Ericaceae-Neotropical Blueberries
James L. Luteyn and Paola Pedraza-Peñalosa
The New York Botanical Garden


Plant/Animal Interactions


     General information is herein given from field observations or from literature as to probable pollinators and visitation of neotropical ericads (birds, bees, buterflies, and wasps), dispersal agents, hummingbird flower mites, miscellaneous relationships between Ericaceae and ants, and of epiphytic and parasitic relationships; literature references are provided at the end. It must be mention that this section is not complete yet and more information on the Ericaceae/Animal relationship should be searched in the specialized literature.

Probable pollinators and visitors of Neotropical Ericaceae

Plants visited by Hummingbirds (Trochilidae):

Arbutus peninsularis:  Basilinna xantusii (Mexico;  Schuchmann, 1999)

Bejaria racemosa:  Colibri coruscans (Colombia;  Melampy, 1987)

Cavendishia bracteata:  Lampornis calolaema (Costa Rica;  Wm. Busby, pers. comm.);  Adelomyia melanogenys, Chlorostilbon poortmani (Colombia;  Snow & Snow, 1980); Chlorostilbon poortmani, Eriocnemis vestitus, Eriocnemis cupreoventris, Heliangelus amethysticollis, Lafresnaya lafresnayi, Lesbia nuna, Metallura tyrianthina (Clombia;  Snow & Snow, 1980;  plant reported as C. cordifolia); Adelomyia melanogenys, Colibri thalassinus (Ecuador;  Bleiweiss 1103).  [NOTE:  the photos of Coeligena helianthea (Schuchmann, 1999,  p. 480) and Heliangelus strophianus (p. 485) visiting C. bracteata were posed and do not reflect actual situations fide K.-L. Schuchmann, pers. comm.]

Cavendishia capitulata:  Lampornis castaneoventris calolaemus  (Costa Rica;  Wm. Busby, pers. comm.)

Cavendishia chiriquiensis var. chiriquiensis: Lampornis castaneoventris castaneoventris (Panama;  Luteyn et al. 15304)

Cavendishia complectens var. complectens: Lampornis calolaema (Costa Rica;  Wm. Busby, pers. comm.)

Cavendishia guatapeensis:  Adelomyia melanogenys, Boissoneaua flavescens, Coeligena torquata, Doryfera ludoviciae (Colombia;  Snow & Snow, 1980)

Cavendishia lactiviscida:  Doryfera ludovicae (Costa Rica;  Wm. Busby, pers. comm.)

Cavendishia melastomoides var. albiflora: Lampornis castaneoventris.  [NOTE: the photo from Schuchmann (1999, p. 474) was posed and does not reflect an actual situation (Schuchmann, pers. comm.)]

Cavendishia melastomoides var. melastomoides: Lampornis calolaema (Costa Rica;  Wm. Busby, pers. comm.)

Cavendishia pubescens Coeligena torquata, Doryfera ludoviciae (Colombia;  Snow & Snow, 1980);  Amazilia franciae, Amazilia tzacatl (Colombia;  Snow & Snow, 1980)

Ceratostema peruvianum:  Coeligena lutitiae (male) (Ecuador;  Bleiweiss 1010, GH)

Disterigma acuminatum Eriocnemis nigrivestis (male) (Ecuador;  Bleiweiss 1141)

Disterigma aff. acuminatum:  Eriocnemis nigrivestis (male) (Ecuador;  Bleiweiss 1140)

Disterigma alaternoides (or possibly D. cryptocalyx):Adelomyia melanogenys, Boissoneaua  flavescens, Doryfera ludoviciae, Heliangelus amethysticollis (Colombia;  Snow & Snow, 1980); Metallura tyrianthina (Colombia;  Snow & Snow, 1980)

Disterigma empetrifolium:  Chalcostigma stanleyi (F. Ortíz, pers. comm.)

Disterigma humboldtii:  Aglaeocercus coelestris (Ecuador; Sperling & Bleiweiss 5053, QCA)

Disterigma stereophyllum Ocreatus underwoodii incommodus [NOTE: this photo from Schuchmann (1999, p. 490) was posed and does not reflect an actual situation fide K.-L. Schuchmann, pers. comm.]

Gaultheria bracteata Metallura tyrianthina (Peru;  Schuchmann, 1999;  plant reported as  G. rufescens)

Gonocalyx pterocarpus:  Lampornis calolaema (Costa Rica;  Wm. Busby, pers. comm.)

Macleania bullata Coeligena torquata, Doryfera ludoviciae (Colombia;  L. Navarro, pers. comm.)

Macleania coccoloboides:  Eriocnemus sp. (Ecuador;  bird det. by site by Peggy Stern)

Macleania cordifolia:  Adelomyia melanogenys, Colibri thalassinus (Ecuador;  Bleiweiss 1102); Lesbia nuna (Ecuador;  Bleiweiss 1105)

Macleania macrantha:  Coeligena torquata (female), Coeligena lutitiae (female), Eriocnemis  nigrivestis (male; this bird is considered CRITICALLY ENDANGERED.  Cites II) (Ecuador; Bleiweiss 1146)

Macleania pentaptera: Amazilia tzacatl, Ocreatus underwoodii (male), Phaethornis longeumareus, Urosticte benjamini (female) (Ecuador;  photos by M. Krause and bird dets. by  K.-L. Schuchmann)

Macleania rupestris:  Eriocnemis luciani, Eriocnemis mosquera (Ecuador;  Bleiweiss 1125);  Eriocnemis mosquera, Eriocnemis vestitus (Ecuador; Bleiweiss 1014, GH)

Macleania talamancensis one species of hummingbird (Costa Rica;   J. L. Luteyn, pers. observ.)

Oreanthes sperlingii:  Eriocnemis luciani (Ecuador; Sperling & Bleiweiss 5141)

Psammisia falcata:  Coeligena prunellei, Doryfera ludoviciae (Colombia;  Snow & Snow, 1980)

Psammisia incana Heliangelus exortis (male) (Ecuador; Bleiweiss & Young 1008, GH)

Psammisia penduliflora:  Amazilia franciae, Amazilia tzacatl (Colombia;  Snow & Snow, 1980)

Psammisia ramiflora:  Doryfera ludovicae veraguensis (Costa Rica;  Wm. Busby, pers. comm.;  also determination of plant by Luteyn from photo in Schuchmann, 1999, p. 507.  Although this photo was probably posed, the situation actually does exist.)

Psammisia sodiroi:  Ocreatus underwoodii (female), Phaethornis syrmatophorus, Phaethornis yaruqui, Thalurania colombica (male and female) (Ecuador;  photos by M. Krause and bird dets. by K.-L. Schuchmann

Psammisia ulbrichiana:  Phaethornis yaruqui (Ecuador;  source unknown)

Satyria meiantha:  Lampornis calolaema (male) (Costa Rica;  Wm. Busby, pers. comm.)

Satyria warszewiczii:  Lampornis calolaema (Costa Rica;  Wm. Busby, pers. comm.)

Satyria sp. nov. "panamensis":  Lampornis c. castaneoventis (Panama; Luteyn et al. 15296)

Thibaudia andrei:  Adelomyia melanogenys, Aglaiocercus coelestis, Boissonneaua jardini, Coeligena wilsoni (Ecuador; Bleiweiss 1057)

Thibaudia floribunda:  Eriocnemis nigrivestis (male;  this bird is considered CRITICALLY  ENDANGERED.  Cites II) (Ecuador;  Bleiweiss 1139)

Thibaudia inflata:  Urosticte benjamini (female) (Ecuador;  photo by M. Krause and bird dets. by K.-L. Schuchmann)

Thibaudia mellifera:  visited by Metallura hummingbird (Peru;  Boeke & Boeke 3003)

Thibaudia rigidiflora Coeligena prunellei (Colombia;  Snow & Snow, 1980)

Vaccinium poasanum Selasphorus scintilla (Costa Rica;  del Hoyo et al., 1999, p. 507)

Hummingbirds visiting Ericaceae for "food and feeding" (from Schuchmann 1999, unless otherwise stated)

Adelomyia melanogenys (Speckled Hummingbird):  Psammisia

Aglaiocercus coelestis (Violet-tailed Sylph):  Ericaceae

Agyrtria franciae (Andean Emerald): Cavendishia, Psammisia

Androdon aequatorialis (Tooth-billed Hummingbird):  Ericaceae

Augastes geoffroyi (Wedge-billed Hummingbird): Cavendishia, Psammisia

Basilinna xantusii (Xantus's Hummingbird): Arbutus peninsularis

Boissonneaua flavescens (Buff-tailed Coronet): Cavendishia, Disterigma

Calliphlox bryantae (Magenta-throated Woodstar): Cavendishia capitulata, plant species determined by Luteyn from slide by M. and P. Fogden)  [NOTE: this photo was possibly staged (K.-L. Schuchmann, pers. comm.) and may not represent a natural situation.]

Campylopterus falcatus (Lazuline Sabrewing):  Ericaceae

Campylopterus largipennis (Grey-breasted Sabrewing):  Ericaceae

Chaetocercus bombus (Little Woodstar):  Cavendishia, Psammisia, and others.  The bird is considered ENDANGERED (Cites II).

Chalcostigma heteropogon (Bronze-tailed Thornbill): Gaultheria, Pernettya

Chalybura urochrysia (Bronze-tailed Plumeleteer):  Ericaceae

Chlorostilbon alice (Green-tailed Emerald):  Ericaceae

Chlorostilbon stenurus (Narrow-tailed Emerald):  Ericaceae

Coeligena bonapartei (Golden-bellied Starfrontlet): Cavendishia, Macleania

Coeligena coeligena (Bronzy Inca): Cavendishia

Coeligena eos (Golden Starfrontlet): Cavendishia, Macleania

Coeligena helianthea (Blue-throated Starfrontlet): Bejaria, Macleania.  [NOTE: The photo of this hummingbird visiting Cavendishia bracteata (reported as C. cordifolia, see Schuchmann, 1999, p. 480), was posed and does not reflect an actual situation fide K.-L. Schuchmann, pers. comm.]

Coeligena inca (Gould's Inca):  Ericaceae

Coeligena lutetiae (Buff-winged Starfrontlet): Cavendishia

Coeligena prunellei (Black Inca): Psammisia falcata, Thibaudia rigidiflora

Coeligena torquata (Collared Inca): Cavendishia guatapeensis, C. pubescens

Coeligena wilsoni (Brown Inca): Cavendishia, Macleania, Psammisia, Satyria

Damophila julie (Violet-bellied Hummingbird):  Ericaceae

Doryfera ludovicae (Green-fronted Lancebill): Cavendishia, Macleania, Satyria  [The photo of this hummingbird visiting Psammisia ramiflora (plant determined by Luteyn) in    Schuchmann (1999, p. 507), is posed and does not reflect an actual situation fide K.-L. Schuchmann, pers. comm.]

Doryfera johannae (Blue-fronted Lancebill): Psammisia, Satyria

Elvira cupreiceps (Coppery-headed Emerald):  Ericaceae

Eriocnemis cupreoventris (Coppery-bellied Puffleg): Cavendishia, Pernettya

Eriocnemis derbyi (Black-thighed Puffleg):  Ericaceae

Eriocnemis nigrivestis (Black-breasted Puffleg):  Disterigma, Macleania macrantha, Thibaudia floribunda.  This bird is considered CRITICALLY ENDANGERED (Cites II).

Eriocnemis vestitus (Glowing Puffleg):  Ericaceae

Eupherusa eximia (Stripe-tailed Hummingbird):  Ericaceae

Eupherusa nigriventris (Black-bellied Hummingbird):  Ericaceae

Goethalsia bella (Pirre Hummingbird):  Ericaceae

Goldmania violiceps (Violet-capped Hummingbird): Psammisia

Heliangelus amethysticollis (Amethyst-throated Sunangel):  Cavendishia, Disterigma

Heliangelus exortis (Tourmaline Sunangel): Psammisia incana (Naskrecki & Colwell, 1998).

Heliangelus strophianus (Gorgeted Sunangel): Cavendishia bracteata [NOTE:  this photo from Schuchmann (1999, p. 485) was posed and does not represent an actual situation fide K.-L. Schuchmann, pers. comm.]

Heliodoxa imperatrix (Empress Brilliant):  Ericaceae

Heliodoxa jacula (Green-fronted Brilliant):  Ericaceae

Heliodoxa schreibersii (Black-throated Brilliant):  Ericaceae

Heliothryx barroti (Purple-crowned Fairy): Cavendishia

Klais guimeti (Violet-headed Hummingbird): Cavendishia

Lafresnaya lafresnayi (Mountain Velvetbreast):  Ericaceae

Lampornis castaneoventris (Variable Mountain-gem):  Ericaceae (Satyria sp., detetmination of  bird by  B. Riggs, plant voucher Luteyn et al. 15296)  [NOTE: the photo of this hummingbird visiting Cavendishia melastomoides var. albiflora Luteyn seen in Schuchmann (1999, p. 474) was posed and does not reflect an actual situation fide K.-L. Schuchmann, pers. comm.]

Lampornis hemileucus (White-bellied Mountain-gem):  Cavendishia, Thibaudia

Lesbia nuna (Green-tailed Trainbearer): Cavendishia bracteata (reported as C. cordifolia)

Mellisuga minima (Vervain Hummingbird): Vaccinium

Metallura baroni (Violet-throated Metaltail): .  The bird is considered VULNERABLE (Cites II).

Metallura theresiae (Coppery Metaltail):  Macleania

Metallura tyrianthina (Tyrian Metaltail):  Ericaceae [including Gaultheria bracteata (reported as  G. rufescens)]

Metallura williami (Viridian Metaltail):  Ericaceae

Microchera albocoronata (Snowcap): Cavendishia

Ocreatus underwoodii (Booted Racquet-tail): Macleania pentaptera, Psammisia sodiroi [NOTE:  the photo of this hummingbird visiting Disterigma stereophyllum in Schuchmann (1999, p. 490) was posed and does not represent an actual situation (Schuchmann, pers. comm.)]

Opisthoprora euryptera (Mountain Avocetbill):  Ericaceae

Panterpe insignis (Fiery-throated Hummingbird): Vaccinium

Phaethornis yaruqui (White-wiskered Hermit):  Ericaceae

Phlogophilus harterti (Peruvian Piedtail):  Ericaceae

Phlogophilus hemileucurus (Ecuadorian Piedtail): Psammisia

Polyerata rosenbergi (Purple-chested Hummingbird): Macleania, Psammisia

Polytmus milleri (Tepui Goldenthroat):  Ericaceae

Ramphomicron dorsale (Black-backed Thornbill):  Ericaceae

Ramphomicron microrhynchum (Purple-backed Thornbill):  Ericaceae

Selasphorus flammula (Volcano Hummingbird): Vaccinium

Selasphorus scintilla (Scintillant Hummingbird): Vaccinium poasanum  [NOTE: this photo in Schuchmann (1999, p. 507) was probably posed and does not represent an actual situation (Schuchmann, pers. comm.]

Thalurania colombica (Purple-crowned Woodnymph): Psammisia sodiroi

Thalurania fannyi (Green-crowned Woodnymph):  Ericaceae

Thalurania furcata (Fork-tailed Woodnymph):  Ericaceae

Thalurania ridgwayi (Mexican Woodnymph):  Ericaceae.  This bird is considered VULNERABLE (Cites II).

Topaza pella (Crimson Topaz):  Ericaceae

Urochroa bougueri (White-tailed Hillstar): Cavendishia, Psammisia

Urosticte benjamini (Purple-bibbed Whitetip): Thibaudia inflata

Other hummingbirds that visit Ericaceae flowers in Ecuador include:  Coeligena torquata, Eriocnemis luciani, Lesbia victoriae, and Metallura tyrianthina (F. Ortíz, pers. comm.).

Plants visited by nectar-robbing birds

Cavendishia bracteata:  Diglossa albilatera, Diglossa carbonaria (Colombia;  Snow & Snow, 1980)

Disterigma acuminatum:  Diglossa lafresnayii (Ecuador; Bleiweiss 1141)

Disterigma aff. acuminatum:  Diglossa lafresnayii (Ecuador; Bleiweiss 1140)

Disterigma humboldtii:  Diglossa duidae (Venezuela; S. Renner, pers. comm.)

Macleania bullata:  Diglossa cyanea (Colombia;  L. Navarro, pers. comm.)

Psammisia falcata Diglossa albilatera (Colombia;  Snow & Snow, 1980).  At this site Aglaiocercus kingi also took nectar, but "illegitimately" through pierce holes (Snow & Snow, 1980).

Psammisia penduliflora:  Coereba flaveola (Colombia;  Snow & Snow, 1980)

Thibaudia rigidiflora:  Diglossa albilatera (Colombia;  Snow & Snow, 1980)

Plants visted by Bees:


Bejaria aestuans:  "honey bees" (Ecuador;  Clemants, 1995)

Bejaria racemosa:  Apis melifera (Ecuador;  Clemants, 1995)

Bejaria sprucei:  Centris flavifrons (oil-collecting bee) (Venezuela;  Clemants, 1995)

Cavendishia lebroniae:  black or dark brow, very large bumble bee-like (Ecuador;  Luteyn, pers. observ.)

Cavendishia martii:  Trigona (Trigona) hyalinata var. amazonensis (Peru;  Luteyn & Lebrón-Luteyn 6368 and pers. observ.)

Disterigma alaternoides (or possibly D. cryptocalyx):  (Colombia;  Snow & Snow, 1980)

Disterigma humboldtii:  pollinated by an oil-collecting bee Centris sp. (subg. Melanocentris) (Venezuela;  S. Renner, pers. comm.)

Pernettya prostrata:  Bombus ephippiatus (Costa Rica;  R. Heithaus, 1983)

Psammisia dolichopoda:  Trigona sp. (Ecuador; Madison 3931)

Plants visited by Butterflies:

 Cavendishia pubescens one species of butterfly (Colombia;  J. L. Luteyn, pers. observ.)

Macleania bullata:  one species of butterfly (Colombia;  L. Navarro, pers. comm.)

Plants visited by Wasps:


Bejaria sprucei:  Scolia sp. (Venezuela;   Clemants, 1995)

Cavendishia pedicellata:  Polybia diguetana (social wasp) (Colombia;  Luteyn et al. 7308 and pers. observ.)


Ericaceae whose fruits are eaten Birds

Cavendishia bracteata:  Diglossa caerulescens (E Andes of Colombia;  D. Snow, pers. comm.)

Cavendishia engleriana var. engleriana Chlorospingus semifuscus, Euphonia xanthogaster (Colombia; C. Restrepo, unpublished)

Cavendishia tarapotana var. gilgiana:  Euphonia xanthogaster, Semnornis ramphastinus (Colombia; C. Restrepo, unpublished)

Diogenesia tetranda:  Semnornis ramphastinus (Colombia; C. Restrepo, unpublished)

Gaultheria glomerata fruits eaten by birds (Ecuador;  herbarium specimen source unknown)

Macleania stricta:  Chlorospingus semifuscus, Euphonia xanthogaster (Colombia;  C. Restrepo, unpublished)

Psammisia cf. columbiensis:  Euphonia xanthogaster, Masius chrysopterus (Colombia;  C. Restrepo, unpublished)

Satyria grandifolia:  Atlapetes brunneinucha, Euphonia xanthogaster, Lipaugus cryptolophus, Mionectes striaticollis, Myadestes ralloides, Pipreola riefferi, Semnornis ramphastinus (Colombia; C. Restrepo, unpublished)

Sphyrospermum cordifolium:  Euphonia xanthogaster, Pipreola riefferi (Colombia;  C. Restrepo, unpublished)

Thibaudia inflata:   fruit eaten by the bird "pava silvestre" (Ecuador;  Cerón M. 6900)

The two most important dispersal agents of Ericaceae at Monteverde (Costa Rica) include the Emerald Toucanets and Black-faced Solitaires (G. Murray, pers. comm.).

Miscellaneous relationships between Ericaceae and Ants

Cavendishia colombiana: ants eating fruits, Acromyrmex (A.) hystrix (Colombia; Luteyn & Lebrón-Luteyn 6995 and pers. observ.)

Cavendishia lebroniae:  ants eating fruits, black, non-biting (Ecuador;  Luteyn & Lebrón-Luteyn 5759)

Satyria sp.:  ants carrying corollas to nest (Panama;  J. L. Luteyn, pers. observ.)

Thibaudia martiniana:  feeding on nectar (Ecuador; Luteyn & Borchsenius 13337) & J. L. Luteyn, pers. observ.)

Hummingbird flower Mites

      Colwell (1973, 1985) described in detail the fascinating story about mites that spend their lives within the flowers of certain hummingbird flowers, such as Ericaceae.  These same mites, however, are dispersed from flower to flower within the nasal passages of hummingbirds.  In 1998, Naskrecki and Colwell monographed mites of the genera Tropicoseius and Rhinoseius (Acari: Mesostigmata: Ascidae) and found that rhinoseius mites apparantly evolved within the flowers of neotropical Ericaceae in montane habitats.  The following is a list of those mites and the ericads they live in (country of origin and herbarium voucher collection are cited in parentheses).

Rhinoseius androdon: Cavendishia coccinea (Colombia, Luteyn 4787)

Rhinoseius antioquiensis: Cavendishia lindauiana (Colombia, Luteyn & Lebrón-Luteyn 7166); Cavendishia palustris (Ecuador, Luteyn 14587)

Rhinoseius haplophaedia: Anthopterus verticillatus (Ecuador, Luteyn 5650); Cavendishia isernii var. pseudospicata (Ecuador, Luteyn 5674); Disterigma pentandrum (Ecuador,  Luteyn 14791); Macleania coccoloboides x M. cf. ericae (Ecuador, Luteyn 8474); Macleania cf. ericae (Ecuador, Luteyn 14789); Macleania stricta (Ecuador, Luteyn 13339)

Rhinoseius luteyni: Psammisia incana (Ecuador, Luteyn 13393)

Rhinoseius nadachowskyi: Cavendishia ruiz-teranii (Venezuela, Luteyn et al. 6099);  Ceratostema reginaldii (Ecuador, Luteyn 14708); Macleania puberula (Ecuador, Luteyn 14712);  Plutarchia angulata (Colombia, Luteyn 5020 & 6786);  Sphyrospermum buxifolium (Ecuador, Luteyn 14400);  Thibaudia floribunda (Ecuador, Luteyn 14787)

Rhinoseius pastorae: Ceratostema peruvianum (Ecuador, Luteyn 14697)

Rhinoseius richardsoni: Cavendishia nobilis var. capitata (Ecuador, Dorr & Barnett 5907);  Cavendishia cuatrecasasii (Ecuador, Luteyn 5677); Cavendishia subamplexicaulis (Colombia, Luteyn 7065); Cavendishia sp. (Costa Rica, Colwell s.n.);  Macleania pentaptera    (Ecuador, Luteyn 14719); Macleania rupestris (Costa Rica, Colwell s.n.);  Macleania stricta (Colombia, Luteyn et al. 12414);  Psammisia sp. (Colombia, Luteyn 12327); Vaccinium floccosum (Panama, Luteyn 3756)

Rhinoseius rafinskii: Vaccinium cf. floribundum (Ecuador;  plant reported as V. corymbosum  a species that does not occur in S. America)

Rhinoseius ucumariensis: Vaccinium mathewsii (Peru, Luteyn 11382)

Tropicoseius steini: Ceratostema peruvianum (Ecuador, Luteyn 14697) 

Epiphyte hosts

     The evergreen, coriaceous leaves of many Ericaceae provide an excellent substrate for mosses, hepatics, lichens, and other fungi.  Also glands and glandular areas on the stems, leaf surfaces, and inflorescences are often covered by fungal mycelia that probably derive nutrients from the exudates. Fungal mycelia of the inoperculate discomycete Lachnum sp. (Hyaloscyphaceae; G. Samuels and P. Johnston, pers. observ.) have been found growing on the lower leaf surfaces of Gaultheria strigosa var. strigosa (syn. G. arachnoidea).  In fact, G. arachnoidea was characerized by its "unique pale arachnoid indumentum of young branchlets and lower leaf surfaces" (Smith, 1933).  The "unique" arachnoid indumentum is only the fungal mycelia.
     The fungus Ophiostoma sp. nov. grows on the type collection of Orthaea paniculata in Mexico (C. Rogerson, pers. comm.).
     The ascomycete fungus Gibbera sphyrospermi (Stigmateaceae) was described from Sphyrospermum cordifolium (G. Samuels, pers. comm.).


     Members of the plant family Loranthaceae frequently parasitize Ericaceae.  Among them are Dendrophthora costaricensis subsp. poasensis on Cavendishia bracteata in Costa Rica, D. chryostachya on Cavendishia sp. from Peru, and D. lindeniana on C. bracteata from Colombia (Kuijt, 1961).


Clemants, S. E.  1995.  Bejaria.  Fl. Neotrop. Monogr. 66: 54-106

Colwell, R. K.  1973.  Competition and coexistence in a simple tropical community.  American Nat. 107: 737-760.

Colwell, R. K.  1985.  Stowaways on the hummingbird express.  Natural History 7/85: 57-63.

Hiethaus, R.  1983.   Bombus ephippiatus (Chiquizá de Montaña, Bumblebee).  Pp. 700-701. In:  Janzen, D. H. (ed.), Costa Rican natural history.  University of Chicago Press, Chicago.

Kuijt, J.  1961.  A revision of Dendrophthora (Loranthaceae).  Wentia 6: 1-145.

Luteyn, J. L.  1976.  A revision of the Mexican-Central American species of Cavendishia (Vacciniaceae).  Mem. New York Bot. Gard. 28: 1-138.

Luteyn, J. L.  1983.  Cavendishia.  Fl. Neotrop. Monogr. 35: 1-290.

Melampy, M. N.  1987.  Flowering phenology, pollen flow and fruit production in the Andean shrub Befaria resinosa.  Oecol. 73: 293-300.

Middleton, D. J.  1991.  Ecology, reproductive biology and hybridization in Gaultheria L. Edinburgh J. Bot. 48(1): 81-89.

Naskrecki, P. & R. K. Colwell.  1998.  Systematics and host plant affiliations of hummingbird flower mites of the genera Tropicoseius Baker & Yunker and Rhinoseius Baker & Yunker (Acari: Mesostigmata: Ascidae).  Monographs Thomas Say Publications in Entomology.     Entomological Society of America,

Santesson, R.  1952.  Foliicolous lichens. I.  Symb. Bot. Upsal. 12: 1-590.

Schuchmann, K.-L.  1999.  Hummingbird chapter, in J. del Hoyo, A. Elliott & J. Sargatal (eds.), Handbook of the Birds of the World. Vol. 5.  Barn-owls to Hummingbirds.  Lynx Edicions, Barcelona.

Smith, A. C.  1933.  Studies of South American plants, III.  New Ericaceae and Vacciniaceae.  Bull. Torrey Bot. Club 60: 99-121.

Snow, D. W. & B. K. Snow.  1980.  Relationships between hummingbirds and flowers in the Andes of Colombia.  Bull. British Mus. (Nat. Hist.), Zool. ser., 38(2): 105-139.