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Cover Photograph: Native plant community at Grotto Beach, San Salvador Island, . Photograph © Carol Landry. CARIBBEAN NATURALIST Board of Editors James D. Ackerman, Department of Biology, University of at Río Piedras, USA Alfonso Aguilar-Perera, Department of Marine Biology, Universidad Autónoma de Yucatán, Wayne J. Arendt, International Institute of Tropical Forestry, Luquillo, Puerto Rico, USA Rüdiger Bieler, Field Museum of Natural History, Chicago, IL, USA Leo Douglas, Department of Geography/Geology, University of the West Indies, Mona, Robert Erdman, Department of Biological Sciences, Gulf Coast University, Fort Myers, FL, USA Keith Goldfarb, Eagle Hill Institute, Steuben, ME, USA ... Editor-in-Chief Grizelle González, International Institute of Tropical Forestry, San Juan, Puerto Rico, USA Gary R. Graves, Department of Vertebrate Zoology, Smithsonian Institution, Washington, DC, USA S. Blair Hedges, Department of Biology, Pennsylvania State University, University Park, PA, USA Julia A. Horrocks, Dept. of Biological and Chemical Sciences, Univ. of the West Indies, Cave Hill Campus, Barbados Scott Jones, Smithsonian Institution, Caribbean Coral Reef Ecosystems, Carrie Bow Cay, Heather Judkins, Department of Biological Sciences, University of South Florida, St. Petersburg, FL, USA Craig A. Layman, Department of Biological Sciences,Florida International University, North Miami, FL, USA John Leavengood, Department of Entomology, University of Kentucky, Lexington, KY, USA Antonio A. Mignucci-Giannoni, Manatee Conservation Center, Inter American University, Bay- amón, Puerto Rico, USA Gregg Moore, Department of Biological Sciences, Jackson Estuarine Laboratory, University of New Hampshire, Durham, NH, USA Robert Powell, Department of Biological Sciences, Avila University, Kansas City, MO, USA Chris Rimmer, Vermont Center for Ecostudies, Norwich, VT, USA Armando Rodríguez-Durán, Dean for Research, Inter American University, Bayamón, Puerto Rico, USA Noris Salazar Allen, Smithsonian Tropical Research Institute, Panama Inés Sastre de Jesus, Biology Department, University of Puerto Rico at Mayagüez, USA J. Angel Soto-Centeno, American Museum of Natural History, Division of Mammalogy, New York, NY, USA Christopher Starr, Department of Life Sciences, University of the West Indies, St. Augustine, Trini- dad and Tobago David W. Steadman, Florida Museum of Natural History, Gainesville, FL, USA Kathleen Sullivan Sealey, Department of Biology, University of Miami, Coral Gables, FL, USA Jarrod M. Thaxton, Department of Biology, University of Puerto at Mayagüez, USA Jason M. Townsend, Department of Wildlife, Fish and Conservation Biology, University of Califor- nia-Davis, USA ... Managing Editor Jill Weber, Eagle Hill Institute, Steuben, ME, USA ... Production Editor Byron Wilson, Department of Life Sciences, University of the West Indies at Mona, Kingston, Jamaica Graham A. J. Worthy, Department of Biology, University of Central Florida, Orlando, FL, USA Joseph M. Wunderle, International Institute of Tropical Forestry, University of Puerto Rico at Río Píedras, USA

The Caribbean Naturalist (ISSN # 2326-7119) is published by the Eagle Hill Institute, PO Box 9, 59 Eagle Hill Road, Steuben, ME 04680-0009. Phone 207-546-2821, FAX 207-546-3042. E-mail: [email protected]. Webpage: www.eaglehill.us/cana. Copyright © 2014, all rights reserved. Periodical postage paid in Steuben, ME and additional mailing offices. Published quarterly. Special issue proposals are welcome. On-line secure subscription ordering: rate per year for Caribbean subscribers - $15 regular, $10 students, $60 organizations; for Non-Caribbean subscribers - $20 regular, $15 students, $80 organizations. Journal subscription exchanges are considered. Authors: submission guidelines are available at www.eaglehill.us/cana. Co-published journals: The Northeastern Natu- ralist (ISSN 1092-6194 [print], ISSN 1938-5307 [online]) and the Southeastern Naturalist (ISSN 1528-7092 [print], ISSN 1938-5412 [online]), journals with separate Boards of Editors. The Eagle Hill Institute is a tax exempt 501(c)(3) nonprofit corporation of the State of Maine (Federal ID # 010379899). 20142014 CARIBBEANCaribbean Naturalist NATURALIST No. 9:1–16No. 9 C.L. Landry, N.B. Elliott, A.J. Finkle, and L.B. Kass

Plant-Pollinator Interactions in Bahamian Coastal Communities

Carol L. Landry1,*, Nancy B. Elliott2, Adam J. Finkle2,3, and Lee B. Kass4

Abstract - In light of increased pressure from direct anthropogenic effects and climate change, coastal conservation has become even more important. Here we report on observed visiting flowers on 30 plant in 14 families during two time periods on San Salvador Island, The Bahamas. An understanding of community pollination dynamics is necessary for successful conservation efforts in coastal ecosystems because most plant species rely on pollinators for cross-pollination; thus, identification of potential pollinators for each plant species is a necessary first step. A total of 41 animal species representing four and two orders were observed visiting flowers. The relative importance of individual species to the community was estimated based on the number of interaction partners and whether the species was active during both time periods.

Introduction Coastal ecosystems are dynamic, sculpted by wind and water. They have great ecological and economic importance because they provide significant ecosystem services: they stabilize coastal sediments (Kumara et al. 2010, Laurance et al. 2011) and build dunes (Miller et al. 2010), buffer adjacent inland communities against storm surge (Gedan et al. 2011, Krauss et al. 2009, Miller et al. 2010), and act as nurseries for marine fishes (Barbier et al. 2011, Beck et al. 2001). Coastal ecosystems are vulnerable to a number of anthropogenic and natural disturbances, including hu- man economic development (Dahdouh-Guebas et al. 2004, Ellison and Farnsworth 1996), sea-level rise associated with climate change (Krauss et al. 2010, Kumara et al. 2010), and hurricanes (Franklin et al. 2006; Landry 2011, 2013; Middleton 2009; Smith et al. 2009; Tanner et al. 1991), as well as interactions between disturbance types (e.g., sea-level rise and hurricanes [Ross et al. 2009], sea-level rise and agricul- ture [Eslami-Andargoli et al. 2010]). This study is important because investigation of the reproductive ecology of organisms in coastal communities, including the effects of disturbance on pollination mutualisms (Ashman et al. 2004; Benedek et al. 2007; Horvitz et al. 2010; Rathcke 2000, 2001), is critical for successful conservation, res- toration, or mitigation efforts (Kearns et al. 1998). Many plant species rely on animal pollinators for successful reproduction, and the lack of consistent pollination services from these animals can result in reduced

1Department of Evolution, Ecology, and Organismal Biology, The Ohio State University, Mansfield, OH 44906, USA.2 Department of Biology, Siena College, Loudonville, NY 11211, USA. 3Current address - Sustainability Science Program, University of Massachusetts, Am- herst, MA 01003, USA. 4L.H. Bailey Hortorium, Department of Plant Biology, Cornell Uni- versity, Ithaca, NY 14853, USA. *Corresponding author - [email protected]. Manuscript Editor: Kathleen Sullivan-Sealey 1 2014 Caribbean Naturalist No. 9 C.L. Landry, N.B. Elliott, A.J. Finkle, and L.B. Kass seed production due to pollination limitation (Campbell 1987; Jurgens et al. 2009; Rathcke 2000, 2001). Further reductions in seed or fruit set due to inbreeding depression can occur in autogamously selfing species (Bellusci et al. 2009, Ka- meyama and Kudo 2009), although increased selfing rates do not always strengthen inbreeding depression in plant populations (Geiger et al. 2010, Landry and Rathcke 2007), and the effects can be mitigated over time due to strong selective filters that favor selfing (Bodbyl-Roels and Kelly 2011). While most plant species are general- ists that can interact with a number of mutualist partners, if pollination services are absent or severely reduced for a prolonged period by a catastrophic disturbance, then plant species that rely exclusively on animal pollinators for seed set could disappear from that community (Kearns et al. 1998). Generalist pollinators can take advantage of floral resources from other plant species (Roubik and Villanueva- Gutiérrez 2009), but pollinators that specialize on only a few plant species may not be able to re-establish viable populations in the community. Further, a subset of the and pollinators in the tropics and sub-tropics are active year-round, but there are also species that are only active for part of the year. The species that are active year-round may interact with one another, but they are also likely to interact with species that are temporally restricted in their activities. Thus, species that are active at different times of the year may facilitate one another indirectly through shared mutualist partners (Lazaro et al. 2009, Waser and Real 1979). Most plant species in The Bahamas rely on animals for cross-pollination, and many have generalist flowers that may be visited by a number of pollinator species. However, some plant species have flowers specialized for a subset of pollinators. The purpose of this study was to identify potential pollinators of plants that occur in four coastal communities on San Salvador Island, The Bahamas: beach-fore- dune, Coccothrinax-shrub, shrub-thicket, and coppice-thicket communities. These communities form a successional series in the coastal ecosystem, from the early- successional beach-foredune community to the late-successional coppice-thicket community. Throughout this paper, we use the term “plant community” to describe a discrete assemblage of interactive plant species that occurs within a successional series, which is appropriate in systems that are strongly influenced by disturbance (sensu Franklin and Steadman 2010). Further, we consider early and late-stage thickets as separate communities (shrub-thicket and coppice-thicket, respectively) because the dominant species in each assemblage are different, and the difference in the vertical structure of each community can influence the animal species assem- blages in that community (Sutton et al. 1983). Here we report on the assemblage of flower visitors observed on plants in coastal communities on San Salvador Island at two distinctly different times of year: July (early in the rainy season) and December (at the beginning of the dry season).

Field-Site Description This study was conducted at five locations on San Salvador Island, The Baha- mas (Fig. 1): East Beach (E), Grahams Harbour (GH), Grotto Beach (GB), Rocky 2 2014 Caribbean Naturalist No. 9 C.L. Landry, N.B. Elliott, A.J. Finkle, and L.B. Kass

Point (R), and Sandy Hook (S). At each location, we surveyed in plant communities within 200 m of the shoreline, including beach-foredune (E, GH, S), Coccothrinax- shrub (R, GB, S), shrub-thicket (E, GH, GB, S), and coppice-thicket (E, GH, S). The strength of the disturbance felt by these communities differs depending on dis- tance from the ocean and local topography. The effect of the disturbance generally decreases with distance from the ocean and/or altitude, but locations that are on the inland side of dune or rock formations may be more protected.

Methods We performed field work in 2010 during two time periods: 18–27 July (≈100 hours of observation) and 2–10 December (≈110 hours of observation). We iden- tified all animal-pollinated plant species in flower during each time interval and recorded and identified all their floral visitors. We made all observations between 8 am and 6 pm. We confirmed plant identifications using Kass (2009) or Correll and Correll (1982), and verified all plant names and family affiliations through the Tropicos.org Missouri Botanical Gardens website (2013). Insect identifications were either confirmed directly by N.B. Elliott, by comparison of to previ- ously identified specimens held in the Gerace Research Centre’s Repository, or by using Elliott et al. (2009). Insect voucher specimens are deposited in the GRC Repository’s research collection.

Figure 1. Map of The Bahamas archipelago and San Salvador Island (inset), with study loca- tions indicated. Location codes: GH = Grahams Harbour, E = East Beach, S = Sandy Hook, GB = Grotto Beach, and R = Rocky Point. Scale shows latitude in degrees North. Image modified from Robinson and Davis (1999) and GinkgoMaps Project (2014). 3 2014 Caribbean Naturalist No. 9 C.L. Landry, N.B. Elliott, A.J. Finkle, and L.B. Kass Results We observed a total of 37 plant species in 15 families flowering in one to three community types during the two time periods (Table 1). Twenty plant species in 12 families were flowering during July at one or more location, while 30 species in 14 families were in flower during December; of these, 13 species were flowering dur- ing both time periods.

Table 1. Phenological data for plants flowering in July and/or December 2010, and the communities within which the plants were observed. community types: B-f = beach-foredune, C-s = Coccothrinax- shrub, S-t = Shrub-thicket, and C-t = Coppice-thicket. Code: X = plants were in flower; P = plants were present in community. Flowering time Community type Family/Species Jul Dec B-f C-s S-t C-t Aizoaceae Sesuvium portulacastrum (L.) L. X P − − − umbellata Jacq. X − − P P Cynanchum bahamense (Griseb.) Gillis X − P P − luteum (L.) B.F. Hansen and Wunderlin X − P P − alba DC. X − P P − Borrichia arborescens (L.) DC. X P − − − Gundlachia corymbosa (Urb.) Britt. X − P P − Wedelia bahamensis (Britt.) Schulz ex Urb. X − − P P Bourreria baccata Raf. X X − − P P Cordia bahamensis Urb. X X − − − P Heliotropium gnaphalodes L. X X P − − − Cakile lanceolata (Willd.) O.E. Schutz X X P − − − Convolvulaceae Merremia dissecta (Jacq.) Hall. f. X − − P − Jacquemontia cayensis Britt. X − P P − Euphorbiaceae Croton discolor Willd. X − P P P Croton linearis Jacq. X X − − P P Fabaceae Calliandra haemotomma (Brit.) Benth. X − − P − rosea (Sw.) DC. X P P − − Chamaecrista lineata (Sw.) Green X X − P P − Leucaena leucocephala (Lam.) de Wit. X − − P − Pithecellobium keyense Britt. Ex Britt.and Rose X − − P − Stylosanthes hamata (L.) Taub X − − P − Goodeniaceae Scaveola taccada (Gaertn.) Roxb. X X P − P − 4 2014 Caribbean Naturalist No. 9 C.L. Landry, N.B. Elliott, A.J. Finkle, and L.B. Kass In July, we observed 96 animals representing 21 species in six orders, four of insects and two of , visiting the flowers of 14 species in 10 plant families (Table 2), although some individuals may have made multiple visits to the same plants. In December, 470 animals representing 35 species in four orders, three of insects and one of birds, were observed visiting the flowers of 26 species in 14 plant families (Table 3); again, some individuals may have visited the same plants multiple times. Only plant species that received floral visitors during our observations are included on the visitor tables. Most animal visitors were insects, primarily hymenopterans and lepidopterans, although two bird visitors (one evelynae [] and one Coereba flaveola [Ba- nanaquit]) were observed in July, and several Bananaquits were observed in De- cember. In July, insects from 13 families in four orders were observed probing flowers. The insects represented one family of (Coleoptera), two of (Diptera), five of bees and wasps (), and five of (Lepi- doptera). In December, insects from 15 families in three orders were observed probing flowers; they represented two families of flies, seven of bees and wasps, and six of butterflies and . All dipteran, hymenopteran, and lepidopteran families represented in July were also represented in December, but not neces- sarily by the same species.

Table 1, continued. Flowering time Community type Family/Species Jul Dec B-f C-s S-t C-t Malvaceae Corchorus hirsutus L. X − − P − Melochia tomentosa L. X − − P P Waltheria bahamensis Britt. X − − P −

Passifloraceae cupraea L. X − − P − Passiflora pectinata Griseb. X − P P − L. X X − P P − Casasia clusiifolia (Jacq.) Urb. X X − P P − Erithalis diffusa Correll X X P P P − Erithalis fruticosa L. X X − P P − littoralis Sw. X − P P − Solanaceae Solanum bahamense L. X X − P P − Surianaceae Suriana maritima L. X P − A − Verbenaceae Lantana involucrata L. X X − P P − Stachytarpheta jamaicense L. X − − P −

5 2014 Caribbean Naturalist No. 9 C.L. Landry, N.B. Elliott, A.J. Finkle, and L.B. Kass

Ve (1 sp.) 1 1 So (1 sp.)

2 2 Ru (2 spp.) 1 1 1 1 1 Pa (3 spp.)

1 Go (1 sp.) 1 Fa (1 sp.) , Ru = Rubiaceae, So = Solanaceae, and

Plant families visited (# species) 1 1 1 Eu (1 sp.) 1 Br (1 sp.)

1 1 Bo (2 spp.)

1 As (1 sp.) Xylocopa cubaecola Lucas Coelioxys sp. Megachile alleni Mitchell signata (L.) Myzinum apicale Cresson And Salt Polistes bahamensis picturatus Beq. Agraulis vanillae insularis Maynard undetermined species Centris versicolor (Fabr.) Leptotes cassius theonus (Lucas) hegesia Cramer Kricogonia lyside Godart Phoebis agarithe antillia Brown

Megachilidae

Sphecidae Tiphiidae Vespidae Heliconiidae Hesperiidae Hymenoptera Apidae Lycaenidae Pieridae Table Animals 2. observed Table visiting flowers, and the number of species in each plant family that received floral visits from the animal during July 2010. Undetermined species were observed visiting flowers but were not collected, and could not be further identified. Plant As Families: = Asteraceae, Bo = Boraginaceae, Br = Brassicaceae, Eu Euphorbiaceae, Fa Fabac eae, Go Goodeniaceae, Pa Ve = Verbenaceae. = Ve Visitor Order/Family/Species Visitor

6 2014 Caribbean Naturalist No. 9 C.L. Landry, N.B. Elliott, A.J. Finkle, and L.B. Kass

1 Ve

So

2 1 1 1 2 1 Ru

1 Pa

Go

1 1 Fa

Plant families visited

1 1 1 Eu

Br

1 1 1 Bo

As Calliphlox evelynae (Bourcier) flaveola (L.) Coereba Oxacis sp. (F.) Ligyra cereberus Poecilanthrax lucifer (F.) undetermined species undetermined species Chrysanthrax maculipennis Scarbrough and Davidson

Apodiformes Trochilidae Passeriformes Emberizidae Coleoptera Oedemeridae Syrphidae Table 2, continued. Table Visitor Order/Family/Species Visitor Diptera

7 2014 Caribbean Naturalist No. 9 C.L. Landry, N.B. Elliott, A.J. Finkle, and L.B. Kass -

1 1 1 1 1 1 Ve (2) 1 Su (1) P a s s i f l o r a

1 So (1) 1 2 1 1 3 3 1 Ru (4)

Pa (1) 1 2 1 1 2 Ma (2)

1 1 (1) Go 1 1 1 1 1 1 1 2 2 Fa (4)

1 1 1 1 1 Eu (2) Plant families visited (# species) 1 1 1 1 Co (1)

1 1 1 1 1 Br (1) 1 2 1 Bo (2)

1 1 1 As (3)

1 1 1 1 1 1 1 1 1 Ap (1) Myzinum sp. Beq. And Salt Pachodynerus cubensis bahamensis Beq. Pachodynerus scrupeus Zavattari Pachodynerus sp. And Salt Polistes bahamensis picturatus Beq. fidelissima vagrans Bates Bates heros Empyreuma Eunomia latenigra Butler Cerceris watlingensis Elliott and Salbert Cerceris Stictia signata (L.) Campsomeris trifasciata (Fabr.) Megachile bahamensis Mitchell Megachile alleni Mitchell Dialictus sp. Xylocopa cubaecola Lucas Agapostemon columbi Roberts Centris versicolor (Fabr.)

Tiphiidae Vespidae Lepidoptera Arctiidae

Sphecidae

Scoliidae

Megachilidae

Hymenoptera Halictidae Apidae

Table Animals 3. observed Table visiting flowers, and the number of species in each plant family that received floral visits from the animal= Apocynaceae. during December, Ap Families: Plant identified. further be not could and collected, not were but flowers visiting observed were species Undetermined 2010. Asteraceae, Bo = Boraginaceae, Br Brassicaceae, Eu Euphorbiaceae, Fa Fabaceae, Go Goodeniaceae, Ma Malvaceae, Pa As = ceae, So = Solanaceae,Su = Surianaceae, and Ve = Verbenaceae. = Ve ceae, So = Solanaceae,Su Surianaceae, and Visitor Order/Family/Species Visitor

8 2014 Caribbean Naturalist No. 9 C.L. Landry, N.B. Elliott, A.J. Finkle, and L.B. Kass

1 2 1 1 1 Ve

Su

1 1 So

2 1 2 1 1 2 1 1 Ru

1 1 1 1 Pa

1 Ma

Go

1 1 1 1 1 1 1 1 1 Fa

1 1 1 Eu Plant families visited (# species)

1 Co

1 1 1 1 1 Br

1 2 Bo

2 2 1 1 1 1 As

Ap Coereba flaveola (L.) Coereba Chrysanthrax maculipennis Scarbrough and Davidson (F.) Ligyra cereberus Undetermined species Ocyptamus sp. (Wied.) Palpada albifrons Agraulis vanillae insularis Maynard Hylephila phyleus Drury sp. Chlorostrymon Leptotes cassius theonus (Lucas) Clench Strymon acis armouri Euptoieta hegesia Cramer Junonia genoveva (Cramer) Memphis intermedia (Witt) Ascia monuste eubotea Latreille chamberlaini (Butler) Eurema Kricogonia lyside Godart Phoebis agarithe antillia Brown

Passeriformes Emberizidae

Visitor Order/Family/Species Visitor Table 3, continued. Table Bombyliidae Syrphidae

Heliconiidae

Hesperiidae Lycaenidae

Nymphalidae

Pieridae Diptera

9 2014 Caribbean Naturalist No. 9 C.L. Landry, N.B. Elliott, A.J. Finkle, and L.B. Kass

Discussion In general, fewer plant species were in flower during late July relative to ear- ly December, and fewer floral visitors were observed in July than in December as well, probably due to climate conditions. Shaklee (1996) reported that on San Salvador, July is relatively dry and has the highest mean monthly temperature and mean monthly maximum of the year (28 ºC and 32 ºC, respectively), where- as December follows the wettest months of the rainy season and has a mean monthly temperature and mean monthly minimum that are among the lowest (23 ºC and 18 ºC, respectively). The rainy season on San Salvador is characterized by occasional storms, not daily showers (Shaklee 1996). Species that were not active in July may have high water requirements for the initiation of flowering (plants) or larval development (insects) that are not met until the peak of the rainy season (October). Hymenopterans and lepidopterans were the most speciose pollinator groups in either season, but some dipteran and coleopteran visitors were also observed, along with a few Bahama Woodstar and Bananaquit individuals. In general, pollinators are scarce on San Salvador, so additional observations are necessary to identify all the pollinators of each plant species. Most of the plants were generalists with respect to floral visitors, although a few species appeared to be more specialized, and several plant species exhibited characteristics associated with bee, , and bird pollination syndromes described by Howe and Wetsley (1988). Rathcke et al. (2005) noted that Corchorus hirsutus exhibits some traits asso- ciated with bee-visited flowers in that the plant has yellow, radially symmetrical flowers that lack a floral tube. However, the flowers also open late in the day and do not produce nectar (Rathcke et al. 2005; C.L. Landry, pers. observ.), traits which are not associated with bee flowers. During our December observations, three of the five species visitingC. hirsutus were bees. On Inagua, the halictid bee Agapos- temon columbi was the most frequent visitor and the only bee observed (Rathcke et al. 2005); A. columbi visited C. hirsutus on San Salvador as well, but Megachile alleni (Megachilidae) was more frequent (Landry et al., in press). All members of the Fabaceae are animal-pollinated, and many provide both nectar and pollen resources. Large bees like Centris versicolor and Xylocopa cubaecola (Apidae) visited the brightly colored single flowers of Chamaecrista lineata and , whereas apids, halictids, and megachilids vis- ited the globe-shaped inflorescences of Pithecelobium keyense and Leucaena leucocephala. We observed 16 visitor species to P. keyense, including six lepidopterans. The most common visitor to P. keyense during our study was , which we observed 22 times. In a previous study on San Salvador during December and January in 2002–2003, M. alleni was the most common visitor to P. keyense, making 13 visits to the plant (Elliott et al. 2011). Observations made by Elliott and Smith (2003) on the southern islands of Ack- lins, Mayaguana, and Great Inagua during the winter months of 2000 showed that many of the same insect species visited P. keyense. However, there were some 10 2014 Caribbean Naturalist No. 9 C.L. Landry, N.B. Elliott, A.J. Finkle, and L.B. Kass differences in visitor species on different islands because of geographic distribu- tions of the species occurring there. A southern species of Campsomeris, C. baha- mensis Bradley, occurred on Acklins and Great Inagua instead of C. trifasciata, which is found in the central and northern Bahamas, including San Salvador. An endemic species of Pachodynerus, P. linda Menke, was the only member of the found on Mayaguana, and a southern subspecies of a Hispaniolan species in the same genus, P. tibialis barbouri Bequaert, visited P. keyense on Great Ina- gua (Elliott and Smith 2003, Menke 1986). Although Rathcke et al. (2001) noted that Bourreria baccata exhibits many characteristics of Howe and Westley’s (1988) butterfly-pollination syndrome, we saw Xylocopa cubaecola and Campsomeris trifasciata visiting this species, as well as Agraulis vanillae and Bananaquits. Miller et al. (1992) reported visits to B. bac- cata by nine species of butterflies on Crooked, Acklins, and Mayaguana islands, as well as visits by A. vanillae to Lantana involucrata and Stachytarpheta jamaicen- sis (both Verbenaceae), which have short tube-shaped flowers with nectar. A number of plants with tubular flowers may be visited by birds. Bananaquits visited Bourreria baccata (Boraginaceae) during both seasons of our observations, and it also visited Erithalis diffusa and (Rubiaceae) in Decem- ber. Bananaquits, , and warblers have also been observed visiting Ernodea species in the Bahamas and Puerto Rico (Negron-Ortiz and Hickey 1996). Despite its floral structure, E. littoralis was frequently visited by the wasp Camp- someris trifasciata, although Elliott (1990) reported that female wasps slit the flo- ral tube from the top to its base before feeding, a behavior that we also observed. Negron-Ortiz and Hickey (1996) also reported that some species of Ernodea were visited by C. trifasciata nassauensis Bradley on Great Abaco and Puerto Rico, but they did not describe damage to the floral tube. Whether or not splitting the floral tube has any effect on seed or fruit set is not known. Many of the animal species we observed visited flowers of several species, al- though during our December observations, they rarely belonged to the same plant family. In December, we observed 35 species visiting an average of 3.5 species of plants in an average of 1.7 families; only eight of the pollinator species were ob- served on a single plant species. In contrast, 12 of the 21 animal species observed in July were recorded on the flowers of a single plant species. The difference could be due to fewer species offering floral resources in July or to specialization; additional studies are needed in order to determine if the insects visit the flowers of other species at different times of the year. A number of Hymenoptera species were active during both time periods and visited a wide range of floral types. Campsomeris trifasciata (Scoliidae) visited 14 species from 10 different families. Two species of bees were frequent flower visitors: Xylocopa cubaecola (Apidae) visited a total of nine different species from seven families, and Megachile al- leni (Megachilidae) visited 11 species in nine families. Stictia signata (Specidae) was observed on species in ten different plant families in December, although we observed only one visit in July. Several species of Lepidoptera were common plant visitors during the December observations. Agraulis vanillae (Heliconiidae) 11 2014 Caribbean Naturalist No. 9 C.L. Landry, N.B. Elliott, A.J. Finkle, and L.B. Kass visited 10 species in seven plant families, while Strymon acis and Leptotes cas- sius (both Lycaenidae) visited seven and six species, respectively. During the July observations, two of these three butterflies each made one floral visit, and S. acis was not seen. The diurnal arctiid moths, Empyreuma heros and Composia fidelissima,visited flowers of three and four species, respectively, in the same number of families in December. Based on these preliminary data, a number of plant and animal species appear to be important to the pollination dynamics of these communities. Erithalis diffusa and Croton discolor were both flowering during July and December at multiple locations. Erithalis diffusa attracted seven floral visitor species in July and 12 in December, with only one animal species (Ligyra cereberus) visiting E. diffusa flowers during both time intervals. Croton discolor attracted eight species in July and seven in December, again with one animal species (Megachile alleni) observed on C. discolor flowers during both intervals. Two other plant species (Cordia baha- mensis and Erithalis fruticosa) attracted at least six floral visitor species in July. In December, at least 10 animal species were attracted to the flowers of five additional plant species: Cakile lanceolata, Cynanchum bahamense, Ernodea littoralis, Lan- tana involucrata, and Pithecellobium keyense. Xylocopa cubaecola was observed visiting flowers during both time periods; individuals visited four plant species in July and six in December. The only plant species visited by X. cubaecola during both time periods was Scaevola taccada. Oxacis beetles were seen visiting the flowers of five different plant species in July. In December, six additional insect species visited the flowers of at least five plant species; these insects were Agraulis vanillae, Campsomeris trifasciata, Leptotes cassius, Megachile alleni, Stictia signata, and Strymon acis. Additional studies to determine the effects of direct and indirect interactions on pollination networks within coastal plant communities are warranted, given the importance of these communities (Miller et al. 2010), the diversity of mutualist interactions (Bronstein 1994), the variation in the strength of those interactions (Bascompte et al. 2006), the insular nature of island systems (MacArthur and Wil- son 1967), and the changes in the strength and direction of interactions in stochastic environments (Ashman et al. 2004, Horvitz et al. 2010). These preliminary data will be included in ongoing studies of coastal community pollination networks on islands of The Bahamas.

Acknowledgments We thank The Bahamas Environment, Science, and Technology (BEST) Commission of the Ministry of the Environment for permission of conduct research in The Bahamas. Further, we thank the director and staff of the Gerace Research Centre, College of The Bahamas, for logistic support and assistance on San Salvador Island, and the editor and two anonymous reviewers for comments that have improved this manuscript. The material included in this paper was first presented in 2011 at the 14th Symposium on the Natural History of the Bahamas and was published in the proceedings of that conference by the Gerace Research Centre. A significant portion of the funding for this work was provided by The Ohio State University-Mansfield.

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