Regional Variation in Caribbean Dry Forest Tree Species Composition

Home , Bucida, Bursera simaruba, Coccoloba, Drypetes, Metopium toxiferum, Oxandra lanceolata

Plant Ecol DOI 10.1007/s11258-015-0474-8

Regional variation in Caribbean dry forest tree species composition

Janet Franklin . Julie Ripplinger . Ethan H. Freid . Humfredo Marcano-Vega . David W. Steadman

Received: 8 November 2014 / Accepted: 14 April 2015 Ó The Author(s) 2015. This article is published with open access at Springerlink.com

Abstract How does tree species composition vary in using clustering, ordination, and indicator species relation to geographical and environmental gradients analysis at regional and local scales. Regionally, the in a globally rare tropical/subtropical broadleaf dry largest cluster group included over half of all plots and forest community in the Caribbean? We analyzed data comprised plots from all three island groups. Indicator from 153 Forest Inventory and Analysis (FIA) plots species were native Bursera simaruba (Burseraceae) from Puerto Rico and the U.S. Virgin Islands (USVI), and Metopium toxiferum (Anacardiaceae). Species along with 42 plots that we sampled in the Bahamian composition was similar to dry forests throughout the Archipelago (on Abaco and Eleuthera Islands). FIA region based on published studies. Other groups we data were collected using published protocols. In the identiﬁed at the regional scale consisted of many Bahamian Archipelago, we recorded terrain and Puerto Rico and USVI plots that were dominated by landscape variables, and identiﬁed to species and non-native species, documenting the widespread na- measured the diameter of all stems C5 cm at 1.3 m ture of novel ecosystems. At the local scale the height in 10 m radius plots. All data were analyzed Bahamian data clustered into two main groups corre- sponding largely to the two islands sampled, a pattern consistent with the latitudinal aridity gradient. Ba- Communicated by Lesley Rigg. hamian dry forests share previously undocumented compositional similarity with native-dominated dry Electronic supplementary material The online version of this article (doi:10.1007/s11258-015-0474-8) contains supple- forests found throughout the Caribbean, but they lack mentary material, which is available to authorized users.

J. Franklin (&) H. Marcano-Vega School of Geographical Sciences and Urban Planning, USDA Forest Service, Southern Research Station, Arizona State University, Tempe, AZ 85287, USA International Institute of Tropical Forestry, 1201 Calle e-mail: [email protected] Ceiba, Jardıń Botańico Sur, San Juan, PR 00926, USA e-mail: [email protected] J. Ripplinger School of Life Sciences, Arizona State University, D. W. Steadman Tempe, AZ 85287, USA Florida Museum of Natural History, University of Florida, e-mail: [email protected] P. O. Box 117800, Gainesville, FL 32611, USA e-mail: dws@flmnh.ufl.edu E. H. Freid Bahamas National Trust/Leon Levy Native Plant Preserve, P.O. Box N-4105, Nassau, Bahamas e-mail: [email protected]

123 Plant Ecol extensive post-disturbance novel dry forests dominat- The broadleaf forest found in the Bahamian Archi- ed by non-native trees found in the Greater Antilles. pelago, known locally as ‘‘coppice’’(Correll and Correll 1982), has been called dry evergreen forest (Smith and Keywords Bahamas Bahamian archipelago Vankat 1992), lowland tropical or subtropical seasonal Á Á Broadleaf subtropical evergreen forest Forest evergreen forest (Areces-Mallea et al. 1999) or Á community Greater Antilles Novel ecosystems Caribbean dry forest (Dunphy et al. 2000). We use Á Á Á Tropical seasonal evergreen forest Vegetation ‘‘dry forest’’ in this paper for simplicity. Bahamian dry Á classification West Indies forest is a globally rare and endangered ecosystem, Á supporting endemic and endangered species (WWF Worldwide Report 2001 http://worldwildlife.org/ ecoregions/nt0203). It has been identified as an impor- Introduction tant conservation target, threatened by clearing and degradation (Sullivan-Sealey et al. 2002; Wunderle and Understanding regional variation in plant community Waide 1993). Little is known, however, about its rela- composition is required in an era of global change tionship to other dry forests in the region, or composi- because ecosystem processes and services depend on tional variation within the archipelago, where dry forest taxon-specific responses to climate change, land use has been studied only within single islands (Currie et al. change, novel species combinations, and altered 2005a; Franklin and Steadman 2013; Freid and Kerwin disturbance regimes (Sala et al. 2000; Tylianakis 1998; Helmer et al. 2010; Smith and Vankat 1992). et al. 2008). Describing plant communities also serves This study begins to fill the gap in our knowledge of important purposes in conservation planning. Plant regional variation in Caribbean dry forest composition communities are used to set conservation targets for by addressing the following research questions: (1) terrestrial ecosystems (Keith et al. 2013; Murdoch What are the broad regional patterns of variation in et al. 2010), and as a ‘‘coarse filter’’ (Noss 1987) for Caribbean dry forest composition? (2) Is there local estimating patterns of biodiversity to assess gaps in variation in dry forest composition within the less- conservation coverage (Butterfield et al. 1994). Major studied Bahamian islands that is geographically or ongoing efforts aim to develop vegetation classifica- environmentally structured? tions using standardized protocols and quantitative We addressed the first question by analyzing Forest methods (Franklin et al. 2012); a global classification Inventory and Analysis (FIA) data available from the does not yet exist, however, and there is great U.S. Department of Agriculture (USDA) Forest Ser- unevenness in the geographical distribution of quan- vice for Puerto Rico and the U.S. Virgin Islands titative plant community data (Dengler et al. 2011). (USVI) in conjunction with comparable plot data we Janzen (1988) declared that tropical dry forests are collected on Bahamian islands. These data range the most threatened tropical forest ecosystems, a across two-thirds of the latitudinal extent and half of statement that remains valid (Gillespie et al. 2012; the longitudinal extent of the West Indies (Fig. 1). Miles et al. 2006). Patterns of diversity and compo- Results of this regional analysis are compared to sition of Neotropical dry forests have been broadly published descriptions of dry forests elsewhere in the described (Gentry 1995), and ecological attributes of Caribbean. We expected that composition would vary Antillean (Lugo et al. 2006) and south Florida, US among major island groups in our sample because of (Gillespie 2006) dry forests have been reviewed. differences in island size and elevation, as well as Although dry forests have been studied on individual geological substrate. We then addressed the second islands within the Caribbean region, a comprehensive, question by comparing our Bahamian dry forest data quantitative analysis of Caribbean dry forest compo- from two islands—Abaco and Eleuthera. These sition is lacking. Furthermore, dry forest communities islands are separated by deep water, and together span in some parts of the Caribbean, such as the Bahamian more than 2° of latitude and nearly 2° of longitude, Archipelago, are not well studied (Carey et al. 2014). almost half of the linear extent of the archipelago. We The Bahamian Archipelago is divided politically into expected to discover latitudinal variation in species The Commonwealth of the Bahamas and The Turks composition related to the north–south gradient of and Caicos Islands. increasing aridity and warmth.

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Fig. 1 The Caribbean study region showing the Bahamian Abaco and Eleuthera. Plots are clustered and cannot be Archipelago (The Bahamas ? Turks and Caicos Islands), Cuba, discerned at this scale. For example, 8 plots were located in a Jamaica, Hispaniola, and Puerto Rico (Greater Antilles), the large patch of forest on east-central Abaco in the area known as U.S. Virgin Islands, and the southern portion of Florida, US. Wilson City Inset shows with the number of vegetation plots for locations on

Materials and methods the locations of the dry forest plots were extracted from WorldClim (worldclim.org; accessed 22 Sep Study area 2014); temperatures for those locations range from an average high of 31 °C in Aug to average low of 20 °C FIA data came from the USVI and the Commonwealth in Jan, with average annual precipitation *1200 mm of Puerto Rico (the main island of Puerto Rico and its (range 900–1900 mm). Although some error is delib- outlying islands), and published vegetation studies erately added to the reported locations of FIA plots also provided descriptions of dry forest on the large owing to privacy regulations (O’Connell et al. 2014)), islands of the Greater Antilles (Jamaica, Cuba, Puerto this merely added a small amount of uncertainty to our Rico and Hispaniola), and in south Florida, US descriptive climate statistics. Many of the dry forests (Fig. 1). Dry forests of Puerto Rico and throughout found today in Puerto Rico and the USVI are the Greater Antilles tend to occur at low elevations and dominated by young small-diameter trees. Further- often on limestone (Lugo et al. 2006). The USVI are more, naturalized, introduced species are an important volcanic, low, hilly islands at the eastern margin of the component of forests regenerating on abandoned Greater Antilles. FIA dry forest plots were located cropland and pastures (Brandeis and Turner 2013a, near the southwestern (leeward) coast of the island of b; Molina Colo´n et al. 2011). Puerto Rico and nearby islands of Mona, Vieques, and The Bahamian Archipelago consists of limestone Culebra, and at lower elevations of St. Croix, St. John from indurated calcareous sand deposited on massive, and St. Thomas islands in the USVI. Climate data for shallow carbonate banks (Sealey 1994). Abaco (also

123 Plant Ecol called Great Abaco), which lies on the Little Bahama Data collection Bank, is the second largest island in the archipelago (1244 km2), has a maximum elevation of 44 masl, and Forest plot data for Puerto Rico (sampled between 2001 extends *120 km N–S (Fig. 1). Climate data were and 2009) and the USVI (2004–2009) were acquired not available for Abaco but at nearby Freeport, Grand from the USDA Forest Service’s Forest Inventory and Bahama (268300N) temperatures range from an aver- Analysis (FIA) Program (http://www.fia.fs.fed.us/). FIA age high of 30 °C in Aug to average low of 18 °C in plots from Puerto Rico have been used to develop bio- Jan, with annual precipitation *1450 mm. Abaco at mass and volume equations for dry forest (Brandeis et al. 26–278N experiences more winter cold fronts than 2006) and those from the USVI have been used in a Eleuthera (Sealey 1994). The most extensive vegeta- study of bird habitat on St. John (Steadman et al. 2009). tion community on Abaco is pine woodland (Areces- We extracted data for 184 plots classified as ‘‘sub- Mallea et al. 1999), dominated by Pinus caribaea var. tropical dry forest.’’ Of these, 31 plots were excluded bahamensis, endemic to the Bahamian Archipelago from our analyses (leaving 153) because their multi- and found on only six of its islands (Sullivan-Sealey variate distance from another plot was zero, precluding et al. 2002). Dry forest occupies about 10 % of the some multivariate analyses. These were plots with only land area of Abaco. Eleuthera (457 km2, maximum a few trees, all of a single species. For example, 25 of the elevation 60 masl) lies on the Great Bahama Bank, is excluded plots contained only a single species of a non- about 120 km long NW–SE, and in places is\1.5 km native leguminous tree. In FIA plots, dbh (diameter at wide (Fig. 1). Climate data were not available for breast height) is measured at 1.37 m height on the trunk Eleuthera but at nearby Nassau, New Providence and the height of each tree is measured. Trees C12.5 cm (25840N) temperatures range from an average high of dbh are sampled in 4 clustered circular subplots of 32 °C in Aug to average low of 21 °C in Jan, with 168 m2 each (total sampling area 672 m2) whose cen- *1400 mm annual precipitation. Upland habitats on ters are 36.6 m apart; trees 2.5 to \12.5 cm dbh are Eleuthera are dominated by broadleaf dry forest and sampled in 4 circular microplots of 13.46 m2 (total scrub. Abaco and Eleuthera, lying on different sampling area 53.84 m2) nested within subplots (Gray Bahamian Banks, would have each been part of much et al. 2012; O’Connell et al. 2014)). We extracted data larger islands and closer together when these shallow only for trees C 5.0 cm dbh for comparability with our banks were exposed at lower sea-levels during much Bahamian data. of the Pleistocene (Steadman and Franklin 2015). In the Bahamian Archipelago, we established 20 Bahamian dry broadleaf woody vegetation is subplots on Abaco and 22 plots on Eleuthera. Plot divided into ‘‘short’’ or ‘‘low’’ (\4.6 m) scrub versus locations were geographically stratified in order to ‘‘tall’’ or ‘‘high’’ ([4.6 m) forest that differs primarily sample all major patches of inland dry forest on Abaco in stature, because of variation in site quality or time and to span the extent of Eleuthera (Fig. 1). Random since disturbance (Currie et al. 2005b), and into sampling was not possible because access was chal- coastal (‘‘whiteland’’) versus inland (‘‘blackland’’) lenged by rugged terrain, lack of trails, and land forest. Coastal broadleaf woody vegetation occurs on ownership. For efficiency, plots often were located in sands near the coast and tends to be short. Inland nearby pairs ([500 m apart) or, in the case of Abaco, broadleaf woody vegetation is found on limestone on transects associated with bird surveys (Franklin and with greater soil development, is typically taller, and Steadman 2013). Sampling took place on 7 Jan and 6–9 differs somewhat in species composition from the May 2009; 9 Jan, 12–17 Mar, 4–6 May, and 30 Nov-3 coastal form (Correll and Correll 1982; Smith and Dec 2012; and 5–10 Dec 2013. We did not observe any Vankat 1992). Our study focuses on inland, tall forest, large scale canopy gaps created by the hurricanes that although these forms grade into one another in many affected the islands during this time frame. locations. Bahamian dry forest is considered to be Bahamian plots were 10 m radius (314 m2), a plot resilient to hurricanes that occur frequently there size adequate for forest community analysis (Otyp- (Morrison and Spiller 2008), although anthropogenic kova´ and Chytry 2006). Within each plot, the disturbances include clearing, grazing, and fire diameters of all stems C 5 cm diameter were mea- (Franklin and Steadman 2013; Larkin et al. 2012; sured at 1.3 m height on the trunk (‘‘dbh’’). Basal area Miller 2007; Sullivan-Sealey et al. 2002). per tree was summed to yield species basal area. Trees

123 Plant Ecol were identified to the greatest level of taxonomic Scaling (NMDS) (Clarke 1993) was used to visualize resolution possible. A few individuals not in fruit or continuous variation in species composition among flower could not be identified to species level, and plots and the distinction among groups identified by were identified to genus or family level or left as clustering. Detrended correspondence analysis was run unknowns in our analyses (Appendix S2). Tree height in parallel, with the results examined for congruence was measured using an Impulse laser rangefinder (Økland 1996). This sequence of clustering and (Laser Technology Inc., Centennial, CO, US) based on ordination was carried out twice—first for the 153 the average height of five overstory trees, chosen FIA plots together with the 42 Bahamian plots in order evenly among all directions from plot center. Slope to analyze regional variation in forest composition, and angle and aspect were measured with a clinometer and then for the Bahamian plots alone in order to examine compass, and percent rock exposed in the plot was local gradients of composition within the Bahamian estimated visually. Plot locations were recorded with a Archipelago. Garmin Global Positioning System (GPSMAP 60CSx, Additionally, Canonical Correspondence Analysis Garmin International Inc., Olathe, KS). Plot elevation (CCA), a constrained ordination method (ter Braak and distance from the nearest road and coastline were 1987), was conducted for the 42 Bahamian plots in measured using Google Earth Pro (Google Inc., Menlo order to identify local scale compositional variation Park, CA). Plot and environmental data are provided explained by environmental variables. These variables in Supplemental Material Appendices S1 and S2 and described geographical location (latitude, longitude), archived in VegBank (www.vegbank.org). While FIA terrain (slope, aspect, percent rock), landscape (dis- plots differed from the Bahamian plots in plot size and tance from coast, distance from road), and stand (very slightly) height of dbh measurement, the multi- structure (average tree height). Statistical analyses variate analysis methods we used are based on relative were conducted using R (R Development Core Team species importance and are robust to plot-size differ- 2012); vegan (Oksanen et al. 2011) was used for ences far greater than those found in our study PERMANOVA, NMDS, DCA and CCA, and labdsv (Otypkova´ and Chytry 2006). (Roberts 2012) was used for Indicator Species analysis. Nomenclature follows Correll and Correll Data analysis (1982) for Bahamian species, with updates from Acevedo-Rodrı´guez and Strong (2012), and for FIA Analyses of species composition were based on data, the USDA, NRCS PLANTS database (2014). patterns of relative abundance described by species Importance Value (average of relative basal area and relative density per plot). Analyses were also conduct- Results ed using relative basal area for comparison. Clustering, based on Jaccard distance and average linkage (Peet Regional patterns and Roberts 2013), was calculated from the plot-by- species Importance Value matrix in order to detect Summaries of the regional plot data are given in subgroups of plots sharing distinct assemblages of Table 1. Over 75 % of FIA dry forest plots in Puerto species. Permutational Multivariate Analysis of Vari- Rico and USVI were found below 100 masl. Multivari- ance or PERMANOVA (Anderson 2001), a non- ate analyses that used relative basal area versus parametric test based on permuting observations to Importance Value yielded almost identical results and obtain a probability of difference among groups, was so only those based on Importance Value are reported used to evaluate the significance of compositional here. The combined FIA and Bahamian plots included differences among groups defined by clustering. Indi- 193 taxa. Based on clustering the regional data and cator Species analysis (Dufreˆne and Legendre 1997) retaining 13 clusters (PERMANOVA R = 0.073, was used to identify species that have strong asso- P = 0.001 based on 999 permutations), the largest ciation (high relative abundance within the group) and group comprised more than half ofall plots,including all fidelity (occurring in many plots within but few outside 42 Bahamian plots, almost half of the Puerto Rican plots the group) with groups defined by clustering. Uncon- (45 of 91), and one quarter (16 of 62) of the plots from strained ordination with Non-metric Multidimensional USVI (Group 1; Table 2). Indicator species (Indicator

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Table 1 Summary of dry forest plot data and forest structure (see Appendices S1 and S2), for trees C 5 cm diameter. See based on FIA plots from Puerto Rico and the U.S. Virgin text for description of plot layout in the respective datasets Islands (USVI), and our data from the Bahamian Archipelago Variable Puerto Rico USVI Bahamas

Number of plots 91 62 42 Number of species 107 63 70 Number of trees 1299 975 4140 Elevation range of plots (masl) 0–281 0–202 10–47 Tree height (m) 6.6 ± 2.1 (2.4–15.8) 6.8 ± 2.8 (0.9–20.1) 7.7 ± 2.0 (3.5–10.9) Stand density (trees ha-1) 1232 ± 886 (15–4288) 2185 ± 2784 (15–15 216) 3138 ± 948 (1305–4934) Basal area (m2 ha-1) 8.8 ± 9.8 (0.4–90.3) 12.5 ± 13.2 (0.2–74.2) 27.3 ± 9.7 (7–50) Density and basal area of FIA plots were calculated using the area of forested subplots and microplots. (Some FIA subplots fall on non-forested land owing to the systematic sampling scheme)

Species Value [ 0.48 and P B 0.2) for this group were based on 999 permutations). Two large clusters accom- Bursera simaruba and Metopium toxiferum, with Coc- modated most plots from Abaco or Eleuthera. Group B1 coloba diversifolia also abundant (Table 2). Although included 13 of 20 plots from Abaco, whereas Group B3 the NMDS plot suggests some differences in dry forest contained 17 of 22 plots from Eleuthera (and 2 from composition among the Bahamian Archipelago, Puerto Abaco). The six additional ‘‘clusters’’ each included only Rico and USVI (Fig. 2), the plots in this large group are one to three plots from Abaco or Eleuthera with much more similar to each other than they are to other distinctive species composition (Fig. 3). dry forest plots in the FIA sample. Dry forest on both Abaco (Group B1) and Eleuthera Group 3 (55 plots from Puerto Rico and USVI; (Group B3) was dominated by Coccoloba diversifolia, Fig. 2) was dominated by Leucaena leucocephala and with Metopium toxiferum, Sideroxylon foetidissimum, Prosopis pallida, both non-native and naturalized Bursera simaruba, Exothea paniculata and Guapira disturbance indicators, and Group 12 (13 plots from obtusata also dominant or abundant (Table 3). The USVI) was dominated by Melicoccus bijugatus (an Group B1 plots were distinguished from those of Indicator Species), also non-native, naturalized and Group B3 by greater abundance of M. toxiferum and S. considered invasive (Table 2). Group 12 was also foetidissimum, and the presence of Swietenia ma- dominated by Trema micrantha, a small shade-intol- hagoni. Group B3 also had four minor species that erant tree, and included Tamarindus indica, a planted were absent from Group B1 (Table 3). and naturalized fruit tree. This group appears to Other groups identified by clustering included only represent forests with a legacy of human activity one to three plots of distinctive species composition, (clearing, agriculture). All other groups identified by such as three Abaco plots dominated by Eugenia clustering included less than 10 plots (five of them axillaris (Group B2; Fig. 3), a rocky hilltop Eleuthera with only one plot), and therefore are outliers of plot dominated by Guaiacum sanctum and Lysiloma unique species composition, or perhaps forest types latisilliquum (B5), an Eleuthera plot distinguished by that have not been adequately sampled. Notably, as the palm Pseudophoenix sargentii (B7), and another with Groups 3 and 12, 10 of these remaining 24 plots Eleuthera plot (B8) with a diverse set of indicator were dominated by non-native, potentially invasive species (Schaefferia frutescens, Stenostomum lucida, leguminous trees (Acacia farnesiana, Pithecellobium Erythroxylum areolatum, Guettarda krugii and Bour- dulce) that usually establish in thickets and near roads. reria succulenta). The first two axes of the constrained ordination, Local patterns CCA, carried out for the Bahamian data had eigenvalues of 0.3706 and 0.2406 (variance explained 0.0915 Average linkage clustering of the local data (42 Bahami- and 0.0594), suggesting that the first axis represents a an plots), retaining eight clusters, resulted in significantly fairly strong gradient and the second axis a somewhat different groups (PERMANOVA R = 0.102, P = 0.001 weaker one. For comparison, unconstrained

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Table 2 Summary of dominant species with average Impor- Bahamian Archipelago (Bah)—42 plots, Puerto Rico (PR)—91 tance Value (IV) [ 0.05, and characteristic species with plots—and the U.S Virgin Islands (USVI)—62 plots. Only Indicator Species Value (ISV) [ 0.48 (and P B 0.2), for shown are three groups with [ 10 plots, but these groups groups of plots resulting from clustering 195 plots from the include 171 (88 %) of all plots analyzed Species (Family) Average IV ISV P

Group 1 103 plots: Bah 42; PR 45; USVI 16 Bursera simaruba (Burseraceae) 0.301 0.7574 0.001 Coccoloba diversifolia (Polygonaceae) 0.098 Metopium toxiferum (Anacardiaceae) 0.076 0.4854 0.186 Group 3 55 plots: PR 28; USVI 27 Leucaena leucocephala (Fabaceae) (I)a 0.208 Prosopis pallida (Fabaceae)a 0.156 Bourreria succulenta (Boraginaceae) 0.092 Pisonia subcordata (Nyctaginaceae) 0.073 Swietenia mahagoni (Meliaceae)b 0.051 Group 12 13 plots USVI Melicoccus bijugatus (Sapindaceae) (I)a 0.309 0.5847 0.181 Trema micrantha (Cannabaceae) 0.246 Eugenia rhombea (Myrtaceae) 0.077 Tamarindus indica (Fabaceae)a 0.064 Bourreria succulenta (Boraginaceae) 0.057 a Non-native, naturalized species in the Caribbean; I, species considered invasive in Puerto Rico and USVI (Rojas-Sandoval and Acevedo-Rodrı´guez 2015) b Non-native and naturalized in Puerto Rico ordination, DCA, had eigenvalues of 0.4762 and instead we found remarkable similarities in native- 0.2992. CCA axes 1 and 2 were positively correlated dominated Caribbean dry forest at the regional level with latitude, distance from roads, and tree height, and when data from the Bahamian Archipelago, Puerto negatively correlated with longitude, elevation, and Rico, and USVI were analyzed. Most plots fell into a slope (Table 4). Rockiness was negatively correlated single region-wide dry forest community type with CCA1 and positively with CCA2. Group B1 plots dominated by native species Bursera simaruba and were at higher latitude and more negative longitude Metopium toxiferum. A geographically structured (north and west, because we used the convention of compositional difference is the dominance of Coc- negative longitude values west of Greenwich), at coloba diversifolia in the Bahamian Archipelago and greater distances from roads, and with taller canopies its absence from most FIA dry forest plots in Puerto (Figs. 3 and 4). Group B3 sites were at somewhat Rico and the USVI. Interestingly, C. diversifolia was higher elevations, on steeper, rockier slopes, and at recorded frequently in other FIA plots from Puerto greater longitude and lower latitude (further east and Rico and the USVI classiﬁed as subtropical moist south; Appendix S1). forest, whereas C. microstachya was recorded frequently in FIA dry forest plots, suggesting niche displacement for these congeners (Silvertown 2004). Discussion Comparison with published descriptions of dry forest communities provides additional evidence for Regional patterns region-wide compositional similarities (Table 5). Greater Antillean dry forests, especially those on We expected to see variation in composition among limestone substrates, have similar species composition the island groups analyzed in this study because of to our Group 1 forests, although when found at higher differences in island size, elevation, and substrate; elevations those forests tend to be more diverse and

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Bahamas and Lugo 2006), was dominated by Coccoloba Puerto Rico diversifolia, and shared other species with our Group USVI Group 3 1. Of the limestone dry forests surveyed in Jamaica by Kelly et al. (1988), their Round Hill sites at *300 masl (annual precipitation *1200 mm) had different dominants, but nonetheless shared many species and genera with the plots analyzed in our study

NMDS2 (Table 5). A mature dry forest on limestone was described by Roth (1999) in the Cordillera Central of the Dominican Republic (Hispaniola) at *300–

Group 1 500 masl (1000 mm strongly seasonal annual rainfall). The most abundant species featured those that -0.4 -0.2 0.0 0.2 0.4 dominated our Group 1 (Bursera simaruba, Coc- coloba diversifolia and so forth) as well as some not -0.4 -0.2 0.0 0.2 0.4 0.6 found in the Bahamian Archipelago (Oxandra lanceo- NMDS1 lata; Table 5). In Cuba, seasonal evergreen forest on Fig. 2 Ordination diagram showing the distribution of 153 limestone-derived soils is similar in composition to plots from the Bahamian Archipelago (Bah), Puerto Rico (PR), these other dry forests and to our Group 1, although it and U.S. Virgin Islands (VI) on first two dimensions (NMDS1, lacks Coccoloba diversifolia; instead it is co-dominat- NMDS2) derived from unconstrained Non-metric Multidimen- sional Scaling (NDMS; stress = 27.0 %). Symbols indicate the ed by Sideroxylon foetidissimum, S. salicifolium, island groups. Ellipses indicate the 95 % confidence ellipse for Roystonea regia, and Bursera simaruba (Table 5), the two largest groups (Groups 1 and 3; see Table 1). Note the with many other associated tree species (Areces- main group (Group 1) with plots from all three regions (see Mallea et al. 1999; Borhidi 1991). The absence of Table 1) Swietenia mahagoni from Greater Antillean dry forests on islands where it is native may be a result of historic selective logging (Rodan et al. 1992). S. mahagoni is introduced and naturalized in Puerto Rico and the USVI. Other studies have noted floristic similarities among dry forests on limestone in the Greater Antilles (Trejo- Torres and Ackerman 2002). Gillespie (2005) also recorded many of the important species identified in

NMDS2 our analyses in subtropical dry forest sites on mainland

Grp B1 southern Florida and the Florida Keys (Table 5). Taken Grp B2 together, our quantitative analysis and comparison Grp B3 Grp B4 with published studies suggest a region-wide dry forest Grp B5 Grp B6 type for south Florida, the Bahamian Archipelago and Grp B7 the Greater Antilles. Comparable plot data from Grp B8

-0.4 -0.2 0.0 0.2 0.4 throughout the region, as well as from dry forest in -0.2 0.0 0.2 0.4 the Lesser Antilles, Central America and northern NMDS1 South America, would allow a more detailed quanti- Fig. 3 Ordination diagram showing the distribution of 42 plots tative classification of variation within the region to be from Abaco and Eleuthera on first two dimensions (NMDS1, used as a basis for conservation planning. NMDS2) based on unconstrained Non-metric Multidimensional Notably, the other two dry forest groups identified Scaling (NDMS; stress = 18.8 %). Symbols indicate the 8 groups (Grp B#) of plots defined by clustering (see text) in our region-wide analysis were dominated by non- native species. An extensive group found in Puerto taller, with greater basal area. A mature dry forest at Rico and the USVI is dominated by non-native *100 masl on limestone in Puerto Rico, in the well- Leucaena leucocephala and Prosopis pallida, and studied Guańica Biosphere Reserve (Molina Coloń one found only in the USVI is dominated by non-

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Table 3 Average basal Species (Family) Group B1 BA (m2 ha-1) Group B3 BA (m2 ha-1) area (BA; m2 ha-1) for indicator and dominant Coccoloba diversifolia (Polygonaceae) 9.47 5.19 species in the two large Metopium toxiferum (Anacardiaceae) 8.40 0.93 groups of Bahamian plots deﬁned by clustering: B1 Sideroxylon foetidissimum (Sapotaceae) 7.21 0.34 (13 plots from Abaco) and Bursera simaruba (Burseraceae) 2.80 7.01 B3 (19 plots—17 from Exothea paniculata (Sapindaceae) 2.54 1.15 Eleuthera, 2 from Abaco) Swietenia mahagoni (Meliaceae) 2.52 – Guapira obtusata (Nyctaginaceae) 2.28 1.53 Eugenia confusa (Myrtaceae) 2.08 0.03 Ficus aurea (Moraceae) 2.54 1.15 Sideroxylon salicifolium (Sapotaceae) 0.97 0.88 Hypelate trifoliata (Sapindaceae) 0.81 0.01 Eugenia axillaris (Myrtaceae) 0.53 0.25 Nectandra coriacea (Lauraceae) 0.46 0.06 Acacia choriophylla (Fabaceae) 0.41 0.55 Amyris elemifera (Rutaceae) 0.36 0.14 Piscidia piscipula (Fabaceae) 0.35 0.58 Guapira discolor (Nyctaginaceae) 0.21 0.29 Tabebuia bahamensis (Bignoniaceae) 0.17 0.08 Bourreria succulenta (Boraginaceae) 0.14 0.04 Thouinia discolor (Sapindaceae) 0.09 0.02 Reynosia septentrionalis (Rhamnaceae) 0.07 0.45 Basal Area values for Eugenia foetida (Myrtaceae) 0.01 0.04 Indicator Species (P B 0.2) Gymnanthes lucida (Euphorbiaceae) – 0.09 shown in italics, and for dominant species (Basal Pithecellobium keyense (Fabaceae) – 0.03 Area [ 5m2 ha-1) in bold. Guettarda kruggii (Rubiaceae) – 0.02 Other species included are Guaiacum sanctum (Zygophyllaceae) – 0.02 those mentioned in the text

Table 4 Correlation of CCA1 CCA2 r2 Pr([ r) environmental site variables with the first two Latitude (Lat); decimal degrees 0.83331 0.55281 0.5714 0.001 constrained ordination axes Longitude (Lon); decimal degrees -0.70789 -0.70633 0.6355 0.001 (CCA1, CCA2) based on Canonical Correspondence Elevation; masl -0.81107 -0.58495 0.1217 0.076 Analysis (CCA) of the 42 Percent rock (rock); percent ground cover -0.92501 0.37995 0.5930 0.001 Bahamian plots Slope; degrees -0.50202 -0.86485 0.1453 0.070 Distance from coast; meters -0.50041 -0.86579 0.0145 0.721 Distance from road; meters 0.70285 0.71134 0.3246 0.018 Average tree height (TreeHt); meters 0.88145 0.47227 0.4872 0.001 native Melicoccus bijugatus (Table 1). Tropical dry USVI but not on Bahamian islands. These two groups, forests are particularly susceptible to invasion by non- as well as many of the remaining dry forest plots that native species owing to extensive human disturbance did not cluster into large groups, were dominated by including land clearing and fire (Fine 2002). These naturalized species considered to be invasive (Rojas- novel ecosystems (Lugo and Helmer 2004; Martinuzzi Sandoval and Acevedo-Rodrı´guez 2015) and consti- et al. 2013; Molina Coloń et al. 2011) with a legacy of tute post-disturbance communities (Lugo et al. 2006). human activity are widespread in Puerto Rico and the This finding reinforced concerns about the

123 Plant Ecol

composition may also be related to disturbance history 4 (Larkin et al. 2012), although dry forest on both islands is subjected to similar natural and human disturbance regimes, and plots were located in tall, 2 Lat closed canopy forests, e.g. mid- to late-successional. TreeHt Published studies on dry forest composition in the Rock Bahamian Archipelago can be used to more fully

CCA2 describe local patterns of variation. Previous studies,

Grp B1 focused on single islands, described forest communi- Grp B2 ties with varying combinations of the same dominant Grp B3 Grp B4 species found in this study. On North Andros (Great -2 0 Lon Grp B5 Bahama Bank), Smith and Vankat (1992) quantita- Grp B6 Grp B7 tively defined three inland dry forest communities that, Grp B8 as in our findings, were distinguished only by relative -6 -4 -2 0 2 abundance of the dominant species C. diversifolia, M. CCA1 toxiferum, Exothea paniculata and/or B. simaruba. Fig. 4 Ordination diagram showing the distribution of 42 plots They found the compositional gradient to be weakly from Abaco and Eleuthera on first two dimensions (CCA1, related to elevation (while ours was not). E. H. Freid CCA2) based on constrained ordination using Canonical (unpublished data) sampled dry forest at Clifton Correspondence Analysis (CCA). Symbols indicate the 8 groups National Heritage Park on New Providence. The forest (Grp B#) of plots defined by clustering (see text). 95 % confidence ellipses are shown for Grp B1 and Grp B3. Vectors there was dominated by Coccoloba sp. and Nectandra show the correlation of environmental variables with con- coriacea, with greater abundance of B. simaruba strained ordination axes, vector length indication strength of nearer the coast. Subdominants included Eugenia correlation (shown for variables with P = 0.001; defined in axillaris, E. foetida, Tabebuia bahamensis and S. Table 3) foetidissimum. This forest is somewhat more similar to Group B1 (Abaco) than to B3 (mostly Eleuthera) in conservation status of dry forest (Janzen 1988; Gille- spite of being on the Great Bahama Bank. spie et al. 2012). Although several geographical and environmental variables measured for our Bahamian plots were Local patterns correlated with species composition, only rockiness correlated with composition independently of the At the local scale on Bahamian islands, we found some geographical factors that differed systematically be- evidence for the expected variation in species com- tween the two islands. Distance from the coast was not position correlated with the north to south gradient of correlated with species composition, but this may have increasing aridity and warmth. Some species recorded been a result of the sampling focused on the inland in plots on Eleuthera but not Abaco (Guaiacum ‘‘blackland’’ form of dry forest. sanctum, Pithecellobium keyense) are abundant in the dry forests of the warmer, more arid southern Vegetation classification for conservation Bahamian islands (JF, EHF and DWS, personal planning observation). Most species characterizing these groups were found on both islands, however, and the These striking similarities and overlap in dominant subtle differences between them were mainly in species composition in our regional-scale analysis patterns of species dominance (Table 2). The taller suggest that a more comprehensive, quantitative re- canopies recorded in plots on Abaco may indicate that gion-wide treatment of Caribbean dry forest is needed. dry forest, restricted to limestone ridges on this pine In addition to geographical variation in community island, occurs in protected and productive sites, while composition, temporal dynamics (succession following on Eleuthera dry forest is the dominant terrestrial plant disturbance) also need to be better understood (Franklin community and was sampled across a wide range of 2007; Larkin et al. 2012). The Nature Conservancy’s site conditions. Differences in canopy height and Guide to Caribbean Vegetation Types (Areces-Mallea

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Table 5 Overview of Caribbean dry forest composition based on this study and published literature Species (Family) BAa PR, VIa PRb JMc DRd CUe FLf

Coccoloba diversifolia (Polygonaceae) X (smf) X X X X Bursera simaruba (Burseraceae) X X X X X X X Metopium toxiferum (Anacardiaceae) X X (sp) X Sideroxylon foetidissimum (Sapotaceae) X X X X Exothea paniculata (Sapindaceae) X (smf) X X X Swietenia mahagoni (Meliaceae) X (x) (smf) X Guapira obtusata (Nyctaginaceae) X X X (sp) Guapira fragrans (Nyctaginaceae) X X X Guapira discolor (Nyctaginaceae) X X Eugenia sp. (Myrtaceae) X X X X X X Sideroxylon salicifolium (Sapotaceae) X X X X X Hypelate trifoliata (Sapindaceae) X X X X (x) Nectandra coriacea (Lauraceae) X (smf) X X X Amyris elemifera (Rutaceae) X X X (sp) X Piscidia piscipula (Fabaceae) X X X Bucida buceras (Combretaceae) X X X Gymnanthes lucida (Euphorbiaceae) X X X X X X Erythroxylum rotundifolium (Erythroxylaceae) X X (sp) Guettarda krugii (Rubiaceae) X X X (sp) Erythroxylum areolatum (Erythroxylaceae) X X X Erythroxylum confusum (Erythroxylaceae) X X Krugiodendron ferreum (Rhamnaceae) (x) X (smf) X X X X Acacia choriophylla (Fabaceae) X Acacia scleroxyla (Fabaceae) X Oxandra lanceolata (Annonaceae) X Roystonea regia (Arecaceae) (x) X (x) a This study b Molina Coloń and Lugo (2006) c Kelly et al. (1988) Round Hill site d Roth (1999) e Borhidi (1991) in Areces-Mallea et al. (1999) Sideroxylon foetidissimum—Sideroxylon salicifolium—Roystonea regia Forest (includes many other species not listed here) f Gillespie (2005) Small (x) indicates species is a minor component of dry forest; (smf) indicates species is commonly recorded in FIA plots classified as subtropical moist forest; (sp) indicates another species of this genus was important Species indicated by X were noted to be abundant, dominant or characteristic of dry forest in our study or in previously published studies of from the Bahamian Archipelago (BA), Puerto Rico (PR), U.S. Virgin Islands (VI) Jamaica (JM), Cuba (CU), Hispaniola (DR, Dominican Republic), or Florida, US (FL). Species in same order as in Table 3 (in order of their importance in Bahamian forests) with additional species included with congeners or at the end et al. 1999) was developed to establish regional Bahamian Archipelago (Reynosia septentrionalis— conservation priorities. While this classification is Sideroxylon americanum—Pithecellobium keyense— comprehensive in its treatment of rain forest, it did not Jacquinia keyensis Forest), synonymized with ‘‘white- describe the Greater Antillean dry forests of Hispaniola land’’ (e.g., coastal) coppice (Correll and Correll 1982); and Jamaica summarized in Table 5. Furthermore, it no inland Bahamian dry forest was described by Areces- included only a single dry forest Alliance for the Mallea et al. (1999).

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Our study begins to address this lack of quantitative Areces-Mallea AE, Weakley AS, Li X, Sayre RG, Parrish JD, understanding of Caribbean dry forest communities, Tipton CV, Boucher T (1999) A guide to Caribbean vegetation types: preliminary classification system and although we note that data are lacking from the descriptions. Nature Conservancy, Washington southeastern half of the Bahamian Archipelago, a gap Borhidi A (1991) Phytogeography and vegetation ecology of that we hope to fill. More comprehensive analysis of Cuba. Akademiai Kiado, Budapest dry forest associations at both regional and local scales Brandeis TJ, Turner JA (2013a) Puerto Rico’s forests, 2009. Resour Bull. SRS-191. U.S. Department of Agriculture could be accomplished by collaboration among Forest Service, Southern Research Station, Asheville vegetation scientists working in the Caribbean (for an Brandeis TJ, Turner JA (2013b) U.S. Virgin Islands’ forests, example, see Franklin et al. 2013). Such analyses could 2009. Resour Bull. SRS-196. U.S. Department of Agri- help define Caribbean dry forest types at the mid-level culture Forest Service, Southern Research Station, Asheville of established vegetation classification hierarchies; Brandeis TJ, Delaney M, Parresol BR, Royer L (2006) Devel- e.g., our regional Group 1 may contribute to vegetation opment of equations for predicting Puerto Rican sub- class descriptions at the Macrogroup level of the U.S. tropical dry forest biomass and volume. Forest Ecol National Vegetation Classification System (Franklin Manage 233:133–142 Butterfield BR, Csuti B, Scott JM (1994) Modeling vertebrate et al. 2012). Local analyses will help to define classes at distribution for gap analysis. In: Miller RI (ed) Mapping the lower levels in a hierarchy, such as Alliances or diversity of nature. Chapman and Hall, London, pp 53–68 Associations (e.g., our groups B1 and B3), that form Carey E, Gape L, Manco BN, Hepburn D, Smith R, Knowles L, the basis of conservation planning. A regionally Knowles D, Daniels M, Vincent M, Freid E, Jestrow B, Griffith MP, Calonje M, Meerow A, Stevenson D, Fran- coherent classification is needed to guide conservation cisco-Ortega J (2014) Plant conservation challenges in the management of this widespread yet threatened forest Bahama Archipelago. Bot Rev 80:265–282 community in an era of global change. Clarke KR (1993) Non-parametric multivariate analyses of changes in community structure. J Ecol 18:117–143 Acknowledgments This research was sponsored by the U.S. Correll DS, Correll HB (1982) Flora of the Bahama archipelago. National Science Foundation (Grant BCS-1118340 to JF and A.R.G. Gantner Verlag K.-G., Fl-9490 Vaduz BCS-1118369 to DWS) and Arizona State University (JF). For Currie D, Wunderle JM, Ewert DN, Anderson MR, Davis A, permissions and cooperation on Abaco and Eleuthera, we thank Turner J (2005a) Habitat distribution of birds wintering in The Bahamas National Trust, The National Museum of The Central Andros, The Bahamas: implications for manage- Bahamas (Antiquities, Monuments and Museums Corporation), ment. Caribb J Sci 41:75–87 Friends of the Environment, the Leon Levy Native Plant Preserve, Currie D, Wunderle JM, Ewert DN, Davis A, McKenzie Z C. Adair, M. and N. Albury, E. Carey, M. Daniels, A. and S. (2005b) Winter avian distribution and relative abundance Knowles, D. Knowles, P. Maillis, O. Patterson, K. Tinker, and K. in six terrestrial habitats on southern Eleuthera, The Ba- Williams. B. Trapido-Lurie produced Fig. 1. The USDA Forest hamas. Caribb J Sci 41:88–100 Service (USFS) FIA program for Puerto Rico and the U.S. Virgin Dengler J, Jansen F, Glockler F, Peet RK, De Caceres M, Chytry Islands is jointly funded by USFS-Southern Research Station-FIA M, Ewald J, Oldeland J, Lopez-Gonzalez G, Finckh M, and the USFS International Institute of Tropical Forestry. Mucina L, Rodwell JS, Schaminee JHJ, Spencer N (2011) The Global Index of Vegetation Plot Databases (GIVD): a Open Access This article is distributed under the terms of the new resource for vegetation science. J Veg Sci 22:582–597 Creative Commons Attribution 4.0 International License (http:// Dufreˆne M, Legendre P (1997) Species assemblages and indi- creativecommons.org/licenses/by/4.0/), which permits unrest- cator species: the need for a flexible asymmetrical ap- ricted use, distribution, and reproduction in any medium, pro- proach. 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123 Supporting*Information*to*the*paper*Franklin,*J.*et*al.*Regional*variation*in*Caribbean*dry*forest*composition.*Plant*Ecology Appendix*S1.*Plot*by*species*basal*area*data*and*Plot*by*environmental*variables*and*other*plot*attributes*for*42*Bahamian*plots. Sum*of*Basal*Area*(m^2)*for*each*species*in*each*plot,*with*three*versions*of*plot*numbers*per*plot*(the*following*three*rows) Plots 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 A01 A02 A03 A04 A05 A06 A07 A08 A09 A10 A11 A12 A13 A14 A15 A16 A17 A18 A19 A20 E01 E02 E03 E04 E05 E06 E07 E08 E09 E10 E11 E12 E13 E14 E15 E16 E17 E18 E19 E20 E21 E22 Accepted_taxon_name Ab8MC02DBHAb8MC04DBHAb8MC06DBHAb8MC08DBHAbCHER03DBHAbCHER05DBHAbCHER07DBHAbGPDBH02 AbGPDBH04 AbIH01DBH AbIH04DBH AbIH05DBH AbWCDBH02AbWCDBH04AbWCDBH05AbWCDBH08AbWCDBH10AbWCDBH11AbWCDBH13AbWCDBH16El_GHDBH21 El_GHDBH22 El_GPDBH19 El_GPDBH20 El_GTDBH17 El_GTDBH18 El_JCDBH15 El_JCDBH16 El_LHDBH13 El_LHDBH14 El_LPDBH11 El_LPDBH12 El_PCBHDBH07El_PCDBH08 El_PondDBH05El_PondDBH06El_RSCOPDBH01El_RSDBH02 El_WBDBH03El_WBDBH04El_WindDBH09El_WindDBH10 Acacia*choriophylla 0.03390483 0.00761679 0.00363168 0 0 0 0 0.01473249 0.03038703 0.01793063 0 0.01600954 0 0.00395919 0 0 0 0 0.0057821 0.02411563 0.00212372 0.04038435 0.03159497 0.03366135 0.05058274 0.03106954 0.03866276 0.0065455 0 0.00826631 0.02470232 0.02497171 0.01866105 0.03811456 0.01515346 0.01614542 0 0.01232524 0.02218984 0 0 0.01627736 Amyris*elemifera 0 0 0.00220618 0 0 0.00204282 0.01405469 0 0 0 0.00363168 0 0.02100938 0.00196349 0.02626212 0.00246301 0.00650388 0.00477836 0 0.01530033 0 0.04455953 0 0 0.00342119 0 0.00451525 0 0 0 0.00311724 0.04658271 0 0 0 0 0 0 0.05801888 0.00674028 0.02260531 0.00490167 Stenostomum*lucidum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.02444471 0 0 0 0 0 0 0 0 0 0 0 0 Bourreria*succulenta 0.02444628 0 0.00418538 0 0 0.01205899 0 0 0 0.01056831 0 0 0 0 0 0 0 0 0 0 0 0.00693977 0 0 0 0 0 0 0.00395919 0.08115041 0 0.00624941 0 0 0 0 0 0 0.01128459 0.0058088 0.00384845 0 Bursera*simaruba 0 0.04697541 0.04687409 0.04861061 0.26997803 0.24158042 0.13600022 0.1689932 0.31071032 0.00467547 0.00515299 0.14838122 0.13368015 0.02144449 0 0.13860302 0.07177355 0.15242759 0.0949899 0 0.84757506 0.24277893 0.09126398 0.10329391 0.16134421 0.31996309 0.19133618 0.09383301 0.02198013 0.33926266 0.0364825 0.04596617 0.28578259 0.33894929 0.05628708 0.00501791 0.11987286 0.22111846 0.00974678 0.04786134 0.43203853 0.19580352 Canella*winterana 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00568 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Coccoloba*cf*kruggii 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00650388 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Coccoloba*diversifolia 0.1681057 0.45779485 0.29987105 0.58117137 0.03517874 0.11914951 0 0.24566056 0.15671508 0.09492 0.19506368 0.20969642 0.18617848 0.07295243 0.13163733 0.2228424 0.28516291 0.13959184 0.22891274 0.10668604 0.41204466 0.41894281 0.18864463 0.32208209 0.16418499 0.27332383 0.09265413 0.04215071 0 0 0.13067679 0.11654671 0.15461492 0.03579763 0.05587789 0.01100892 0.14642873 0.20029521 0.09745448 0.13301256 0.10630041 0.20776513 Coccoloba*swartzii 0 0 0 0.02164241 0 0 0 0.05718322 0.01015755 0 0.00554177 0.00212372 0 0.01246897 0 0 0 0 0 0.00204282 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Coccothrinax*argentata 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.0066476 0 0 0 0.01192548 0.04732099 0 0 0.02269799 0 0 0 DBH03_unkB 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00754296 0 0 0 DBH03_unkC 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00301907 0 0 0 Drypetes*diversifolia 0 0 0 0 0 0 0 0 0 0 0 0 0.01038688 0.03998851 0.01177468 0.02354465 0 0 0 0 0 0 0 0 0 0 0 0.05468801 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Drypetes*lateriflora 0 0 0.14214595 0 0 0.00680547 0 0 0 0 0 0.00606327 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Erithalis*fruiticosa 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00273397 0 0 0 0 0 0 0 0 0 0 0 0 0 Erythroxylum*areolatum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.01800995 0 0 0 0 0 0 0 0 0 0 0 0 Eugenia*axillaris 0.00255176 0.03421506 0.13858889 0.02494423 0.02842668 0.06801778 0.03054254 0.00458201 0 0 0 0 0 0 0 0 0 0.00311724 0.00229022 0.00237583 0.02102902 0 0 0.00301907 0.00237583 0 0 0 0 0 0 0 0 0 0 0 0.01479139 0.00622113 0 0.00212372 0.0076749 0 Eugenia*confusa 0 0 0 0 0 0.00264208 0 0.02973201 0.00693977 0 0 0 0 0.02435832 0.34616238 0.07620633 0.35905232 0 0 0.01588938 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00352565 0.00586299 Eugenia*foetida 0 0 0 0 0.00502419 0 0.0049323 0 0 0 0 0 0 0 0.00458201 0 0 0 0 0 0 0 0.00196349 0 0 0.00363168 0 0 0.00594467 0.01344993 0 0 0 0 0 0 0 0.00212372 0 0 0.00738981 0.00363168 Eugenia*rhombea 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.004249 0 0 0 0.00537605 0.00453646 Exostema*caribaeum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00292246 0.03009251 0 0 0 0 0 0 0 0 0 0 0 0 Exothea*paniculata 0.05662716 0.02369073 0.05330886 0.08515515 0.12637203 0.11604248 0.05391283 0.02197307 0 0.06108979 0.13903578 0.13469803 0.02924899 0.02907463 0.11682709 0.04297145 0.03887639 0.11384337 0.02192359 0.00623606 0.14833724 0.02260138 0 0 0 0 0 0.01693474 0 0 0 0.01129951 0 0.00255176 0 0 0.1076136 0.08264738 0.00273397 0.05940118 0.01165687 0.05318948 Ficus*americana 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.02216706 0 0 0.01998837 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Ficus*aurea 0 0.09079195 0 0 0 0.03255394 0.31016604 0 0 0 0 0.07720457 0.04058542 0 0 0 0 0.16786851 0.1214256 0 0 0 0 0 0.09002383 0.16426981 0.0174696 0.05888047 0 0 0 0 0 0.75122014 0 0 0 0 0 0 0 0 Ficus*citrifolia 0 0 0.00622113 0 0 0 0 0 0.00708821 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00196349 0 0 0 0 0 0 0 0 0 0 0 0 0 0.02124814 Guaiacum*sanctum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.10893699 0.14018481 0 0 0 0 0 0 0 0 0 0 0 0.00985203 Guapira*discolor 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.02507382 0 0.06026277 0.00490167 0.04472289 0.0092025 0 0.02086801 0.03172221 0.03428417 0.0064002 0.00738666 0.04698641 0 0 0 0 0 0 0 0 0 0 0.01817253 0 Guapira*obtusata 0.00849486 0 0.00255176 0 0.01593179 0.07951757 0 0 0.02573041 0.01288759 0.06992394 0 0.08830224 0.09703115 0.13146455 0.02934559 0.13226172 0.09351335 0.10009813 0.04260624 0 0.12466772 0 0.01527912 0.02494501 0.00554177 0 0 0 0 0.14344971 0.17061426 0 0.00738981 0.05852232 0.01525321 0.18412152 0.20756014 0.01946137 0.08750742 0.09782754 0.07111853 Guettarda*kruggii 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.01162624 0 0 0 0.00273397 0 0 0 0 0 0 0 0 0 0 0 0 Guettarda*scabra 0 0 0 0 0 0 0 0.02932596 0 0 0 0 0 0.0127148 0 0 0 0 0 0.0100955 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Guettarda*sp. 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00311724 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Gymnanthes*lucida* 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.03181096 0.01237315 0 0.00196349 0.00237583 0 0 0 0 0 0.00466919 0 0 0 0 0 0.00246301 0 Hypelate*trifoliata 0 0 0 0 0 0 0.03110253 0 0 0 0 0 0.10861655 0 0 0 0.09025867 0 0.00608212 0.12560155 0 0 0 0 0 0 0 0 0 0 0 0.00508074 0 0 0 0 0 0.00759087 0 0 0 0 Jacquinia*keyensis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00384845 0 0 0 0 0 0 0 0 0 0 0 Krugiodendron*ferreum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00636172 0 0 0.00418538 0.00212372 0.02686452 0 0 0 0.01564983 0 0 0 0 0 0 0 0 0 0 0 0 0 0.0033183 0.03197275 0.02811645 Leucaena*leucocephala 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00229022 0 0 0 0.00424743 0 0 0 0 0 0 0 0 0.02233121 0 0 0 0 0 0 0 0 Chionanthus*sp. 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00237583 0 0 0 0 0 0 0 0 0 0 Lysoloma*latisilliquum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.07943432 0.01651298 0 0 0 0 0 0 0 0 0 0 0 0 Lysoloma*sabicu 0 0 0 0 0 0 0 0.02296581 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.0138929 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Manilkara*bahamensis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.01101834 0 0 0 0 0 0 0 0 0 0 Manilkara*jaimiqui*subsp.*emarginata 0 0 0.00255176 0 0 0 0 0 0 0 0.00594467 0 0.03279506 0 0 0 0 0 0.07128739 0.00738038 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00220618 0 0 0 0 0 0 0 Manilkara*zapota 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.02488453 0.00477836 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Sideroxylon*foetidissimum 0.16533167 0.22171693 0.06637316 0.30507745 0.09175092 0.07728233 0.18230333 0.02918537 0.01122647 0.29408581 0.02188824 0.27912477 0.26773101 0.08357415 0.14540928 0.11023839 0.11810336 0.18017176 0.27052153 0.17450354 0 0 0 0 0 0 0 0 0 0 0.00264208 0 0 0.02490967 0 0 0.02257704 0.00352565 0.11303519 0.09990806 0 0 Maytenus*buxifolia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.01039159 0 0 0 0 0 0 0 0 0 0 0 0 0 Metopium*toxiferum 0.19764764 0.11238253 0.20480968 0.03905703 0.01056831 0.01168986 0 0.18284604 0.16973225 0.50211012 0.60255382 0.18069562 0.0864919 0.27637431 0 0.04987588 0.03976075 0.08603558 0.09594966 0.11785204 0 0 0.01243206 0 0.03297334 0.01767459 0.11930581 0.01694888 0 0 0.01138591 0.11523117 0.00613317 0.00950331 0.0306305 0.10783508 0.02758709 0.02768526 0.11392662 0.43431696 0.00981747 0.06285144 Myricanthes*fragrans 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00909647 0.00801577 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Nectandra*coriacea 0 0.0071856 0 0 0 0 0 0.02480128 0.03331578 0 0 0 0.02212072 0.01072303 0.01003895 0.04572035 0.01313892 0.02684096 0.00985752 0.01779554 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 PCBHDBH07_UnkB 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00465662 0.00301907 0 0 0 0 0 0 0 0 Persea*palustrus 0.00662012 0 0 0 0 0 0 0 0 0 0 0.00686516 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Picramnia*pentandra 0 0 0 0 0 0 0.00441865 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.0067929 0 0 0 0 0 0 0 0 Pilosocereus*millspaughii 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.08647227 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Piscidia*piscipula 0 0 0 0 0.1192791 0.09406706 0.04178001 0 0 0 0.0232352 0.05139406 0 0.06721589 0 0 0 0 0 0 0.04240597 0.01611086 0 0.03556201 0 0 0 0 0 0.09729818 0 0 0 0 0 0 0.01097279 0.00465662 0 0 0.00196349 0.02268856 Pithecellobium*keyense 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00408721 0 0.00196349 0 0.00798042 0.00416968 0 0.00400631 0.00466605 0 0 0 0 0 0 0 0 0 0 0.00237583 0 Prunus*myrtifolia 0 0 0 0 0 0.00212372 0 0 0 0 0 0 0 0.00430084 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Pseudophoenix*sargentii 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.32266957 0 0 0 0 0 0 0 0 0 0 Reynosia*septentrionalis 0 0 0.00292246 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.03801324 0.06849137 0 0.00220618 0.00636172 0 0 0.03039645 0 0.25870208 0.01651298 0 0 0.02511544 0.01851732 0 0.11806409 0.00883651 0 0.00866686 0 Roystonea*regia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.02516806 0 0 0 Schaefferia*frutescens 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.01209905 0 0 0 0 0 0 0 0 0 0 0 0 Sideroxylon*salicifolium 0 0 0 0.13202689 0.12695636 0.0944951 0.10682898 0.0040715 0 0.03040352 0.04802313 0.00770082 0.01038688 0.05330493 0.00220618 0.05212526 0 0 0.03486144 0.00237583 0.08871065 0 0.00264208 0.01796204 0 0 0 0.00796236 0 0 0 0.11399416 0 0 0.01082356 0 0 0.026043 0.00363168 0.01431387 0.03935077 0.11227493 Simarouba*glauca 0.0033183 0 0 0 0 0 0 0 0 0.00430084 0.00282743 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.02373943 0 0 0 0 0 0 0 0 0 Swietenia*mahagoni 0.16503401 0.2041413 0.14039452 0 0 0 0 0.09997953 0 0 0 0.02707187 0 0.02138245 0 0.18985728 0 0.1832246 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Tabebuia*bahamensis 0 0 0.00863623 0 0 0.01199223 0 0.00465662 0.01736828 0 0.01913857 0 0 0 0 0 0.01724733 0 0 0.00636172 0.00321699 0 0 0 0.00908941 0.00301907 0 0 0 0.00220618 0.03145438 0 0.00212372 0.00196349 0.00196349 0.00301907 0 0 0.00204282 0.00876582 0 0 Thouinia*discolor 0 0.01197339 0.01474898 0 0 0 0 0.00246301 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00255176 0.00441393 0 0 0.00301907 0.00519698 0.00384845 0 0 0 0 0 0 0.00624077 0 0.00282743 0 Thrinax*morrisii 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.0427916 0.0056745 0 Trema*lamarckiana 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00282743 0 0 0 0 0 0 0 0 Zanthoxylum*fagara 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00196349 0 0.0140539 0 0 0 0 0 0 0 0.00292246 0 0 0 0 0 0 0 0 Ziziphus*taylori 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00255176 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Zuelania*guidonia* 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00475166 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 zz_AbWCDBH04_UnkB 0 0 0 0 0 0 0 0 0 0 0 0 0.01020703 0.00717853 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 zz_CHER05_A 0 0 0 0 0.00968631 0.00596981 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 zz_CHER07_F 0 0 0 0 0 0 0.00282743 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 zz_CHER07_G 0 0 0 0 0 0 0.01056831 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 zz_RUBIACEAE 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00229022 0 0 0 0 zz_UnkLP 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.00395919 0 0 0 0 0 0 0 0 0 0

Environmental*variables*and*other*attributes*of*each*plot

PlotNo PlotID PlotName Island Lat*(decimal*degrees)Lon*(decimal*degrees)Elevation*(masl)Rickiness*(%) Slope*(degrees)Aspect*(degrees)Distance*from*Coast*(m)Distance*from*Road*(m)Average*Tree*Height*(m)Stems/plot(314m^2)BA*m2/plot(314m^2) 1 A01 Ab8MC02DBHAbaco 25.9722741 d77.197373 13 0 0 na 727 275 9 86 0.83208231 2 A02 Ab8MC04DBHAbaco 25.9754574 d77.197091 12 5 0 na 649 573 9.3 73 1.21848453 3 A03 Ab8MC06DBHAbaco 25.9789839 d77.197035 16 10 5 180 693 931 9.3 136 1.14002175 4 A04 Ab8MC08DBHAbaco 25.98239 d77.197145 13 1 3 230 682 1307 9.4 91 1.23768514 5 A05 AbCHER03DBHAbaco 26.3089389 d77.005716 23 0 3 210 376 10 9.3 82 0.83915246 6 A06 AbCHER05DBHAbaco 26.3157628 d77.005943 17 0 15 210 570 29 9.9 106 0.97803116 7 A07 AbCHER07DBHAbaco 26.3210488 d77.005059 17 0 7 200 470 33 7.6 50 0.92943783 8 A08 AbGPDBH02 Abaco 26.1069463 d77.18851 15 1 0 na 1082 66 10.1 99 0.94315166 9 A09 AbGPDBH04 Abaco 26.1062361 d77.184519 13 1 0 na 646 63 6.4 106 0.77937115 10 A10 AbIH01DBH Abaco 26.1372891 d77.191612 11 0 0 na 809 20 8.1 115 1.03297207 11 A11 AbIH04DBH Abaco 26.1419403 d77.187367 19 0 0 na 384 42 8.4 124 1.1419609 12 A12 AbIH05DBH Abaco 26.1454034 d77.188245 12 0 0 na 459 12 7.9 139 1.14702907 13 A13 AbWCDBH02Abaco 26.3730238 d77.007806 22 1 0 na 156 140 10.8 82 1.04774069 14 A14 AbWCDBH04Abaco 26.3714481 d77.007853 17 1 0 na 299 273 10.1 80 0.84731353 15 A15 AbWCDBH05Abaco 26.3724739 d77.005175 16 1 0 na 386 367 9.9 88 0.93204457 16 A16 AbWCDBH08Abaco 26.369427 d77.005776 14 1 3 230 657 580 10.9 77 1.03396246 17 A17 AbWCDBH10Abaco 26.366274 d77.005974 17 1 0 na 620 770 9.6 97 1.18327987 18 A18 AbWCDBH11Abaco 26.3755546 d77.005522 21 5 0 na 167 198 8.5 61 1.17648697 19 A19 AbWCDBH13Abaco 26.3764057 d77.00412 19 1 3 135 196 362 10.2 87 1.08397029 20 A20 AbWCDBH16Abaco 26.3730269 d77.002569 14 0 0 na 268 553 6.4 83 0.74817044 21 E01 ElGHDBH21 Eleuthera 25.1755333 d76.212633 25 3 15 210 1085 68 8.52 149 1.5747589 22 E02 ElGHDBH22 Eleuthera 25.1757167 d76.212433 30 2 18 210 1118 97 8.64 123 1.03067321 23 E03 ElGPDBH19 Eleuthera 25.1324187 d76.138971 18 5 0 na 1109 27 5.08 121 0.40623508 24 E04 ElGPDBH20 Eleuthera 25.1310684 d76.141876 10 10 3 290 1431 23 6.86 134 0.53478658 25 E05 ElGTDBH17 Eleuthera 25.3829057 d76.536391 42 40 0 na 468 16 5.5 118 0.6071696 26 E06 ElGTDBH18 Eleuthera 25.383429 d76.53645 47 30 0 na 508 17 6.06 132 0.94272126 27 E07 ElJCDBH15 Eleuthera 25.3414565 d76.377065 12 40 0 na 379 18 5.16 91 0.59328377 28 E08 ElJCDBH16 Eleuthera 25.3397524 d76.376732 11 40 0 na 317 24 4.14 75 0.30827088 29 E09 ElLHDBH13 Eleuthera 24.6328123 d76.157827 24 60 3 130 850 46 4 44 0.28046859 30 E10 ElLHDBH14 Eleuthera 24.6322009 d76.157034 21 0 0 na 775 15 6.56 102 0.84038318 31 E11 ElLPDBH11 Eleuthera 25.1872442 d76.213376 17 1 5 160 483 32 5 136 0.65830605 32 E12 ElLPDBH12 Eleuthera 25.1849463 d76.213458 20 0 8 10 745 144 7.76 142 1.0169209 33 E13 ElPHDBH07 Eleuthera 25.5528258 d76.712789 14 1 3 45 950 61 5.58 85 0.49571149 34 E14 ElPCDBH08 Eleuthera 25.5576464 d76.697728 14 10 15 45 163 200 9 41 1.24829272 35 E15 ElPondDBH05Eleuthera 25.1606526 d76.205892 19 10 12 65 439 13 4.16 85 0.2731746 36 E16 ElPondDBH06Eleuthera 25.1607863 d76.206255 20 15 20 165 444 21 3.5 44 0.22411789 37 E17 ElRSDBH01 Eleuthera 24.812569 d76.174478 20 5 0 na 815 30 7.7 72 0.63821401 38 E18 ElRSDBH02 Eleuthera 24.8120586 d76.175662 21 5 0 na 616 30 8.14 155 0.92214699 39 E19 ElWBDBH03 Eleuthera 24.916249 d76.165836 12 0 0 na 2071 32 6.08 101 0.52703157 40 E20 ElWBDBH04 Eleuthera 24.9154377 d76.166936 15 0 0 na 2216 47 7.31 141 0.94586992 41 E21 ElWindDBH09Eleuthera 25.0688457 d76.132603 22 0 0 na 815 41 9.7 85 0.82152814 42 E22 ElWindDBH10Eleuthera 25.0701321 d76.133522 24 5 0 na 706 30 9.08 102 0.82011835 Supporting*Information*to*the*paper Franklin,*J.*et*al.*Regional*variation*in*Caribbean*dry*forest*composition.*Plant*Ecology Appendix*S2.*List*of*all*tree*taxa*sampled*in*Bahamian*plots*(Appendix*S1) SpCode Species_in_raw_data Accepted_taxon_name Accepted_author Accepted_family Acacho Acacia*choriophylla Acacia*choriophylla Benth. Fabaceae Amyelm Amyris*elemifera Amyris*elemifera L. Rutaceae Antluc Antirhea*lucida Stenostomum*lucidum (Sw.)*C.F.*Gaertn. Rubiaceae Bouova Bourreria*ovata Bourreria*succulenta Jacq. Boraginaceae Bursim Bursera*simaruba Bursera*simaruba (L.)*Sarg. Burseraceae Canwin Canella*winterana Canella*winterana (L.)*Gaertn. Canellaceae Cocarg Coccothrinax*argentata Coccothrinax*argentata (Jacq.)*L.H.*Bailey Arecaceae Cocdiv Coccoloba*diversifolia Coccoloba*diversifolia Jacq. Polygonaceae Cockru Coccoloba*cf*kruggii Coccoloba*krugii Lindau Polygonaceae Cocswa Coccoloba*swartzii Coccoloba*swartzii Meisn. Polygonaceae Drydiv Drypetes*diversifolia Drypetes*diversifolia Krug*&*Urb. Putranjivaceae Drylat Drypetes*lateriflora Drypetes*lateriflora (Sw.)*Krug*&*Urb. Putranjivaceae Erifru Erithalis*fruiticosa Erithalis*fruticosa L. Rubiaceae Eryare Erythroxylum*areolatum Erythroxylum*areolatum L. Erythroxylaceae Eugaxi Eugenia*axillaris Eugenia*axillaris (Sw.)*Willd. Myrtaceae Eugcon Eugenia*confusa Eugenia*confusa DC. Myrtaceae Eugfoe Eugenia*foetida Eugenia*foetida Pers. Myrtaceae Eugrho Eugenia*rhombea Eugenia*rhombea (O.*Berg)*Krug*&*Urb. Myrtaceae Exocar Exostema*caribaeum Exostema*caribaeum (Jacq.)*Roem.*&*Schult. Rubiaceae Exopan Exothea*paniculata Exothea*paniculata (Juss.)*Radlk. Sapindaceae Ficame Ficus*americana Ficus*americana Aubl. Moraceae Ficaur Ficus*aurea Ficus*aurea Nutt. Moraceae Ficcit Ficus*citrifolia Ficus*citrifolia Mill. Moraceae Guadis Guapira*discolor Guapira*discolor (Spreng.)*Little Nyctaginaceae Guaobt Guapira*obtusa Guapira*obtusata (Jacq.)*Little Nyctaginaceae Guasan Guaiacum*sanctum Guaiacum*sanctum L. Zygophyllaceae Guekru Guettarda*kruggii Guettarda*krugii Urb. Rubiaceae Guesca Guettarda*scabra Guettarda*scabra (L.)*Vent. Rubiaceae Guesp Guettarda*sp. Guettarda L. Rubiaceae Gymluc Gymnanthes*lucida* Gymnanthes*lucida Sw. Euphorbiaceae Hyptri Hypelate*trifoliata Hypelate*trifoliata Sw. Sapindaceae Jackey Jacquinia*keyensis Jacquinia*keyensis Mez Primulaceae Krufer Krugiodendron*ferreum Krugiodendron*ferreum (Vahl)*Urb. Rhamnaceae Leuleu Leucaena*leucocephala Leucaena*leucocephala (Lam.)*de*Wit Fabaceae Linosp Linociera*sp. Chionanthus*sp. L. Oleaceae Lyslat Lysoloma*latisilliquum Lysiloma*latisiliquum (L.)*Benth. Fabaceae Lyssab Lysiloma*sabicu Lysiloma*sabicu Benth. Fabaceae Manjai Manilkara*jaimiqui*subsp.*emarginata Manilkara*jaimiqui*subsp.*emarginata (L.)*Cronquist Sapotaceae Manzap Manilkara*zapota Manilkara*zapota (L.)*P.*Royen Sapotaceae Masfoe Mastichodendron*foetidissimum* Sideroxylon*foetidissimum Jacq. Sapotaceae Maybux Maytenus*buxifolia Maytenus*buxifolia (A.*Rich.)*Griseb. Celastraceae Mettox Metopium*toxiferum Metopium*toxiferum (L.)*Krug*&*Urb. Anacardiaceae Myrfra Myricanthes*fragrans Myrcianthes*fragrans (Sw.)*McVaugh Myrtaceae Neccor Nectandra*coriacea Nectandra*coriacea (Sw.)*Griseb. Lauraceae Perpal Persea*palustrus Persea*palustris (Raf.)*Sarg. Lauraceae Picpen Picramnia*pentandra Picramnia*pentandra Sw. Picramniaceae Pilosp Pilocereus*sp. Pilosocereus*millspaughii (Britton)*Byles*&*G.D.*Rowley Cactaceae Pispis Piscidia*piscipula Piscidia*piscipula (L.)*Sarg. Fabaceae Pitkey Pithecellobium*keyense Pithecellobium*keyense Britton Fabaceae Prumyr Prunus*myrtifolia Prunus*myrtifolia (L.)*Urb. Rosaceae Psesar Pseudophoenix*sargentii Pseudophoenix*sargentii H.*Wendl.*ex*Sarg. Arecaceae Reysep Reynosia*septentrionalis Reynosia*septentrionalis Urb. Rhamnaceae Royreg Roystonea*regia Roystonea*regia (Kunth)*O.F.*Cook Arecaceae Schfru Schaefferia*frutescens Schaefferia*frutescens Jacq. Celastraceae Sidsal Sideroxylon*salicifolium Sideroxylon*salicifolium (L.)*Lam. Sapotaceae Simgla Simarouba*glauca Simarouba*glauca DC. Simaroubaceae Swimah Swietenia*mahagoni Swietenia*mahagoni (L.)*Jacq. Meliaceae Tabbah Tabebuia*bahamensis Tabebuia*bahamensis (Northr.)*Britton Bignoniaceae Thodis Thouinia*discolor Thounia*discolor Grisleb. Sapindaceae Thrmor Thrinax*morrisii Thrinax*morrisii H.*Wendl. Arecaceae Trelam Trema*lamarckiana Trema*lamarckiana (Roem.*&*Schult.)*Blume Cannabaceae Zanfag Zanthoxylum*fagara Zanthoxylum*fagara (L.)*Sarg. Rutaceae Ziztay Ziziphus*taylori Ziziphus*taylorii (Britton)*M.C.*Johnst. Rhamnaceae Zuegui Zuelania*guidonia* Zuelania*guidonia (Sw.)*Britton*&*Millsp. Salicaceae uWC04B zz_AbWCDBH04_UnkB uCH05A zz_CHER05_A uCH07F zz_CHER07_F uCH07G zz_CHER07_G uRUB zz_RUBIACEAE uLP zz_UnkLP