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We conducted an ornithological survey of the Colombian slope of Cerro Tacarcuna, the highland region adjacent to the 'Darién Gap' on the Colombia / Panama border, and one of the most poorly known and threatened regions in the world. We present novel data on distribution, habitat, breeding biology and vocalisations for 27 species, including the first confirmed records in Colombia of Ochraceous Wren Troglodytes ochraceus and Beautiful Treerunner Margarornis bellulus, and the first records in the Darién highlands of Black-headed Antthrush Formicarius nigricapillus, Scaly-throated Foliage-gleaner Anabacerthia variegaticeps, Yellow-throated Chlorospingus Chloropingus flavigularis hypophaeus and, based on previously overlooked specimens, report the first confirmed records for Colombia of Sooty-faced Finch Arremon crassirostris. In addition, we collected the first Colombian specimens of Violet-capped Hummingbird Goldmania violiceps, Bare-shanked Screech Owl Megascops clarkii, Tacarcuna Tapaculo Scytalopus panamensis and Varied Solitaire Myadestes coloratus. For several subspecies endemic to the region, we collected the first or second specimens for Colombia. Finally, we discuss the elevational ranges of Darién endemic species and subspecies, which are mostly concentrated above 600 m. The Darién highlands remain poorly studied and threats to their conservation are increasing. Therefore, effective measures are needed, particularly in Colombia, where the sole protected area in the region currently covers forests only below 600 m.
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Luis Miguel Renjifo et al. 46 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club
New bird distributional data from Cerro Tacarcuna,
with implications for conservation in the Darién
highlands of Colombia
by Luis Miguel Renjifo, Augusto Repizo, Juan Miguel Ruiz-Ovalle,
Sergio Ocampo & Jorge Enrique Avendaño
Received 25 September 2016; revised 20 January 2017; published 13 March 2017
         


the world. We present novel data on distribution, habitat, breeding biology and
          
of Ochraceous Wren Troglodytes ochraceus and Beautiful Treerunner Margarornis
bellulus   
Formicarius nigricapillus, Scaly-throated Foliage-gleaner Anabacerthia variegaticeps,
Yellow-throated Chlorospingus Chloropingus avigularis hypophaeus and, based on

of Sooty-faced Finch Arremon crassirostris      
     Goldmania violiceps, Bare-
shanked Screech Owl Megascops clarkii, Tacarcuna Tapaculo Scytalopus panamensis
   Myadestes coloratus. For several subspecies endemic to the

          
        
         
are needed, particularly in Colombia, where the sole protected area in the region
currently covers forests only below 600 m.

isolated massifs of moderate elevation, at the junction between Central and South America
           

as a route for or barrier to the exchange of Central and South American faunas during the



  
          
represent separate evolutionary lineages    et al. 1984,
Miller et al et al. 2012). Some authors have
proposed the existence of isolated forest refuges on the slopes of several massifs in the

et al. 1992)

         
          
Luis Miguel Renjifo et al. 47 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club
              
et al. 2014, Hruska et al 
    
1959, 1967a,b, Bran-Castrillón et al        

to massifs such as Cerro Pirre, Alturas de Nique, Cerro Tacarcuna and its spur Cerro Malí
 et al. 1984, Robbins et al    
1989, Hruska et al
(Angehr et al. 2004), Serranía de Majé (Angehr & Christian 2000) and the foothills of Cerro
Piña (Miller et al. 2011), the last two of which are both isolated massifs north of the Serranía
 

et al. to Alto Barrigonal
(c            
and Pearman      
          
c
and by JMR-O & T. Walschburger in November 2008 to the headwaters of the río Tanelita
c
Unguía, dpto. Chocó et al. 2014). Olaciregui et al. (2016) visited the Serranía de

 

that several bird species will be found on the Colombian side, representing range extensions

America.
Here, we present the results of an expedition to the Colombian slope of Cerro Tacarcuna
in August 2010. We present a list of bird species recorded, several of which represent
noteworthy range extensions, or additional records of poorly known species (e.g. endemics)


Methods
Three main mountain ranges run parallel from the eastern Isthmus of Panama to
   
ranges from its northern outlier, Cerro Sapo, south to the headwaters of the río Jaqué, and
is connected to the Cordillera de Juradó and Altos de Aspavé. To the east, this serranía is
separated from the Serranía de Pirre by c
and Juradó (Angehr et al. 2004). The Serranía de Pirre (1,550 m) extends further south to

        et al. 2004). Nearly 50 km of lowlands
              et
al. 1985). This range rises to 1,875 m at Cerro Tacarcuna and extends north to the Serranía
   
             


 et al. 2014).
Luis Miguel Renjifo et al. 48 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club


The area lies in the headwaters of the río Bonito, on the east slope of the Cerro Tacarcuna
(Fig. 1). We accesed Cuchilla del Lago by opening a trail in primary forest upslope from
the left bank of the río Bonito (c.450 m), close to a property currently occupied by the Toro
family. Based on our observations, the transition from wet to cloud forests on this part of
Cerro Tacarcuna starts at c



Chocó (Poveda-M. et al. 2004).
Three observers made visual and aural records along two transects, between 05.45 h and
c.
   
          
trail we opened to reach the campsite. Simultaneously, two of us undertook c.288 mist-net
  
Figure 1. Map of
north-western South
America and eastern
Panama showing
the location of the
   
geographical features
mentioned in the
text. Land above
600 m is indicated in
grey to black. Circles
represent historical
and recent localities,
including our study
site in the Serranía del
    
  
municipality Acandí,
dpto. Chocó; 2 = Alto
Barrigonal (1,400 m),
municipality Acandí,
   
Cuchilla del Lago (1,150
m), Corregimiento de
Balboa, municipality
Unguía, dpto. Chocó;
4 = headwaters of the
  
1,400 m), Comunidad
 
Unguía, dpto. Chocó;
and 5 = headwaters of
the río Tigre (1,250 m),
municipality Unguía,
dpto. Chocó.
Luis Miguel Renjifo et al. 49 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club

digital recorders with Sennheiser ME-67 shotgun microphones. Recordings are deposited





Results


data or range extensions are presented for 27 species. Most (16) were expected to occur
on the Colombian slope of Cerro Tacarcuna, given that they have been recorded on the
Panamanian slope of Cerro Tacarcuna and adjacent Cerro Malí. However, seven species


Another four records correspond to rare and poorly known species in Colombia, but
previously known from Cerro Tacarcuna.
BARRED HAWK Morphnarchus princeps
  
 (Wetmore 1965, Robbins et al. 1985) and
 et al.  
 (Hilty & Brown 1986).
RUSSET-CROWNED QUAIL-DOVE Geotrygon goldmani
Recorded by voice almost daily at 1,150 m near our campsite, but apparently in smaller
        G. violacea   

Tacarcuna (1,450 m; Wetmore 1968), and recently the Serranía de Jungurudó (Angehr et al.
2004) and Cerro Piña (c.1,100 m; Miller et al. 2011). In Colombia, it is known only from the
headwaters of the río Cutí on Cerro Tacarcuna and the río Juradó (c.90 m) in Chocó (Hilty
& Brown 1986).
VIOLACEOUS QUAIL-DOVE Geotrygon violacea

Cerro Pirre above Cana and El Real, and Cerro Sapo in Panama (Wetmore 1968, Ridgely &

Serranía de San Lucas (Hilty & Brown 1986, Salaman et al. 2002).
BARE-SHANKED SCREECH-OWL Megascops clarkii




          
showed moderate body, wing and tail moult, with relatively large testes (left 5.0 × 1.7 mm).
Stomach contents included parts of beetles (Coleoptera). M. clarkii is rare in Panama, with
            
Luis Miguel Renjifo et al. 50 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club
(1,450 m), the crest of the Cerro Tacarcuna ridge (1,460 m, in both Panama and Colombia)
 
et al. 1985).
VIOLET-CAPPED HUMMINGBIRD Goldmania violiceps
     
   

at the río Tanelita (c.1,400 m), c
 (Wetmore

of the río Cutí at Cerro Tacarcuna near the Panama border (Hilty & Brown 1986).
RED-HEADED BARBET Eubucco bourcierii
            

Tacarcurna (c.et al
 (Hilty & Brown 1986). Recorded in the
northern Central Andes, Serranía de San Lucas and the adjacent West Andes (Salaman et al.

           
Hummingbird Goldmania violiceps; (B) male White-bibbed Manakin Corapipo leucorrhoa 
Myadestes coloratusChlorospingus avigularis
Luis Miguel Renjifo et al. 51 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club
ZELEDON’S ANTBIRD Myrmeciza zeledoni

Dysithamnus puncticeps, Chestnut-backed Antbird Poliocrania exsul, White-cheeked Antbird
Gymnopithys leucaspis, Ocellated Antbird Phaenostictus mcleannani and Black-headed
Formicarius nigricapillus. On 6 August 2010, an adult male was foraging with a

in Panama on Cerro Pirre (c cet
al
ridges such as Serranía de Abibe (1,500 m), Serranía de Baudó (c.900 m), the Chocó lowlands
and adjacent West Andes et al. 2016).
TACARCUNA TAPACULO Scytalopus panamensis


alone, mainly at ground level up to c.0.5 m. Most were on slightly steep and humid slopes,
               


  
gonads (left testis 1.8 × 0.4 mm), but a fresh incubation patch and symmetric moult on its

that S. panamensis 
 
S. chocoensis            
Jungurudó (c (Angehr et al
  
1997). The vocalisations of S. panamensis   
 
184866, 184868). The call of S. panamensis 


& Schulenberg (1997) suggested some similarity between the calls of S. panamensis and S.
chocoensis  S.
 Scytalopus 
adjacent Western Andes of Colombia, from left to right: Nariño Tapaculo S. vicinior 
Pisones, 8 km north-west Jegüadas, municipality Mistrató, dpto. Risaralda, 6 June 1992; Tacarcuna Tapaculo
S. panamensis
August 2010; and Chocó Tapaculo S. chocoensis
Altaquer, municipality Barbacoas, dpto. Nariño, 11 October 2009 (J. E. Avendaño)
Luis Miguel Renjifo et al. 52 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club
chocoensis
nn
of Nariño Tapaculo S. vicinior  S. panamensis in its mean
n = 7) and note shape (e.g. length of stroke ‘tails’). A
natural song of S. panamensis 


second, n = 2). Note shape is similar to that of some individuals of S. chocoensis 
  S. vicinior  
S. chocoensis, but slower than S. vicinior. Thus, S. panamensis appears to be a vocal and
S.
vicinior and S. chocoensis
herein are detailed in Appendix 2.
BLACK-HEADED ANTTHRUSH Formicarius nigricapillus
         
above 1,000 m at our study site. Previously recorded in the Chocó lowlands at Nuquí north


locality in Panama is Nusagandi, western San Blas 
F. analis   

  
 
(Sullivan et al. 2009), plus an aural record below 800 m at our study site suggests possible
contact in the foothills of Cerro Tacarcuna, where the species possibly replace each other


Figure 4. Calls of Scytalopus
S. panamensis
municipality Unguía, dpto. Chocó; (B) Chocó Tapaculo S. chocoensis
et al. 2007), Tambito Nature Reserve, municipality El Tambo, dpto. Cauca, Colombia; (C) Nariño Tapaculo S.
vicinioret al. 2007), Las Nubes Nature Reserve, municipality Jericó,
dpto. Antioquia. Note the distinctive frequency bandwidth and note shape in S. panamensis. Spectrograms
were created using Syrinx v2.6h (Burt 2006) applying the same parameters except for adjusting brightness
to improve note resolution.
Luis Miguel Renjifo et al.  Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club
Figure 5. Primary songs of Scytalopus          
(A) Tacarcuna Tapaculo S. panamensis  
Corregimiento de Balboa, municipality Unguía, dpto. Chocó, Colombia; (B) Chocó Tapaculo S. chocoensis
natural song (LNS 40016, T. S. Schulenberg), El Placer, prov. Esmeraldas, Ecuador; (C) S. chocoensis natural
S. vicinior natural song
 et al 
Colombia.
Luis Miguel Renjifo et al. 54 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club
SPOTTED WOODCREEPER Xiphorhynchus erythropygius
   
Previously known in Panama from Cerro Pirre (1,580 m), Cerro Malí (1,450 m; Robbins
et al. 1985, Wetmore 1972) and Serranía de Majé (c.
Adjacent records in Colombia are from the Serranía del Baudó, ríos Atrato and Truandó
            (Hilty &
Brown 1986), Serranía de Abibe (Olaciregui et al. 2016), the northern Central Andes (Cuervo
et al
BROWN-BILLED SCYTHEBILL Campylorhamphus pusillus
           

(760 m; Wetmore 1972, Robbins et al. 1985) and Serranía de Majé (c
& Christian 2000). In Colombia, known from Serranía de Abibe (Olaciregui et al. 2016), the
northern West Andes at Frontino (Hilty & Brown 1986) and the northern Central Andes
(Cuervo et al
the Tacarcuna massif.
SCALY-THROATED FOLIAGE-GLEANER Anabacerthia variegaticeps
Cranioleuca
erythrops and several Tangara species. The nearest localities to our record are Cerro Flores
            
    
of the West Andes, Colombia (Olaciregui et al. 2016). Taxonomy of Middle and South
American populations of this species is controversial, with some authors suggesting that
A. v. temporalis of the West Andes should be recognised as a separate species from Middle
American populations (Remsen et al  
resolve species limits in the group.
BEAUTIFUL TREERUNNER Margarornis bellulus

Its presence in Colombia was based on mainly habitat continuity and geographic proximity
c
et al. 2002)
and recently Serranía de Majé (1,150 m; Angehr et al
   
       
(Remsen et al. 2016).
RUFOUS-BROWED TYRANNULET Phylloscartes superciliaris

the Panamanian side of Cerro Tacarcuna and Cerro Malí (1,060 m; Wetmore 1972, Ridgely

Andes (Cuervo et al    et al. 2002, Cuervo et al. 2008a) and
Serranía de San Lucas 
Santander (Hilty & Brown 1986).
Luis Miguel Renjifo et al. 55 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club
PIRATIC FLYCATCHER Legatus leucophaius

          et al. 1985), the Chocó
lowlands and northern West Andes (Hilty & Brown 1986, Olaciregui et al. 2016). This is a
widespread species of lowland habitats that has possibly colonised the region due to the

WHITE-BIBBED MANAKIN Corapipo leucorrhoa
              
           
             
Colombia are the humid northern slopes of the West and Central Andes (Hilty & Brown
      C. altera, which also occurs on Cerro

 (Wetmore 1972) and Serranía de Jungurudó (Angehr et al. 2004),
             (Hilty & Brown
1986). These taxa appear to be separate species based on their sympatry in this region,

 et al. 2016). However, lek
behaviour has been studied only in C. altera (Hilty & Brown 1986, Remsen et al. 2016).
OCHRACEOUS WREN Troglodytes ochraceus
  
They searched for insects within dense mosses and bromeliads, climbing vine tangles and
          
mainly of tanagers and furnariids, although the wren also foraged and vocalised in the

sound-recorded. These match recordings from Chiriquí province in Panama (Fig. 6) and
T. solstitialis, which occurs in the West
Andes. This wren ranges from Costa Rica to Panama where it has been recorded on Serranía
de Majé (ccet
al. 1985), and Cerro Malí (c.1,460 m; Wetmore et al
 et al. 2016, Remsen et
al. 2016).
VARIED SOLITAIRE Myadestes coloratus




with an immature collected on 29 August 1996 on Serranía de Majé (Angehr & Christian
 
río Tanelita (c

   et al. 1984), and recently
from Serranía de Majé (cc.1,000 m) (Angehr &
Christian 2000, Angehr et al

Luis Miguel Renjifo et al. 56 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club
Angehr et al. 2004), populations of M. coloratus on Cerro Pirre are genetically distinct from
those of Serranía de Majé (Miller et al. 2007).
BLACK-AND-YELLOW TANAGER Chrysothlypis chrysomelas
   

Chlorospingus species,
          
          et
al. 1984) and Serranía de Majé (Angehr & Christian 2000). Our records represent the third

c.1,400
m), on the east slope of Cerro Tacarcuna et al. 2014); and one observed by J.
Zuluaga-Bonilla on Cerro La Nevera (cc
January 2007 (Fig. 1).
Figure 6. Songs and calls of Troglodytes          
Ochraceous Wren T. ochraceus   
Chiriquí, Panama; (B) T. ochraceus
de Balboa, municipality Unguía, dpto. Chocó, Colombia; (C) Mountain Wren T. solstitialis natural song (BSA
et al. 2007), Ibanasca, Corregimiento de Juntas, municipality Ibagué, dpto. Tolima, Colombia;
T. ochraceus
(E) T. ochraceus            
municipality Unguía, dpto. Chocó, Colombia; (F) T. solstitialis
Frontino, dpto. Antioquia, Colombia.
Luis Miguel Renjifo et al. 57 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club
BLUE-AND-GOLD TANAGER Bangsia arcaei
Singles recorded daily at 1,150 m accompanying midstorey and subcanopy mixed-species
         Myrmotherula
schisticolor, Red-faced Spinetail, Olivaceous Woodcreeper Siasomus griseicapillus,
Ochraceous Wren, Speckled Tanager Tangara guata, Bay-headed Tanager T. gyrola,
Silver-throated Tanager T. icterocephala, Black-and-yellow Tanager, Yellow-throated
Chlorospingus Chlorospingus avigularis and Tawny-capped Euphonia Euphonia anneae.

            
Brewster in western San Blas et al. 1984). Our record

in November 2008 between 1,250 m and 1,400 m at río Tanelita, on the east slope of Cerro
Tacarcuna et al. 2014).
BAY-HEADED TANAGER Tangara gyrola

de Majé, Cerro Sapo, Cerro Pirre, Cerro Malí, Cerro Tacarcuna (Wetmore et al. 1984) and
Serranía de Jungurudó (Angehr et al. 2004). This record extends the species’ range to the
Colombian slope of the Cerro Tacarcuna. Adjacent records from Colombia are from Serranía
de Abibe, the West Andes and northern Central Andes (Hilty & Brown 1986, Cuervo et al.
2008b, Olaciregui et al. 2016).
EMERALD TANAGER Tangara orida

(Robbins et al. 1985), Cerro Tacarcuna (Wetmore et al. 1984) and Serranía de Jungurudó
(Angehr et al. 2004); and Serranía de Abibe, south to El Carmen de Atrato in the West Andes
of Colombia (Sullivan et al. 2009, Olaciregui et alT. f.
auriceps, whereas the nominate occurs on the Caribbean slope from Costa Rica to Panama.

is unknown, although it has been considered to represent an intermediate population
et al. 1984).
TACARCUNA CHLOROSPINGUS Chlorospingus tacarcunae

            
accompanied by Yellow-throated Chlorospingus C. avigularis, which was more abundant.
Both Chlorospingus  
Cryptopipo holochlora, Scale-
crested Pygmy Tyrant Lophotriccus pileatus, Rufous-browed Tyrannulet, Ochraceous Wren,
Slate-throated Redstart Myioborus miniatus, Silver-throated Tanager, Black-and-yellow
           
begging for food from an adult on 6 August 2010, which is consistent with previous
observations of nest construction and immatures that suggest the species’ breeding season
probably begins in late February and extends to the middle of the year (Wetmore et al. 1984,


cfet al.
2011) of a species whose presence in the country had been suggested based on Panamanian
 (Wetmore et al. 1984,

Luis Miguel Renjifo et al. 58 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club
YELLOW-THROATED CHLOROSPINGUS Chlorospingus avigularis
 

  C. f.
hypophaeus
eyes and breast of C. f. marginatus
C. f. hypophaeus, was collected by JMR-O
in November 2008 at 1,400 m on the río Tanelita, east slope of Cerro Tacarcuna. These

time. Its nearest locality is Cerro Brewster in western San Blas  
In Colombia the species occurs north to the upper río Sinú (Hilty & Brown 1986) and was

CHESNUT-CAPPED BRUSH FINCH Arremon brunneinucha
Fairly common in the understorey above 1,000 m. Previously recorded on most massifs

(1,400 m) (Wetmore et al. 1984), Serranía de Majé and Serranía de Jungurudó (Angehr &
Christian 2000, Angehr et al. 2004). Adjacent Colombian records are from the West Andes
(Hilty & Brown 1986), the northern Central Andes (Cuervo et al. 2008b) and Serranía de San
Lucas (Salaman et al. 2002).
SOOTY-FACED FINCH Arremon crassirostris
et al. 2011, Remsen et al. 2016).
Although we did not record the species, we report here two specimens collected in the
foothills of Cerro Tacarcuna, at Alto Barrigonal, in June 1980. Both specimens, a male and




JMR-O & T. Walschburger at río Tanelita (1,408 m), on the east slope of Cerro Tacarcuna.
These records remove any doubts concerning the species’ presence in Colombia and South
America (Remsen et al. 2016).
HEPATIC TANAGER Piranga ava
             
         (Angehr & Christian 2000)
and Cerro Pirre (Robbins et al
of the West Andes, in Serranía de Abibe south (Hilty & Brown 1986, Olaciregui et al. 2016).
Discussion

      
        
       

         
species reported in the region (Fig. 8A). In addition, four species previously known from
         
(Scaly-throated Foliage-gleaner and Ochraceous Wren) inhabit cloud forests of the West
Luis Miguel Renjifo et al. 59 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club
Andes and the Chiriquí highlands, respectively, whereas the remaining species (Black-

ranges such as San Blas and Serranía del Baudó. These range extensions, plus another also
presented here, are indicative of the poorly known avifauna of the region.
           
        
           
     Phaethornis guy coruscans, Catharus f. fuscater, Basileuterus
tristriatus tacarcunae) and foothills (Schiornis veraepacis acrolophites, Chlorospingus avigularis
hypophaeus          
taxonomic, geographic and temporal representation of several poorly known species in bird
collections (Cuervo et al
Tapaculo song, and documented the vocalisations on the Colombian side of endemic taxa
          
Chlorospingus.
             
Although this inventory is probably far from complete, the combination of visual and
         
assemble a representative inventory swiftly. These complementary methodologies have
Figure 7. Lateral views of specimens of Sooty-faced Finch A. crassirostris
slope of Cerro Tacarcuna, compared to a specimen of Olive Finch A. castaneiceps (C) from the West Andes of

   
(S. Sierra)
Luis Miguel Renjifo et al. 60 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club
 
forests            

Tacarcuna range. For example, Robbins et al. (1985) recorded 186 species on Cerro Pirre at

m).
          
             

          
eight species are currently considered nationally threatened in Colombia (Renjifo et al.
          
above 600 m, based on the midpoints of each species’ range (Fig. 8A). Exceptions are the
rare Spiny-faced Antshrike Xenornis setifrons and Yellow-green Tyrannulet Phylloscartes
avovirens. For highland species, 15 of 16 reach their lower elevational limit at or above 600

             

only below 600 m.



   
     
             
              

subspecies, 21 of them mainly found above 600 m (Fig. 8B). However, endemism could be

 
Figure 8. Elevational and geographic distribution of 29 restricted-range species (A) and 24 restricted-range

their elevational midpoint (black dots). Note that most taxon ranges are concentrated above 600 m as shown
by the vertical dashed line. Taxon names are followed by one or more superscripts indicating geographic
distribution. 1  2Central: Serranía de Pirré,
    
et al. (1984), Robbins et al. (1985), Hilty & Brown (1986),
et al. (2004).
Luis Miguel Renjifo et al. 61 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club
could potentially represent distinctive species (e.g. Basileuterus tristriatus tacarcunae;

et al.
2014).

           
plantations), which have accelerated in recent decades in Panama and Colombia (Rangel-C.
2004, Angehr et al         

change because of the small ranges and isolation of their endemic populations. Indeed,
most of those species whose ‘climate envelope’ is projected to disappear in Colombia due to

          
is projected to disappear by 2050 et al 
will encourage further biological exploration of the region, and more importantly, draw
            


the preservation of one of the most interesting biogeographical regions in the Neotropics

Acknowledgements

              
Córdoba-Córdoba and C. A. Medina facilitated access to, and provided photographs of, specimens at the


     

References:
Guía sonora de los Andes colombianos


range in eastern Panama. Bull. Brit. Orn. Cl
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isolated mountain range in eastern Panama. Bull. Brit. Orn. Cl
   Biodiver. Conserv.

                
Orn. Neotrop

(accessed 27 April 2014).

Río Atrato (Turbo, Antioquia). Orn. Colombiana
Burt, J. 2006. Syrinx version 2.6h. www.syrinxpc.com.
      
species is Arremon torquatusBiol. J. Linn. Soc

llamado a fortalecer las colecciones ornitológicas. Orn. Colombiana
        
records from the northern sector of the Western Andes of Colombia. Bull. Brit. Orn. Cl
      
Central of the Colombian Andes, with implications for the biogeography of northwestern South
America. Condor
Luis Miguel Renjifo et al. 62 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club
. A., Palacio, A., Botero,
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           
Biota Colombiana
Chapman, F. M. 1917. The distribution of bird-life in Colombia. Bull. Amer. Mus. Nat. Hist
Orn. Neotrop. 12:

        Myrmeciza immaculata, with a new
subspecies from the Central Andes of Colombia. Bull. Brit. Orn. Cl

Lucas, an isolated mountain range in northern Colombia. Bull. Brit. Orn. Cl
Basileuterus
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                
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
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          
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             Scytalopus tapaculos
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     
in Rangel-C., J. O. (ed.) Colombia diversidad biótica
                 in
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


  Libro rojo
de aves de Colombia. Instituto de Investigación de Recursos Biológicos Alexander von Humboldt &
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  A guide to the birds of Panama with Costa Rica, Nicaragua and Honduras.
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in
Neotropical ornithology.
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Aves del Parque Nacional Natural Los Katíos
 Bangsia arcaei y Chrysothlypis chrysomelas
             Orn.
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Colombia ’98 expedition to Serranía de los Churumbelos: preliminary report.
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              
Ecosphere
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Orn. Columbianae
Wetmore, A. 1965. The birds of the Republic of Panama
Wetmore, A. 1968. The birds of the Republic of Panama
Wetmore, A. 1972. The birds of the Republic of Panama
Proc. Biol. Soc. Wash
Wetmore, A., Pasquier, R. F. & Olson, S. L. 1984. The birds of the Republic of Panama, pt. 4. Smithsonian
.
Addresses             
       
          
         
   
   


Appendix 1
                

individuals captured in parenthesis. *Colombian near-endemic (Chaparro et al  
Remsen et al. (2016).
Family / English name Scientic name Foothills Study site
TINAMIDAE
 Crypturellus soui A
CRACIDAE
 Penelope purpurascens
 Crax rubra
ODONTOPHORIDAE
 Rhynchortyx cinctus S
Luis Miguel Renjifo et al. 64 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club
Family / English name Scientic name Foothills Study site
ACCIPITRIDAE
Barred Hawk Morphnarchus princeps 
Black Hawk-Eagle Spizaetus tyrannus A A
White Hawk Pseudastur cf. albicollis
COLUMBIDAE
Short-billed Pigeon Patagioenas nigrirostris A
 Geotrygon goldmani*AT
 Geotrygon violacea 
CUCULIDAE
Squirrel Cuckoo Piaya cayana A
STRIGIDAE
Bare-shanked Screech Owl Megascops clarkii 
 Ciccaba virgata AT
TROCHILIDAE
White-tipped Sicklebill Eutoxeres aquila C (1)
Stripe-throated Hermit Phaethornis striigularis
 Phaethornis guy 
Tooth-billed Hummingbird Androdon aequatorialis*
 Haplophaedia aureliae*CS (2)
 Heliodoxa jacula C (1)
 Chalybura urochrysia C (2)
Crowned Woodnymph Thalurania colombica
Rufous-tailed Hummingbird Amazilia acatl
 Goldmania violiceps CS (2)
TROGONIDAE
Slaty-tailed Trogon Trogon massena AT
Black-throated Trogon Trogon rufus A
MOMOTIDAE
Broad-billed Motmot Electron platyrhynchum 
Rufous-capped Motmot Baryphthengus rucapillus A
CAPITONIDAE
Red-headed Barbet Eubucco bourcieri
RAMPHASTIDAE
Black-mandibled Toucan Ramphastos ambiguus 
Emerald Toucanet Aulacorhynchus prasinus 
PICIDAE
Olivaceus Piculet Picumnus olivaceus A
Smoky-brown Woodpecker Picoides fumigatus 
PSITTACIDAE
 Pyrilia pyrilia* A
Mealy Parrot Amazona farinosa AT
Red-and-green Macaw Ara chloropterus A
THAMNOPHILIDAE
Black-crowned Antshrike Thamnophilus atrinucha A
Plain Antvireo Dysithamnus mentalis 
Spot-crowned Antvireo Dysithamnus puncticeps
Slaty Antwren Myrmotherula schisticolor 
Chestnut-backed Antbird Poliocrania exsul 
Zeledon’s Antbird Haeria zeledoni 
Bicoloured Antbird Gymnopithys bicolor 
Luis Miguel Renjifo et al. 65 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club
Family / English name Scientic name Foothills Study site
Ocellated Antbird Phaenostictus mcleannani 
RHINOCRYPTIDAE
Tacarcuna Tapaculo Scytalopus panamensis*
FORMICARIIDAE
 Formicarius analis A
 Formicarius nigricapillus 
FURNARIIDAE
Olivaceus Woodcreeper Siasomus griseicapillus
Long-tailed Woodcreeper Deconychura longicauda
Plain-brown Woodcreeper Dendrocincla fuliginosa 
Wedge-billed Woodcreeper Glyphorhynchus spirurus 
 Xiphorhynchus erythropygius 
Brown-billed Scythebill Campylorhamphus pusillus 
Streak-headed Woodcreeper Lepidocolaptes souleyetii A
 Xenops minutus
Scaly-throated Foliage-gleaner Anabacerthia variegaticeps
Lineated Foliage-gleaner Syndactyla subalaris 
Beautiful Treerunner Margarornis bellulus*
Red-faced Spinetail Cranioleuca erythrops
TYRANNIDAE
Rufous-browed Tyrannulet Phylloscartes superciliaris
Olive-striped Flycatcher Mionectes olivaceus 
Scale-crested Pygmy Tyrant Lophotriccus pileatus 
White-throated Spadebill Platyrinchus mystaceus 
Smoke-coloured Pewee Contopus fumigatus A
Tufted Flycatcher Mitrephanes phaeocercus 
 Myiarchus tuberculifer 
PIPRIDAE
White-bibbed Manakin Corapipo leucorrhoa*
 Cryptopipo holochlora 
TITYRIDAE
 Schiornis stenorhyncha*
TROGLODYTIDAE
Scaly-breasted Wren Microcerculus marginatus A A
Ochraceous Wren Troglodytes ochraceus 
White-headed Wren Campylorhynchus albobrunneus*
Sooty-headed Wren Pheugopedius spadix*
White-breasted Wood Wren Henicorhina leucosticta A
 Henicorhina leucophrys 
Song Wren Cyphorhinus cf. phaeocephalus A
TURDIDAE
 Myadestes coloratus 
Slaty-backed Nightingale-Thrush Catharus fuscater 
THRAUPIDAE
Blue-and-gold Tanager Bangsia arcaei
Speckled Tanager Tangara guata
 Tangara fucosa
Bay-headed Tanager Tangara gyrola
Emerald Tanager Tangara orida
Silver-throated Tanager Tangara icterocephala
Luis Miguel Renjifo et al. 66 Bull. B.O.C. 2017 137(1)
© 2017 The Authors; Journal compilation © 2017 British Ornithologists’ Club
Family / English name Scientic name Foothills Study site
Black-and-yellow Tanager Chrysothlypis chrysomelas
Bananaquit Coereba aveola C (1)
EMBERIZIDAE
Tacarcuna Chlorospingus Chlorospingus tacarcunae 
Yellow-throated Chlorospingus Chlorospingus avigularis 
Chestnut-capped Brush Finch Arremon brunneinucha 
CARDINALIDAE
Hepatic Tanager Piranga ava
PARULIDAE
 Myiothlypis fulvicauda 
Three-striped Warbler Basileuterus tristriatus 
Slate-throated Redstart Myioborus miniatus 
FRINGILLIDAE
Tawny-capped Euphonia Euphonia anneae 
Appendix 2
Sound-recordings used in vocal comparisons of Scytalopus tapaculos (see text). Sound collections: BSA =

Library. Songs: Tacarcuna Tapaculo S. panamensis     
S. chocoensis  S. vicinior  
Calls: S. panamensis   S. chocoensis        
S. vicinior
... Such landscape heterogeneity has been identified as a driver for bird speciation in the Neotropics (Smith et al., 2014), and is thus important to highlight in distribution maps. Our new maps also identify other hotspots of richness in relatively unexplored areas such as the Serranía de San Lucas, the Darién region (Renjifo et al., 2017) and Santa Marta Woodstar (Chaetocercus astreans). These species are geographically restricted but include widespread elevation and habitat breadth, and current methods exclude suitable lower elevation habitats from the artificial circular shape distribution presented by Lumbierres, Dahal, Soria, et al. (2022) (Lumbierres, Dahal, Soria, et al., 2022). ...
... Despite having the highest richness of bird species in the country, the Amazon shows the largest extent of uncertainty in the country when multiple species are analysed. The findings align with results from prior studies that examined information gaps for different taxa in Colombia (Suárez-Castro et al., 2021), and they can thus guide concerted national efforts to survey these regions (Renjifo et al., 2017). For instance, recent expeditions in the uncertainty hotspot of the lower interfluvial zone of the Putumayo-Caquetá rivers (Figure 5b), provided new distributional information for poorly known species in northwestern Amazonia (Carantón-Ayala, 2023;Socolar et al., 2022). ...
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... For these species, behavioral data is in need. A few studies have focused on Z. goldmani, including inventories at Tacarcuna, Pirre, Altos de Quia, Nique, Jurado, and other highlands in Darien and at the Colombian border, which contain nearly untouched mountain forest ecosystems (Renjifo et al., 2017). Studies with the genus Zentrygon and Geotrygon have examined behaviour (vocalization, courtship) (Tubaro & Mahaler, 1998;Johnson and Weckstein 2011;Donegan and Salaman, 2012;Renjifo et al., 2017;Baptista et al., 2020), genetic and evolutionary adaptations (Brooks, 2012;Lapiedra et al., 2013). ...
... A few studies have focused on Z. goldmani, including inventories at Tacarcuna, Pirre, Altos de Quia, Nique, Jurado, and other highlands in Darien and at the Colombian border, which contain nearly untouched mountain forest ecosystems (Renjifo et al., 2017). Studies with the genus Zentrygon and Geotrygon have examined behaviour (vocalization, courtship) (Tubaro & Mahaler, 1998;Johnson and Weckstein 2011;Donegan and Salaman, 2012;Renjifo et al., 2017;Baptista et al., 2020), genetic and evolutionary adaptations (Brooks, 2012;Lapiedra et al., 2013). The aims for this study were to build preliminary list of wild-dove species and obtain behavioural information that can help as baseline for monitoring and conserve the wild-doves from Darien. ...
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... Analysis of new bird records is important for evaluating extinction risks, predicting the future changes in ecological communities, and undertaking actions to preserve species of conservation concern (Chen et al., 2011;Li et al., 2013;Liu et al., 2013;Renjifo et al., 2017). A recent meta-analysis has shown that the number of new bird records in China has increased yearly during the past two decades (Chen et al., 2021). ...
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Birds are sensitive to environmental changes and can drive range shifts rapidly due to their high mobility. Though previous studies have examined the associations between species traits and range shifts, whether species traits could still explain heterogeneity in shift directions remains poorly explored. Here, we compiled new bird records of China from 2000 to 2019 and analyzed species traits associated with apparent shift directions. We collected 350 provincial-level new records of birds belonging to 67 families of 22 orders. Of these, 32 are threatened, with 3 critically endangered, 11 endangered, and 18 vulnerable. Provinces in western China (i.e., Yunnan and Xizang) had relatively higher species richness of new recorded birds; this pattern was also reflected in the phylogenetic diversity we observed. In addition, provinces in northern China (i.e., Tianjin, Shandong, and Beijing) had relatively higher richness-controlled phylogenetic diversity. Phylogenetic overdispersion of new recorded bird communities was observed in 61.29% of provinces (19 of 31). The main shift directions indicated by new bird records were northward (with nearly 50% of birds moving NW, N and NE). Migration, hand-wing index (HWI), body mass, and range size are the four key factors that most significantly influence the shift directions in bird species, suggesting that bird movement toward newly suitable areas varies with species-specific traits. Together, these results demonstrate the importance of considering species ecological traits when predicting shift directions of birds.
... Serranía de Majé, Panama, and along the Serranía del Darién from Cerro Azul in the west, to Cerro Tacarcuna in the east (Wetmore et al. 1984, Cuervo 2013, Renjifo et al. 2017. ...
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Bird inventories of the isolated Serranía de San Lucas, northern Colombia, yielded range extensions for several foothill or highland birds of the adjacent Central Andes: Pavonine Cuckoo Dromococcyx pavoninus, Lyre-tailed Nightjar Uropsalis lyra, White-tipped Sicklebill Eutoxeres aquila, Masked Trogon Trogon personatus, Golden-headed Quetzal Pharomachrus auriceps, Smoky-brown Woodpecker Picoidesfumigatus, Crimson-mantled Woodpecker Colaptes rivolii, Redfaced Spinetail Cranioleuca erythrops, Western Woodhaunter Hyloclistes virgatus, Buff-fronted Foliage-gleaner Philydor rufum, Spotted Woodcreeper Xiphorhynchus erythropygius, Slaty Antwren Myrmotherula schisticolor, White-crowned Tapaculo Scytalopus atratus, Rufous-browed Tyrannulet Phylloscartes superciliaris, Ornate Flycatcher Myiotriccus ornatus, White-winged Becard Pachyramphus polychopterus, Brown-capped Vireo Vireo leucophrys, Orange-billed Nightingale-Thrush Catharus aurantiirostris, Black-billed Thrush Turdus ignobilis, Golden Tanager Tangara arthus and Yellow-throated Bush Tanager Chlorospingus flavigularis. M. schisticolor and S. atratus show small differences from Central Andes populations and require further research. Range extensions for lowland species (some based on older specimens) include Ornate Hawk-Eagle Spizaetus ornatus, Scaled Dove Scardafella squammata, Vermiculated Screech Owl Megascops 'guatemalae' centralis, Central American Pygmy Owl Glaucidiutn griseiceps, Spot-tailed Nightjar Hydropsalis maculicaudus, Green-and-rufous Kingfisher Chloroceryle inda, Sooty Grassquit Tiaris fulighwsus and Black-faced Grassquit T. bicolor. The threatened Chestnut-bellied Hummingbird Amazilia castaneiventris and Recurve-billed Bushbird Clytoctantes alixii (both last collected in the 1940s in the region) were recorded. Tens of new elevational records for typically lowland species were made. San Lucas woodnymphs are intermediate between Thalurania fannyi and T. colombica, raising questions as to whether two species are involved. Nominate Lineated Foliage-gleaner Syndactyla subalaris (of the West Andes) occurs in the northern Central Andes including San Lucas. Various highland species recorded at similar elevations in the Central Andes are apparently absent, perhaps due to lower humidity, island effects and ecological release of lowland species. The highest elevations of San Lucas remain unknown but occupy very small areas. Further work will doubtless yield additional new records, but is complicated by security issues and land-use conflicts. Anti-personnel landmines do not provide a viable 'defence' against deforestation in the long term due to habitat disturbance associated with their clearance.