Heliconia

Heliconia
Heliconia latispatha inflorescences
Scientific classification
Kingdom:	Plantae
(unranked):	Angiosperms
(unranked):	Monocots
(unranked):	Commelinids
Order:	Zingiberales
Family:	Heliconiaceae Vines[1]
Genus:	Heliconia L.
Synonyms[2]
Bihai Mill. Heliconiopsis Miq.

Heliconia mariae inflorescence

Heliconia, derived from the Greek word helikonios, is a genus of flowering plants in the Heliconiaceae. Most of the species are native to the tropical Americas, but a few are indigenous to certain islands of the western Pacific and Maluku.^[2] Many species of Heliconia are found in rainforests or tropical wet forests of these regions. Several species are widely cultivated as ornamentals, and a few are naturalized in Florida, Gambia and Thailand.^[3] Common names for the genus include lobster-claws, wild plantains or false bird-of-paradise. The last term refers to their close similarity to the bird-of-paradise flowers (Strelitzia). Collectively, these plants are also simply referred to as heliconias.

The Heliconia are a monophyletic genus in the family Heliconiaceae, but was formerly included in the family Musaceae, which includes the bananas (e.g., Musa, Ensete; Judd et al., 2007). However, the APG system of 1998, and its successor, the APG II system of 2003, confirm the Heliconiaceae as distinct and places them in the order Zingiberales, in the commelinid clade of monocots.

These herbaceous plants range from 0.5 to nearly 4.5 meters (1.5–15 feet) tall depending on the species (Berry and Kress, 1991). The simple leaves of these plants are 15–300 cm (6 in-10 ft). They are characteristically long, oblong, alternate, or growing opposite one another on non-woody petioles often longer than the leaf, often forming large clumps with age. Their flowers are produced on long, erect or drooping panicles, and consist of brightly colored waxy bracts, with small true flowers peeping out from the bracts. The growth habit of heliconias is similar to Canna, Strelitzia, and bananas, to which they are related.The flowers can be hues of reds, oranges, yellows, and greens, and are subtended by brightly colored bracts. The plants typically flower during the wet season. These bracts protect the flowers; floral shape often limits pollination to a subset of the hummingbirds in the region (Gilman and Meerow, 2007).

Uses and ecology

Heliconias are grown for the florist's trade and as landscape plants. These plants do not grow well in cold, dry conditions. They are very drought intolerant, but can endure some soil flooding. Heliconias need an abundance of water, sunlight, and soils that are rich in humus in order to grow well. These flowers are grown in tropical regions all over the world as ornamental plants (Ong, 2007). The flower of H. psittacorum (Parrot Heliconia) is especially distinctive, its greenish-yellow flowers with black spots and red bracts reminding of the bright plumage of parrots.

Several cultivars and hybrids have been selected for garden planting, including:

• H. psittacorum × H. spathocircinata, both species of South America, mainly Brazil
• H. × rauliniana = H. marginata (Venezuela) × H. bihai (Brazil)
• H. chartacea cv. 'Sexy Pink'

Most commonly grown landscape Heliconia species include Heliconia augusta, H. bihai, H. brasiliensis, H. caribaea, H. latispatha, H. pendula, H. psittacorum, H. rostrata, H. schiediana, and H. wagneriana.

Heliconias are an important food source for forest hummingbirds, especially the hermits (Phathornithinae), some of which – such as the rufous-breasted hermit (Glaucis hirsuta) – also use the plant for nesting. The Honduran white bat (Ectophylla alba) also lives in tents it makes from heliconia leaves.

Characteristics

Heliconia psittacorum

Heliconia rostrata inflorescence

Closeup of Heliconia vellerigera inflorescence

H. psittacorum × H. spathocircinata cv. Golden Torch

Heliconia bourgaeana Petersen

Leaf

Leaves in different positions on the plant have a different absorption potential of sunlight for photosynthesis when exposed to different degrees of sunlight (He et al. 2006).

Flower

Flowers produce ample nectar that attracts pollinators, most prevalent of which are hummingbirds (Bruna et al. 2004)

Seeds

Fruits are blue-purple when ripe and primarily bird dispersed (Uriarte et al. 2011). Post dispersal seed survival was studied and had many results (Hoii and Lulow 2006). The study showed that seed size was not a determinant of post dispersal seed survival. The highest amount of seed predation came from mammals.

Heliconia and bats

Pollination

Although Heliconia are almost exclusively pollinated by hummingbirds, some bat pollination has been found to occur. Heliconia solomonensis is pollinated by the macroglosine bat (Melonycteris woodfordi) in the Solomon Islands. Heliconia solomonensis has green inflorescences and flowers that open at night, which is typical of bat pollinated plants. The macroglosine bat is the only known nocturnal pollinator of Heliconia solomonensis (Kress 1985).

Habitat

Many bats use Heliconia leaves for shelter. The Honduran white bat, Ectohylla alba, utilizes five species of Heliconia to make diurnal tent shaped roosts. The bat cuts the side veins of the leaf extending from the midrib causing the leaf to fold like a tent. This structure provides the bat with shelter from rain, sun, and predators. In addition, the stems of the Heliconia leaves are not strong enough to carry the weight of typical bat predators, so shaking of the leaf alerts roosting bats to presence of predators (Timm 1976). The bats Artibeus anderseni and A. phaeotis form tents from the leaves of Heliconia in the same manner as the Hondurian white bat (Timm 1987). The Neotropical disk-winged bat, Thyroptera tricolor, has suction disks on the wrists which allow it to cling to the smooth surfaces of the Heliconia leaves. This bat roosts head up in the rolled young leaves of Heliconia plants (Findley 1974).

Insects and Heliconia

Heliconias provide shelter for a diverse range of insects within their young rolled leaves and water-filled floral bracts. Insects that inhabit the rolled leaves often feed upon the inner surfaces of the leaf, such as beetles of the family Chrysomelidae. In bracts containing small amounts of water, fly larvae and beetles are the dominant inhabitants. In bracts with greater quantities of water the typical inhabitants are mosquito larva. Insects living in the bracts often feed on the bract tissue, nectar of the flower, flower parts, other insects, microorganisms, or detritus in the water contained in the bract (Siefert 1982). Almost all species of Hispini beetles that use rolled leaves are obligate herbivores of plants of the order of Zingiberales, which includes Heliconia. These beetles live in and feed from the rolled leaf, the stems, the inflorescences, or the unfurled mature leaves of the Heliconia plant. In addition, these beetles deposit their eggs on the leaf surface, petioles of immature leaves, or in the bracts of the Heliconia (Strong 1977). Furthermore, some wasp species such as Polistes erythrocephalus build their nest on the protected underside of large leaves. ^[4]

Hummingbirds and Heliconia

Hummingbirds are the main pollinators of flowers in the genus Heliconia (order Zingiberales: family Heliconiaceae) in many locations. The concurrent diversification of hummingbird pollinated taxa in the order Zingiberales and the hummingbird family (Trochilidae: Phaethorninae) 18 mya supports that these radiations have significantly influenced one another through evolutionary time (Bleiweiss 1998; Kress and Specht 2005). At La Selva research station in Costa Rica it was found that specific species of Heliconia have specific hummingbird pollinators (Stiles 1975). These hummingbirds can be organized into two different groups: hermits and non-hermits. Hermits are a subfamily of Phaethornithinae, consisting of the genera Anopetia, Eutoxeres, Glaucis, Phaethornis, Raphodon, and Threnetes (McGuire et al. 2008). Non-hermits are a paraphyletic group within the Trochilidae, comprising several clades (McGuire 2008). Hermits are generally traplining foragers, where individuals visit a repeated circuit of high-reward flowers instead of holding fixed territories (Stiles 1975; Dobkin 1984). Non-hermits are territorial over their Heliconia clumps, causing greater self-pollination (Stiles 1975). Hermits tend to have long curved bills while non-hermits tend to possess short straight bills, a morphological difference that likely spurred the divergence of these groups in the Miocene era (Graham et al. 2009; Temeles et al. 2010). Characteristics of Heliconia flowers that select for either hermit or non-hermit pollinator specificity are degree of self-compatibility, flowering phenology, nectar production, color, and shape of flower (Kress and Specht 2005; Meléndez-Ackerman et al. 2005; Graham et al. 2009). The hummingbird itself will choose the plants its feeds from based on its beak shape, its perch on the plant, and its territory choice (Linhart 1973).

It was found that hummingbird visits to the Heliconia flower did not affect its production of nectar (Feinsinger 1983). This may account for the flowers not having a consistent amount of nectar produced from flower to flower.

Different Heliconia species have different flowering seasons. This suggests that the species compete for pollinators. It was found that many species of Heliconia, even the newly colonized species, all had many pollinators visit (Feinsinger 1978).

Species

Species accepted by Kew Botanic Gardens^[3]

1. Heliconia abaloi -Colombia
2. Heliconia acuminata - South America
3. Heliconia adflexa - S Mexico, Guatemala, Honduras
4. Heliconia aemygdiana - South America
5. Heliconia albicosta - Costa Rica
6. Heliconia angelica - Ecuador
7. Heliconia angusta - SE Brazil
8. Heliconia apparicioi - Ecuador, Peru, NW Brazil
9. Heliconia arrecta - Colombia
10. Heliconia atratensis - Colombia
11. Heliconia atropurpurea - Colombia, Panama, Costa Rica
12. Heliconia aurantiaca - S Mexico, Central America
13. Heliconia auriculata - Bahia
14. Heliconia badilloi - Colombia
15. Heliconia barryana - Chiriquí
16. Heliconia beckneri - Costa Rica
17. Heliconia bella - Panama
18. Heliconia berriziana - Colombia
19. Heliconia berryi - Napo
20. Heliconia bihai - West Indies, N South America
21. Heliconia bourgaeana - S Mexico, Central America
22. Heliconia brachyantha - Panama, Colombia, Venezuela
23. Heliconia brenneri - Ecuador
24. Heliconia burleana - Colombia, Ecuador, Peru
25. Heliconia caltheaphylla - Costa Rica
26. Heliconia caquetensis - Colombia
27. Heliconia carajaensis - Pará
28. Heliconia caribaea - West Indies
29. Heliconia carmelae - Colombia
30. Heliconia chartacea - N South America
31. Heliconia chrysocraspeda - Colombia
32. Heliconia clinophila - Costa Rica, Panama
33. Heliconia colgantea - Costa Rica, Panama
34. Heliconia collinsiana - S Mexico, Central America
35. Heliconia combinata - Colombia
36. Heliconia cordata - Colombia, Ecuador
37. Heliconia crassa - Guatemala
38. Heliconia cristata - Panama
39. Heliconia cucullata - Costa Rica, Panama
40. Heliconia curtispatha - Colombia, Ecuador, Central America
41. Heliconia danielsiana - Costa Rica, Panama
42. Heliconia darienensis - Colombia, Panama
43. Heliconia dasyantha - Suriname, French Guiana
44. Heliconia densiflora - Trinidad, N South America
45. Heliconia dielsiana - NW South America
46. Heliconia donstonea - Colombia, Ecuador
47. Heliconia episcopalis - South America
48. Heliconia estherae - Colombia
49. Heliconia estiletioides - Colombia
50. Heliconia excelsa - Napo
51. Heliconia farinosa - SE Brazil, NE Argentina
52. Heliconia faunorum - Panama
53. Heliconia fernandezii - Antioquia
54. Heliconia × flabellata - Ecuador
55. Heliconia foreroi - Colombia
56. Heliconia fragilis - Colombia
57. Heliconia fredberryana - Imbabura
58. Heliconia fugax - Peru
59. Heliconia gaiboriana - Los Ríos
60. Heliconia gigantea - Colombia
61. Heliconia gloriosa - Peru
62. Heliconia gracilis - Costa Rica
63. Heliconia griggsiana - Colombia, Ecuador
64. Heliconia harlingii - Ecuador
65. Heliconia hirsuta - Central + South America, Trinidad
66. Heliconia holmquistiana - Colombia
67. Heliconia huilensis - Colombia
68. Heliconia ignescens - Costa Rica, Panama
69. Heliconia imbricata - Costa Rica, Panama, Colombia
70. Heliconia impudica - Ecuador
71. Heliconia indica - Papuasia, Maluku
72. Heliconia intermedia - Colombia
73. Heliconia irrasa - Costa Rica, Panama, Nicaragua
74. Heliconia julianii - N South America
75. Heliconia juruana - Ecuador, Peru, NW Brazil
76. Heliconia kautzkiana - Espírito Santo
77. Heliconia lanata - Solomon Islands
78. Heliconia lankesteri - Costa Rica, Panama
79. Heliconia lasiorachis - Colombia, Peru, NW Brazil
80. Heliconia latispatha - from S Mexico to Peru
81. Heliconia laufao - Samoa
82. Heliconia laxa - Colombia
83. Heliconia lentiginosa - Antioquia
84. Heliconia librata - S Mexico, Central America
85. Heliconia lingulata - Peru, Bolivia
86. Heliconia litana - Imbabura
87. Heliconia longiflora - Colombia, Ecuador, Central America
88. Heliconia longissima - Colombia
89. Heliconia lophocarpa - Costa Rica, Panama
90. Heliconia lourteigiae - South America
91. Heliconia lozanoi - Colombia
92. Heliconia luciae - B Amazonas
93. Heliconia lutea - Panama
94. Heliconia luteoviridis - Colombia
95. Heliconia lutheri - Ecuador
96. Heliconia maculata - Panama
97. Heliconia magnifica - Panama
98. Heliconia × mantenensis - Minas Gerais
99. Heliconia marginata - N South America, S Central America
100. Heliconia mariae - NW South America, Central America
101. Heliconia markiana - Ecuador
102. Heliconia marthiasiae - S Mexico, Central America
103. Heliconia meridensis - Colombia, Venezuela
104. Heliconia metallica - N South America, Central America
105. Heliconia monteverdensis - Costa Rica
106. Heliconia mooreana - Guerrero
107. Heliconia mucilagina - Colombia
108. Heliconia mucronata - Venezuela, NW Brazil
109. Heliconia mutisiana - Colombia
110. Heliconia nariniensis - Colombia, Ecuador
111. Heliconia necrobracteata - Panama
112. Heliconia × nickeriensis - Suriname, French Guiana
113. Heliconia nigripraefixa - Colombia, Ecuador, Panama
114. Heliconia nitida - Colombia
115. Heliconia nubigena - Costa Rica, Panama
116. Heliconia nutans - Costa Rica, Panama
117. Heliconia obscura - Ecuador, Peru
118. Heliconia obscuroides - Colombia, Ecuador, Peru
119. Heliconia oleosa - Colombia
120. Heliconia ortotricha - Colombia, Ecuador, Peru
121. Heliconia osaensis - Colombia, Central America
122. Heliconia paka - Fiji
123. Heliconia paludigena - Ecuador
124. Heliconia papuana - New Guinea
125. Heliconia pardoi - Ecuador
126. Heliconia pastazae - Ecuador
127. Heliconia peckenpaughii - Napo
128. Heliconia pendula - Guiana, Fr Guiana, NE Brazil
129. Heliconia penduloides - Peru
130. Heliconia peteriana - Ecuador
131. Heliconia × plagiotropa - Ecuador
132. Heliconia platystachys - NW South America, S Central America
133. Heliconia pogonantha - NW South America, S Central America
134. Heliconia pruinosa - Peru
135. Heliconia pseudoaemygdiana - Rio de Janeiro
136. Heliconia psittacorum - N South America, Panama, Trinidad
137. Heliconia ramonensis - Costa Rica, Panama
138. Heliconia × rauliniana - Venezuela
139. Heliconia regalis - Colombia, Ecuador
140. Heliconia reptans - Colombia
141. Heliconia reticulata - NW South America, S Central America
142. Heliconia revoluta - Colombia, Venezuela, NW Brazil
143. Heliconia rhodantha - Colombia
144. Heliconia richardiana - NE South America
145. Heliconia rigida - Colombia
146. Heliconia riopalenquensis - Ecuador
147. Heliconia rivularis - São Paulo
148. Heliconia robertoi - Colombia
149. Heliconia robusta - Peru, Bolivia
150. Heliconia rodriguensis - Venezuela
151. Heliconia rodriguezii - Costa Rica
152. Heliconia rostrata - Colombia, Ecuador, Peru, Bolivia
153. Heliconia samperiana - Colombia
154. Heliconia sanctae-martae - Sierra Nevada de Santa Marta
155. Heliconia sanctae-theresae - Antioquia
156. Heliconia santaremensis - Pará
157. Heliconia sarapiquensis - Costa Rica, Panama
158. Heliconia scarlatina - Colombia, Panama, Peru
159. Heliconia schiedeana - Mexico
160. Heliconia schumanniana - Colombia, Ecuador, Peru, N Brazil
161. Heliconia sclerotricha - Ecuador
162. Heliconia secunda - Costa Rica, Nicaragua
163. Heliconia sessilis - Panama
164. Heliconia signa-hispanica - Colombia
165. Heliconia solomonensis - Solomon Islands, Bismarck Archipelago
166. Heliconia spathocircinata - South America, Panama, Trinidad
167. Heliconia spiralis - Colombia
168. Heliconia spissa - S Mexico, Central America
169. Heliconia standleyi - Ecuador, Peru
170. Heliconia stella-maris - Colombia
171. Heliconia stilesii - Costa Rica, Panama
172. Heliconia stricta - N South America
173. Heliconia subulata - South America
174. Heliconia tacarcunae - Panama
175. Heliconia talamancana - Costa Rica, Panama
176. Heliconia tandayapensis - Ecuador
177. Heliconia tenebrosa - Colombia, NE Peru, NW Brazil
178. Heliconia terciopela - Colombia
179. Heliconia thomasiana - Panama
180. Heliconia timothei - NE Peru, NW Brazil
181. Heliconia titanum - Colombia
182. Heliconia tortuosa - S Mexico, Central America
183. Heliconia trichocarpa - Costa Rica, Panama, Colombia
184. Heliconia tridentata - Colombia
185. Heliconia triflora - B Amazonas
186. Heliconia umbrophila - Costa Rica
187. Heliconia uxpanapensis - Veracruz
188. Heliconia vaginalis - Costa Rica, Panama, Colombia, Ecuador
189. Heliconia vellerigera - Ecuador, Peru
190. Heliconia velutina - Colombia, Ecuador, Peru, NW Brazil
191. Heliconia venusta - Colombia, Ecuador
192. Heliconia villosa - Venezuela
193. Heliconia virginalis - Ecuador
194. Heliconia wagneriana - Central America, N South America, Trinidad
195. Heliconia willisiana - Pichincha
196. Heliconia wilsonii - Costa Rica, Panama
197. Heliconia xanthovillosa - Panama
198. Heliconia zebrina - Peru

• 

  Heliconia rostrata, in a botanical garden, Costa Rica

• 

  Heliconia wagneriana in Florida

• 

  Heliconia sp. in tropical rain forest at Sierra del Escambray, Cuba

• 

  Heliconia sp. in tropical rain forest at Sierra del Escambray, Cuba

• 

  Heliconia stricta (Dwarf Jamaican leaf). Location: Maui, Kula Ace Hardware and Nursery

See also

• National Tropical Botanical Garden, designated a conservation center by the Heliconia Society International

External links

• Heliconias of Peru (pdf file with scientific name and photos) in Spanish
• Heliconiaceae in L. Watson and M.J. Dallwitz (1992 onwards). The families of flowering plants: descriptions, illustrations, identification, information retrieval. Version: 27 April 2006. http://delta-intkey.com.
• Monocot families (USDA)
• NCBI Taxonomy Browser
• links at CSDL

References

1. Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III" (PDF). Botanical Journal of the Linnean Society 161 (2): 105–121. doi:10.1111/j.1095-8339.2009.00996.x. Retrieved 2013-06-26.
2. Kew World Checklist of Selected Plant Families
3. Kew World Checklist of Selected Plant Families, genus Heliconia
4. "N esting hab'its and nest symbionts of Polistes erythrocephalus Latreille (Hymenoptera V'espidae) in Costa Rica" (PDF). Biology. . Retrieved 2014-10-14. line feed character in |title= at position 87 (help); External link in |publisher= (help)

1. Berry, Fred and Kress, John. Heliconia Identification Guide. Smithsonian Institution Press, 1991. Print.

2. Bleiweiss, R. 1998. Tempo and mode of hummingbird evolution. Biological Journal of the Linnean Society 65(1): 63-76.

3. Bruna, E. M., W. J. Kress, F. Marques, and O. F. da Silva. 2004. Heliconia acuminata reproductive success is independent of local floral density. Acta Amazonica 34(3): 467-471

4. Dobkin, D. S. 1984. Flowering patterns of long-lived "Heliconia" inflorescences: implications for visiting and resident nectarivores. "Oecologia" ""64""(2): 245-254.

5. Ong, Chong Ren. "Heliconia Basics." Green Culture Singapore. N.p., March 2007. Web. 6 Nov 2011. <http://greenculturesg.com/articles/mar07/mar07_heliconia.htm>.

6. Feinsinger, Peter. 1978. Ecological interactions between plants and hummingbirds in a successional tropical community. Ecological Monographs 48: 269-287.

7. Feinsinger, Peter. 1983. Variable nectar secretion in a Heliconia species pollinated by hermit hummingbirds. Biotropica 15: 48-52.

8. Findley, J.S. and Wilson, D.E. (1974). “Observations on the Neotropical disk-winged bat, Thyroptera tricolor spix.” Journal of Mammalogy 55(3): 563-571.

9. Gilman, Edward, and Alan Meerow. "Heliconia spp. Heliconia." University of Florida IFAS Extension. N.p., 01-05-2007. Web. 25 Sept 2011. <http://edis.ifas.ufl.edu/fp249>.

10. Graham, C. H., J. L. Parra, C. Rahbek, and J. A. McGuire. 2009. Phylogenetic structure in tropical hummingbird communities. "Proceedings of the National Academy of Sciences of the United States of America" ""106"": 19673-19678

11. He, J., C. Chee. C. Goh. ‘Photoinhibition’ of Heliconia under natural tropical conditions: the importance of leaf orientation for light interception and leaf temperature. Plant, Cell, and Environment 19: 1238-1248.

12. Hoii, Karen, Megan Lulow. 2006. Effects of species, habitat, and distance from edge on post-dispersal seed predation in a Tropical Rainforest. Biotropica 29: 459-468.

13. Judd, Walter, et all. Plant Systematics: A phylogenetic approach. 3rd. Sunderland: Sinauer Associates, Inc., 2007. 78, 230, 304, and 367.

14. Kress, W. J. (1985). "Bat Pollination of an Old World Heliconia." Biotropica 17(4): 302-308.

15. Kress, W. J. and C. D. Specht. 2005. Between Cancer and Capricorn: phylogeny, evolution and ecology of the primarily tropical Zingiberales. "Biologiske Skrifter" ""55"": 459-478.

16. Linhart, Yan. 1973. Ecological and behavioral determinants of pollen dispersal in hummingbird- pollinated Heliconia. The American Naturalist 107. 511- 523.

17. McGuire, J. A., C. C. Witt, J. V. Remsen Jr., R. Dudley, and D.L. Altshuler. 2008. A higher-level taxonomy for hummingbirds. "Journal of Ornithology" ""150"": 155-165.

18. Meléndez-Ackerman, E. J., P. Speranza, W. J. Kress, L. Rohena, E. Toledo, C. Cortés, D. Treece, M. Gitzendanner, P. Soltis, and D. Soltis. 2005. Microevolutionary processes inferred from AFLP and morphological variation in Heliconia bihai (Heliconiaceae). "International Journal of Plant Sciences" ""166""(5): 781-794.

19. Seifert, R.P. 1982. Neotropical Heliconia insect communities. The Quarterly Review of Biology 57: 1-28.

20. .Stiles, Gary. 1975. Ecology, flowering phenology, and hummingbird pollination of some Costa Rican "Heliconia" species. "Ecology" ""56"": 285-301.

21. Strong Jr., Donald R. "Insect Species Richness: Hispine Beetles of the Heliconia Latispatha." Ecology. 58. (1977): n. page. Print.

22. Temeles, E. J., J. S. Miller, and J. L. Rifkin. 2010. Evolution of sexual dimorphism in bill size and shape of hermit hummingbirds (Phaethornithinae): a role for ecological causation. "Philosophical Transactions of the Royal Society" ""365"":1053-1063.

23. Timm, R.W. and Mortimer, J. (1976). “Selection of Roost sited by Honduran White Bats, Ectophylla Alba (Chiroptera: Phyllostomatidae).” Ecology 57(2):385-389.

24. Timm, R.W. and Patterson, B.D. (1987). “Tent Construction by bats of the genera Artibeus and Uroderma.” Fieldiana: Zoology 29: 188-212.

25. Uriarte, M. Anciães, M. T.B. da Silva, P. Rubim, E. Johnson, and E. M. Bruna. 2011. Disentangling the drivers of reduced long-distance seed dispersal by birds in an experimentally fragmented landscape. Ecology 92(4): 924-937.