Marine Micropaleontology 82–83 (2012) 38–45

Contents lists available at SciVerse ScienceDirect

Marine Micropaleontology

j ourna l homepage: www.e lsev ie r .com/ locate /marmic ro

SHE Analysis for Biozone Identification among foraminiferal sediment assemblageson reefs and in associated sediment around St. Kitts, Eastern Caribbean Sea, and itsenvironmental significance

Brent Wilson a,⁎, Kate Orchard b, Jason Phillip c

a Petroleum Geoscience Programme, Department of Chemical Engineering, University of the West Indies, St. Augustine, Trinidad and Tobagob The St. Christopher National Trust., The National Museum, P. O. Box 888, Bay Road, Basseterre, Saint Kitts and Nevisc Dive Caribbean, P.O. Box 2247, Basseterre, Saint Kitts and Nevis

⁎ Corresponding author. Tel.: +1 868 662 2002x3676E-mail address: brent.wilson@sta.uwi.edu (B. Wilson

0377-8398/$ – see front matter © 2011 Elsevier B.V. Alldoi:10.1016/j.marmicro.2011.10.001

a b s t r a c t

a r t i c l e i n f o

Article history:Received 21 January 2011Received in revised form 14 October 2011Accepted 19 October 2011Available online 25 October 2011

Keywords:SHEBIHurricanesAmphistegina gibbosaAsterigerina carinataDiscorbis roseaArchaias angulatus

Sixteen nearshore (≤1 m water depth) sediment samples, termed Sample Set 1, were collected from theeastern and south-western coasts of the South East Peninsula (SEP) of St. Kitts. Twelve samples termed Sam-ple Set 2 were taken from nearby fringing (6–17 m) and offshore (~18–25 m) reefs. The sample sets wereanalysed separately using SHE Analysis for Biozone Identification (SHEBI), α and β diversities and comple-mentarity. SHEBI is a statistical technique that determines abundance biozones (ABs) using the entire vectorof species abundances in samples, species richness S, the information function H, and the equitability index E.Alpha diversity expresses the diversity within an AB, while β diversity quantifies diversity changes betweenadjacent ABs. Complementarity assesses the permeability of AB boundaries.SHEBI divided Sample Set 1 into five ABs but did not distinguish separate bays. Beta diversities reflected thehighest and lowest α diversities at Mosquito Bay and South Friar's Beach\Frigate Bay respectively. Comple-mentarity was highest between two ABs in South Friar's Bay, indicating a close relationship between shore-line sediment and material marginally farther offshore. It was lowest between Whitehouse and South Friar'sBays. The assemblages along this transect were rich in Discorbis rosea.For Sample Set 2, samples were accumulated for SHEBI by depth. SHEBI distinguished four ABs (three on thefringing reefs) characterised by differing percentages of Asterigerina carinata, Archaias angulatus and Amphis-tegina gibbosa. Comparison between Sample Sets 1 and 2 indicates that during storms sediment moves shore-ward from the fringing reefs. Foraminifera show that there is less movement of sediment>63 μm from theshoreline (Sample Set 1) to the fringing and deeper reefs.

© 2011 Elsevier B.V. All rights reserved.

1. Introduction

Foraminifera have a moderate to high post-mortem preservationpotential (Martin and Liddell, 1991) and abound in all marine envi-ronments. They are particularly diverse in tropical reefal environ-ments (Murray, 2006) such as those in the Caribbean Sea, which isbordered to the north and east by islands that are encircled by coralreefs (Fig. 1). The nearshore benthonic foraminifera around theislands of the Caribbean were first studied by d'Orbigny (1839) andhave since been reported in >100 papers (Culver and Buzas, 1982).However, the majority of these studies were qualitative, reportingspecies presence or absence or giving only brief comments regardingabundance (e.g., Brasier, 1975a, 1975b; Radford, 1976a, 1976b, 1998).Such information is of limited use in environmental work.

.).

rights reserved.

Martin and Liddell (1988) quantitatively examined the foraminif-era along two transects across reefs in Discovery Bay, Jamaica, fromthe back-reef (1–4 m) to the deep fore-reef (75 m) zones. Theyfound that fore-reef zones were poorly delineated in Q-mode clusteranalyses of species abundances obtained from traditional countingprocedures (300 foraminifera from a size fraction>0.125 mm), butwere able to discern a very detailed zonation using alternative pick-ing procedures. However, Gischler et al. (2003) used this alternativepicking procedure and found cluster analysis to distinguish four ben-thonic foraminifera associations on three reefs off Belize, CentralAmerica, sampled depths ranging from 0 m (beach) to 40 m (fore-reef). Fore-reef samples contained abundant Amphistegina gibbosaand Asterigerina carinata, while high-energy, marginal-reef areaswere dominated by encrusting Homotrema rubrum. Platform-interiors contained abundant Miliolina (especially Archaias angulatus,Quinqueloculina sp., and Triloculina sp). Elphidium poeyanum was in-dicative of low-energy or deep-lagoonal regimes. Sediment transportwas discerned using abundant occurrences of tests outside their orig-inal habitats. Gischler et al. (2003) concluded, however, that

Fig. 1. The Caribbean Sea and St. Kitts.

a

b

c

Fig. 2. a. The South East Peninsula (SEP) of St. Kitts showing the locations of bays sam-pled and sediment colours. b. The SEP showing the locations of samples (prefixed “SK:”in text) from the shoreline Sample Set 1 and the prevailing current direction. c. St. Kittsshowing the locations of reefs and of samples (prefixed “SK:” in text) from the reefalSample Set 2.

39B. Wilson et al. / Marine Micropaleontology 82–83 (2012) 38–45

sediment redeposition did not significantly alter characteristic taxo-nomic compositions.

Li et al. (1997) used foraminifera to trace sediment transport bystorms around Grand Cayman, in the western Caribbean. They con-cluded that A. angulatus dominates in the inner lagoon, A. gibbosaand A. carinata on the fore-reef terraces and Discorbis rosea on theupper reef terrace. These they termed “tracer species” and postulatedthat they can be used to detect sediment movement due to storms. Liet al. (1997), however, excluded juvenile A. gibbosa and A. angula-tusb0.345 mm from their analyses because they were poorly pre-served (although presumably still recognisable).

Buzas andHayek (1996) introduced a technique called SHE analysisthat resolves biodiversity into three components: species richness S,diversity as measured using the information function H (=−Σpi·lnpi,where pi is the proportional abundance of the ith species), and even-ness measured using the equitability index E (=eH/S). Decompositionof the equitability index shows that lnE=H− lnS. From this Buzas andHayek (1998) developed SHE Analysis for Biozone Identification(SHEBI), in which lnS, H and lnE are recalculated as samples are accu-mulated and the number of specimens N increases. SHEBI delimitsabundance biozones (ABs), an AB being an area in which the abun-dance of a particular species or group of species differs significantlyfrom that in adjacent ABs. An AB differs from a biofacies in being de-fined using the entire assemblage, rather than merely dominant spe-cies (Buzas, written communication, 2008). An AB boundary is drawnwhere there is a break in slope on a graph of lnE against lnN. SHEBI isthus not subject to any distortions that may arise from the exclusionof juveniles as practised by Li et al. (1997). It has two advantagesover cluster analysis: it is incapable of grouping samples separated bysamples belonging to another AB and it allows ready and rapid assess-ments of changes in diversity across AB boundaries.

In this paper, SHEBI is for the first time applied to foraminiferal as-semblages in samples from reefs and backreef areas, using samplescollected around the South East Peninsula (SEP) of St. Kitts, West In-dies (Fig. 2). It employs sample sets that differ from transects in notbeing linear. Comparisons of assemblages in the different ABs areused to detect sediment transport during storms.

2. The study area

St. Kitts consists of an amalgamated series of Plio-Pleistocene vol-canic cones on the St. Eustatius Bank of the inner arm of the LesserAntilles double island arc (Martin-Kaye, 1969). This study concen-trates especially on the waters around the SEP, which is relativelyundeveloped compared to the remainder of the island and has onlybeen readily accessible since the opening of a highway in 1990, at

40 B. Wilson et al. / Marine Micropaleontology 82–83 (2012) 38–45

which time a beach bar was constructed at Mosquito Bay. Theremaining distal Cockleshell, Major's and Whitehouse Bays wereundeveloped at the time of sampling, although concern was alreadybeing expressed about the possible deleterious effects of land clear-ance and sedimentation on reef health (Smith et al., 1997). Mora(2008, fig. 1) has more recently detected negative impacts on fishes,corals and macroalgae throughout the Caribbean Sea (including St.Kitts) from agricultural land use, coastal development, overfishingand climate change, although it is not clear how severe these wereat the time of sampling. At that time the more proximal South Friar'sBay and Frigate Beach were popular with tourists.

The Caribbean Sea around St. Kitts is microtidal (Kjerfve, 1981),oligotrophic (Mendoza et al., 2009) and supports a rich growth ofcorals and coralline algae. Consequently, shoreline sediment aroundthe island consists of silici-carbonates that range from pale yellowishbrown (Munsell 10YR 6/2) to greyish orange pink (5YR 7/2; Fig. 2a).Adey and Burke (1976) suggested that three reef tracts occur alongthe windward (in the case of St. Kitts, the NE) coasts of the Lesser An-tilles: fringing (or bench), bank barrier and shelf edge. However,available maps show only fringing reefs (e.g., CCA, 1990). One of us(JP) identified, and collected sediment samples from, two reef tractsoff the SW coast of the island: a fringing reef tract close in shore(depths of 6–17 m) and a tract farther offshore (depths~25 m) thatWilkinson (2004) termed “offshore bank reefs”; the latter may beanalogous to the mid shelf reefs around the Cayman Islands recordedby Li et al. (1997) (Fig. 2). The platform around the SEP supports lushseagrass meadows (CCA, 1990) with a rich epiphytal foraminiferalfauna (Wilson, 1998). Currents in the microtidal area flow from SEto NW.

3. Materials and methods

Sixteen samples, numbered SK:I through SK:X (Fig. 2b), were col-lected from nearshore sediment (water depth≤1 m) between thefringing reefs and the SEP shoreline by scraping a sample jar overthe sediment surface to recover the top 1 cm (samples SK:A−SK:H,collected from the SEP north coast, are not considered here). Mostwere taken in 2006, but sample SK:L* (distinguished with an asterisk)was collected from Mosquito Bay in 1993. Sample SK:R was takenfrom the shoreline at the SE end of South Friar's Beach, while theremaining samples from there were taken from deeper (~1 m)water. Twelve sediment samples (SK:1 to SK: 12) were taken by adiver in 2010 from the fringing and offshore bank reefs south of St.Kitts and from a shipwreck (the River Tawe) (Fig. 2c). Sample SK:11was recovered a few metres from a hot spring off Camps Village.Most samples were taken adjacent to quadrats used to estimate per-centage bare sand versus percentage live coral\sponge cover andcoral\sponge health. The corals were generally in fair to excellenthealth but covered b10% of the quadrat.

Nearshore sediment being typically devoid of live foraminifera(Brasier, 1975b), the sampleswere not stained to distinguish live forami-nifera. However, a few specimens of A. carinata and A. angulatus from thecoral reefs were greenish brown from enclosed algal symbionts, and sowere alive at the time of collection (Pamela Hallock, written communi-cation, October 2010). The assemblages are thus regarded as comprisingtotal (live+dead) faunas.

The samples were washed over a 63 μm mesh to remove silt andclay, and then dried over a gentle heat. As implied by Hofker(1976), the proportion of sediment comprising foraminifera variesspatially around Caribbean islands. Approximately equal volumes ofsediment from each sample were picked clean of benthonic forami-nifera such that the number of specimens recovered varied from sam-ple to sample but was generally >250. Species were identified usingCushman (1921, 1922, 1923, 1929, 1930, 1931), Brooks (1973),Hofker (1956, 1964, 1971, 1976, 1980), Todd and Low (1971) andSchnitker (1971), and the number of specimens per species counted.

Point (sample) diversities were measured using species richness Sand the information function H. Point dominance was determinedusing the equitability function E.

The data set was examined using SHEBI (Buzas and Hayek, 1998,2005; Hayek and Buzas, 2010), which Wilson (2008b) has shownyields different results according to the direction taken along thetransect. Previous SHEBIs of foraminiferal associations have usuallyproceeded from shallow water to deep (Buzas et al., 2007a, 2007b).However, as samples SK:I−SK:X, were all collected from ≤1 m andform a transect along shore, it was not possible to include all samples(i.e., the nearshore SK:I−SK:X and the reefal SK:1−SK:12) in a sin-gle SHEBI. Instead, two sets of samples were analysed separatelyand then compared. The nearshore samples SK:I−SK:X, here termedSample Set 1, were for SHEBI accumulated along the direction of theprevailing surface current. The reefal samples SK:1−SK:12, whichare here termed Sample Set 2 despite being geographically scatteredalong the island's SW coast, were accumulated in order of increasingdepth. SHEBI was conducted using the freeware PAST programme ofHammer and Harper (2005).

It is challenging when using SHEBI to decide how significant abreak in slope must be to warrant drawing an AB boundary (Buzasand Hayek, 1998). An AB boundarywas for this study drawnwhereverthere was:

1. an increase in lnE between two samples. This is the easiest ABboundary to recognise and this type of break can indicate an ABcontaining one sample only.

2. a break in slope after which the new trend in lnE versus lnN wasmaintained for several samples. This can distinguish an AB withas few as two samples.

Whittaker (1972) distinguished two categories of diversity: inven-tory diversity, which calculates the diversity of associations withinsamples (point diversity), habitats (α diversity) or the combined eco-systems within a region (γ diversity); and differentiation diversity,which examines the change in diversity between samples (pattern di-versity) or between habitats (β diversity). Ecologists usually expressinventory diversity in terms of S only, as when Hunter (2002) termedγ diversity “geographic-scale species diversity”. This, however, is mis-leading as it gives equal weight to abundant and rare species (Hayekand Buzas, 2010). For each AB, the final value of cumulative H is usedhere as the index for α diversity (Jost, 2006; Wilson et al., 2010;Wilson, 2011; Wilson and Costelloe, 2011), which is expressed interms of the effective number of species SE (=eH). The between hab-itat or β diversity between successive ABs along the two transectswas calculated using the method described by Wilson (2011) andWilson and Costelloe (2011), where the dimensionless β=SE2/SE1and SE1 and SE2 are the effective number of species in successive ABsalong the transect. The statistical significance of β was determinedusing the t-test developed for the raw information function H byHutcheson (1970), in which

t ¼ H1−H2ð Þ= var:H1 þ var:H2ð Þ1=2

and

df ¼ var:H1 þ var:H2ð Þ2=½ð var:H12Þ=N1 þ ð var:H2

2Þ=N2�

where

var:H ¼ ½Σpi lnpið Þ2–ðΣpi· lnpiÞ2�=N−½ S−1ð Þ=2N2�:

Each AB boundary was placed into one of three categories followingWilson and Costelloe (2011). At Type 0β AB boundaries,Hutcheson's (1970) t-test does not reveal any difference in raw Hfor the α diversities and β is ~1. At the other two AB boundary

a

b

c

d

Fig. 3. SHE Analysis for Biozone Identification (SHEBI) for the shoreline Sample Set 1. a.SHEBI using all samples from Sample Set 1, showing point of inflection at boundary ofabundance biozone 1 (AB1) after the first three samples. b. SHEBI for Sample Set 1 sam-ples after the exclusion of samples in AB1, showing the AB2\AB3 boundary after threesamples. c. Plot showing delineation of AB3 (three samples). d. Plot showing lnE vs. lnNfor the final two abundance biozones (AB4\AB5).

41B. Wilson et al. / Marine Micropaleontology 82–83 (2012) 38–45

types, Hutcheson's (1970) t-test is significant: β>1 at Type +1βboundaries, while βb1 at Type −1β boundaries.

The β diversities presented here compare α diversities in adjacentABs but not assemblage compositions. Two ABs with different speciescompositions but similar vectors of species abundances will be sepa-rated by a Type 0β boundary. Complementarity sensu Vane-Wright etal. (1991) assesses the difference between two ABs in terms of the as-semblages contained. This study quantifies complementarity usingWhittaker's (1972) percentage similarity index CP, for which

CP ¼ 100–0:5Σ pi2–pi1j j;

where pi1 and pi2 are the percentage abundances of the ith species inthe lower and upper ABs respectively. Strayer et al. (2003) found thatspatial ecological boundaries are either permeable or impermeable tospecies' migrations, depending on the difference in the environmen-tal milieu across the boundary. Values of CP give an indication of per-meability. Where two assemblages are identical such that pi1=pi2and CP=100, the AB boundary is completely permeable. Where thebarrier is completely impermeable, CP=0.

4. Results

4.1. Sample Set 1

The sixteen samples in Sample Set 1 yielded 4619 benthonic fora-minifera in 81 species. Total recovery was dominated by D. rosea(20.2%) with subdominant Quinqueloculina auberiana (14.0%), A.angulatus (13.0%) and A. carinata (11.1%). Also common were Penero-plis proteus (8.2% of total recovery), Discorbis auberii (4.3%) and A. gib-bosa (4.2%). The highest and lowest point (per sample) values of S, Hand E are given in Table 1, with sample locations. Cockleshell Bay (SK:N) with the lowest H, had the highest percentage (49.7%) of D. rosea,which formed a mean of 19.9% of the recovery per sample along Sam-ple Set 1. In contrast, Major's Bay (SK:O) had the highest species rich-ness, the lowest percentage of D. rosea (2.8%) but the highest of A.angulatus (31.0%, mean 13.2%).

The lowest and highest point values of H equated to 6.3 and 26.2effective species respectively. Pattern diversity, measured using SE2/SE1 and where 1.0 indicates no change in diversity between samples,attained a minimum of 0.52 between SK:L* and SK:M, (Mosquito Bayand Cockleshell Bay), for which it indicates a decrease in diversity. Itattained a maximum of 2.88 between SK:N and SK:O (Cockleshell Bayand Major's Bay), indicating an increase in diversity between thesebays. These data reflect the low diversity in Cockleshell Bay.

SHEBI was commenced at Mosquito Bay, for which bay sample SK:L* (collected 1993) was included as the last sample. It detected fiveABs (Fig. 3), although the boundaries between themwere mostly sub-tle. AB1 contained the three samples from Mosquito Bay collected in2006, while AB2 contained SK:L* fromMosquito Bay and the two sam-ples from Cockleshell Bay. AB3 consisted of the one sample fromMajor's Bay and the two from Whitehouse Bay, while AB4 containedtwo samples from SE South Friar's Beach. The final AB contained theremaining sample from South Friar's Bay and four from FrigateBeach. Thus, SHEBI grouped some bays together but drew an ABboundary part way along South Friar's Beach. Therewas no correlation

Table 1Highest and lowest point (sample) measures along Sample Set 1 (shoreline). S=spe-cies richness, H=information function, E=equitability index.

Measuretype

Highest Lowest

Sample Value Location Sample Value Location

S SK:O 50 Major's Bay SK:X 20 Frigate BayH SK:I 3.27 Turtle Beach SK:N 1.84 Cockleshell BayE SK:I 0.61 Turtle Beach SK:Q 0.29 Whitehouse Bay

between sediment colour and ABs.While the foraminifera in AB1–AB3were pristinely preserved, those at South Friar's Beach\Frigate Bay(AB4 and AB5) were smooth and highly polished, perhaps due to anovergrowth of aragonite crystals (cf. Tucker and Wright, 1990, fig.3.10).

Alpha andβdiversities are listed in Tables 2 and3. Alphadiversitywashighest at Mosquito Bay (AB1) and lowest at South Friar's Beach\FrigateBay (AB5). Beta diversities ranged between 0.53 (AB1\AB2) and 1.18(AB2\AB3), but only those between AB1\AB2 and AB3\AB4 are statistical-ly significant, forming Type −1β boundaries. The other two AB bound-aries were of Type 0β.

Complementarity was greatest between AB4\AB5, at South Friar'sBay, indicating little change in the assemblages between these ABs.It was lowest across the AB3\AB4 boundary, between Whitehouseand South Friar's Bays. Table 4 lists species that change in abundanceby >1% between adjacent ABs. The following species (in alphabeticalorder) changed by >5% across any AB boundary: A. angulatus, A.

Table 4Species for which the percentage abundance changes by >1% across the AB boundaries,Sample Set 1 (shoreline).

Species AB1\AB2 AB2\AB3 AB3\AB4 AB4\AB5

Amphistegina gibbosa 1.8 −2.2 1.8 −1.4Archaias angulatus 3.4 20.3 −8.6 −3.9Asterigerina carinata −7.4 −2.8 7.6 −3.5Borelis pulchra −0.4 1.9 −2.2 0.4Discorbis auberii 2.6 −3.4 −5.7 0.2Discorbis granulosa 2.7 −2.4 −0.6 0.4Discorbis rosea 17 −12.5 −11.8 7.1Elphidium discoidale −2.8 −0.1 2.4 −1.5Hauerina bradyi −0.7 1.4 −1.4 0Nodobaculariella mexicana 0.2 0.2 1.9 2Peneroplis proteus 1.2 0.8 3.9 1.6Quinqueloculina auberiana 8.4 −3.4 17.2 1.2Quinqueloculina candeiana 0.1 0 1.8 −1.7Quinqueloculina lamarckiana −2.3 1.2 −1.9 1.5Quinqueloculina venezuelaensis 0.1 −0.8 0 1.2

Table 2Alpha diversities for abundance biozones along Sample Set 1 (shoreline). N=numberof specimens, S=species richness, H=information function, SE=effective number ofspecies, STG=percentage of assemblage as Sediment Tracer Group.

Abundancebiozone

No. ofsamples

N S Alpha (asH)

Var. H Alpha (asSE)

STG

AB1 3 811 63 3.16 0.0138 23.5 23.8AB2 3 877 45 2.53 0.0091 12.5 21.6AB3 3 787 65 2.7 0.0116 14.8 36.8AB4 2 5841 32 2.32 0.0106 10.2 32AB5 5 1563 40 2.4 0.0042 11 29

42 B. Wilson et al. / Marine Micropaleontology 82–83 (2012) 38–45

carinata, D. rosea, Q. auberiana and Textularia candeiana. The two larg-est changes were for A. angulatus, which increased by 20.3% betweenAB2\AB3, and D. rosea, which increased by 17.0% across the AB1\AB2boundary. Complementarity and β diversity were not significantlycorrelated along Sample Set 1.

Siphonaperta agglutinans 0 0.3 1.2 0.6Siphonaperta bidentata −0.1 0.1 2.5 −1.7Textularia candeiana −5.4 −1.1 1.1 −0.2Triloculina bicarinata −0.2 0 1.1 −1

Table 5Highest and lowest point (sample) measures along Sample Set 2 (reefs). S=species

4.2. Sample Set 2

The twelve samples from Sample Set 2 yielded 3539 benthonic fo-raminifera in 133 species. Five species each formed >5% of the totalrecovery: A. gibbosa (16.5%), A. carinata (13.9%), A. angulatus (7.9%),Peneroplis carinatus (6.7%) and P. proteus (6.2%). D. rosea and Q.auberiana, the two most abundant species from Sample Set 1, bothformed ≤2% of the total recovery from Sample Set 2. A. gibbosa wasmore abundant in the deeper part of Sample Set 2 and formed67.0% of the recovery from the deepest sample. P. proteus was at itsmost abundant at ≤6 m, while P. carinatus formed >7% of each sam-ple between 10 and 17 m.

The highest and lowest point values of S, H and E along Sample Set2 are given in Table 5. Brimstone Shallow Reef (SK:12, 27 m) had thelowest H, E and S and the highest percentage of A. gibbosa, whileFriar's Reef (SK:5, 10 m) had the highest H. The vent area offCamps Village had the highest S and low percentages of A. carinata,A. angulatus, P. carinatus and P. proteus.

The lowest and highest point diversities equated to 5.1 and 31.0effective species respectively. Pattern (between point) diversityattained a minimum of 0.17 between SK:11 (vent area, Camps Village18 m) and SK:12 (Brimstone Shallow Reef, 27 m) and were indicativeof a decrease in diversity. It attained a maximum of 1.64 betweenSK:10 (Old Road Reef, 17 m) and SK:11 (vent area, Camps Village,18 m). This reflects a high diversity (H=3.41, SE=30.1) at the ventarea off Camps Village relative to adjacent samples along the transect.

SHEBI was commenced at the shallowest sample and detectedfour ABs numbered AB:i through AB:iv (Fig. 4). Boundaries betweenABs occurred at ~8, 14, and 17 m. Alpha and β diversities are listedin Tables 6 and 7. Alpha diversity differed little between ABi throughABiii, but was markedly lower in ABiv. Thus, β ranged between 0.68(ABiii\ABiv) and 1.03 (ABi\ABii), but only that between ABiii\ABivwas statistically significant, indicating a Type −1β boundary. The re-mainder are Type 0β boundaries.

Complementarity was greatest across the ABii\ABiii boundary andlowest between ABiii and ABiv. Table 8 lists species that change inabundance by >1% between adjacent ABs. Only four species changed

Table 3Beta diversities and complementarities along Sample Set 1 (shoreline). t= t-test,df=degrees of freedom, CP=complementarity. Asterisk indicates significant t-test.

AB boundary Beta diversity t df CP

AB1\AB2 0.53 4.17* 1591 60.3AB2\AB3 1.18 1.18 1612 64.28AB3\AB4 0.69 2.53* 1353 54.72AB4\AB5 1.08 0.66 1072 86.38

by >5% across any AB boundary along Sample Set 2. From shallowestwater to deepest, these changes reflect the following:

1. P. proteus was at its most abundant on the shallower parts of thefringing reefs (ABi),

2. A. carinata increased only marginally across the ABi\ABii boundaryand thereafter decreased markedly, indicating a penchant forfringing reefs (ABi and ABii),

3. P. carinatus increased across the ABi\ABii and decreased across theABiii\ABiv boundary, reflecting its concentration the deeper partsof fringing reefs (ABii and ABiii),

4. A. angulatus changed little across the shallower boundaries but de-creased markedly between ABiii and ABiv. This reflects a fringingreef habitat,

5. A. gibbosa showed persistent positive changes across all bound-aries, reflecting its preference for deeper water but especially foroffshore bank reefs (ABiv).

4.3. Sample Set 1 versus Sample Set 2

Sample Set 1 (b1 m) contained five ABs and Sample Set 2 con-tained four. Beta diversity was used to document differences betweenthe ABs from Sample Set 1 and the shallowest water ABi from SampleSet 2. Beta ranged from 1.11 (between AB1\ABi) and 2.55 (AB5\ABi;Table 9). That all β diversities were positive suggests that diversity in-creased with depth. Only four of the β diversities were statisticallysignificant (the odd man out being the AB1\ABi boundary) but theywere indicative of Type +1β boundaries. Complementarities werelow (44.5–58.7: Table 9) reflecting especially decreases in the per-centage abundances of D. rosea and Q. auberiana, and increases in A.carinata and P. proteus, between the ABs in Sample Set 1 and ABi.

richness, H=information function, E=equitability index.

Measuretype

Highest Lowest

Sample Value Depth Location Sample Value Depth Location

S SK:11 66 18 Ventarea

SK:12 30 27 BrimstoneShallow

H SK:5 3.44 10 Friar'sReef

SK:12 1.63 27 BrimstoneShallow

E SK:5 0.5 10 Friar'sReef

SK:12 0.17 27 BrimstoneShallow

a

b

c

Fig. 4. SHE Analysis for Biozone Identification (SHEBI) for the reefal Sample Set 2. a.SHEBI using all samples from Sample Set 2, showing point of inflection at the boundaryfor the first abundance biozone (ABi) after the first four samples. b. SHEBI for SampleSet 2 samples after the exclusion of samples in ABi, showing the ABii\ABiii boundaryafter three samples. c. Plot showing delineation of the final two abundance biozones(ABiii\ABiv).

Table 7Beta diversities and complementarities along Sample Set 2 (reefs). t= t-test, df=degrees of freedom, CP=complementarity. Asterisk indicates significant t-test.

AB boundary Beta diversity t df CP

ABi\ABii 1.03 0.19 1858 61.19ABii\ABiii 0.88 0.78 1723 71.39ABiii\ABiv 0.68 2.16* 1261 50.94

Table 8Species for which the percentage abundance changes by >1% across the AB boundaries,Sample Set 2 (reefs).

Species ABi\ABii ABii\ABiii ABiii\ABiv

Amphistegina gibbosa 6.8 10 25.6Archaias angulatus 0.5 −0.4 −8.3Asterigerina carinata 2.5 −7.1 −11.4Borelis pulchra 0.5 0.7 −1.4Cibicidoides pseudoungerianus 0 0.6 1.2Cyclorbiculina compressa −2.2 −0.9 0.2Discorbis rosea −2.8 1.4 −1.9Elphidium discoidale 1.9 0.9 −3.2Eponides antillarum 1.3 0.4 2.1Hauerina ornatissima 0.5 0 1.3Hauerina pacifica −0.5 −0.2 1.8

43B. Wilson et al. / Marine Micropaleontology 82–83 (2012) 38–45

5. Discussion and conclusions

The ABs detected here reflect sediment assemblages stratified bydepth, with a particularly pronounced progression from dominantD. rosea (nearshore sediment) through A. carinata (fringing reefs) toA. gibbosa (offshore bank reefs). P. proteus, abundant in the shallowerparts of the fringing reefs (ABi), was replaced by the more disc-likeand delicate P. carinatus in their deeper parts (ABii\ ABiii).

The total assemblage preserved at a site is controlled by twocauses:

1. the composition of the live assemblage, which in the shallowwaterof the eastern Caribbean is predominantly epiphytal (Brasier,1975a; Steinker, 1977; Steinker and Clem, 1984), and

Table 6Alpha diversities for abundance biozones along Sample Set 2 (reefs). N=number ofspecimens, S=species richness, H=information function, SE=effective number ofspecies.

Abundancebiozone

No. ofsamples

N S Alpha (asH)

Var. H Alpha (asSE)

ABi 4 1231 92 3.26 0.01 26ABii 3 869 86 3.29 0.0144 26.8ABiii 3 856 74 3.16 0.0134 23.6ABiv 2 583 75 2.77 0.0192 16

2. post-mortem effects. The latter consists of specimen loss throughdissolution and abrasion (Martin and Liddell, 1991; Kotler et al.,1992) and transport during storms (Coulbourn and Resig, 1975;Debenay, 1988; Li et al., 1997, 1998) and by other means such asfish predation (Lipps, 1988; Debenay et al., 2010).

It is not possible to assess the effects of dissolution and abrasionfrom the data presented here. However, some comments can bemade regarding sediment transport.

Previous studies indicate that the dominance of D. rosea in thenearshore Sample Set 1 reflects its abundance on associated nearshorevegetation and that these specimens have been little transported.Wilson and Ramsook (2007) examined live, epiphytal foraminiferalfaunas at b1 m water depth around Nevis, ~3 km south of St. Kitts.The live, epiphytal fauna on seagrass rhizomes in a pristine bay was44% D. rosea, 13% D. auberii and 10% Tretomphalus bulloides. However,Li et al. (1997) found D. rosea to dominate sediment assemblages onthe upper reef terrace around the Cayman Islands. The low abun-dances of D. rosea along the reefal Sample Set 2 (maximum 3.6%,ABi) compared to Sample Set 1 (minimum 9.4%, AB4, shoreline atSouth Friar's Beach; maximum 33.8%, AB2, Mosquito and CockleshellBays) suggests that there is limited offshore transport of nearshoresediment onto the shallower parts of the fringing reefs. This isreflected in the low complementarities between Sample Sets 1 and2. It is possible that nearshore sediment is instead transported throughchutes between the fringing reefs and deposited between them andthe offshore banks (cf. Li et al., 1997).

The three other tracer species recorded by Li et al. (1997; A.carinata, A. angulatus and A. gibbosa) were recovered more or less

Nodobaculariella mexicana 0.5 0.6 −1.9Peneroplis carinatus 7.4 −0.4 −8.2Peneroplis pertusus −0.3 0.4 −1.2Peneroplis proteus −16 0.9 −1.5Quinqueloculina auberiana −1.4 −0.6 −0.2Quinqueloculina lamarckiana −0.7 −1.1 −1.4Quinqueloculina sp. D. Wilson 1998 1 −0.8 0.1Quinqueloculina venezuelaensis −2.5 0 −1.1Rosalina subaraucana 0.9 −1 0.9Sahulia cf. conica 0 0 1.2Siphonaperta agglutinans −1.2 1.8 0.2Siphonaperta bidentata −2.2 0.9 −0.4Siphonina pulchra 0.1 0.9 1.5Sorites obiculus 0.1 1.6 −0.9Textularia candeiana 3.2 −2.4 −0.6

Table 9Beta diversities and complementarities between abundance biozones along Sample Set1 (shoreline) and the shallowest abundance biozone along Sample Set 2 (reefs). t= t-test, df=degrees of freedom, CP=complementarity. Asterisk indicates significant t-test.

AB boundary Beta t df CP

AB1\ABi 1.11 0.66 1792 54.4AB2\ABi 2.08 5.30* 2077 44.5AB3\ABi 1.76 3.83* 1846 48.5AB4\ABi 2.55 4.79* 518 58.7AB5\ABi 2.37 7.63* 2076 55.9

44 B. Wilson et al. / Marine Micropaleontology 82–83 (2012) 38–45

abundantly from the nearshore Sample Set 1, but none of them livescommonly in the nearshore (b1 m) area. Wilson and Ramsook(2007) recorded only 47 A. gibbosa, 6 A. angulatus and 13 A. carinataamong 11,150 live foraminifera recovered from epiphytal communi-ties in waterb1 m around Nevis. Their presence in Sample Set 1thus indicates shoreward transport of sediment.

Most sediment transported shoreward from the reefs to the near-shore areas, being characterised especially by A. carinata and A. angu-latus, was derived from the shallower parts of the fringing reefs. A.carinata was along Sample Set 1 abundant in Mosquito Bay (AB1,17.0%) and South Friar's Bay+Frigate Beach (AB4, 14.3%; AB5,10.8%) reflecting a concentration here of material derived from theshallower parts of the fringing reefs (ABi\ABii). The high concentra-tion of A. angulatus along the shoreline at South Friar's Beach (AB4,18% of recovery) reflects the manner in which this species has beenconcentrated here by wave action, it forming only ~6.5% of the recov-ery elsewhere along South Friar's Bay+Frigate Beach (AB5) and innearby Whitehouse and Major's Bays (AB3). A. gibbosa, which domi-nated the offshore bank reefs, was generally a small component ofsamples from Sample Set 1 (mean, 4.2%, s.d. 1.7%), where its abun-dance differed little from the shallow fringing reefs (ABi, 3.9%). Thissuggests that there was limited sediment transport into nearshoreareas from the deeper-water, offshore bank reefs (ABiv). This doesnot mean, however, that sediment from offshore bank reefs is nottransported towards shore; it might be deposited in the area betweenthe platform edge and the fringing reefs.

Percentage abundances along Sample Set 1 for A. carinata+A. gib-bosa+A. angulatus (here termed a sediment tracer group) range be-tween 21.6 and 36.6% (Table 2). It might be argued that the highestabundances between Major's Bay and Frigate Beach (AB3–AB5), arerelated to the proximity of fringing reefs. However, this is unlikely:fringing coral reefs occur close to the other bays also.

There is evidence of considerable seaward transport of sedimentfrom the reefs. Wilson (2008a, 2011) reported that foraminiferal as-semblages in bathyal sediment around St. Kitts contains up to 56%allochthonous, shallow water specimens, with especially abundantA. carinata, Peneroplis bradyi, P. proteus and A. gibbosa. He recordedonly a single D. rosea, however. Thus, the data presented here donot support Wilson's (2008a) assertion that the allochthonous ele-ment in bathyal sediment comprises “predominantly back-reef spe-cies” that were displaced “on floating marine-plant debris.” Rather,the approximately equal abundances of A. carinata and A. gibbosa sug-gest that the allochthonous component comprises reefal species. Itmay be that such sediment is transported during hurricanes and trop-ical storms (cf. Li et al., 1997), of which the NE Caribbean Sea was im-pacted by >60 between 1871 and 1986 alone (Lugo et al., 2000).

Hallock et al. (2003) noted that environmental change at a sitewill, if it persists long enough, be reflected in the sediment assem-blage, although it will initially be reflected by a regime shift in thelive, epiphytal fauna (e.g., Wilson and Ramsook, 2007). There issome evidence of environmental change around the SEP. The ABs inSample Set 1 frequently grouped samples from more than one bay.Thus, sample SK:L*, collected from Mosquito Bay in 1993, groupedwith samples taken from Cockleshell Bay in 2006. However, three

other samples collected fromMosquito Bay in 2006 formed a separateAB. (This remained so when SK:L* was placed as the first sample fromMosquito Bay, rather than the last, during the accumulation of sam-ples from SHEBI.) This reflects a decrease in D. rosea in sediment atthis site between 1993 and 2006 accompanied by an increase in α di-versity. Wilson and Ramsook (2007) having shown that (a) D. rosea isindicative of sites that are relatively unaffected by organic matter en-richment and (b) organic matter enrichment in the Caribbean in-creases epiphytal foraminiferal diversity, it is concluded that thischange in the sediment assemblage at Mosquito Bay reflects the im-pact of the beach bar on organic matter concentrations. There hadnot by 2006, however, been any noticeable change in the proportionof opportunists within the sediment (cf. Hallock et al., 2003).

Foraminifera living on fringing coral reefs are a considerable com-ponent of nearshore sediment (Hallock, 2000). The continued healthof foraminiferal communities is thus vital to the integrity of beachesaround St. Kitts. This study provides a baseline survey of the forami-nifera on reefs around St. Kitts and Nevis. Given that total sedimentassemblages react to environmental change (albeit more slowlythan live ones), future surveys incorporating the sites studied herehave the potential to contribute to the monitoring and managementof coral reefs around St. Kitts and Nevis.

Acknowledgement

Sample Set 2 was collected by JP as part of a United Nations GlobalEnvironment Fundproject entitled “Towards the SustainableMonitoringand Management of Coral Reefs in St. Kitts and Nevis,”while JacquelineWilson aided in the collection of Sample Set 1. A contribution from theResearch and Publications Fund of the University of the West Indies isgratefully acknowledged. Thoughtful and thorough reviews by RonaldLewis and Ronald Martin improved the manuscript greatly while clear-ing up some misunderstandings on the part of BW.

Appendix A. Supplementary data

Supplementary data to this article can be found online at doi:10.1016/j.marmicro.2011.10.001.

References

Adey, W.H., Burke, R., 1976. Holocene bioherms (algal ridges and bank-barrier reefs) ofthe eastern Caribbean. Geological Society of America Bulletin 87, 95–109.

Brasier, M.D., 1975a. Ecology of Recent sediment-dwelling and phytal foraminiferafrom the lagoons of Barbuda, West Indies. Journal of Foraminiferal Research 5,42–62.

Brasier, M.D., 1975b. The ecology and distribution of Recent foraminifera from the reefsand shoals around Barbuda, West Indies. Journal of Foraminiferal Research 5,193–210.

Brooks, W.W., 1973. Distribution of Recent foraminifera from the southern coast ofPuerto Rico. Micropaleontology 19, 385–416.

Buzas,M.A., Hayek, L.-A.C., 1996. Biodiversity resolution: an integratedapproach. BiodiversityResearch 3, 40–43.

Buzas, M.A., Hayek, L.-A.C., 1998. SHE analysis for biofacies identification. Journal ofForaminiferal Research 28, 233–239.

Buzas, M.A., Hayek, L.-A.C., 2005. On richness and evenness within and between communi-ties. Paleobiology 31, 199–220.

Buzas, M.A., Hayek, L.-A.C., Culver, S.J., 2007a. Community structure of benthic forami-nifera in the Gulf of Mexico. Marine Micropaleontology 65, 43–53.

Buzas, M.A., Hayek, L.-A.C., Hayward, B.W., Grenfell, H., Sabaa, A.T., 2007b. Biodiversityand community structure of deep-sea foraminifera around New Zealand. Deep-SeaResearch I 1641–1654.

Caribbean Conservation Association, 1990. Country Environmental Profile: St. Kitts andNevis. Island Resources Foundation, St. Thomas, US Virgin Islands. 276 pp.

Coulbourn, W.T., Resig, J.M., 1975. On the use of benthic foraminifera as sedimenttracers in a Hawaiian bay. Pacific Science 29, 99–115.

Culver, S.J., Buzas, M.A., 1982. Distribution of Recent benthic foraminifera in the Caribbeanregion. Smithsonian Contributions to Marine Science 14, 1–382.

Cushman, J.A., 1921. Foraminifera from the north coast of Jamaica. Proceedings of theU. S. National Museum 59 (2360), 47–82.

Cushman, J.A., 1922. Shallow-water foraminifera of the Tortugas region. Carnegie Institutionof Washington 17, 3–85.

Cushman, J.A., 1923. The foraminifera of the Atlantic Ocean, Part 4: Lagenidae. UnitedStates National Museum Bulletin 104 (4), 1–228.

45B. Wilson et al. / Marine Micropaleontology 82–83 (2012) 38–45

Cushman, J.A., 1929. The foraminifera of the Atlantic Ocean, Part 6: Miliolidae. UnitedStates National Museum Bulletin 104 (6), 1–129.

Cushman, J.A., 1930. The foraminifera of theAtlantic Ocean, Part 7: Nonionidae, Camerinidae,Peneroplidae and Alveolinellidae. United States National Museum Bulletin 104 (7),1–179.

Cushman, J.A., 1931. The foraminifera of the Atlantic Ocean, Part 8: Rotaliidae, Amphis-teginidae, Calcarinidae, Cymbalopoerttidae, Globorotaliidae, Anomalinidae, Pla-norbulinidae, Rupertiidae, and Homotremidae. United States National MuseumBulletin 104 (8), 1–179.

Debenay, J.-P., 1988. Recent foraminifera tracers of oceanic water movements in thesouthwestern lagoon of New Caledonia. Palaeogeography, Palaeoclimatology,Palaeoecology 65, 59–72.

Debenay, J.-P., Sigura, A., Justine, J.-L., 2010. Foraminifera in the diet of coral reef fishfrom the lagoon of New Caledonia: predation, digestion, dispersion. Revue deMicropaleontologie 54, 87–103.

d'Orbigny, A., 1839. Foraminifères. In: de La Sagra, R. (Ed.), histoire physique, politiqueet naturelle de l'ile de Cuba. Arthus Bertrand, Paris, France, pp. 1–224.

Gischler, E., Hauser, I., Heinrich, K., Scheitel, U., 2003. Characterization of depositionalenvironments in isolated carbonate platforms based on benthic foraminifera,Belize, Central America. Palaios 18, 236–255.

Hallock, P., 2000. Larger foraminifers as contributors to carbonate beach sands. In:Magoon, O.T., Robbins, L.L., Ewing, L. (Eds.), Carbonate Beaches 2000. AmericanSociety of Civil Engineers, Key Largo, Florida, U.S.A., pp. 97–98.

Hallock, P., Lidz, B., Cockey-Burkhard, E.M., Donnelly, K.B., 2003. Foraminifera as bioindica-tors in coral reef assessment and monitoring: the FORAM Index. Environmental Moni-toring and Assessment 81, 221–238.

Hammer, Ø., Harper, D.A.T., 2005. Paleontological Data Analysis. Wiley-Blackwell,Oxford, UK. 368 pp.

Hayek, L.-A.C., Buzas, M.A., 2010. Surveying Natural Populations: Quantitative Tools forAssessing Biodiversity. Columbia University Press, New York, USA. 590 pp.

Hofker, J., 1956. Foraminifera dentata: foraminifera of Santa Cruz and Thatch Island,Virginia Archipelago, West Indies. Spolia Zoologica Musei Hauniensis 15, 1–237.

Hofker, J., 1964. Foraminifera from the tidal zone in the Netherlands Antilles and otherWest Indian islands. Studies on the Fauna of Curacao and other Caribbean Islands21, 1–119.

Hofker, J., 1971. The foraminifera of Piscadera Bay, Curacao. Studies on the Fauna ofCuracao and other Caribbean Islands 35, 1–94.

Hofker, J., 1976. Further studies on Caribbean foraminifera. Studies on the Fauna of Curacaoand other Caribbean Islands 40, 1–252.

Hofker, J., 1980. The foraminifera of the Saba Bank Expedition 1972 (Cigar Cruises 34, 35).ZoologischeVerhandelingenUitgegevendoorhet RijksmuseumvanNatuurlijkeHistoirete Leiden 177 73 pp.

Hunter, M., 2002. Fundamentals of Conservation Biology. Blackwell Science, Inc., Malden,Massachussetts, USA. 555 pp.

Hutcheson, K., 1970. A test for comparing diversities based on the Shannon formula.Journal of Theoretical Biology 29, 151–154.

Jost, L., 2006. Entropy and diversity. Oikos 113, 363–375.Kjerfve, B., 1981. Tides of the Caribbean Sea. Journal of Geophysical Research 86 (C5),

4243–4247.Kotler, E.,Martin, R.E., Liddell,W.D., 1992. Experimental analysis of abrasion anddissolution

resistance of modern reef-dwelling foraminifera; implications for the preservation ofbiogenic carbonate. Palaios 7, 244–276.

Li, C., Jones, B., Blanchon, P., 1997. Lagoon-shelf sediment exchange by storms—evidencefrom foraminiferal assemblages, east coast of Grand Cayman, British West Indies.Journal of Sedimentary Research 67, 17–25.

Li, C., Jones, B., Kalbfleisch, W.B.C., 1998. Carbonate sediment transport pathways basedon foraminifera: case study from Frank Sound, Grand Cayman, British West Indies.Sedimentology 45, 109–120.

Lipps, J.H., 1988. Predation on foraminifera by coral reef fish; taphonomic and evolu-tionary implications. Palaios 3, 315–326.

Lugo, A.E., Rogers, C.S., Nixon, S.W., 2000. Hurricanes, coral reefs and rainforests: resis-tance, ruin and recovery in the Caribbean. Ambio 29, 106–114.

Martin, R.E., Liddell, W.D., 1988. Foraminiferal biofacies on a North Coast Fringing Reef(1–75 m), Discovery Bay, Jamaica. Palaios 3, 298–314.

Martin, R.E., Liddell, W.D., 1991. Taphonomy of foraminifera in modern carbonateenvironments: implications for the formation of foraminiferal assemblages. In:Donovan, S.K. (Ed.), The Processes of Fossilization. Belhaven Press, London, pp.170–193.

Martin-Kaye, P.H.A., 1969. A summary of the geology of the Lesser Antilles. OverseasGeology and Mineral Resources 10, 172–206.

Mendoza, W.G., Zika, R.G., Corredor, J.E., Morrel, J., Ko, D.S., Mooers, C.N., 2009. Develop-mental strategy for effective sampling to detect possible nutrientfluxes in oligotrophiccoastal reef waters in the Caribbean. Journal of Operational Oceanography 2, 35–47.

Mora, C., 2008. A clear human footprint in the coral reefs of the Caribbean. Proceedingsof the Royal Society B 275, 767–773.

Murray, J.W., 2006. Ecology andApplicationsof Benthic Foraminifera. CambridgeUniversityPress, Cambridge, UK. 438 pp.

Radford, S.S., 1976a. Depth distribution of Recent foraminifera in selected bays, TobagoIsland, West Indies. Revista Espanola de Micropaleontología 8, 219–238.

Radford, S.S., 1976b. Recent foraminifera from Tobago Island, West Indies. RevistaEspanola de Micropaleontología 8, 193–218.

Radford, S.S., 1998. Foraminifera from the Southern Caribbean–Atlantic Province. In:Ali, W., Paul, A., Young On, V. (Eds.), 3rd Geological Conference of the GeologicalSociety of Trinidad and Tobago and the 14th Caribbean Geological Conference,Port-of-Spain, Trinidad, pp. 163–178.

Schnitker, D., 1971. Distribution of foraminifera on the North Carolina continentalshelf. Tulane Studies in Geology and Paleontology 8, 169–215.

Smith, A.H., Rogers, C.S., Bouchon, C., 1997. Status of western Atlantic coral reefs in theLesser Antilles. In: Lessios, H.A., Macintyre, I.G. (Eds.), Proceedings of the 8th Inter-national Coral Reef Symposium. Smithsonian Tropical Research Institute, PanamaCity, pp. 351–356.

Steinker, D.C., 1977. Foraminiferal studies in tropical carbonate environments: SouthFlorida and Bahamas. Florida Science 40, 42–45.

Steinker, D.C., Clem, K.V., 1984. Some nearshore foraminiferal assemblages from phytalsubstrates and bottom sediments, Bermuda. The Compass 61, 98–115.

Strayer, D.L., Power, M.E., Fagan, W.F., Pickett, S.T.A., Belnap, J., 2003. A classification ofecological boundaries. BioScience 53, 723–729.

Todd, R., Low, D., 1971. Foraminifera from the Bahama Bank west of Andros Island. USGeological Survey Professional Paper 683-C, 1–22.

Tucker, M.E., Wright, V.P., 1990. Carbonate Sedimentology. Blackwell Science, Oxford,UK. 482 pp.

Vane-Wright, R.I., Humphries, C.J., Williams, P.H., 1991. What to protect? Systematicsand the agony of choice. Biological Conservation 55, 235–254.

Whittaker, R.H., 1972. Evolution and measurement of species diversity. Taxon 21,213–251.

Wilkinson, C., 2004. Status of Coral Reefs of theWorld, 2004, Volume2. Australian Instituteof Marine Science, Townsville, Queensland, Australia, p. 557.

Wilson, B., 1998. Epiphytal foraminiferal assemblages on the leaves of the seagrassesThalassia testudinum and Syringodium filiforme. Caribbean Journal of Science 34,131–132.

Wilson, B., 2008a. Late Quaternary benthonic foraminifera in a bathyal core from theLeeward Islands, Lesser Antilles, NE Caribbean Sea. Journal of Micropalaeontology27, 177–188.

Wilson, B., 2008b.Using SHEBI (SHEAnalysis for Biozone Identification): to proceed from thetop down or the bottom up? A discussion using twoMiocene foraminiferal successionsfrom Trinidad, West Indies. Palaios 23, 636–644.

Wilson, B., 2011. Αlpha and beta diversities of Late Quaternary bathyal benthonic fora-miniferal communities in the NE Caribbean Sea. Journal of Foraminiferal Research41, 40–47.

Wilson, B., Costelloe, A., 2011. Abundance biozone boundary types and characteristicsdetermined using beta diversity: an example using Pleistocene benthonic forami-nifera in DSDP Hole 148, eastern Caribbean Sea. Palaios 26, 152–159.

Wilson, B., Ramsook, A., 2007. Population densities and diversities of epiphytal forami-nifera on nearshore substrates, Nevis, West Indies. Journal of Foraminiferal Re-search 37, 213–222.

Wilson, B., Jones, B., Birjue, K., 2010. Paleoenvironmental interpretations based on fora-miniferal abundance biozones, Mayo Limestone, Trinidad, West Indies, includingalpha and beta diversities. Palaios 25, 158–166.