.00133.x Lidz BH, Rose PR (1989) Diagnostic foraminiferal assemblages of Florida Bay and adjacent shallow waters: a comparison. Bull Mar Sci 44:399418 Loeblich AR Jr, Tappan H (1988a) Foraminiferal genera and their classiWcation. Reinhold, New York, 970 pp Loeblich AR Jr, Tappan H (1988b) Foraminiferal genera and their classiWcationplates. Reinhold, New York, 212 pp Martin RE (1986) Habitat and distribution of the foraminifer Archaias angulatus (Fichtel and Moll) (Miliolina, Soritidae), northern Florida Keys. J Foraminiferal Res 16:201206 Martin RE, Liddell WD (1988) Foraminiferal biofacies on a north coast fringing reef (175 m), Discovery Bay, Jamaica. Palaios 3:298314. doi:10.2307/3514659 Martin RE, Liddell WD (1989) Relation of counting methods to taphonomic gradients and information content of foraminiferal sediment assemblages. Mar Micropal 15:6789. doi:10.1016/03778398(89)90005-4

Conclusions Four benthic foraminiferal associations were identiWed on Banco Chinchorro. The Homotrema assemblage characterizes the windward platform margin, (2) the Archaias-Homotrema assemblage is found at the leeward margin and platform interior patch reefs, (3) the Quinqueloculininae-Archaias-Rosalina assemblage is characteristic of the western platform, and (4) the ArchaiasQuinqueloculininae assemblage of the eastern platform interior. Environmental factors that may be used to explain the variation in foraminiferal diversity and distribution on Banco Chinchorro include exposure to waves and currents and plant/algal growth on the substrate. Mean grain-size and sorting of foraminiferal tests, and the occurrence of some taxa outside their typical habitats in Chinchorro suggest sediment transport to some degree at platform margins. Overall, sediment redeposition does not appear to be an important factor that potentially distorts foraminiferal distribution.
Acknowledgements We are grateful to Chevron Overseas Petroleum, especially T. Lomando, who graciously Wnanced the expedition to Banco Chinchorro. H. Legarre assisted during sediment sampling. G. Garcia and J. Perez helped with the diYcult task of exporting the samples. I. Hauser, K. Heinrich, U. Scheitel, and S. Schultz gave helpful advice during the study. W. Schiller assisted during SEM sessions. We thank the editor A. Freiwald for his thoughtful comments that improved this paper.

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