Flora 255 (2019) 80–85

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Flora
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Bird visitation to a high Andean Anthurium (Araceae) in Eastern Ecuador T

a,⁎ b,1 c d
Robert Bleiweiss , Francisco Sornoza Molina , Efraín Freire , Thomas B. Croat
a
Department of Integrative Biology and the Zoological Museum, Room 441, Birge Hall, 430 Lincoln Drive, University of Wisconsin, Madison, WI, 53706, USA
b
Fundación Jocotoco, Lizardo García E9-104 y Andrés Xaura, Quito, Ecuador
c
Herbario Nacional del Ecuador, Museo Ecuatoriano de Ciencias Naturales del Instituto Nacional de Biodiversidad, Av. Río Coca E6-115, Quito, Ecuador
d
P. A. Schulze Curator, Missouri Botanical Garden, P.O. Box 299, St. Louis, MO, 63166-0299, USA

ARTICLE INFO ABSTRACT

Edited by Timotheus van der Niet The limited information available on aroid reproductive systems suggests that most species are visited and
Keywords: pollinated by arthropods, especially bees, beetles, and flies. However, hummingbirds and passerine birds have
Cloud forest been reported to visit some Colombian Anthurium that retain many typical aroid traits. Observations in upper
Hummingbird montane cloud forests (2700 m) on the eastern slope of the Andes in northern Ecuador revealed a third
Flowerpiercer Anthurium species visited by birds. Male-phase inflorescence features that seemed to favor bird visitation in-
Pollen cluded a robust and all red inflorescence, and copious fluid secretions that appeared to have nutritive value. Over
Red coloration a weeklong period, at least four species of hummingbird and one passerine bird, a flowerpiercer, were observed
Thrips
to pick up pollen when they fed on the fluid secretions during repeated visits to a male-phase inflorescence. The
bird-plant interaction depended on species-specific foraging methods, rank in the local dominance hierarchy,
and diel activity patterns. In addition, several arthropod taxa were documented as Anthurium visitors for the first
time. We consider the potential for a bird or a mixed pollination system of this Anthurium.

1. Introduction
The conserved reproductive biology of Araceae raises fundamental
issues about the role of pollinators in the extraordinary diversification
of this plant family (Croat, 1980; Gibernau, 2003; Henriquez et al.,
2014). The characteristic aroid inflorescence comprises a large
sheathing bract (spathe) that supports or partly encloses a floral spike
(spadix) along which are arrayed numerous tiny white to green flowers
that often emit volatile attractants (Gibernau, 2003). Field studies
support inferences from this morphology that the main aroid pollinators
are typically arthropods (Croat, 1980, Gibernau 2003, 2011, 2016;
Gibernau et al., 2010), which contrasts with the diversity of pollinators
recorded in other taxon-rich and widespread plant families (Gorelick,
2001; van der Niet and Johnson, 2012). However, the pollination sys-
tems of only a tiny fraction of aroid species have been studied in any
detail. Notably, recent investigations suggest that even subtle variations
on the basic aroid inflorescence form constitute specializations for very
different arthropod pollinators, distinguishing species adapted for pol-
lination by bees, beetles, or flies (Croat, 1980; Gibernau, 2011;
Gibernau et al., 2010; Prieto and Cascante-Marín, 2017). Thus, the true
diversity of aroid visitors and pollination systems is probably
underestimated.
In this regard, the possible association of aroids with bird visitors is
of particular interest. Although the various forms of aroid reproductive
structures are atypical for bird-pollinated plants, aroid flowers secrete
nectar-like fluids indicating potential pollination by animals with
higher energy requirements than arthropods (Daumann, 1931; Vogel,
1983; Kraemer and Schmitt, 1999). Indeed, reports by early naturalists
that birds sometimes visit these plants (reviewed in Brown, 1988) have
been confirmed by two more recent reports. In the Andean foothills of
western Colombia, the hummingbird Aglaiocercus coelestis was photo-
graphed sitting on the red spathe of an unidentified Anthurium
(Salaman, 1996). Elsewhere in the same Andean range, eight species of
hummingbird and two species of passerine bird (Diglossa albilatera
flowerpiercers, Anisognathus flavinucha tanagers) were observed to visit
male-phase inflorescences of Anthurium sanguineum, a large epiphytic
species with erect green spadices subtended by bright red spathes
(Kraemer and Schmitt, 1999). This system revealed important new
details regarding bird-aroid interactions by documenting that the bird
species accumulated pollen loads during their frequent visits to feed on
the copious, sugared secretions produced by the male-phase spadices of
this aroid. Thus, use of aroids by birds appears to extend beyond mere

Corresponding author.
⁎

E-mail addresses: rebleiwe@wisc.edu (R. Bleiweiss), franciscosornoza@ecociencia.org (F. Sornoza Molina), efrain.freire@biodiversidad.gob.ec (E. Freire),
thomas.croat@mobot.org (T.B. Croat).
1
Present address: Fundación EcoCiencia, San Ignacio E12-143 y Humboldt Edf. Carmen Lucía, Quito, Ecuador.

https://doi.org/10.1016/j.flora.2019.02.010
Received 10 August 2018; Received in revised form 9 February 2019; Accepted 20 February 2019
Available online 27 March 2019
0367-2530/ © 2019 Elsevier GmbH. All rights reserved.
R. Bleiweiss, et al. Flora 255 (2019) 80–85

Fig. 1. Female (A) and incipient male (B, C) phases of the Guango Anthurium inflorescence Typically, a recently opened inflorescence had an erect spathe (A, arrow 1)
oriented parallel to a spadix that produced small droplets of stigmatic fluid over its entire length. Pollen and fluid production began at the base of the spadix (B, arrow
1; C arrow 2), by which time the spathe was often bent over (B, arrow 2). Both hymenoptera (A, arrow 2) and thrips (C, arrow 1; barely visible in A and B) were
present at these and later inflorescence phases.

casual associations in some cases. 2. Materials and methods

While conducting fieldwork in upper montane cloud forest on the
eastern slope of the Andes in northern Ecuador, several hummingbird
and one flowerpiercer species were observed visiting a distinctive
Anthurium with mature male-phase inflorescences in which both the
spathe and spadix were entirely red. Inflorescence form therefore dif-
fered from the other two aroids visited by birds, providing a third ex-
ample of the association. We report field observations of the birds and
arthropods visiting the Ecuadorian Anthurium, present evidence that
birds picked up pollen during their frequent visits to feed on the copious
fluid secretions produced by male-phase inflorescences, describe simi-
larities and differences in foraging methods among bird species, and
discuss the possibility that this Anthurium employs a bird or a mixed
bird-arthropod pollination system.
2.1. Study organism and study site
Reviews of information provided in the Flora of Ecuador, the
Herbario Nacional del Ecuador (QCNE), and our own electronic data-
base (Croat, Lucid Anthurium Key that includes more than 1550 spe-
cies), indicate that the Anthurium is a distinctive new species belonging
to the section Cardiolonchium. The plant is hereafter simply referred to
as the “Guango Anthurium”, pending its formal description. The plant
was discovered at 2700 m in humid montane cloud forest at Guango
Lodge (0°22′46.56″ S, 78°4′36.12″ W), which is located approximately
4.8 km west of the larger town of Cuyuja, in Napo Province Ecuador.
Visitors to the plants were recorded during a total of 35 person-hours
spread out from January 25 to 31, 2016. During the study period, days
were clear and virtually cloudless and without rain except for one

81
R. Bleiweiss, et al.
afternoon shower on January 28th.
2.2. Plant growth
The Guango Anthurium grew as a hemi-epiphyte in the forest sub-
canopy. Two plants were discovered, located approximately 10 m apart,
and their different flowering phases indicated protogynous develop-
ment typical of the genus. The dry weather suggested that fluids de-
tected on the surface of spadices were secretions produced by the plant
rather than condensate or rain. When discovered, one plant had one
male-phase inflorescence producing pollen and fluid, one unopened and
one incompletely opened inflorescence, and two fruiting structures. The
other plant had five unopened inflorescences, two more in the female
phase, and five others in various stages of fruiting. All colors related to
inflorescence development were assessed visually.
2.3. Animal visitation
Observations centered on the plant (hereafter, the “focal plant”)
with the male-phase inflorescence, especially at the male-phase in-
florescence itself (hereafter, the “focal inflorescence”). An inventory of
visitors was conducted both visually (including with binoculars), and
through digital imaging (for micro-fauna) during the 35 h spent
watching visitors. All images of visitors were captured with a Canon
f5.6 55–250 mm zoom lens on a Canon EOS Digital Rebel XTi camera
set to ISO 400, AI Servo AF Mode, auto white balance, and sRGB color
space. All photos were taken by Robert Bleiweiss.
3. Results
3.1. Plant growth
Inflorescence structures of the Guango Anthurium were large and
robust (spathe: 192.5 mm, N = 4; spadix: 184.5 mm, N = 4). Prior to
unfurling, each inflorescence closely resembled a slender red candle,
complete with a wick-like tip. The spathe remained red after separating
from the spadix (Fig. 1A). The spadix appeared greenish-orange in-
itially, and produced small droplets of stigmatic fluid along its entire
length (Fig. 1A). Pollen production began at the spadix base (Fig. 1B),
at which time the spathe either reflexed (Fig. 1B) or remained parallel
to the inflorescence as a crescent-shaped bract (Fig. 2). Over time, the
male-phase spadix turned bright red and produced copious amounts of
fluid, especially in a transition zone where new male-phase flowers
were opening (Figs. 1C; 2). Reddening encompassed the entire spadix as
male-phase flowers opened apically. Flower sexual phases did not ap-
pear to overlap on a spadix, but fruiting began before the end of the
male phase. Coincident with fruiting, the spadix continued to elongate
and gradually lost red coloration.
3.2. Bird visitation
Bird visitors to the focal plant included four hummingbird species
(Trochilidae) and a passerine bird, the flowerpiercer Diglossa cyanea
(Thraupidae) (Table 1). The focal inflorescence was located within the
feeding territory of the hummingbird Boissonneaua matthewsii. The
latter always won contests against the other bird visitors, which in-
cluded both other hummingbirds [Heliangelus exortis (10 times),
Aglaiocercus kingi (2 times), Adelomyia melanogenys (1 time)] and a
flowerpiercer [Diglossa cyanea (1 time)]. Social dominance by Boisson-
neaua appeared to dictate choice of inflorescence and foraging method
by all species. Compared to other bird visitors, Boissonneaua was ob-
served to feed only at the most productive, focal inflorescence, and to
forage at the spadix only from its perch on the spathe (Fig. 2A). Bois-
sonneaua took some time to settle on its perch; its feeding bouts pro-
ceeded uninterrupted, but it often interrupted visits made by sub-
ordinate species to the focal plant. The remaining species typically
82
Flora 255 (2019) 80–85
perched directly on the spadix and began to feed immediately, either at
the focal inflorescence (Heliangelus, Diglossa; Fig. 2B) or at other in-
florescences (Adelomyia and Aglaiocercus) on the focal plant that were
weighted down to a horizontal position by ripening fruit, and partly
hidden from the resident Boissonneaua by vegetation and woody
growth. Infrequently, hummingbird visitors hovered at the focal in-
florescence, either to (apparently) inspect (Boissonneaua) or feed (He-
liangelus) off the spadix (Table 1).
3.3. Fluid production and bird foraging behaviors
Inspection of the focal inflorescence over time and space indicated
the presence of extensive fluid secretions (Fig. 3A) as early as first light
(˜7 AM). The amount of fluid was greatest in the transition zone
(Fig. 3A) of nascent, male-phase flowers located between regions with
older (more basal), and unopened (more apical) male-phase flowers
(Fig. 3A). Failure to exclude visitors from feeding made estimates of
production rates difficult; however the amount of fluid appeared only
slightly lower early and late, suggesting relatively constant production
during most of the day. Hummingbird visits to the focal plant increased
through the morning, and stopped after noon (Table 2). Diglossa visited
at low frequencies throughout the day, suggesting more opportunistic
patterns.
3.4. Plant-animal interactions
Pollen and fluid uptake by birds was supported in several ways.
Birds arrived at the focal plant with clean bills, but picked up pollen on
various bill surfaces during their visits (Fig. 3B–E). Digital images and
video clips indicated that birds used their tongues to feed on fluids
secreted by the male-phase spadix of the focal inflorescence (Fig. 3B,
D). Moreover, the amounts of fluid on this spadix appeared to diminish
over the course of the main feeding activities during the morning hours
(compare Fig. 3A, C). Regardless of social status, bird visitors to the
focal inflorescence appeared to feed mainly in the transition zone,
where both nascent pollen and fluid production was greatest (Figs. 2
and 3). However, bird visitors sometimes also scraped their bills side-
ways along the long axis of the spadix, suggesting that they were re-
moving as much of the fluid as they could during a visit (especially
characteristic of Diglossa; Fig. 3D). Either feeding method resulted in
the deposition of considerable quantities of pollen, usually at or near
the tip of the bird’s bill (Fig. 3). The birds’ tongues also presumably
contacted male-phase flowers, so pollen itself cannot be entirely ruled
out as a protein and nutrient source, particularly because the birds were
molting at the time of flowering. Species that fed while perched on the
spadix (see above) may have also picked up pollen on their plumage
and feet (Fig. 2).
3.5. Arthropod visitation
Birds were recorded as more frequent visitors than arthropods to the
focal plant (Tables 1 and 2). The latter comprised an unidentified wasp
(Hymenoptera) with a distinctive pigmentation pattern on its wings
(Fig. 1A), and numerous thrips (Thysanoptera). Both of these ar-
thropods visited or were present along the spadix at various develop-
mental phases (Table 1; Fig. 1). However, arthropod foraging on pollen
or fluids produced by the spadix could not be determined.
4. Discussion
Upper montane cloud forests are among the least studied Andean
environments. Our observations add to a few earlier reports (summar-
ized in Kraemer and Schmitt, 1999) that aroids are important plants for
the activities of nectar-feeding birds in these habitats. While much re-
mains to be learned about the nature of these bird-aroid associations,
our observations suggest that montane aroids can be important
R. Bleiweiss, et al. Flora 255 (2019) 80–85

Fig. 2. Typical feeding methods adopted by the hummingbird Boissonneaua matthewsii (A) and the flowerpiercer Diglossa cyanea (B) at the Guango Anthurium. All
visitors focused on the fluid-rich transition zone, where male-phase flowers (white) were beginning to open. However, feeding tactics and strategies differed among
bird visitors (see Results). Shown is the focal inflorescence, which is in the male phase.

resources for resident nectarivores. We were unable to measure the
sugar content and caloric reward of the fluid secretions present on the
surface of female-phase and male-phase spadices. However, persistent
visits by birds to male-phase inflorescences, and evidence that birds
used their tongues to feed on inflorescence secretions rather than on
other potential resources there (arthropods), suggests that the fluids
most likely had nutritive value. Moreover, Kraemer and Schmitt (1999)
measured appreciable glucose (up to 12% in standing crop) in the male-
phase spadix secretions of the related A. sanguineum. Pending evidence
that the male-phase inflorescences of the Guango Anthurium secrete
fluids that also qualify as nectar, we tentatively conclude that these
fluids are at least nutritious for birds. Thus, our observations are con-
sistent with the suggestion of Kraemer and Schmitt (1999) that the
presence of nutritive fluids accessible on the surface of an exposed
spadix encourages visits to certain aroids by a diverse assortment of
birds. Moreover, the associations of birds specifically with the genus
Anthurium in all three recent examples of birds feeding at aroids are
unlikely to be coincidental given that many other, often dissimilar aroid
genera also grow in the Andes (Henriquez et al., 2014; Sierra-Giraldo
et al., 2013; Vargas et al., 2004). Although the general morphology of
the Guango Anthurium inflorescence evokes little potential for or-
nithophily in the context of conventional pollination syndromes, we
here consider several lines of evidence that this Anthurium might be
pollinated, at least in part, by its bird visitors.
First, we obtained direct evidence that birds picked up large quan-
tities of pollen on their bills when they visited male-phase in-
florescences of the Guango Anthurium to feed on the fluid secretions.
This basic observation is consistent with a mutualistic relationship in

Table 1
Visitors to inflorescences of the Guango Anthurium.
Inflorescence Phasea Visitor N

Early (female), N=1 Middle (male), N=23 Late (male), N=3

b
Method H ST SX H ST SX H ST SX
Taxonc
Hummingbirdsd
Boissonneaua − − − 2 7 − 1 − − 10
Heliangelus − − − 2 − 4 − − − 6
Adelomyia − − − − − − − − 1 1
Aglaiocercus − − − − − − − 1 1 2
Passerines
Diglossa − − − − − 6 − − − 6
Insects
Hymenoptera − − 1 − − 2 − − − 3
Thysanopterae − − + − − + − − + many

a
Inflorescence phases: early (spadix just unfurling), middle (pollen at spadix base), late (pollen along most of spadix length).
b
Feeding tactics: H (hover), ST (perch on spathe), SX (perch on spadix).
c
Full species names identified in text.
d
Hummingbirds observed feeding in the area but not at the Guango Anthurium included Boissonneaua flavescens, Heliodoxa rubinoides, Coeligena torquata, Ensifera
ensifera, and Metallura tyrianthina.
e
Thrips (Thysanoptera) were present on the spadix at least from the time it was uncovered, through the fluid-producing phase.

83
R. Bleiweiss, et al. Flora 255 (2019) 80–85

Fig. 3. Bird-plant interactions at the Guango Anthurium. Male-phase spadix ontogeny (A) indicates copious fluid production in the transition zone (arrow 2) between
unopened (arrow 1) and opened (arrow 3) male-phase flowers. Cameos of pollen and fluid uptake by the hummingbird Boissonneau matthewsii (B and C) and the
flowerpiercer Diglossa cyanea (D and E) indicate roles of bill and tongue (arrows) in acquisition of these plant resources.

which both bird and plant benefit from their interaction. An important
caveat to our observations is that pollination requires that birds loaded
with pollen also visit female-phase inflorescences; otherwise what we
observed could be interpreted as floral parasitism and a reduction in
fitness of male-phase flowers through loss of reproductive effort
(pollen, nutritive fluids). Although we did not observe unequivocal
visits to any inflorescence in the female phase, stigmatic fluid (Fig. 1A)
could be attractive and just as accessible to mobile and opportunistic
nectar-feeding birds (see also Kraemer and Schmitt, 1999). Thus, pol-
lination by birds remains a likely possibility.
Second, the economics of foraging provide indirect evidence that
birds can be reliable visitors to the Guango Anthurium beyond what we
observed. Boissonneaua’s priority of access to the focal inflorescence
appeared to confer both benefits and costs to all bird visitors. Thus,
territorial defense of this Anthurium by the resident Boissonneaua im-
plied that this species obtained benefits from feeding that outweighed
its costs of interference competition. Similarly, persistent attempts by
subordinate species to visit the plant suggested that they also obtained
foraging benefits that outweighed the costs of attacks by the territorial
resident. At the same time, interspecific social interactions may com-
plicate the dynamics of pollen flow within and between plants. The
strong social dominance expressed by Boissonneaua presumably allowed
this species to be resident around the single most productive male-
phase inflorescence, and to perch uninterrupted on the erect spathe so
that only its bill and tongue contacted the spadix. By comparison,
subordinate species typically made quick landings on the spadix before
departing, but doing so may have allowed pollen to be deposited on the
plumage and feet, not just the bill and tongue. These differences in
feeding strategies and tactic could thereby limit outcrossing by
Boissonneaua, but enhance that role by subordinates.
Third, the Guango Anthurium expressed typical attractants for birds,
such as red coloration and copious nutritive fluid production (Figs. 2

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R. Bleiweiss, et al. Flora 255 (2019) 80–85

Table 2 Declaration of interest

Temporal visitation patterns at inflorescences of the Guango Anthurium.
Time of Daya Visitor N None.

Earlyb Middle Late Acknowledgements

(AM) 7 8 9 10 11 12 1 2 3+ (PM)
Taxonc (Totals)
We extend our gratitude to Irene Bustamante, proprietor of Guango
Hummingbirdsd Lodge, for her permission to conduct this work. She and her fabulous
Boissonneaua – 1 2 5 2 – – – – 10 staff worked tirelessly to make our stay comfortable and successful. We
Heliangelus – – 1 1 4 – – – – 6 thank Sarah Friedrich for her meticulous work on the figures. Timo van
Adelomyia – – 1 – – – – – – 1
der Niet and two anonymous reviewers provided many helpful com-
Aglaiocercus – – 1 1 – – – – – 2
Passerines ments on the manuscript. This research did not receive any specific
Diglossa 1 1 1 – 1 1 – – 1 6 grant from funding agencies in the public, commercial, or not-for-profit
Insectse sectors.
Hymenoptera – – – – 2 – – – 1 3
Hourly N 1 2 6 7 9 1 0 0 2
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