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577
Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 53 (3-4): 577-594, September-December 2005
Birds of a high-altitude cloud forest in Alta Verapaz, Guatemala
Knut Eisermann
1
& Ulrich Schulz
University of Applied Sciences Eberswalde, Fr. Ebertstr. 28, 16225 Eberswalde, Germany.
1 Current address and address of correspondence: Knut Eisermann. P.O. Box 098 Periférico. Guatemala, Ciudad
Guatemala, Centroamérica. Tel.: 502-5906 6479; knut.eisermann@proeval-raxmu.org
Abstract: The Northern Central American Highlands have been recognized as endemic bird area, but little is
known about bird communities in Guatemalan cloud forests. From 1997 to 2001 a total of 142 bird species
were recorded between 2 000 and 2 400 masl in cloud forest and agricultural clearings on Montaña Caquipec
(Alta Verapaz, Guatemala). The bird community is described based on line transect counts within the for-
est. Pooling census data from undisturbed and disturbed forest, the Gray-breasted Wood-Wren (Henicorhina
leucophrys) was found to be the most abundant species, followed in descending order by the Common Bush-
Tanager (Chlorospingus ophthalmicus), the Paltry Tyrannulet (Zimmerius vilissimus), the Yellowish Flycatcher
(Empidonax flavescens), the Ruddy-capped Nightingale-Thrush (Catharus frantzii), and the Amethyst-throated
Hummingbird (Lampornis amethystinus). Bird communities in undisturbed and disturbed forest were found to
be similar (Sørensen similarity index 0.85), indicating low human impact. Of all recorded species, ~27% were
Nearctic-Neotropical migratory birds. The most abundant one was the Wilson’s Warbler (Wilsonia pusilla).
The Montaña Caquipec is an important area for bird conservation, which is indicated by the presence of four
species listed in the IUCN Red List (Highland Guan Penelopina nigra, Resplendent Quetzal Pharomachrus
mocinno, Pink-headed Warbler Ergaticus versicolor, Golden-cheeked Warbler Dendroica chrysoparia), and
42 Mesoamerican endemics, of which 14 species are endemic to the Central American Highlands. The results
presented here will be useful as baseline data for a long-term monitoring. Rev. Biol. Trop. 53(3-4): 577-594.
Epub 2005 Oct 3.
Key words: Bird community, cloud forest, conservation, Guatemala, Nearctic-Neotropical migrants, spe-
cies richness.
Tropical rainforests have been labeled
as some of the most endangered biomes on
a global scale (Dobson 1996). While public
interest is mainly focused on lowland rain-
forests (Wilson 1992, Primack 1993, Dobson
1996, Primack and Corlett 2005), the overall
area of cloud forests in the world has been
more greatly reduced than lowland forests in
the last few decades (Doumenge et al. 1995,
Hamilton et al. 1995). Between 1990 and
2000, Guatemala had an annual deforestation
rate of 1.7% of remaining forests, equaling
485 km
2
(FAO 2003).
Humid broadleaf forests within the conden-
sation zone, which is usually above 1 000 m,
are known as cloud forests (Richards 1996,
Received 03-III-2005. Corrected 18-VII-2005. Accepted 28-VII-2005.
Brown and Kappelle 2001). Characteristic
northern Mesoamerican cloud forest birds, like
the Resplendent Quetzal (Pharomachrus mocin-
no) and the Slate-colored Solitaire (Myadestes
unicolor), range in altitude from ~1 000-
3 000 m. At the lower range limit, these species
are associated with many lowland rainforest spe-
cies with an upper range limit at 1 500-1 800 m
(Howell and Webb 1995). High species turnover
rates have been observed along an altitudinal
transect in the Sierra Madre (Navarro 1992),
and there is almost no similarity in bird species
composition between lowland broadleaf forests
and cloud forests above 2 000 m (pers. obs.),
which can therefore be considered as the habitat
of cloud forest specialists.
578
Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 53 (3-4): 577-594, September-December 2005
Although the Guatemalan highlands have
previously been included in an endemic bird
area (Bibby et al. 1992b, Long 1995, Wege and
Long 1995, Stattersfield et al. 1998), there is
no comprehensive description of bird commu-
nities widely available. This study was carried
out as a thesis research (Eisermann 1999) and
aims to increase the knowledge on the structure
of bird communities in high-altitude cloud for-
ests in Guatemala.
In order to decrease human impact on
the cloud forest, local conservation institu-
tions provide agricultural support and alterna-
tive income opportunities to the local people
(Schulz and Unger 2000). Birds have been used
as indicators for the effectiveness of conserva-
tion efforts in biomonitoring programs (Kremen
et al. 1994). In order to measure the impact of
the efforts of several conservation institutions
at Caquipec, a long-term monitoring program
of the avifauna will be implemented based on
the results of this study.
MATERIALS AND METHODS
Study area
The Montaña Caquipec (15°23’N, 90°11’W)
is a karst mountain (Weyl 1980) which is charac-
terized by a rugged topography, located ~45 km
southeast of the city of Cobán, department Alta
Verapaz. The study area includes cloud forest
and open agricultural habitats, at elevation of
2 000 - 2 400 m.
The area is often enveloped in clouds with
an annual precipitation of ~4 000 mm and a
mean annual temperature of 15°C (MAGA
2002). Primary vegetation is an evergreen
humid broadleaf forest, with canopy heights
between 25 and 40 m. The tree community
is species rich, including different species of
the laurel family (Lauraceae), oaks (Quercus
spp.), and yellowwood (Podocarpus sp.). The
forest canopy on ridges is dominated by oaks
(Quercus spp.). A low-canopy (5 m) elfin
forest (Cavendishia sp. and other Ericaceae)
is found in small patches on ridges most
exposed to wind. Treeferns (Dicksoniaceae
and Cyathaceae), various shrubs (e.g.
Gesneriaceae, Melastomataceae) and small
palms (Chamaedorea spp.) dominate the for-
est understory in most areas. Due to the high
humidity in the forest, most of the trunks and
branches are heavily covered (<15 cm deep)
with mosses and liverworts (Bryophyta), ferns
(Filicatae), orchids (Orchidaceae), bromeliads
(Bromeliaceae) and other epiphytes of the fam-
ilies Ericaceae, Campanulaceae, Piperaceae,
and Araceae.
The Montaña Caquipec has been mostly
deforested, leaving a remnant forest fragment
of ~12 km² on the least accessible slopes
of the mountain peaks. Within this fragment
there are clearings of up to 75 ha occupied by
small indigenous settlements. The residents of
Caquipec belong to the Maya Q’eqchi’ eth-
nic group. They are subsistence farmers who
mainly depend on the cultivation of corn (Zea
mayz) and beans (Phaseolus sp.) by means of
slash-and-burn agriculture. After a few sub-
sequent crop cycles, fields are left fallow for
about 5 years, resulting in a large land require-
ment per farmer. Due to this factor and popula-
tion increase, the slash-and-burn agricultural
methods are unsustainable and have resulted
in deforestation of all but the most inaccessible
slopes. People live in traditional settlements
with simple huts dispersed widely in agricultur-
al clearings. Houses are constructed using the
stems of treeferns and timber from the forest.
People also use the forest for a variety of sub-
sistence purposes including firewood, hunting,
medicinal and edible plants, and tourism. The
secondary growth adjacent to cultivated areas
is used for cattle raising, the harvest of a sedge
(Carex sp.) for roof material, and the fruit of
the arrayan-shrub (Myrica cerifera), which
is used to produce a natural wax for candles
(Schulz and Unger 2000).
The forest appears generally undisturbed,
except in some parts where previous tim-
ber extraction created an interrupted canopy.
Considering birds as highly mobile animals,
there is no truly pristine habitat available due to
the small size of the remaining forest area.
579
Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 53 (3-4): 577-594, September-December 2005
Methods
The bird community of the cloud forest
and open habitat was studied for 208 days from
September, 1997 to October, 1998.
A definition of terms used in this report
follows:
Undisturbed forest: Cloud forest with a closed
canopy, without signs of timber extraction.
Disturbed forest: Cloud forest with canopy
gaps caused by selective timber extraction
for subsistence use.
Open habitat: Agricultural clearings and sec-
ondary scrub associated with fallow fields.
Nearctic birds: Nearctic-Neotropical migrants,
transient or wintering species which breed
in the Nearctic region.
Transient: Nearctic species in the process of
migration, present at Caquipec just tempo-
rary during the northern winter.
Vagrant: Occasionally observed species, which
are resident in the region.
Species nomenclature refers to AOU
(1998) and its supplements, names of species
treated in this report are affected by Banks et
al. (2003, 2004). We treat the Central American
form of the Sharp-shinned Hawk (Accipiter
striatus chionogaster) separated from the North
American form (Accipiter striatus striatus).
Two field methods were applied: casual
observations and line-transect counts.
Casual observations
From September, 1997 to February, 1998
qualitative species composition data were col-
lected using the ad libitum method (Lehner
1978). The principal investigator (KE) walked
along existing trails in undisturbed forest, dis-
turbed forest, and open habitat types, between
one hour before sunrise until three hours after
sunset. Birds were identified by voice or visu-
al signs. The principal investigator was able
to familiarize himself with the calls and songs
of all resident and migratory bird species
during the course of the ad libitum surveys,
which was later essential in order to conduct
the audio-visual transect counts. During 208
days of field work, a total effort of 1 250 hr of
survey were conducted throughout the study
site. Eight of the 143 species were added to
the species list through casual observations
made on additional visits from 1999-2001
(Eisermann, pers. observ., P. Kaestner, M.
Schweighöfer, R. Haupt, pers. comm.).
Transect counts
All line-transect sampling was conducted
by the principal investigator, between March
and August 1998 when favorable weather con-
ditions permitted (no rain, calm). Transect sam-
pling methods followed Bibby et al. (1992a).
Nine transects between 200 m and 900 m long
were flagged on an existing trail network (Fig.
1, Table 1). The total length of all transect lines
was 4 850 m, of which 3 680 m sampled undis-
turbed forest and 1 170 m sampled disturbed
forest. All individuals seen or heard within a
perpendicular distance of 30 m from the tran-
sect line were recorded. Sampling was repeated
two to nine times per transect, between 5:30
and 9:00 a.m. The census taker walked at an
average speed of 90 m per 15 min. Total length
Fig. 1. Location of transects (black lines) within the cloud
forest fragment (gray) of the Montaña Caquipec. The inset
map shows the location of the study area in Guatemala.
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Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 53 (3-4): 577-594, September-December 2005
of sampled transects was 31.79 km. Because
the effective strip width along the transects was
60 m, the survey design effectively sampled an
area of 29.1 ha, representing ~2.4% of the total
forest area. Three of the transects crossed both
forest types, but no sampling was conducted
within the distance of 50 m to the habitat edge.
Data Analysis
Species Richness
To predict species richness we applied
a first-order jackknife estimator to the line
transect data (Krebs 1999), using the software
“Biodiversity Professional” (McAleece el al.
1997). Each transect was treated as sample
unit. Calculation is based on presence/absence.
We excluded the records of Nearctic species
from this analysis; so the estimate indicates
species richness of breeding species only.
Dominance
To describe the community we calculated
dominance ranking using the following equa-
tion (modified after Mühlenberg 1993):
with and
where D
i
is the dominance of species i, n
i
is
the mean number of individuals of species i per
100 m transects (across all transects), N is the
sum of all n
i
, t is the number of transects, l is
the total length (in 100 m units) of all counts
along one transect.
The relative abundance values used for this
analysis were based on the frequency of obser-
vation per 100 m of sampling effort (n/100 m)
on the line transects during all months of the
survey (March-August). We assumed no sig-
nificant changes in abundance due to the brief
sampling period, which coincided with the
peak of reproductive activity.
Abundance
We compared species abundance between
undisturbed and disturbed forest using a per-
mutation test for independent samples based
on mean differences. Following Rice (1989)
we adapted significance at the alpha level for
the multiple comparisons at 0.025. For these
calculations we used the statistical package SSS
(Engel 1998). Each transect section representing
a particular forest type was used as a replicate
unit (7 sections in undisturbed and 5 in disturbed
forest, Table 1). The same abundance estimate
used for the dominance ranking analysis was
used for this comparison. In order to describe
ß-diversity between open and forest habitats, as
well as between undisturbed and disturbed for-
est, a Sørensen similarity index was calculated
based on presence-absence data using the fol-
lowing equation (Magurran 1991):
where Cs is the similarity between the two assem-
blages, j is the number of species occurring in
both compared habitats and a and b are the num-
ber of species occurring in habitat a and b.
RESULTS
Inventory and species richness
In the current study, a total of 142 bird
species were recorded on Montaña Caquipec
TABLE 1
Lenght of sampled transects
Transect length in m
Undisturbed forest Disturbed forest
Transect 1 610
Transect 2 470 100
Transect 3 450
Transect 4 600 50
Transect 5 150 330
Transect 6 900
Transect 7 750
Transect 8 240
Transect 9 200
Total 3 680 1 170
581
Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 53 (3-4): 577-594, September-December 2005
at elevations between 2 000 and 2 400 m.
Table 2 shows the relative abundance of spe-
cies obtained from transect counts, and table
3 shows relative abundance values for spe-
cies which were only recorded during ad
libitum sampling. The following bird families
were found to be most species rich: Parulidae
with 22 species, followed by Tyrannidae (13),
Turdidae (12), Trochilidae (9), Emberizidae
(8), and Vireonidae with 7 species.
Ninety-three species (~65%) were record-
ed within the cloud forest (undisturbed and
disturbed). Subtracting from this total the
number of species classified as feeding visi-
tors, vagrants, overhead transients, Nearctic-
Neotropical migrants, and unclassified species,
the remaining 60 species are assumed to be
local breeders. These are classified as perma-
nent residents, summer residents, or probable
altitudinal migrants (Table 2). The predictions
of species richness for resident forest-dwelling
birds based on transect counts, provided a first-
order jackknife value of 74 species.
Composition and habitat use
Of 93 species recorded in the cloud for-
est, ~47% were insectivores, ~18% omni-
vores, ~11% nectarivores, and ~9% carnivores.
Approximately 40% (37 of 93) of all species
observed in forest (disturbed and undisturbed)
were recorded exclusively in this habitat and
~45% (44 of 98) of the species recorded in
open habitat were observed exclusively there.
Fifty six species were recorded in the forest
and in the open habitat as well. The Sørensen
similarity index value comparing the bird com-
munity between both forest types combined
and open habitat was 0.53.
The Sørensen-index value comparing
undisturbed and disturbed forest was 0.85,
indicating a minor difference in species com-
position. When comparisons between the two
forest types were made using the permutation
test, only three species had a significant dif-
ference in abundance (Table 2). The total bird
abundance of all species was not significantly
different between disturbed and undisturbed
forest (permutation test, mean differences -
3.127, p = 0.10, based on 792 permutations).
The most abundant species with
a dominance >5% of the total number of
individuals were (in descending order): Gray-
breasted Wood-Wren (Henicorhina leucoph-
rys), Common Bush-Tanager (Chlorospingus
ophthalmicus), Paltry Tyrannulet (Zimmerius
vilissimus), Yellowish Flycatcher (Empidonax
flavescens), Ruddy-capped Nightingale-Thrush
(Catharus frantzii), and Amethyst-throated
Hummingbird (Lampornis amethystinus).
Migratory birds
Approximately a fourth of all recorded
species (39 of 143 species, see Table 2 and
Table 3) are classified as Nearctic-Neotropical
migrants. Nine of these species were observed
during the whole migratory season and are
considered Nearctic wintering migrants. The
remaining 30 Nearctic species are classified as
transients. The number of Nearctic-Neotropical
migratory species was found to be higher in
open habitat than in the forested habitats (Table
4). Tables 2, 3 and 4 classify migratory spe-
cies based on the pattern of presence over one
year of sampling. The most abundant winter-
ing Neartic species was the Wilson’s Warbler
(Wilsonia pusilla), which was recorded from
beginning of September until mid-May. The
Western Wood-Pewee (Contopus sordidulus)
was the only summer resident, recorded from
February to July.
Mixed-species flocks
We observed 30 mixed-species flocks
during casual observations and recorded the
associated species. Of 93 species recorded
for combined forest types, 23 species (~25%)
were observed foraging in mixed-species flocks
(Table 5). Twelve species were wintering or tran-
sient Nearctic birds. The number of individuals
detected in a single flock ranged from 3 to 30
containing a range of 2 to 6 species. The nucleus
species was always the Common Bush-Tanager
(Chlorospingus ophthalmicus), no other species
582
Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 53 (3-4): 577-594, September-December 2005
TABLE 2
Status, conservation status, and abundance of bird species recorded during transect counts in the cloud forest of Montaña Caquipec. Species in descending order
of relative abundance. Species with significant different abundance in disturbed and undisturbed forest are marked in bold
Abundance / Dominance
Status
1
CS
2
Undisturbed and disturbed forest
Undisturbed
forest
Disturbed
forest
Permutation test
n
3
n/100m
4
Dom. (%) n
3
n/100m.
4
Dom. (%) n
3
n/100m
4
Dom. (%) md
5
P
Total lenght of transects (km) 31.79 22.78 9.01
Maximum unrepeated length of transect (km) 4.85 3.68 1.17
Number of recorded species 63 56 54
Number of recorded individuals 5158 3458 1700
TOTAL 16.826 100 15.927 100.00 18.949 100.00
Gray-breasted Wood-Wren (Henicorhina leucophrys)
r 651 2.127 12.64 440 2,000 12,64 211 2.324 12.26 0.324 0.29
Common Bush-Tanager (Chlorospingus ophthalmicus)
r 620 1.961 11.65 443 1,943 12,28 177 2.204 11.63 0.261 0.55
Paltry Tyrannulet (Zimmerius vilissimus)
r 447 1.262 7.50 317 1,157 7,31 130 1.514 7.99 0.357 0.35
Yellowish Flycatcher (Empidonax flavescens)
r e 321 1.054 6.27 238 1,118 7,07 83 0.931 4.91 0.187 0.25
Ruddy-capped Nightingale-Thrush (Catharus frantzii)
r e 317 1.048 6.23 209 0,990 6,26 108 0.965 5.09 0.025 0.91
Amethyst-throated Hummingbird (Lampornis amethystinus)
r e 333 0.994 5.91 210 0,835 5,28 123 1.401 7.39 0.566 <0.02
Rufous-browed Wren (Troglodytes rufociliatus)
r E, e 192 0.702 4.17 137 0,714 4,51 55 0.778 4.10 0.064 0.82
Golden-browed Warbler (Basileuterus belli)
r e 174 0.675 4.01 105 0,659 4,16 69 0.695 3.67 0.037 0.89
Chestnut-capped Brush-Finch (Buarremon brunneinucha)
r 162 0.528 3.14 101 0,463 2,92 61 0.627 3.31 0.164 0.14
Tufted Flycatcher (Mitrephanes phaeocercus)
r 126 0.474 2.81 72 0,444 2,80 54 0.693 3.66 0.250 0.19
Band-tailed Pigeon (Patagioenas fasciata)
sr 119 0.454 2.70 79 0,486 3,07 40 0.353 1.86 0.133 0.41
Black Robin (Turdus infuscatus)
sr e 118 0.451 2.68 82 0,499 3,16 36 0.379 2.00 0.120 0.49
Spot-crowned Woodcreeper (Lepidocolaptes affinis)
r 117 0.423 2.52 80 0,437 2,76 37 0.513 2.71 0.076 0.52
Garnet-throated Hummingbird (Lamprolaima rhami)
r e 128 0.411 2.44 93 0,390 2,46 35 0.413 2.18 0.024 0.85
Mountain Robin (Turdus plebejus)
r e 119 0.386 2.30 77 0,355 2,25 42 0.495 2.61 0.140 0.21
Slate-colored Solitaire (Myadestes unicolor)
r e 114 0.370 2.20 82 0,377 2,38 32 0.401 2.12 0.024 0.85
Brown-backed Solitaire (Myadestes occidentalis)
r e 91 0.310 1.84 52 0,230 1,45 39 0.331 1.75 0.101 0.45
Wilson’s Warbler (Wilsonia pusilla)
*** 85 0.283 1.68 58 0,286 1,81 27 0.334 1.76 0.047 0.43
Green-throated Mountain-gem (Lampornis viridipallens)
sr E, e 111 0.282 1.67 86 0,299 1,89 25 0.295 1.56 0.004 0.99
583
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Abundance / Dominance
Status
1
CS
2
Undisturbed and disturbed forest
Undisturbed
forest
Disturbed
forest
Permutation test
n
3
n/100m
4
Dom. (%) n
3
n/100m.
4
Dom. (%) n
3
n/100m
4
Dom. (%) md
5
P
Mountain Trogon (Trogon mexicanus)
r 75 0.248 1.47 47 0,227 1,43 28 0.225 1.19 0.002 0.97
Barred Parakeet (Bolborhynchus lineola)
r 60 0.238 1.41 35 0,186 1,18 25 0.248 1.31 0.062 0.77
Black-throated Jay (Cyanolyca pumilo)
r E, e 58 0.229 1.36 33 0,200 1,26 25 0.252 1.33 0.052 0.58
Cinnamon-bellied Flowerpiercer (Diglossa baritula)
r e 62 0.195 1.16 46 0,225 1,42 16 0.160 0.84 0.066 0.37
Elegant Euphonia (Euphonia elegantissima)
sr e 74 0.193 1.15 35 0,103 0,65 39 0.306 1.62 0.204 0.18
Hairy Woodpecker (Picoides villosus)
r 58 0.187 1.11 34 0,152 0,96 24 0.216 1.14 0.064 0.35
Resplendent Quetzal (Pharomachrus mocinno)
r R, e 54 0.168 1.00 36 0,154 0,98 18 0.234 1.24 0.080 0.30
Pink-headed Warbler (Ergaticus versicolor)
sr E, R, e 24 0.157 0.94 21 0,204 1,29 3 0.104 0.55 0.100 0.51
Emerald Toucanet (Aulacorhynchus prasinus)
r e 59 0.150 0.89 36 0,113 0,71 23 0.187 0.99 0.074 0.41
Crescent-chested Warbler (Parula superciliosa)
r e 37 0.136 0.81 27 0,137 0,87 10 0.094 0.50 0.043 0.49
Ruddy Foliage-gleaner (Automolus rubiginosus)
r 43 0.121 0.72 34 0,137 0,87 9 0.078 0.41 0.059 0.20
Blue-crowned Chlorophonia (Chlorophonia occipitalis)
sr e 35 0.097 0.58 18 0,060 0,38 17 0.179 0.94 0.119 0.13
Spotted Woodcreeper (Xiphorhynchus erythrophygius)
sr 30 0.093 0.55 12 0,039 0,25 18 0.335 1.77 0.296 <0.002
Slate-throated Redstart (Myioborus miniatus)
r 25 0.071 0.42 18 0,084 0,53 7 0.125 0.66 0.041 0.59
Hutton’s Vireo (Vireo huttoni)
r 15 0.043 0.26 6 0,018 0,11 9 0.053 0.28 0.035 0.35
Chestnut-collared Swift (Streptoprocne rutila)
f 11 0.032 0.19 4 0,016 0,10 7 0.073 0.39 0.058 0.12
Tawny-throated Leaftosser (Sclerurus mexicanus)
sr 11 0.031 0.19 8 0,030 0,19 3 0.054 0.28 0.023 0.75
White-faced Quail-Dove (Geotrygon albifacies)
r e 9 0.026 0.16 5 0,014 0,09 4 0.046 0.24 0.031 0.21
Western Wood-Pewee (Contopus sordidulus)
sr 7 0.025 0.15 5 0,020 0,13 2 0.021 0.11 0.001 1
Highland Guan (Penelopina nigra)
r E, R, e 8 0.023 0.14 5 0,017 0,11 3 0.019 0.10 0.002 0.87
Black-capped Swallow (Notiochelidon pileata)
r E, e 4 0.016 0.09 3 0,015 0,10 1 0.012 0.06 0.003 1
White-eared Hummingbird (Hylocharis leucotis)
sr 4 0.015 0.09 4 0.049 0.26 0.049 <0.02
Violet Sabrewing (Campylopterus hemileucurus)
sr e 6 0.014 0.08 4 0,011 0,07 2 0.010 0.05 0.001 0.84
Rufous-collared Robin (Turdus rufitorques)
sr E, e 4 0.013 0.08 4 0.024 0.13 0.024 0.15
TABLE 2 (Continued)
Status, conservation status, and abundance of bird species recorded during transect counts in the cloud forest of Montaña Caquipec. Species in descending order
of relative abundance. Species with significant different abundance in disturbed and undisturbed forest are marked in bold
584
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TABLE 2 (Continued)
Status, conservation status, and abundance of bird species recorded during transect counts in the cloud forest of Montaña Caquipec. Species in descending order
of relative abundance. Species with significant different abundance in disturbed and undisturbed forest are marked in bold
Abundance / Dominance
Status
1
CS
2
Undisturbed and disturbed forest
Undisturbed
forest
Disturbed
forest
Permutation test
n
3
n/100m
4
Dom. (%) n
3
n/100m.
4
Dom. (%) n
3
n/100m
4
Dom. (%) md
5
P
Northern Flicker (Colaptes auratus) r 6 0.013 0.08 5 0,015 0,10 1 0.005 0.03 0.010 0.58
Canada Warbler (Wilsonia canadensis) ** 3 0.011 0.07 1 0,004 0,03 2 0.014 0.07 0.010 0.42
Barred Forest-Falcon (Micrastur ruficollis) r 3 0.011 0.06 2 0,008 0,05 1 0.009 0.05 0.002 1
Blue-throated Motmot (Aspatha gularis) r E, e 4 0.010 0.06 2 0,005 0,03 2 0.010 0.05 0.005 0.73
Magnificent Hummingbird (Eugenes fulgens) sr 4 0.009 0.05 3 0,008 0,05 1 0.005 0.03 0.003 1
Scaly-throated Foliage-gleaner (Anabacerthia variegaticeps) sr 3 0.008 0.05 1 0,004 0,03 2 0.027 0.14 0.023 0.42
Sharp-shinned Hawk (Accipiter striatus chionogaster) sr E, e 3 0.007 0.04 3 0,011 0,07 0.011 0.47
Rufous-browed Peppershrike (Cyclarhis gujanensis) sr 1 0.005 0.03 1 0.009 0.05 0.009 0.42
Rose-throated Becard (Pachyramphus aglaiae) r 2 0.005 0.03 2 0.010 0.05 0.010 0.42
Blue-headed Vireo (Vireo solitarius) *** 1 0.004 0.02 1 0,005 0,03 0.005 1
White-tipped Dove (Leptophila verreauxi) v 1 0.004 0.02 1 0.080 0.42 0.080 0.42
Bushy-crested Jay (Cyanocorax melanocyaneus) v E, e 1 0.003 0.02 1 0,004 0,03 0.004 1
Green Violet-ear (Colibri thalassinus) sr 1 0.003 0.02 1 0,004 0,03 0.004 1
Warbling Vireo (Vireo gilvus) *** 1 0.003 0.02 1 0.005 0.03 0.005 0.42
Wine-throated Hummingbird (Atthis ellioti) sr E, e 1 0.002 0.01 1 0,003 0,02 0.003 1
Hooded Grosbeak (Coccothraustes abeillei) sr e 1 0.002 0.01 1 0,003 0,02 0.003 1
Mountain Elaenia (Elaenia frantzii) sr 1 0.002 0.01 1 0,003 0,02 0.003 1
Eastern Wood-Pewee (Contopus virens) ** 1 0.002 0.01 1 0,003 0,02 0.003 1
Black-throated Green Warbler (Dendroica virens) ** 1 0.002 0.01 1 0.022 0.12 0.022 0.42
Olive-sided Flycatcher (Contopus cooperi) ** 1 0.002 0.01 1 0,003 0,02 0.003 1
1 Status: r-resident, sr-summer resident or altitudinal migrant, f-feeding visitor, v-vagrant, ***=Nearctic wintering migrant; **=Nearctic transient; *=Nearctic overhead transient.
2 Conservation status (CS): E=endemic to the Northern Central American Highlands (Stattersfield et al. 1998), e=endemic to Mesoamerica (Stotz et al. 1996), R=listed in the
global Red List (BirdLife International 2004).
3 n - sample size.
4 n/100m-relative abundance in n/100 m of transect (mean across all transects) (60 m strip width).
5 mean differences in 792 permutations.
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TABLE 3
Status and number of observations of bird species not recorded during transect counts
Number of observations in
Status
1
CS
2
Cloud forest Openland
Overhead
transient
Great Blue Heron (Ardea herodias)
*1
Cattle Egret (Bubulcus ibis)
v3
Upland Sandpiper (Bartramia longicauda)
** 1
Broad-winged Hawk (Buteo platypterus)
** 1
Red-tailed Hawk (Buteo jamaicensis)
r >10 >10
Gray Hawk (Asturina nitida)
t2
Hook-billed Kite (Chondrohierax uncinatus)
v1 1
Sharp-shinned Hawk (Accipiter striatus striatus)
*1
Swallow-tailed Kite (Elanoides forficatus)
t1
American Kestrel (Falco sparverius)
t1
Black Vulture (Coragyps atratus)
t4
Turkey Vulture (Cathartes aura)
t1
Plain Chachalaca (Ortalis vetula)
sr
e 2
Buffy-crowned Wood-Partridge (Dendrortyx leucophrys)
re 1 4
Mourning Dove (Zenaida macroura)
** 1
Northern Pygmy-Owl (Glaucidium gnoma)
r11
Fulvous Owl (Strix fulvescens)
r E, e >10
Mottled Owl (Ciccaba virgata)
sr >10
Unspotted Saw-whet Owl (Aegolius ridgwayi)
ve 1
Whip-poor-will (Caprimulgus vociferus)
r e >10 >10
Black Swift (Cypseloides niger)
? 1 flock 1 flock
White-collared Swift (Streptoprocne zonaris)
f >10
Vaux’s Swift (Chaetura vauxi)
? 1 flock
White-throated Swift (Aeronautes saxatilis)
? 1 flock
Sparkling-tailed Hummingbird (Tilmatura dupontii)
ve 1
Golden-fronted Woodpecker (Melanerpes aurifrons)
v1
Yellow-bellied Sapsucker (Sphyrapicus varius)
*** 7
White-throated Flycatcher (Empidonax albigularis)
ve
1
Least Flycatcher (Empidonax minimus )
** 1
Hammond’s Flycatcher (Empidonax hammondii)
*** >10 >10
Buff-breasted Flycatcher (Empidonax fulvifrons)
ve 3
Yellow-bellied Flycatcher (Empidonax flaviventris)
** 1 1
Acadian Flycatcher (Empidonax virescens)
** 1
Plumbeous Vireo (Vireo plumbeus)
v1
Brown-capped Vireo (Vireo leucophrys)
v1
Philadelphia Vireo (Vireo philadelphicus)
** 3 2
Tree Swallow (Tachycineta bicolor)
*** 2 flocks
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Number of observations in
Status
1
CS
2
Cloud forest Openland
Overhead
transient
Cliff Swallow (Petrochelidon pyrrhonota)
** 3
Barn Swallow (Hirundo rustica)
** 3
Plain Wren (Thryothorus modestus)
r e >10
House Wren (Troglodytes aedon)
r >10
Eastern Bluebird (Sialia sialis)
v >10
Orange-billed Nightingale-Thrush (Catharus aurantiirostris)
r >10
Spotted Nightingale-Thrush (Catharus dryas)
v1
Swainson’s Thrush (Catharus ustulatus)
** 2
Hermit Thrush (Catharus guttatus)
** 1
Clay-colored Robin (Turdus grayi)
v1
Blue-and-white Mockingbird (Melanotis hypoleucus)
r E, e >10
Gray Catbird (Dumetella carolinensis)
*** 5
Gray Silky-flycatcher (Ptilogonys cinereus)
r >10 >10
Olive Warbler (Peucedramus taeniatus)
v1
Tennessee Warbler (Vermivora peregrina)
** 3 2
Chestnut-sided Warbler (Dendroica pensylvanica)
** 1
Yellow-rumped Warbler (Dendroica coronata)
*** >10
Townsend’s Warbler (Dendroica townsendi)
*** >10 >10
Golden-cheeked Warbler (Dendroica chrysoparia)
** R 1
Hermit Warbler (Dendroica occidentalis)
** 3
Blackburnian Warbler (Dendroica fusca)
** 3
Black-and-white Warbler (Mniotilta varia)
** 10
MacGillivray’s Warbler (Oporonis tolmiei)
** 2
Red-faced Warbler (Cardellina rubrifrons)
** 1
Rufous-capped Warbler (Basileuterus rufifrons)
v8
Gray-crowned Y
ellowthroat (Geothlypis poliocephala)
ve 1
Northern Waterthrush (Seiurus noveboracensis)
** 1
Louisiana Waterthrush (Seiurus motacilla)
** 3
Blue-black Grassquit (Volatinia jacarina)
v1
Yellow-faced Grassquit (Tiaris olivaceus)
sr 4
White-collared Seedeater (Sporophila torqueola)
sr >10
White-naped Brush-Finch (Atlapetes albinucha)
r >10
Lincoln’s Sparrow (Melospiza lincolnii)
** 1
Rufous-collared Sparrow (Zonotrichia capensis)
r >10
Rusty Sparrow (Aimophila rufescens)
ve 1
Grayish Saltator (Saltator coerulescens)
v1
Rose-breasted Grosbeak (Pheucticus ludovicianus)
** 3
TABLE 3 (Continued)
Status and number of observations of bird species not recorded during transect counts
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Number of observations in
Status
1
CS
2
Cloud forest Openland
Overhead
transient
Great-tailed Grackle (Quiscalus mexicanus)
r >10
Melodious Blackbird (Dives dives)
ve 1
Yellow-backed Oriole (Icterus chrysater)
v3
Baltimore Oriole (Icterus galbula)
** 1
Black-capped Siskin (Carduelis atriceps)
v E, e 2
Black-headed Siskin (Carduelis notata)
sr >10
1 Status: r-resident, sr-summer resident or altitudinal migrant, f-feeding visitor, v-vagrant, t-overhead transient, ?-status
unknown, ***=Nearctic wintering migrant; **=Nearctic transient or vagrant; *=Nearctic overhead transient.
2 Conservational status (CS): E=endemic to the Northern Central American Highlands (Stattersfield et al. 1996);
e=endemic to Mesoamerica (Stotz et al. 1996), R=listed in the global Red List (BirdLife International 2004).
TABLE 3 (Continued)
Status and number of observations of bird species not recorded during transect counts
TABLE 4
Status of bird species recorded in Caquipec from 1997-2001
1
No. of species
Study area Forest Openland
Neotropical species (n=101):
Permanent resident (recorded during the whole year) 45 38 29
Summer resident and probable Teilzieheraltitudinal migrant (recorded only
in summer months, or during the whole year with exception of short periods)
26 22 17
Feeding visitor (highly mobile, aerial species with few records) 2 2 2
Vagrant (in Guatemala resident species with few records in Caquipec) 23 7 18
Overhead transient (records of high overflying species without
observed relationship to the habitat)
5--
Nearctic species (n=39):
Nearctic wintering migrant (recorded during the whole migratory season) 9 7 8
Nearctic transients
(recorded during short periods of the migratory season)
27 14 21
Nearctic overhead transient (records of high overflying species without
observed relationship to the habitat)
3--
Status unknown (n=3): 3 3 3
TOTAL 143 93 98
1 Classification of Neotropical and Nearctic species according to Howell and Webb (1995) and DeGraaf and Rappole
(1995), subclassifications using the author’s observations. Resident and Nearctic form of Sharp-shinned Hawk
(Accipiter striatus chionogaster and A. s. striatus) are listed separately.
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TABLE 5
Bird species observed in mixed-species foraging flocks
FH
1
MS
2
Nucleus species:
THRAUPIDAE
Common Bush-Tanager (Chlorospingus ophtalmicus)
Or
Attendant species:
PARULIDAE
Crescent-chested Warbler (Parula superciliosa)
Ir
Tennessee Warbler (Vermivora peregrina)
In
Wilson’s Warbler (Wilsonia pusilla)
In
Canada Warbler (Wilsonia canadensis)
In
Pink-headed Warbler (Ergaticus versicolor)
Ir
Slate-throated Redstart (Myioborus miniatus)
Ir
Golden-browed Warbler (Basileuterus belli)
Ir
Black-throated Green Warbler (Dendroica virens)
In
Blackburnian Warbler (Dendroica fusca)
In
Townsend’s Warbler (Dendroica townsendi)
In
Black-and white Warbler (Mniotilta varia)
In
VIREONIDAE
Hutton’s Vireo (Vireo huttoni)
Or
Blue-headed Vireo (Vireo solitarius)
On
Warbling Vireo (Vireo gilvus)
On
Philadelphia Vireo (Vireo philadelphicus)
On
Tyrannidae
Tufted Flycatcher (Mitrephanes phaeocercus)
Ir
Rose-throated Becard (Pachyramphus aglaiae)
On
FURNARIIDAE
Scaly-throated Foliage-gleaner (Anabacerthia variegaticeps)
In
DENDROCOLAPTIDAE
Spot-crowned Woodcreeper (Lepidocolaptes affinis)
Ir
Emberizidae
Cinnamon-bellied Flowerpiercer (Diglossa baritula)
Nr
FRINGILLIDAE
Blue-crowned Chlorophonia (Chlorophonia occipitalis)
Fr
Elegant Euphonia (Euphonia elegantissima)
Fr
1
Food habits (FH): O=omnivore; I=insectivore, F=frugivore, N=nectarivore (Stiles and Skutch 1989, Howell and
Webb 1995).
2 Migratory status (MS): n=Nearctic migrant, r=resident (DeGraaf and Rappole 1995, Howell and Webb 1995).
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was present in all flocks. Adjoining species were
observed in all forest strata.
DISCUSSION
Inventory and species richness
Species richness depends on various eco-
logical factors. In general, it decreases with
increasing latitude and altitude (Begon et al.
1990). Terborgh (1977) found a peak in spe-
cies richness at the elevation of 1 500 m along
an altitudinal transect on the eastern slope
of the Peruvian Andes, where species rich-
ness decreased above this elevation. Navarro
(1992) recorded a declining species richness
from 680 m to 3 100 m in the Sierra Madre,
Mexico. Therefore we expected species rich-
ness at Caquipec (>2 000 m) would be lower
compared to species richness values of ~100
resident species recorded locally in undis-
turbed cloud forest at 1 500 m (Eisermann,
unpub. data). In the current study we predicted
a species richness of 74 species with jack-
knife estimator, which is 14 species more than
we obtained from transect counts and casual
observations (60 species). Although the spe-
cies accumulation curve from transect counts
indicates that almost all species were detected
(Fig. 2), several rare or vagrant species may
be expected to contribute to species richness.
Comparison with other
cloud forest avifaunas
The distribution of cloud forest species
in northern Central America is known on
a large scale (Howell and Webb 1995), but
information on relative abundance in bird com-
munities is lacking. Currently no published
description of a Guatemalan cloud forest bird
community, treating all species, is widely avail-
able. Komar (2002) provided the first relative
abundance description for a northern Central
American cloud forest bird community at Cerro
Montecristo (14°26’ N 89°21’ W), in northern
El Salvador. An inventory is available for the
El Triunfo cloud forest (15°39’ N 92°48’ W) in
Chiapas, Mexico, without detailed data on rela-
tive abundance (Parker III et al. 1976, Gómez
de Silva et al. 1999). Unpublished data are
available from the Sierra de las Minas (15°05’ N
89°57’ W), department El Progreso, Guatemala
(Robbins and Dowell 1992, Ponciano 1998,
Eisermann 1999, Nájera 2004) and the Montaña
Yalijux (15°23’ N 90°04’ W), Alta Verapaz
in Guatemala (Renner 2003, pers. obs.). The
cloud forests at all sites cover a similar altitudi-
nal range as the Caquipec forest at elevations of
2 000-2 400 m (Montecristo: 2 000-2 350 m,
El Triunfo: 1 870-2 450 m, Sierra de las Minas
and Yalijux: 2 000-2 500 m). To compare the
most common species between the five sites,
we use the species detected by Komar (2002)
at least once in a four-hour period of ‘intensive
searches’ at Montecristo; the species which
were labeled ‘numerous’ in Gomez de Silva
et al. (1999) or ‘common’ in Parker III et
al. (1976) in El Triunfo; the species with a
dominance >1% in Yalijux (Renner 2003); and
the species classified as ‘common’ and ‘fairly
common’ in Sierra de las Minas (Nájera 2004).
From our data from the Montaña Caquipec we
classify species with a relative abundance value
of >0.28 per 100 m of transect as common,
which equals 1 bird per 360 m of transect sam-
pled (4 hours) and is therefore comparable with
the effort of Komar (2002). Five of the most
common species have been recorded at all sites:
Rufous-browed Wren (Troglodytes rufocilia-
tus), Gray-breasted Wood-Wren (Henicorhina
leucophrys), Ruddy-capped Nightingale-Thrush
(Catharus frantzii), Golden-browed Warbler
Fig. 2. Species accumulation during transect counts.
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(Basileuterus belli), and Common Bush-
Tanager (Chlorospingus ophthalmicus). The
Spotted Nightingale-Thrush (Catharus dryas),
classified as common in Montecristo and El
Triunfo, was not recorded or was classified as
vagrant in all of the Guatemalan high-altitude
cloud forest sites (one observation at Caquipec,
P. Kaestner pers. comm., Robbins and Dowell
1992, Renner 2003). In Alta Verapaz, this spe-
cies is common at lower elevations (<1 500 m,
Eisermann, unpub. data). Although the cloud
forest avifauna of northern Central America
(eastern Chiapas, Guatemala, El Salvador and
Honduras) has been described as similar based
on regional species lists (Hernández-Baños
et al. 1995), there are differences between
the five cloud forest sites. Figure 3 shows the
similarity of the bird communities of Caquipec,
Yalijux, Sierra de las Minas, El Triunfo and
Montecristo, taking into account all species
recorded within the forest except Nearctic-
Neotropical migrants and overhead transients.
Yalijux is located ~11 km to the east of
Caquipec. Both sites are connected by a narrow
corridor of cloud forest, and bird communities
are very similar. Higher dissimilarity seems
to result from longer distances between sites.
The Sierra de las Minas is located at a distance
of ~35 km from Caquipec, separated by the
Polochic valley. An interchange between bird
populations of both sites has been observed
on the Resplendent Quetzal (Pharomachrus
mocinno), which moved apparently from the
Sierra de las Minas eastwards to the Sierra de
Chuacús and from there to Caquipec and Yalijux
(Paiz 1996). The Sierra de los Cuchumatanes
is an altitudinal corridor between Caquipec
and El Triunfo, which is ~270 km away. The
high dissimilarity of the bird community at
Montecristo (Fig. 3, ~135 km from Caquipec), is
probably caused by isolation from the Chiapas-
Guatemalan highlands. Current descriptions of
northern Central American cloud forest sites are
insufficient to explain differences in the compo-
sition of the bird communities. All papers pres-
ent data from short-term studies, thus annual or
long-term population fluctuations are unknown.
Long-term monitoring of bird populations and
their habitats is necessary in order to identify
factors that affect populations.
We observed a remarkably high relative abun-
dance of the Amethyst-throated Hummingbird
(Lampornis amethystinus) at Caquipec (0.99
birds per 100 m transect, dominance 5.9%,
Tab. 2). In Montecristo, no hummingbird was
recorded with a dominance >5% of all individu-
als (data from Komar 2002). In South American
cloud forests, hummingbirds have been recorded
within the most abundant species. Poulsen and
Krabbe (1998) recorded three hummingbird spe-
cies at some study sites above 3 000 m with more
than 5% dominance. The highest dominance
was recorded for the Tyrian Metaltail (Metallura
tyrianthina) at the Chaucha site with 11.6%.
Terborgh (1977) has shown that the dominance
of the nectarivorous guild increases toward high-
er elevations and attributed this phenomenon to a
more abundant and more constant supply of food
resources in higher altitudes and a lack of food
resources for other species.
Fig. 3. Cluster analysis based on presence / absence of
bird species in five northern Central American high-alti-
tude cloud forests, performed with software Biodiversity
Professional (McAleece et al. 1997). Data for El Triunfo
from Parker III et al. (1976) and Gómez de Silva et al.
(1999), for Montecristo from Komar (2002); for Sierra
de las Minas from Robbins and Dowell (1992), Ponciano
(1998), Eisermann (1999) and Nájera (2004); for Sierra
Yalijux from Renner (2003) and pers. obs.
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Human Impact
Because there are no historical data avail-
able on the bird community of the Montaña
Caquipec, the human impact on it is difficult
to assess. Thirty nine species were record-
ed exclusively in open habitat and indicate
increased species richness throughout the
whole area, because this kind of habitat did
not exist in the research area before human
settlement. We assume that human activity has
had some negative effects on species richness
through the reduction of forest size and hunt-
ing. The Horned Guan (Oreophasis derbianus),
for example, was not found in the current study.
This endemic cloud forest cracid is very sensi-
tive to disturbance, especially hunting (Howell
and Webb 1995, González-García et al. 2001),
and was still reported for the region of Cobán
until the first half of the 20th century (Hellmayr
and Conover 1942). Unfortunately no exact
geographic data are available, but these reports
are probably from Caquipec, since this moun-
tain range is accessible by road since long and
it is the highest mountain in Alta Verapaz.
Thiollay (1984) and Peres (2000) have dem-
onstrated that even low intensity hunting can
heavily affect game populations and their natu-
ral predators, which could explain the absence
of the Horned Guan, and the low abundance of
the Highland Guan (Penelopina nigra) in the
study area (Table 2).
In other tropical regions, little similarity
was found between commercially exploited for-
est and pristine forest (Newmark 1991, Thiollay
1992, Mason 1996). In Caquipec, the high
degree of similarity between the bird communi-
ties of undisturbed and disturbed forest indicates
that the subsistence use of forest resources by
Q’eqchi’ communities may have low impact on
the overall species composition.
Conservation
The Northern Central Amerian Highland,
which includes the mountains of Chiapas,
Guatemala, El Salvador and Honduras, is
a recognized endemic bird areas (Bibby et
al. 1992b, Stattersfield et al. 1998, Brown
and Kappelle 2001). In the current study, 42
(~30%) of the recorded species are endemic
to Mesoamerica and 14 species (~10%) are
endemic to the Northern Central American
Highlands. Twelve species of the latter group
were recorded in cloud forest (Tables 2 and 3).
Four species recorded at Caquipec are listed
in the Red List of globally threatened spe-
cies (BirdLife International 2004): Highland
Guan (Penelopina nigra: Near Threatened),
Resplendent Quetzal (Pharomachrus mocin-
no: Near Threatened), Pink-headed Warbler
(Ergaticus versicolor: Vulnerable), and the
migratory Golden-cheeked Warbler (Dendroica
chrysoparia: Endangered). The high percentage
species with a restricted range and the presence
of four globally threatened species identify
the Montaña Caquipec as an important area
for bird conservation and emphasize the high
importance of Guatemalan high-altitude cloud
forests for conservation. Conservation efforts
in these forests are insufficient in Guatemala.
Although the Guatemalan system of protected
areas covers almost 30% (31 930 km
2
) of the
country (CONAP 2003), only ~2 235 km
2
are
cloud forests (~7% of protected areas). These
include the Sierra de las Minas, some smaller
areas in the southern volcanic belt, the ‘Biotopo
del Quetzal’ in Baja Verapaz and areas in the
western highlands. Humid montane forests of
the northernmost mountain chain of Guatemala
are underrepresented in the system of protected
areas. In the northern part of the department of
Alta Verapaz there are only 5 protected areas
with cloud forest. Their sizes are rather small,
ranging from 48 to 1 366 ha, for a total of
26 km
2
(CONAP 2003). We assume that such
small areas will not support viable populations
of many bird species. Altitudinal migration has
been reported for many cloud forest bird spe-
cies (Powell and Bjork 1994, Howell and Webb
1995, Paiz 1996, Winker et al. 1997). This has
led to the recognition that an extension of con-
servation activities along the altitudinal gradi-
ent is essential for the effective conservation of
many species. Conservation activities need to
be extended into rural communities (Islebe and
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Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 53 (3-4): 577-594, September-December 2005
Véliz 2001). Various local NGOs have been
active at Caquipec by supporting indigenous
communities with agricultural and ecotour-
ism projects, in order to generate alternative
income and reduce the rate of deforestation
(Schulz and Unger 2000). At present, we are
unable to evaluate the impact of alternative for-
est use, such as tourism, on the habitat quality
of the forest. A long-term monitoring is needed
to be able to evaluate population trends and
human impact. Data provided here will be use-
ful baseline data for further research.
ACKNOWLEDGMENTS
The period of familiarization with the bird
voices would have been much longer without
the invaluable support of Amy Seglund and
Shawn Conner during an introductory field
trip. We are grateful to the staff of BIDAS /
Proyecto Eco-Quetzal (Cobán, Alta Verapaz)
for its technical support and to David Unger for
constructive discussion during the field work.
Thanks to the Q’eqchi’ indigenous people of
the community of Chicacnab for their kind
cooperation. We thank Jürgen Engel for statis-
tical advice. We appreciate critical comments
on the manuscript by an unknown reviewer
and Claudia Avendaño, and the improvements
in English usage made by Erick Baur through
the Association of Field Ornithologists’ pro-
gram of editorial assistance. Special thanks to
Peter, Gisela and Käthe Eisermann for their
financial support.
RESUMEN
Las alturas del norte de Centroamérica han sido reco-
nocidas como región de aves endémicas, pero se conoce
poco sobre las comunidades de aves en bosques nubosos
de Guatemala. De 1997 a 2001 se han detectado 142
especies de aves entre 2 000 y 2 400 msnm en el bosque
nuboso y áreas agrícolas en la Montaña Caquipec (Alta
Verapaz, Guatemala). El patrón de la comunidad de aves
se describe por medio de censos en transectos de línea.
Combinando los datos de censos en bosque prístino y bos-
que perturbado se concluyó que Henicorhina leucophrys
es la especie más abundante, seguida en orden descendente
por Chlorospingus ophthalmicus, Zimmerius vilissimus,
Empidonax flavescens, Catharus frantzii y Lampornis
amethystinus. El índice de Sørensen de 0.85 entre bosque
prístino y bosque perturbado indica un impacto relativa-
mente bajo de la población humana local. El ~27% de todas
las especies encontradas fueron aves migratorias neárticas,
siendo la más abundante Wilsonia pusilla. La Montaña
Caquipec es un área importante para la conservación de
aves, lo cual es indicado por la presencia de cuatro especies
incluidas en la Lista Roja de IUCN (Penelopina nigra,
Pharomachrus mocinno, Ergaticus versicolor, Dendroica
chrysoparia) y de 42 especies endémicas de Mesoamérica,
de las cuales 14 son endémicas de las alturas norteñas de
Mesoamérica. Estos resultados servirán como base para un
monitoreo a largo plazo.
Palabras clave: Aves, bosque nuboso, conservación,
Guatemala, migración, Neartico-Neotrópico migrants,
riqueza de especies.
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