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Patterns of Species Composition and Endemism in the Northern Neotropics: A Case for
Conservation of Montane Avifaunas
Author(s): Luis Miguel Renjifo, Grace P. Servat, Jaqueline M. Goerck, Bette A. Loiselle and John
G. Blake
Source:
Ornithological Monographs,
No. 48, Studies in Neotropical Ornithology Honoring Ted
Parker (1997), pp. 577-594
Published by: for the University of California Press American Ornithologists' Union
Stable URL: http://www.jstor.org/stable/40157554
Accessed: 16-09-2015 15:00 UTC
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PATTERNS OF SPECIES COMPOSITION AND
ENDEMISM IN THE NORTHERN NEOTROPICS:
A CASE FOR CONSERVATION OF
MONTANE AVIFAUNAS
Luis Miguel Renjifo, Grace P. Servat, Jaqueline M. Goerck, Bette
A. LOISELLE, AND JOHN G. BLAKE
Department
of Biology and International
Center
for Tropical Ecology,
University of Missouri-St.
Louis, St. Louis, Missouri
63121-4499, USA
Ornithological Monographs
Volume 48 (1997), pp. 577-594
Abstract. - A review of the composition of five montane avifaunas in northwestern
South America and southern Central America confirmed the distinctness of these com-
munities from adjacent lowland areas. Excluding species that depend on aquatic re-
sources, 1,800 bird species from 52 families were classified according to principal life
zones in this review. There were 1,366 species associated with lowland areas, whereas
877 species occurred in montane areas (i.e., generally above 1,200 or 1,500 m elevation).
Nearly one-half of these montane species are restricted to these high elevations, with the
greatest diversity found within subtropical zones, followed by temperate and p&ramo
zones, respectively. Comparisons with lowland avifaunas revealed that montane com-
munities differed in trophic structure as well as familial composition. Specifically, mon-
tane communities had proportionately more nectarivores than expected by chance. When
compared to randomly generated montane communities, nearly 30% of the families dif-
fered significantly in number of species from that expected if these communities were
randomly assembled; eight families were more species-rich and seven families less spe-
cies-rich in montane areas than expected. Moreover, montane areas had a greater number
and percentage of species with restricted ranges than did lowland areas. Overall, 217
montane species (24.7%) had small geographic ranges; 142 of these were restricted to
one of the five montane regions reviewed here. As evidence of the endangerment of
these montane communities, nearly 10% (82) of the species are listed as threatened or
near-threatened. Montane habitats are under extreme pressure from human activities.
Most urban centers are located in or close to the mountains in the regions reviewed here.
Given the high diversity and singularity of these avifaunas, together with high levels of
habitat alteration, protection of montane ecosystems should become a priority for con-
servation efforts in the Neotropics.
Resumen.
- Una revisi6n de la composici6n de cinco avifaunas de montana en el
Noroeste de Sur America y el sur de Centro America confirm6 la diferencia de estas
comunidades con las de las tierras bajas. Sin incluir especies que dependen de recursos
acu&ticos, 1,800 especies de aves de 52 familias fueron clasificadas de acuerdo a las
principales zonas de vida. 1,366 especies fueron asociadas con areas de tierras bajas, de
las cuales 877 especies se encontraron en areas montanosas (generalmente encima de
1,200-1,500 m de elevacion). Casi la mitad de estas especies de areas montanosas estan
restringidas a estas elevaciones altas, encontr&ndose
la mayor diversidad en zonas sub-
tropicales, seguidas por zonas templadas y de paramo respectivamente. Comparaciones
con avifaunas de tierras bajas revelan que las comunidades de montana difieren en es-
tructura tr6fica asi como en la composici6n de las Familias. Especificamente, las co-
munidades de montana tuvieron proporcionalmente mayor numero de nectarivoros que
los esperados al azar. Se compararon avifaunas de montanas con comunidades ensam-
bladas al azar usando simulaciones de Monte Carlo. En estas comparaciones, cerca del
30% de las familias se diferenciaron significativamente en el numero de especies; ocho
fueron mas ricas y siete fueron menos ricas en areas montanosas de lo esperado. Ademds,
las areas de montana tuvieron un mayor numero y porcentaje de especies con rangos
restringidos que las de tierras bajas. Un total de 217 especies de montana (24.7%) tu-
vieron rangos geogrdficos pequenos; 142 de estas fueron restringidas a una de cinco
regiones de montana que fueron revisadas en este trabajo. Cerca del 10% (82) de estas
especies se encuentran enlistadas como amenazadas o casi-amenazadas, lo que evidencia
el peligro de extinci6n de estas comunidades de montana. Los areas de montana se
encuentran bajo extrema presi6n por actividades humanas. Muchas de los centros urbanos
577
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578 ORNITHOLOGICAL MONOGRAPHS NO. 48
se encuentran localizados en 6 cerca de las montanas que fueron revisadas. La protecci6n
de los ecosistemas de montana debe ser una prioridad para los esfiierzos de conservaci6n
en los Neotr6picos debido a la alta diversidad y singularidad de estas avifaunas, asi como
a los altos niveles de alteraci6n en los habitats.
The avifauna of the Neotropics has long been recognized by biologists as being exceedingly
rich. Colombia, with 1,758 species (Hernandez 1993), alone contains approximately 19% of the
world's bird species, and even the relatively small country of Costa Rica (ca. 50,000 km2)
contains about 9% of the world's avifauna (840 species; Stiles and Skutch 1989). Several factors
promote high diversity in the Neotropics, including historical, geological, biotic, and environ-
mental factors (e.g., Haffer 1967, 1974; Karr 1976; Pearson 1982; Cracraft 1985; Ricklefs and
Schluter 1993). Indeed, the great diversity of habitat types, the complex topographies, and range
of elevations found in Neotropical areas, together with historical events, undoubtedly have been
important influences on patterns of bird species composition and richness.
Many Neotropical regions have, in recent times, experienced considerable deforestation (e.g.,
Houghton et al. 1991; Sayer and Whitmore 1991; Reid 1992; Whitmore and Sayer 1992), and
much attention by conservation groups, media, and the public has been directed to the defor-
estation crisis in Neotropical rain forests (especially the Amazon basin) (e.g., Hecht and Cock-
burn 1990; Miller and Tangley 1991; Myers 1992). Much less attention, however, has been given
to montane forests in the Neotropics (T. Parker, pers. comm.), despite the fact that these forests
harbor many endemic species and have been subject to habitat alteration for centuries (see Long
1993). In fact, recent evidence reveals that forest loss is greatest in countries with high endemism
levels (Balmford and Long 1994). Andean forests, which contain 6.3% of the world's bird species
(Fjeldsa and Krabbe 1990), have only 10 to 27% of forest remaining in the Colombian Andes
(Henderson et al. 1991; Etter 1993). The situation is even more grim in Ecuador, where less
than 4% of cloud forests remain intact (Dodson and Gentry 1991). In contrast, only 6 to 10%
of Amazonia has been deforested, although certain Amazonian regions experience much greater
pressure than others (Skole and Tbcker 1993; see also Balmford and Long 1994).
Here we draw attention to Neotropical montane avifaunas, with the intent of documenting
some of their differences from lowland counterparts and their contribution to the region's bio-
diversity. We focus on montane birds of northwestern South America and southern Central
America, describing general community patterns and taxonomic affiliations of montane and low-
land faunas. Additionally, we summarize patterns of restricted-range species and status of threat-
ened species in this region. Many of the patterns that we report here are well known to orni-
thologists who conduct research in the Neotropics. By quantifying these patterns, however, we
aim to highlight and better document the importance of montane forest avifaunas to biodiversity
conservation.
METHODS
We examined the structure and composition of the regional avifauna of northwestern South
America and southern Central America by compiling information on the distribution, endemism,
body size, and feeding behavior of landbird species using general sources (Mayr and Phelps
1967; Delacour and Amadon 1973; Ridgely 1976; Meyer de Schauensee and Phelps 1978; Stiles
1983; Hilty and Brown 1986; Isler and Isler 1987; Ridgely and Gwynne 1989; Ridgely and
T\idor 1989, 1994; Stiles and Skutch 1989; Fjeldsa and Krabbe 1990; Karr et al. 1990; Willard
et al. 1991; Collar et al. 1992; ICBP 1992; Dunning 1993; Remsen et al. 1993; BirdLife Inter-
national, unpubl. data), supplemented by specific accounts (Fitzpatrick and O'Neill 1986; Graves
1988; Graves and Uribe 1989; Renjifo 1991, 1994; Stiles 1992; Vuilleumier et al. 1992) and
personal observations of the authors. We generally followed the taxonomic classification of
Clements (1991); use of a different classification system (e.g., Sibley and Monroe 1990) would
have led to slightly different results, but would not be expected to change the major conclusions
of this review.
We compiled distributional data for birds over an area of approximately 2,180,000 km2, which
included continental Costa Rica, Panama, Colombia, and Venezuela (Fig. 1). We used geographic
rather than political criteria to separate the five montane and two lowland regions used in this
review. Broad regional boundaries were defined by elevation or habitat discontinuities between
regions. Lowland areas were divided into cis-Andean and trans-Andean regions by the Andes
(cf. Haffer 1967). Cis-Andean encompassed lowland areas east and south of the Andes, including
the Orinoco Basin and a sector of the Amazon Basin; trans- Andean encompassed lowlands north
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CONSERVATION OF MONTANE AVIFAUNAS 579
Fig. 1
. Schematic map showing location of the five montane systems in southern Central America and
northwestern South America used in this review. Stippled areas indicate montane regions. Heavy black lines
indicate coastlines and country borders.
and west of the Andes, including the Pacific and Caribbean drainages of these four countries.
We did not include avifauna of the dry northwestern Costa Rica in this review because it has
strong affinities to northern Central America and differs markedly from avifaunas of southern
Central America (Stiles 1983; Stiles and Skutch 1989; Howell and Webb 1995).
We analyzed the five largest mountain systems of southern Central America and northwestern
South America (Fig. 1). Three of these montane regions
- Chiriqui-Talamanca mountains, Sierra
Nevada de Santa Marta, and Tepuis
- are totally isolated from other mountains. Indeed, Tepuis
are a series of relict table top mountains in the Guiana Shield which are largely isolated from
each other (Mayr and Phelps 1967; Meyer de Schauensee and Phelps 1978). Colombian and
Venezuelan Andes are relatively more connected, but are divided by the Tdchira depression, an
important biogeographic barrier forming a gap of 40 km (elevation from 800-900 m) between
the two systems (Fjeldsa and Krabbe 1990; Hernandez 1990). Although the Colombian Andes
are divided into three distinct Cordilleras separated by two large and deep Inter-Andean valleys
of ca. 500 m and ca. 1,000 m elevation, these Cordilleras merge in the south (Hilty and Brown
1986). We treat the Colombian Andes as a single unit for this analysis, but it should be recog-
nized that the distributions of a number of bird species are restricted to certain regions (e.g.,
Central Andes of Colombia and Ecuador, Eastern Andes of Colombia; Hilty and Brown 1986;
Long 1993). The eastern slope of northern Perija\ as well as the eastern slope of the Tamd massif,
were included in biogeographic Colombian Andes, even though they are politically within Ven-
ezuela. Venezuelan Andes, as used here, encompassed high mountains of the west, as well as
lower coastal Cordilleras. The oldest mountain system in the area is the Tepuis, which has been
above sea level since Precambrian and was uplifted in the Tertiary. In contrast, Sierra Nevada
de Santa Marta and Chiriqui-Talamanca mountain systems in Colombia and southern Central
America, respectively, were uplifted during the Pliocene and Pleistocene (Haffer 1970, 1974).
The highest mountains among these five regions are in Sierra Nevada de Santa Marta (hereafter
Santa Marta), with peaks at nearly 5,800 m; other regions decrease in elevation in the following
order: Colombian Andes, Venezuelan Andes, Chiriqui-Talamanca, Tepuis.
We did not include avifaunas from Cordillera de Caripe and Paria Peninsula in northeastern
Venezuela, Serrania de la Macarena in central Colombia, and Serrania del Darie"n and Cerro
Pirre"
along the border between Colombia and Panama. At least 17 bird species are restricted to
these mountain ranges (Haffer 1974; Robbins et al. 1985; Hilty and Brown 1986; Long 1993;
BirdLife International, unpubl. data). Inclusion of these areas would be valuable, but lack of
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580 ORNITHOLOGICAL MONOGRAPHS NO. 48
specific details regarding elevational distributions would contribute additional sources of error
to this study.
When a species was present in at least one region, we added the following information to the
database: (1) trophic group (fruit or fruit and seeds [FR]; large invertebrates and small vertebrates
[LI]; large invertebrates, small vertebrates, and fruit (i.e., large omnivores) [LO]; small inver-
tebrates [SI]; small invertebrates and fruits (i.e., small omnivores) [SO]; nectar and insects [NI];
nectar, fruit, and insects [NF]; vertebrates [PR]; carrion [CA]; seeds and insects [SE]; herbivore
[HE]; (2) body weight; (3) restricted-range species (i.e., species that occupy a geographic range
<50,000 km2, BirdLife International, unpubl. data); (4) "endemic" species (i.e., species restricted
to a single biogeographic region); (5) minimum and (6) maximum elevations regularly used by
species in each montane region. We limited our analyses to resident and migrant landbirds, and
excluded birds that take primarily aquatic foods.
Species were assigned to only one trophic group based on their primary food habits. If no
data were available on a species' diet and we had no personal knowledge, then we placed the
species in the same category as other members of that genus. Placement of species in either SI
or LI (or SO, LO) categories often was somewhat arbitrary because no information about prey
size was available in any reference that we examined. In those cases, decisions about trophic
group were based on bird's overall size and bill size.
We assigned birds to one of seven size classes based on body weights: ^ 10 g, 10-25 g, 25-
50 g, 50-100 g, 100-300 g, 300-1,000 g, > 1,000 g. We used body weight of a similar-sized
congener when no data were available for a species and average weight when sexes differed.
We used four elevational zones defined by Chapman (1917, 1926, 1931) and followed in
recent treatments of Neotropical avifaunas (Meyer de Schauensee and Phelps 1978; Hilty and
Brown 1986; Fjeldsa and Krabbe 1990). These zones are tropical (equivalent to lowland), sub-
tropical, temperate, and pdramo. We adopt the original terms used by Chapman because of then-
widespread use by later authors and for ease of presentation; we recognize, however, that use of
some terms (e.g., temperate) is misleading. Stiles and Skutch (1989) did not separate the sub-
tropical and temperate avifaunas in the Chiriqui-Talamanca; however, we acknowledge these
avifaunas here based on similar vegetation and elevational zones. In the relatively low lying
Tepuis, a distinction cannot be drawn between temperate and p&ramo
avifaunas (Chapman 1931;
Mayr and Phelps 1967). The actual elevation range of these zones changes among regions (as
well as within regions) because of variation in rainfall, height of mountain range, and local
topography. We assigned species to zones based on their elevational distribution and the char-
acterization of their habitats. We could not control for fine-scale differences in elevational limits
of these zones within regions and, consequently, some errors likely occur in our classification.
We feel, however, that these errors are uncommon given the broad elevational categories used.
Moreover, such misclassifications would tend to obscure rather than overemphasize patterns
described here. It would likely be wrong to assume, however, that these elevational zones control
the distributional limits of most bird species (e.g., Terborgh 1971, 1985), but such zones are
used here to facilitate broad scale comparisons among regions.
The limits for tropical, subtropical, temperate, and paramo were set to < 1,500 m, 1,500-2,500
m, ^2,500 m to timberline (ca. 3,500 m), respectively, for the Colombian and Venezuelan Andes,
and Santa Marta; < 1,200 m (Caribbean slope) or 1,500 m (Pacific slope), 1,200 (or 1,500 m)-
2,000 m, and >2,000 m to timberline (ca. 3,000 m), respectively, for Chiriqui-Talamanca. In the
Tepuis, we defined only two elevational zones: < 1,000 m, tropical; ^1,000 m subtropical.
We compared avifaunas among montane regions, elevational zones, and between montane and
lowland areas. In addition to a general description of montane and lowland avifaunas, we focused
our comparisons on: distribution of species among species-rich families and genera; trophic
group composition; number and composition of restricted-range and "endemic" species; and
size class distribution. Species-rich families and genera for the entire lowland and montane
avifauna (n = 1,800 species) were defined as those families and genera with equal to or more
than 45 and 10 species, respectively. We used Sorensen's similarity coefficient (Pielou 1977) to
evaluate similarity in species composition among regions.
To determine whether montane avifaunas were simply chance assemblages of the combined
lowland and montane avifaunas, we generated random montane communities using Monte Carlo
simulations (n = 100 runs); the total source pool consisted of all bird species in the data set.
Selection of the source pool for simulations can be problematic (e.g., Connor and Simberloff
1978; Diamond and Gilpin 1982; Graves and Gotelli 1983; Harvey et al. 1983; Gilpin and
Diamond 1984; see also review in Wiens 1989) and may greatly affect interpretation of results.
For this study, we used the entire data set (i.e., bird species occurring in lowland or montane
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CONSERVATION OF MONTANE AVIFAUNAS 581
TABLE 1
Numbers of Species, Genera, Families, and Restricted-range (i.e., <50,000 km2) Species
(RRS) Contained in Different Biogeographic Areas and Elevational Zones. Species
Restricted to a Biogeographic Region ("Endemics") Follow RRS in Parentheses. For
"Restricted Lowland" and "Restricted Montane" Areas, Genera and Family Numbers
Represent Those That Have Species Limited to Lowland or Montane Areas, and Do Not
Represent Genera or Families Totally Restricted to Those Areas (See Text for
Description of Regions)
Species Genera Families RRS
Areas
Total lowland 1,366 502 52 148
Restricted lowland 923 387 49 96
Total montane 877 372 48 217
Restricted montane 434 208 38 165
Shared montane-lowland 443 260 44 52
Regions
Cis-Andean 975 445 49 32(16)
Trans-Andean 932 412 46 120(79)
Colombian Andes 634 324 46 95(47)
Venezuelan Andes 350 215 39 37(17)
Chiriqui-Talamanca 253 174 39 51(47)
Santa Marta 224 165 36 21(14)
Tepuis 137 106 28 37(17)
Montane Elevational Zones
Subtropical 807 346 47 186
Temperate 465 241 43 99
Paramo 117 86 26 32
Totals 1,800 596 52 313
areas) and assumed that if communities were assembled randomly, then each of these species
would have an equivalent chance of occurring in montane habitats. This source pool, however,
was biased because it only contained birds that occurred in the countries and montane regions
selected for this study, and thus did not include all bird species found in montane or cis- Andean
and trans-Andean lowlands.
We used chi-square tests for independence to compare distributions of species between areas
with respect to trophic groups, families, or size classes. Area comparisons included (1) lowland
vs. montane areas, (2) among montane regions, and (3) among elevational zones in montane
areas. In all cases, the null hypothesis was that distributions did not differ between areas, among
regions, or among zones. When necessary, categories were excluded or merged if expected cell
frequencies did not meet assumptions of the chi-square test (see results). Differences in number
of species within taxonomic or trophic groups were compared between randomly generated and
observed montane communities using t-tests (Zar 1984:100).
Similarly, chi-square tests were used to examine the composition of restricted-range species
relative to widespread species (i.e., distribution among elevational zones, lowland vs. montane
areas, trophic groups, etc.). To compare number of species within genera between lowland and
montane areas, we used Fisher's Exact Test (Sokal and Rohlf 1981); only genera with 10 or
more total species were examined.
RESULTS
General description of avifauna.
- The avifauna of montane and lowland regions combined
was composed of 1,800 species representing 52 families and 596 genera (Table 1); this is almost
20% of the world's bird species. The total species list contained 673 non-Passeriformes, 590
suboscine Passeriformes, and 537 oscine Passeriformes. Six families (Tyrannidae, Thraupidae,
Trochilidae, Formicariidae, Emberizidae, and Furnariidae) accounted for 51.8% of the 1,800
species (Table 2).
A total of 877 species representing 48 families was present in all montane regions combined;
1,366 species from 52 families were present in lowland areas (Table 1). Within the lowlands,
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582 ORNITHOLOGICAL MONOGRAPHS
NO. 48
TABLE 2
Number of Total and Restricted-range (RRS) Species Contained within Families in
Lowland and Montane Areas of Northwestern South America and Southern Central
America. Total Number of Species for These Families (i.e., Montane/Lowland Combined)
Are Provided. In All Cases Expected Values for Chi-Square Tests Used in Text Follow
Observed Values in Parentheses. Only Families with More than 45 Species Total Are
Included in the Table
Family Lowland Montane RRS Total
Accipitridae 44(40.0) 24(28.0) 0(9.8) 52
Psittacidae 53(48.9) 30(34.1) 14(12.9) 69
Trochilidae 114(133.1) 112(92.9) 51(34.1) 182
Picidae 41 (33.0) 15 (23.0) 7 (9.2) 49
Fumariidae 48(58.3) 51(40.7) 18(15.8) 84
Formicariidae 128(101.3) 44(70.7) 34(29.1) 155
Cotingidae 37(34.7) 22(24.3) 12(8.8) 47
Tyrannidae 178(167.8) 107(117.2) 19(42.9) 229
Parulidae 48(57.1) 49(39.9) 13(12.8) 68
Emberizidae 64(71.2) 57(49.8) 20(18.6) 99
Thraupidae 120(129.5) 100(90.5) 40(34.1) 182
Total 875 611 228 1,216
the cis-Andean region contained slightly more species, genera, and families than did the trans-
Andean. Among montane regions, greater numbers of species, genera, and families were found
in the Colombian Andes, followed in decreasing order by Venezuelan Andes, Chiriqui-Talaman-
ca, Santa Marta, and Tepuis (Table 1). We suggest caution when interpreting and comparing
among regions the species richness values reported here as these regions differ in total land area
(an important determinant of species richness patterns), as well as in a number of environmental
factors (e.g., rainfall, seasonally, etc.). Our purpose here is not to highlight the effects of area
on species richness patterns, but rather compositional differences between montane and lowland
avifaunas. The latter comparisons are particularly important from a conservation perspective,
and that, together with information on number of total and "endemic" species, will facilitate
setting of conservation priorities.
Much of the region's avifauna was restricted to either montane (435 species, 24.2%) or lowland
(923 species, 51.3%) areas; 442 species (24.6%) were found in both areas (Table 1). Four
families, Opisthocomidae, Psophiidae, Burhinidae, and Sylviidae, only occurred in the lowlands.
Both Psophiidae and Opisthocomidae had one species in the cis-Andean region. Burhinidae had
one species in both cis-Andean and trans-Andean regions, whereas, Sylviidae had five and four
species in cis-Andean and trans-Andean regions, respectively.
Three families were restricted to higher elevations: Ptilogonatidae, Alaudidae, and Oxyrun-
cidae. The two species of Ptilogonatidae are only found in Chiriqui-Talamanca highlands. A very
localized distribution in the Colombian Andes of one species of Alaudidae is a marginal isolate
of a widespread northern hemisphere species. Oxyruncidae (Oxyruncus cristatus) was more prob-
lematic because it is found in the subtropical zone, as well as the upper tropical zone. We
regarded it here solely as a montane taxon because it is always associated with mountains in
our study area (note: other classification systems would include this species within the subfamily
Cotinginae; e.g., Sibley and Monroe 1990).
One-half of the bird species present in montane regions were strictly montane; species re-
stricted to montane areas were represented by 209 genera of which 93 were confined to the
mountains. Some of these genera, however, are found at lower elevations in other regions (Sick
1985; Fjeldsa and Krabbe 1990; Willis 1992; Parker and Goerck 1997).
Nearly 20% (170 species) of the bird species found in montane areas were restricted to a
single elevational zone; 127 species and 13 genera were restricted to the subtropical zone, 30
species and nine genera to the temperate zone, and 13 species and nine genera to paramo
(Appendix 1). Two families, Trochilidae (12 genera) and Fumariidae (seven genera), accounted
for most of the genera restricted to a particular elevational zone. The proportion of montane
species restricted to specific elevational zones was notably similar among Chiriqui-Talamanca,
Colombian Andes, and Venezuelan Andes (i.e., 72% subtropical zone, 19% temperate zone, and
9% paramo). Santa Marta diverged from these regions by having a greater proportion of montane
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CONSERVATION
OF MONTANE AVIFAUNAS 583
TABLE 3
Sorensen's Similarity Coefficients Comparing Species Composition between Regions Using
All Species (A) and Species Restricted to Montane Regions (B)
Cis- Trans- Colombian Venezuelan Chiriqui- Santa
Andean Andean Andes Andes Talamanca Marta
A. ALL SPECIES
Trans-Andean 0.56 - - - - -
Colombian
Andes 0.30 0.46 - - - -
Venezuelan
Andes 0.24 0.29 0.61 - - -
Chiriquf-Talamanca 0.18 0.27 0.29 0.32 - -
Santa
Marta 0.21 0.27 0.48 0.53 0.37 -
Tepuis 0.15 0.11 0.16 0.24 0.16 0.24
B. RESTRICTED
MONTANE SPECIES
Venezuelan
Andes - - 0.59 - - -
Chiriquf-Talamanca - - 0.09 0.12 - -
Santa Marta - - 0.29 0.40 0.10 -
Tepuis - - 0.07 0.10 0.05 0.11
species restricted to paramo (43%). The relatively small area but high elevation of Santa Marta
may account for this departure.
The two lowland regions generally shared more species and had a higher similarity coefficient
than did most montane regions (Table 3); Colombian and Venezuelan Andes, however, displayed
the greatest similarity of all pair-wise comparisons, which likely corresponds to their geographic
proximity. Tepuis had the most distinct avifauna as indicated by consistently low similarity
coefficient scores with other regions (Table 3). When similarities in species compositions were
compared among montane regions excluding those taxa that also occur in lowland areas, a
two-fold or greater reduction in similarity coefficients generally occurred (Table 3). To a large
extent, then, species shared among montane regions also are shared with lowland avifaunas.
Composition of lowland and montane avifaunas. - Despite similarities in family composition,
the distribution of species within the most species-rich families (i.e., 11 families with ^45
species) differed between lowland and montane areas (^ = 43.7, P < 0.0001; Table 2). In
particular, montane areas had more Trochilidae and Parulidae, but fewer Picidae and Formica-
riidae than did lowlands when based on expected values (Table 2).
When numbers of species contained within species-rich genera (i.e., >10 species) were com-
pared between lowland and montane areas, seven genera showed significantly different distri-
butions than expected (Table 4); 1.4 of 29 genera would have been expected to show differences
by chance at P < 0.05 significance criteria. Four of the seven genera (Grallaria, Myioborus,
Basileuterus, Atlapetes) were more species-rich in montane areas, whereas three (Pteroglossus,
Myrmotherula, and Myrmeciza) had more species in lowlands (Table 4).
Distribution of species among major trophic categories differed between lowland and montane
areas (x1 = 27.5, P < 0.001; Table 5). Differences in trophic composition were influenced
primarily by nectarivores, as expected given the importance of Trochilidae in montane areas.
Indeed, when nectarivores were removed from the analysis, differences in distribution of species
among trophic groups did not differ significantly between lowland and montane areas (x1 = 10.5,
P > 0.15). In contrast, no significant differences in trophic composition were found either among
the five montane regions (x1 = 28.6, P > 0.4; NF combined with SO due to low numbers; CA
and HE excluded from analysis due to low numbers) or among elevational zones (x1 = 19.5, P
> 0.15; Table 5).
In both montane and lowland areas, the majority of the avifauna occurred in the three smaller
size classes (68% and 69% of lowland and montane birds below 50 g, respectively). There was
a trend for larger size classes to be poorly represented in montane bird communities (x1 = 11.3,
P < 0.08).
Comparison of observed and randomly generated montane avifaunas. - We found that mon-
tane communities were not simply chance assemblages; nearly 30% of bird families (15 of 52
families) had a significantly different number of species in montane communities than expected
if patterns were random (Table 6) (note: 2.6 of 52 families would be expected to show differences
by chance alone using P < 0.05 criteria). Moreover, when the number of species occurring in
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584 ORNITHOLOGICAL
MONOGRAPHS NO. 48
TABLE 4
Number of Species Contained within Genera in Lowland and Montane Areas for Those
Genera with More Than 10 Species. Results of Fisher's Exact Test Are Shown (P
Value); Genera Showing Significant Differences in Number of Species between Lowland
and Montane Areas Are Bold. Also Shown Are the Number and Proportion of Species
with Restricted Ranges (RRS)
Number of Proportion of
Genera Lowland Montane P value RRS species RRS species
Crypturellus 10 3 0.239 5 0.42
Buteo 8 10 0.138 0 0.00
Caprimulgus 7 6 0.381 2 0.20
Phaethornis 12 6 0.345 0 0.00
Amazilia 18 7 0.178 5 0.28
Trogon 12 4 0.221 3 0.23
Pteroglossus 10 0 0.014 2 0.20
Picumnus 10 3 0.239 3 0.30
Xiphorhynchus 10 3 0.239 0 0.00
Synallaxis 8 8 0.266 4 0.31
Thamnophilus 12 3 0.141 2 0.14
Myrmotherula 19 2 0.005 2 0.10
Myrmeciza 14 1 0.011 4 0.28
Grallaria 3 17 0.001 10 0.53
Elaenia 9 8 0.306 1 0.08
Phylloscartes 6 5 0.431 6 0.60
Pachyramphus 9 4 0.385 0 0.00
Hylophilus 12 3 0.141 1 0.08
Catharus 8 8 0.266 1 0.10
Turdus 10 12 0.116 1 0.07
Thryothorus 15 6 0.228 4 0.25
Dendroica 14 10 0.316 0 0.00
Myioborus 1 10 0.002 6 0.60
Basileuterus 8 13 0.038 5 0.31
Atlapetes 3 14 0.001 6 0.40
Sporophila 13 5 0.248 0 0.00
Icterus 10 2 0.141 0 0.00
Euphonia 20 6 0.081 2 0.09
Tangara 26 22 0.147 3 0.08
TABLE 5
Number of Species Contained within Trophic Groups by Area and Elevational Zones.
Only Montane Zones are Shown. See Text for Descriptions of Trophic Groups
Trophic group
SI SO LI LO FR SE PR NI NF
Areas
Lowland 444 242 135 73 214 59 62 114 16
Montane 280 177 60 34 115 37 43 122 6
Montane
Regions
Colombian Andes 199 127 49 24 79 26 38 82 6
Venezuelan Andes 118 64 23 11 47 16 23 42 3
Chiriqui-Talamanca 84 64 13 13 26 18 15 18 0
Santa
Marta 80 41 15 6 23 17 18 18 1
Tepuis 47 31 5 4 17 8 7 17 1
Montane
Elevational Zones
Subtropical 255 167 58 31 109 33 41 104 6
Temperate 161 88 28 15 53 21 30 64 2
Paramo 46 18 2 3 9 7 6 24 1
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CONSERVATION
OF MONTANE AVIFAUNAS 585
TABLE 6
Number of Species Contained within Species Rich Montane Families for Observed and
Randomly Generated Montane Communities (See Text). Mean Number and Standard
Deviation of Species Are Shown for Random Communities (n = 100 Runs). Results of
t-test Comparing Differences Between Number of Species in Observed and Random
Communities Are Shown. Only Families Showing Significant Patterns Are Included
Montane species
Family Observed Random r-value P value
More Species-rich
in Montane Areas:
Odontophoridae 10 5.6 ± 1.8 -2.5 <0.05
Trochilidae 112 88.1 ± 7.3 -3.3 <0.01
Furnariidae 51 41.7 ± 4.4 -2.1 <0.05
Rhinocryptidae 8 4.5 ± 1.4 -2.4 <0.05
Ptiligonatidae 3 1.3 ± 0.8 -2.0 <0.05
Ibrdidae 26 15.0 ± 2.7 -4.1 <0.001
Hirundinidae 13 8.6 ± 2.0 -2.2 <0.05
Parulidae 49 32.8 ± 3.8 -4.3 <0.001
Less Species-rich
in Montane Areas:
Galbulidae 1 5.9 ± 1.7 2.9 <0.01
Bucconidae 4 12.0 ± 2.5 3.2 <0.01
Ramphastidae 8 14.8 ± 2.7 2.5 <0.05
Picidae 15 24.3 ± 3.7 2.5 <0.05
Formicariidae 44 75.6 ± 6.2 5.1 <0.001
Pipridae 9 15.9 ± 2.5 2.8 <0.01
Sylviidae 0 3.1 ± 1.2 2.5 <0.05
trophic groups was compared between observed and randomly generated montane communities,
we found that three of nine trophic groups showed non-random patterns (CA and HE excluded).
Specifically, more small omnivores and nectarivores, and fewer large insectivores were recorded
in the montane community than expected if those communities were randomly assembled subsets
of the region's avifauna (t > 2.0, P < 0.05).
Patterns of restricted-range species. - Based on distributional data made available by BirdLife
International (unpubl. data), 313 species (17.4% of the total) were restricted-range species
(<50,000 km2); most occurred in only a single biogeographical region (142 in montane and 95
in lowland) (Table 1). Restricted-range species occurred in a wide range of families (32 of 52)
and genera (172 of 596), but they were not distributed equally among species-rich families Ot2
= 35.50, P < 0.001) (Table 2). Tyrannidae, as well as Accipitridae, had a low number of small
range species. When these two families were removed from the analysis, the frequency of re-
stricted-range species no longer differed (j2 = 6.3, P > 0.60). Although Terborgh and Winter
(1983) suggested that species with small geographic ranges tend to be of small size, we found
no difference in body size between bird species with restricted ranges and other bird species Ot2
= 10.4, P > 0.1).
Montane areas had both a greater number and percentage of species with small geographic
ranges than did lowland areas (Table 1). In fact, of those birds only occurring in either montane
or lowland areas, more restricted-range species than expected occurred in the montane avifaunas
(X2 = 172.6, P < 0.0001). Moreover, restricted-range species were particularly prevalent among
those genera that were significantly associated with montane areas (Table 4). Such differences,
however, may be artifacts of current classification systems; further analysis may reveal that many
wide-ranging species are actually morphologically similar allospecies (e.g., see Isler et al. 1997).
DISCUSSION
Montane avifaunas of northwestern South America and southern Central America contribute
substantially to the region's diverse avifauna. As this review has confirmed, montane avifaunas
are not simply a random subset of the rich lowland communities. Within the study region, 93
genera and 435 species occur exclusively in the mountains. Genera restricted to montane areas
represent evolutionary lineages that would be lost if montane habitats are not effectively pro-
tected (cf. Humphries et al. 1991; Williams et al. 1991). Moreover, the Colombian and Vene-
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586 ORNITHOLOGICAL
MONOGRAPHS
NO. 48
zuelan Andes and Santa Marta were found to have some of the Neotropics' greatest density of
recently evolved species, and thus may represent critical regions for evolutionary change (Fjeldsa
1994). Consequently, the evidence strongly supports the fact that montane avifaunas are not only
distinct bird communities but also centers of radiation that deserve special attention (see also
Erwin 1991).
One of the most important ecological differences between lowland and highland areas is the
greater importance (i.e., proportionally more species than expected) of nectarivores in montane
avifaunas. In fact, the most species-rich genera restricted to the mountains are nectarivores,
namely Eriocnemis, Heliangelus, and Diglossa. This increased importance of nectarivores in
montane areas also has been documented elsewhere at different geographic scales of analysis
(Stiles 1985; Graham 1990). Indeed, differences between montane and lowland avian commu-
nities match major changes in plant communities in terms of plant family composition and
physiognomy (Gentry 1988, 1992). For example, the greatest number of bird-pollinated plants
occurs in middle elevations, and bird-pollinated families such as Ericaceae and Gesneriaceae
become more important with increasing elevation (i.e., abundance and species richness) (Stiles
1981; Gentry 1988, 1992).
Differences in land surface area may largely account for observed differences in diversity of
lowland and montane bird communities (Table 1). Yet at the scale of our biogeographic units,
montane areas were more distinct from each other than were the two lowland regions; this
highlights the importance of regional diversity at high elevations. The low species overlap among
montane avifaunas calls for a system of protected areas covering a wide range of geographic
locations.
Restricted-range species are of particular concern from a conservation perspective. Given their
small geographic ranges, these species are highly vulnerable to habitat destruction and other
forms of human pressure. Consequently, degree of overlap in distribution of restricted-range
species is considered a useful criterion for assigning conservation priorities. The majority of bird
species with restricted-ranges in the regions reviewed here are found in the mountains (Table 1
;
see also Terborgh and Winter 1983; ICBP 1992; BirdLife International, unpubl. data). Thus, the
greater total number and proportion of restricted-range species in montane avifaunas further
highlight these regions as areas of utmost conservation concern.
As we have attempted to demonstrate in this review, the distinctiveness and richness of the
montane avifauna provide a strong argument for establishment of montane regions as priority
areas for conservation. When taken together with human activities in these regions, however,
the urgency of conservation action becomes substantially more obvious and pressing. The An-
dean region of Colombia, with the highest diversity of montane birds and the greatest number
of "endemics", is considered one of the most human-modified regions in South America (Hilty
1985; Myers 1988). Similarly, major population centers of Costa Rica and Venezuela are located,
to a large extent, in or close to the mountains. Although Panama's population is centered in the
lowlands, deforestation in the Chiriquf Mountains is extensive. Worldwide trends reveal an
alarming correlation between avian endemism patterns and deforestation, and the countries rep-
resented in our study were among those with the highest degrees of endemism and deforestation
rates (see Fig. lb in Balmford and Long 1994). As a result, 82 species of these montane regions
are currently listed as threatened or near threatened (Collar et al. 1992), and as many as 50 are
restricted to the mountains (Appendix 2).
Despite the richness, distinctiveness and endangerment of the montane avifaunas, conservation
of Neotropical mountain ecosystems has been largely neglected. More emphasis towards con-
servation of montane ecosystems is sorely needed.
ACKNOWLEDGMENTS
We all strongly share Ted Parker's enthusiasm for montane birds and his deep concern for
their conservation. We are profoundly indebted to Ted for his willingness to share so freely with
us his immense knowledge of birds, conservation, and natural history. We thank David Wege
and BirdLife International for providing us with unpublished information on the birds with small
geographic ranges for the countries included in this review. This paper would not have been
possible without their help. We thank J. V. Remsen, G. Barrantes, M. Kelsey, and an anonymous
reviewer for their comments and suggestions on the manuscript. Any errors contained within
this paper rest solely with the authors. Due to space limitations, we have not printed a full
appendix here, but this is available from the authors upon request by sending a blank disk.
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CONSERVATION OF MONTANE AVIFAUNAS 587
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590 ORNITHOLOGICAL MONOGRAPHS NO. 48
APPENDIX 1
Species Restricted to a Single Elevation al Zone within Montane Areas. Species of
Genera Restricted to a Particular Elevational Zone Are Indicated by an Asterisk
SUBTROPICAL ZONE
Nothocercus bonapartei; Crypturellus ptaritepui; Crypturellus obsoletus; Penelope perspicax; Odontopho-
rus melanonotus; Odontophorus strophium; Odontophorus columbianus; Odontophorus leucolaemus; Lep-
totila conoveri; Pyrrhura viridicata; Pyrrhura hoematotis; Touit batavica; Amazona dufresniana; Otus col-
ombianus; Otus guatemalae; Nyctibius leucopterus; Caprimulgus whitelyi; Campylopterus hyperythrus\
Campylopterus duidae; Chlorostilbon alice; Polytmus milleri; Eupherusa eximia*; Eupherusa nigriventris*;
Elvira chionura*; Elvira cupreiceps*; Lampornis hemileucus; Adelomyia melanogenys*; Heliodoxa rubino-
ides\ Sternoclyta cyanopectus*; Urochroa bougueri*; Boissonneaua matthewsii; Eriocnemis godini; Erioc-
nemis mirabilis; Urosticte ruficrissa; Schistes geoffroyi*; Selasphorus ardens; Pharomachrus fulgidus; Tro-
gon aurantiiventris; Veniliornis kirkii\ Cranioleuca demissa; Siptornis striaticollis*; Roraimia adusta*;
Premnornis guttuligera*; Premnoplex brunnescens*; Premnoplex tatei*; Margarornis stellatus; Syndactyla
guttulata; Anabacerthia striaticollis; Thripadectes virgaticeps; Thripadectes melanorhynchus; Automolus
roraimae; Sclerurus albigularis; Thamnophilus unicolor; Thamnophilus insignis; Thamnomanes plumbeus;
Myrmotherula behni; Herpsilochmus roraimae; Chamaeza ruficauda; Chamaeza turdina; Grallaria excelsa;
Grallaria alleni\ Grallaria chthonia\ Grallaria bangsi\ Grallaria kaestneri\ Grallaria flavotincta\ Grallaria
hypoleuca; Grallaricula loricata; Pipreola lubomirskii; Pipreola whitelyi; Lipaugus streptophorus; Rupico-
la peruviana; Oxyruncus cristatus*\ Chloropipo flavicapilla; Todirostrum russatum; Phyllomyias zeledoni;
Phyllomyias plumbeiceps; Zimmerius viridiflavus; Elaenia dayi; Elaenia pallatangae\ Mecocerculus minor;
Phylloscartes poecilotis; Phylloscartes nigrifrons; Phylloscartes chapmani; Phylloscartes superciliaris; Pla-
tyrinchus flavigularis; Myiophobus ftavicans; Myiophobus roraimae; Myiophobus pulcher; Contopus lugub-
ris; Myiarchus cephalotes; Cyanolyca cucullata; Troglodytes rufulus; Myioborus castaneocapillus; Myiobo-
rus brunniceps; Myioborus albifacies; Myioborus cardonai; Basileuterus bivittatus; Basileuterus griseiceps;
Melozone biarcuatum*; Melozone leucotis*; Atlapetes gutturalis; Atlapetes flaviceps; Atlapetes fuscooliva-
ceus; Atlapetes albofrenatus; Atlapetes personatus; Pselliophorus luteoviridis; Oreothraupis arremonops*;
Pheucticus chrysopeplus; Chlorospingus parvirostris; Thlypopsis fulviceps; Mitrospingus oleagineus; Pi-
ranga leucoptera; Bangsia melanochlamys; Bangsia aureocincta; Iridosornis porphyrocephala; Euphonia
elegantissima; Euphonia musica; Chlorochrysa nitidissima; Tangara xanthocephala; Tangara rufigenis;
Tangara ruficervix; Tangara labradorides; Tangara cyanotis; Diglossa duidae; Diglossa major; Diglosso-
pis glauca.
TEMPERATE ZONE
Metriopelia melanoptera*; Hapalopsittaca fuertesi; Aegolius ridgwayi; Caprimulgus saturatus; Panterpe
insignis*; Eugenes fulgens*; Aglaeactis cupripennis*; Coeligena orina; Coeligena lutetiae; Ensifera ensi-
fera*; Ramphomicron microrhynchum; Metallura iracunda; Andigena hypoglauca; Veniliornis nigriceps;
Grallaria milleri; Grallaricula lineifrons; Acropternis orthonyx*; Contopus ochraceus; Muscisaxicola ma-
culirostris; Cyanolyca turcosa; Eremophila alpestris*; Urothraupis stolzmanni*; Acanthidops bairdii*; Sal-
tator cinctus; Conirostrum rufum; Conirostrum sitticolor; Hemispingus goeringi; Hemispingus verticalis;
Buthraupis wetmorei; Anisognathus igniventris.
PARAMO
Phalcoboenus carunculatus*; Vanellus resplendens; Chalcostigma heteropogon*; Chalcostigma herrani*;
Oxypogon guerinii*; Cinclodes fliscus*; Cinclodes excelsior*; Leptasthenura andicola*; Asthenes wyatti*;
Asthenes flammulata*; Cnemarchus erythropygius*; Junco vulcani*; Phrygilus unicolor*
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