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Diet composition of birds associated to an urban forest patch in Northern Venezuela/Composicion de la dieta de las aves asociadas a un parche de bosque urbano en el Norte de Venezuela/Composicao da dieta das aves associada a uma trecho de floresta urbana no Norte da Venezuela.


Diet studies are necessary for the understanding of the biological and ecological strategies of birds. In Venezuela, few studies about the diet of birds have been carried out, restricted to dry forests and arid zones (Poulin et al., 1994, Quilarque et al., 2010, Perez et al., 2001). In the neotropics, however, there are abundant studies, mostly based on stomach contents using emetics (Loiselle and Blake, 1990; Perez et al., 2001; Duraes and Marini, 2005; Gaiotti and Pinho, 2013) and on stomach contents of collected birds (Remsen et al., 1993; Rosenberg, 1993; Lopes et al., 2005). The use of emetics has the disadvantage that they may cause accidental death of the studied specimens, while fecal analysis is a non-invasive method that does not endanger birds. Several studies have demonstrated the convenience of feces' studies to determine feeding guilds in birds (Rouges and Blake, 2001; Piratelli and Pereira, 2002; Carvallo Lima, 2008; Navas, 2009). The disadvantage of this technique is that certain food items may be misrepresented due to differences in digestibility (Voiter et al., 2003). Carlisle and Holberton (2006) found that by increasing the number of fecal samples between five and 17, the results obtained are similar to the number of taxa detected in regurgitated samples.

Studies on urban birds are becoming more relevant due to the growth of urban areas and the concomitant increase in anthropic effects on bird biology. In the neotropics, several studies have focused on urban bird communities (Stiles, 1990; Rivera-Gutierrez, 2006; Villegas and Garitano-Zavala, 2010; Carbo-Ramirez and Zuria, 2011; Reis et al., 2012) and many showed that cities retain a considerable diversity of bird species (Clergeau et al., 1998; Marzluff et al., 2001). However, few explore how urban habitats can affect the diet of birds. The majority of studies are restricted to a single species, mainly Falconiformes and owls (Yalden, 1980; Tatner, 1983; Gavett and Wakeley, 1986; Roth and Lima, 2003; Delgado and Calderon, 2007; Hernandez-Munoz and Mancina, 2011). Several studies have reported that certain bird species tend to consume anthropogenic food (Ottoni et al., 2009) such as bread and food scraps (Rollinson et al., 2003).

In Venezuela, there is scarce information about urban birds (Caula et al. 2003; 2010; Seijas et al., 2011; Sanz and Caula, 2014), and none about their diets. Caracas is a city with over five million people and has a great diversity of birds: about 300 species have been recorded that find refuge in many parks and other green areas of city (Aveledo, 1968; Goodwin 1987; Fundacion La Salle, 1997; Caula and Levin, 1999; Levin et al., 2000; FJBC-SCAV, 2006; SCAV, 2007; Sainz-Borgo and Levin, 2012; Sainz-Borgo, 2012, 2014, 2015).

The aim of this work was to study the diet composition of a bird community associated with an urban patch of semi-deciduous forest, using quantitative analysis of their fecal contents.


Study area

The study involved two field visits per month, from September 2013 to July 2014, to a 2ha patch of semi-deciduous, seasonal, premontane forest located at the Experimental Station Arboretum (Instituto de Biologia Experimental, Universidad Central de Venezuela, Caracas, Venezuela, 1100m, 10[degrees]30'36"N - 66[degrees]53' 92"W), located within a residential zone. The study covered the end of 2013 wet season (September-October), the following dry season (December 2013-April 2014) and the beginning of the 2014 wet season (May-July), with precipitation ranging from 500 to 1000mm per year and an temperature between 19 and 27[degrees]C. The vegetation is secondary, consisting of two layers: an upper stratum where trees 8-18m high predominate, while the bottom contains grasses and shrubs, mixed with abundant trees. Tree species are predominantly deciduous, while in the shrub layer there are abundant vines, thorny shrubs and evergreen species (Hokche and Ramirez, 2006). More than 300 plant species of 77 families have been reported, the most abundant families being Asteraceae, Fabaceae, Poaceae, Euphorbiacea, Mimosaceae, Malvaceae and Rubiacea (Lopez and Ramirez, 2004), and xerophilous species of the Cactaceae and Agavaceae families (Ewell et al., 1976). The study area is surrounded by residential buildings and vehicular transit took place during collection.

Data collection

Eight mist nets (12 x 2.8, 36mm mesh) were used along the main path of the study area, placed twice a month from September 2013 to July 2014. The mist nets were opened from 6:30 to 15:30 and checked every 15min. All captured birds were classified according to the South America Classification Committee (SACC) (Remsen et al, 2016).

For feces collection, individual birds were placed during 20min in a plastic container (20 x 30cm) with a mesh at the bottom and topped with black fabric to reduce stress. After that period, the bird was released and feces were collected in a 8ml glass tube containing 4ml of 70% ethanol. The items in the diet were identified to order or family using a stereo-microscope (Wild HeerbruggMSA), in the Ornithology Laboratory, Universidad Simon Bolivar, Caracas, Venezuela,. The plant samples were identified using the Lau (2010) data base.

The relative occurrence (Ro) of each category was calculated as number of samples in which the category occurred divided by the total number of samples; while the relative abundance (Ra) of each category was calculated as a number of diet items of each category divided by the total numbers of diet items in the sample, averaged for all samples (Martini and Duraes, 2005).

Permanova analyses were conducted with the program Primer v5 (Clarke and Gorley, 2006) using the fecal samplings data by bird family, in order to determine if seasonality had some effect on diet. The food items were used as variables and seasons and bird species were considered factors. To check for differences in diet between dry and wet seasons a Chi-square ([chi square]) test was used.


From a total of 227 captures, 60% provided useful fecal samples, belonging to 33 species, 4 orders and 16 families (Table I). Bird families having the largest numbers of fecal samples were Tyrannidae (45 samples), Thraupidae (18), Thamnophillidae (12), Turdidae (12) and Galbulidae (10). The more abundant species were Atalotriccus pilaris (N=28), Coereba flaveola (22), Galbula ruficauda (16), Formicivora grisea (14), Turdus nudigenis (14) and Camptostoma obsoletum (13). We have identified 32 different food items (Tables I, II, Figure 1). Of the fecal samples, 63.5% contained only insects, 14.5% fruit and seeds, 12.4% fruits and insects, and 9.5% mostly seeds.

Captures in the wet season were 120 (53%) and in dry season 107 (47%). Richness was greater during the rainy season than the dry season (28 vs 21 bird species respectively). The number of fecal samples collected in the two seasons differed: in the wet season 87 feces samples were collected, whereas only 45 samples were obtained in dry season. Food items with the highest values of Ro and Ra were Coleoptera, Formicidae, fruit pulp, Diptera, and seed fragments (Table II). Ants were consumed mainly in the dry season, and Coleoptera, Clusia, and Diptera during wet season (Figure 2). Four species (Columbina talpacoti, G. ruficauda, F. grisea and A. pilaris) consumed a higher diversity of diet items in the wet season than in the dry one, whereas three species (Ramphocaenus melanurus, Thryophilus rufalbus and Saltator coerulescens) consumed more items in the dry season (Figure 3, Table I).

The Permanova analysis indicated that there was an interaction between the factors (species x season, P=0.013) in Tyrannidae, showing that there are differences in the diet of species between seasons. In the rest of birds evaluated, no interactions were detected (Permanova, Thraupidae: P=0.052, Formicariidae: P=0.517). Significant differences in birds abundance were found between seasons for frugivorous-insectivorous (N wet season=34, N dry season=14, [chi square] P=0.0035) and for insectivorous (N wet season=21, N dry season=25, [chi square] P=0.028). The frugivorous and granivorous-insectivorous did not show differences in abundance in dry and wet seasons ([chi square] P=0.135 and P=0.739 respectively).

Feeding guilds

In insectivorous birds, the most abundant food items were Coleoptera (found in 85 samples), Formicidae (35) and Diptera (30) (Tables II). The most common bird species was A. pilaris, which consumed the major variety of food items consisting of eleven different families of insects, as well as Clusia and Loranthaceae seeds (Table I). Strict insectivores were F. grisea, R. melanurus, Thamnophilus doliatus, T. rufalbus, Grallaricula ferrugineipectus, P. squamulatus, Synallaxis albescens, Campyloramphus trochilirostris, Cranioleuca subcristata, and Rhodinocihla rosea.

Samples from C. subcristata contained only insects, except in one sample where seed fragments were found during the wet season. In a similar case, one sample of P. squamulatus contained a Loranthacea seed.

In one sample from F. grisea two individuals of the family Phoridae (Diptera) were found. Several items were registered only once: Araneae, Curculionidae and Blatodea, consumed by A. pilaris; Lampyridae by T. aedon; Diplopodae, Homoptera, and stones by F. grisea; and an ant nymph by T doliatus (Table I).

T. nudigenis, C. flaveola, Cychlaris gujanensis, Euphonia trinitatis and Hylophilus flavipes consumed mostly fruits, and in lesser proportions insects. A. pilaris, C. obsoletum and Pitangus sulphuratus were considered mainly insectivorous with occasional fruits consumed. Strict frugivorous were Thraupis episcopus, T. glaucocolpa, S. coerulescens and E. flavogaster.

Clusia (Clusiaceae) was the most frequently fruit consumed, but seeds of Clydemia (Melastomataceae), Palicourea (Rubiaceae), and Lauraceae were also commonly found in fecal samples. Non-identified small flowers were found in samples of C. flaveola, C. fuscatus and T. glaucocolpa. The samples of E. trinitatis contained abundant Loranthaceae seeds, which were also consumed by E. flavogaster, C. obsoletum, C. fuscatus, H. flavipes, and A. pilaris.

In the feces of C. talpacoti, Tiaris bicolor and Cyanocompsa cyanoides seed fragments and arthopods in smaller proportions were found. In the samples from Leptotila verreauxi only seed fragments were found.


The bird community in the study area was dominated by insectivores, similar to what has been observed in other neotropical forests (Terborgh et al., 1990; Poulin et al., 1994; Duraes and Marini, 2005). During the wet season several species, especially insectivorous, consume a greater variety of food items, a fact that has been reported for different birds assemblages (Leck, 1972; Blake and Loiselle, 1991; van Schaik et al., 1993; Poulin et al., 1994; Williams and Middleton, 2007). Also, a marked seasonality was found in several food items. For example, ant consumption is greater in the dry season, while Coleoptera, Diptera and Clusia seeds are more consumed during the wet season, These results are consistent with previous findings about seasonal changes in arthropod availability on deciduous forests (Wolda, 1978; Wolda and Fisk, 1981), as well as with fluctuations in number of insectivorous birds with changes in food availability (Frith and Frith, 1990), Stouffer and Bierregards (1995) found that in an isolated forest patch the abundance of insectivorous birds tended to decrease, However, the Arboretum Experimental Station is connected to other green areas and parks, constituting an ecological pathway through the city, which allows movements of birds between a continuum of green areas; but probably, the connection between patches is not enough to allow the maintenance of diversity of insectivorous present in pristine areas.




Tyrannidae showed the highest diversity in food items, A. pilaris contained the highest number of food items in the feces (as previously reported by Poulin et al., 1994), followed by C. fuscatus (Gaiotti and Pinho, 2013). For Furnariidae, this study provided the first report of seeds consumption by the specie C. subcristata, There are previous reports of other Furnariidae feeding on seeds and fruits (Poulin et al., 1994) but none for this species, The Phoridae consumption by F. grisea were probably incidental while it was eating ants, because they are parasitoids of the insects (Porter et al., 1995; Morrison and Gilbert, 1999; Elizalde and Folgarait, 2011). The consumption of insects like Formicidae and Coleoptera in addition to seeds by C. talpacoti and T. bicolor has been previously reported in other species of these genera (Poulin et al., 1994; Perez et al., 2001); in the case of ants, they were probably ingested by accident while foraging seeds, or as a protein and fat supplement (Gill, 2006). However, none of these individuals were present in the breeding season, when it has been reported that these species consume proteins, Frugivorous-insectivorous birds were more abundant in the wet season, as has been reported in other seasonal habitats (Karr, 1976; Blake and Loiselle, 1991), An exception was P. sulphuratus, with a single capture in the dry season, Between food items, Clusia and Loranthaceae fruit pulp were the most frequently consumed, Frugivorous birds in general tended to consume arthropods, probably as a nutritional supplement (Levey and Martinez Del Rio, 2001).

Leaves were not found in feces of S. striatipectus and S. coerulescens, in contrast to previous reports (RodriguezFerraro et al., 2007; Chatellenaz, 2008; Garcia-Amado et al., 2011). In this study, S. coerulescens consumed only fruit; whereas S. striatipectus only consumed Coleoptera, In the literature, occasional consumption of insects has been reported in S. orenocensis, but its diet is mostly composed of fruits and leaves (Garcia-Amado et al., 2011). The difference may be related to only one sample obtained of S. striatipectus, from which it is difficult to make a generalization of the diet, In the case of S. coerulescens, where five samples of feces were obtained, despite it being a mainly folivorous species, the abundance of fruit in the diet probably corresponds to an opportunistic consumption at a time when this resource was abundant.

C. flaveola consumed a wide variety of items, consistent with previous reports (Skutch, 1954; Gross, 1958; Snow and Snow, 1971; Cruz, 1974; Feinsingerp et al., 1985; Poulin et al., 1994). The abundance of Loranthaceae seeds in E. trinitatis feces is consistent with numerous reports of Euphonia as the principal seed disperser of this family (Restrepo et al., 2002; Ribeiro et al., 2013). Loranthaceae seeds were also found in several Tyrannidae and in H. flavipes, as had been previously reported (Guerra and Marini, 2002).

Communities in disturbed habitats tend to be less diverse than in pristine areas, with differences in composition, richness, and abundance, among others. Urban birds are affected by different causes than in a wild habitats (Sanz and Caula, 2014). For example, studies of biodiversity in urban habitats show a gradie nt where biodiversity declines with increased urbanization (Clergeau et al., 2006; McKinney, 2006), where species distribute themselves according to resource-matching (Sutherland, 1983; Morris, 1994). These differences in the availability of resources affect the diet of the species; for example, in the case of the feeding guilds in temperate regions there was an increase in omnivorous abundance in urban areas, but not so in the neotropics (Lim and Sodhi 2004), where there is greater availability of resources throughout the year. In this sense, in order to understand the dynamics of urban birds feeding in the neotropical region, this study is pioneer generating information about the diet of birds living in urban habitats in Venezuela. The birds in our study area probably consume different food items that the same species would have in pristine areas, due to a limited availability of resources. Therefore, this kind of research, as well as long-term studies of bird communities in urban habitats, are necessary in different neotropical localities in order to understand the processes of adaptation of birds to urban habitats.

Received: 05/30/2015. Modified: 01/22//2016. Accepted: 01/25/2016.

Cristina Sainz-Borgo. Biologist, Universidad Central de Venezuela. Ph.D. in Biological Sciences, Universidad Simon Bolivar, Venezuela. Professor, Universidad Simon Bolivar, Venezuela. Address: Departamento de Biologia de Organismos, Universidad Simon Bolivar, Apartado 89000, Caracas 1080A, Venezuela email:


The author thanks Luis Gonzalo Morales, Emilio A. Herrera, Adriana Rodriguez Ferraro, Pedro A. Borges L. and two anonymous referees for suggestions on the manuscript; Sandra B. Giner F. for fieldwork help; Luis Levin of the E. S. Arboretum for logistical support, Ruben Candia for help on the identification of Phoridae; and Pablo Lau for providing bibliography.


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                              Diet items

Bird species                  Wet season

Columbina talpacoti (G)       Nid seed fragments,
                                Leaf fragments,
                                Arthropod fragments
Leptotila verreauxi (G)       Nid seed fragments

Amazilia tobaci (NI)          Arthropod fragments,

Picumnus squamulatus (I)      Formicidae, Coleoptera

Galbula ruficauda (I)         Coleoptera, Diptera,
                                Insect eggs,
                                Orthoptera, Apidae

Thamnophilus doliatus (I)     Orthoptera, Formicidae,
                                ant nymph

Formicivora grisea (I)        Orthoptera, Formicidae,
                                Hemiptera, Araneae,

Grallaricula                  Coleoptera
  ferrugineipectus (I)

Cranioleuca subcristata (I)   Coleoptera, Nid
                                seed fragments
Synallaxis albescens (I)      Coleoptera, Hemiptera
Campylorhamphus               Formicidae, Coleoptera
  trochilirostris (I)

Atalotriccus pilaris (I)      Orthoptera, Lepidoptera,
                                Curculionidae, Araneae,
                                Blatodea, Apidae,
                                insect eggs,
                                Formicidae, Clusia
                                seeds, Coleoptera,
Cnemotriccus fuscatus (I)     Coleoptera, Diptera,
                                Hemiptera, Orthoptera,
                                Clusia seeds,
                                Loranthaceae seeds,
                                Formicidae, NI seed
Elaenia flavogaster (F)       Seed fragments,
                                Lorantacea seeds,
                                Palichourea seeds
Pitangus sulphuratus (FI)

Lathrotriccus euleri (I)      Formicidae
Camptostoma obsoletum (FI)    Coleoptera, Formicidae,
                                Loranthaceae seeds,
                                Diptera, fruit pulp,
                                Nid seed fragments

Hylophilus flavipes (F)

Cychlaris gujanensis (F)

Thryophilus rufalbus (I)      Formicidae, Coleoptera

Troglodytes aedon (I)         Orthoptera, Formicidae,
                                Hemiptera, Apidae,
                                Diptera, Elateroidea

Ramphocaenus melanurus (I)    Orthoptera, Coleoptera,

Thraupis episcopus (F)

Thraupis glaucocolpa (F)      Coleoptera, flowers
Tiaris bicolor (GI)           Nid seed fragments,
Coereba flaveola (FI)         Coleoptera, Diptera,
                                Flowers, fruit pulp,
                                Clusia seeds,
                                Formicidae, Isoptera,

Incertae sedis
Rhodinocichla rosea (I)       Coleoptera
Saltator coerulescens (F)     Clusia seeds, fruit pulp

Saltator striatipectus (I)    Coleoptera

Cyanocompsa cyanoides (G)     Coleoptera, Nid seed

Parkesia                      Formicidae, Coleoptera
  noveboracensis (I)

Euphonia trinitatis (F)

Turdus nudigenis (FI)         Formicidae, fruit pulp,
                                Nid seed fragments,
                                Clusia seeds

                                                     No. fecal samples

Bird species                  Dry season              Wet      Dry
                                                     season   season

Columbina talpacoti (G)       Nid seed fragments      1(4)     1(2)

Leptotila verreauxi (G)                               1(1)     0(0)

Amazilia tobaci (NI)                                  1(5)     0(2)

Picumnus squamulatus (I)      Formicidae,             1(1)     2(7)

Galbula ruficauda (I)         Coleoptera             9(14)     1(2)

Thamnophilus doliatus (I)     Orthoptera,             1(4)     2(2)
Formicivora grisea (I)        Orthoptera,            4(11)     5(5)
                              Isoptera, Phoridae,
                                Diptera, Insect
                                egg, Hymenoptera,

Grallaricula                                          1(1)     0(0)
  ferrugineipectus (I)

Cranioleuca subcristata (I)                           2(7)     0(1)

Synallaxis albescens (I)                              1(1)     0(0)
Campylorhamphus                                       3(3)     0(3)
  trochilirostris (I)

Atalotriccus pilaris (I)      Coleoptera,            12(12)   8(16)

Cnemotriccus fuscatus (I)     Formicidae,             7(7)     1(1)

Elaenia flavogaster (F)                               4(7)     1(7)

Pitangus sulphuratus (FI)     Formicidae,             0(0)     1(1)
                                fruit pulp
Lathrotriccus euleri (I)                              1(1)     0(0)
Camptostoma obsoletum (FI)    Formicidae,            9(10)     1(1)
                                insect eggs

Hylophilus flavipes (F)       Loranthaceae seeds,     0(0)     1(1)
                                fruit pulp
Cychlaris gujanensis (F)      Clusia seeds,           0(0)     1(1)
                                fruit pulp

Thryophilus rufalbus (I)      Formicidae,             2(3)     1(2)
Troglodytes aedon (I)         Orthoptera,             2(9)     1(5)
                                Hemiptera, insect
                                eggs, Diptera

Ramphocaenus melanurus (I)    Orthoptera,             1(1)     5(5)
                                insect eggs,
                                Diptera, fruit

Thraupis episcopus (F)        Clydemia seed,          0(0)     2(2)
                                Clusia seeds
Thraupis glaucocolpa (F)                              1(1)     0(0)
Tiaris bicolor (GI)           Nid seed fragments      3(5)     3(3)

Coereba flaveola (FI)         Fruit pulp,             5(8)    4(13)

Incertae sedis
Rhodinocichla rosea (I)                               1(1)     0(0)
Saltator coerulescens (F)     Fruit pulp, Clusia      1(2)     4(5)
Saltator striatipectus (I)                            1(2)     0(4)

Cyanocompsa cyanoides (G)                             2(2)     0(0)

Parkesia                                              1(1)     0(0)
  noveboracensis (I)

Euphonia trinitatis (F)       Loranthaceae seeds      0(0)     1(1)

Turdus nudigenis (FI)         Coleoptera, Clusia      9(9)     3(6)
                                seeds, fruit pulp,
                                Nid seed

* Feeding guilds were assigned based on the items found
in the feces. G: granivorous, NI: nectarivorous-insectivorous,
I: insectivorous, FI: frugivorous-insectivorous. In
parenthesis in the columns of fecal samples are the number
of total captures (N). Nid: non identified.


Food item            No. total   No. feces   No. bird    Ro      Ra
                      of item     samples    species

Formicidae              226         35          15      0,26    0,2
Coleoptera              185         83          25      0,61    0,16
Clusia seeds            137         16          9       0,12    0,12
NI Insects              121         30          26      0,88    0,03
Diptera                 108         30          9       0,22    0,09
Insect eggs             74          13          5       0,09    0,06
Loranthaceae seeds      53           8          7       0,06    0,05
Fruit pulp              37          31          9       0,23    0,03
Orthoptera              24          19          9       0,13    0,02
Lepidoptera             16          10          5       0,07    0,01
Apidae                  13           6          4       0,04    0,01
Hymenoptera             11           8          5       0,08    0,01
Hemiptera                8           7          4       0,05    0,01
Vespidae                 6           6          3       0,04    0,01
Leaf fragments           6           6          6       0,01    0,01
Clydemia seeds           6           1          1       0,01    0,01
Palicourea seeds         6           2          2       0,01     0
Isoptera                 4           4          4       0,01     0
Araneae                  3           3          3       0,02     0
Stone                    3           1          1       0,02     0
Flowers                  3           3          3       0,02     0
Lauraceae                3           1          1       0,01     0
Phoridae (Diptera)       2           1          1       0,01     0
Curculionidae            1           1          1       0,03     0
Blatodea                 1           1          1       0,007    0
Ant nymph                1           1          1       0,007    0
Lampyridae               1           1          1       0,007    0
Pollen                   1           1          1       0,007    0
Diplopod                 1           1          1       0,007    0
Homoptera                1           1          1       0,007    0

NI: non identified, Ro: relative occurrence, and
Ra: relative abundance of feeding items found in
the feces of birds from the studied forest patch.
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Title Annotation:texto en ingles
Author:Sainz-Borgo, Cristina
Date:Feb 1, 2016
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