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ON 24(2) 183-200 NEW.Pdf ORNITOLOGIA NEOTROPICAL 24: 183–200, 2013 © The Neotropical Ornithological Society LEK CHARACTERISTICS AND THE STATIC MALE SONG OF THE GREEN VIOLET-EAR (COLIBRI THALASSINUS) DURING A 3- YEAR STUDY IN A TEMPERATE FOREST OF CENTRAL MEXICO Vanessa Martínez-García1, Carlos Lara2, & Juan Francisco Ornelas3 1Doctorado en Ciencias Naturales, Universidad Autónoma de Tlaxcala, México. 2Centro de Investigación en Ciencias Biológicas, Universidad Autónoma de Tlaxcala, Tlaxcala, México. E-mail: [email protected] 3Departamento de Biología Evolutiva, Instituto de Ecología, A.C., Xalapa, Veracruz, Mexico. Resumen. – Características de los leks y el canto estático de los machos del Colibrí oreji violeta- verde (Colibri thalassinus) durante un estudio de 3 años en un bosque templado del centro de México. – Aun cuando los colibríes son el segundo grupo de aves más diverso, el conocimiento sobre la conducta de formación de leks es limitado. Las características del establecimiento de 11 leks del Colibrí oreji violeta-verde (Colibri thalassinus) fueron analizadas en un bosque templado del centro de México, desde el año 2009 al 2011. En cada lek contamos el número de machos que lo conformaba, el tipo de percha utilizada para los despliegues vocales, y medimos la distancia entre las perchas adyacentes. Asimismo, verificamos la permanencia espacial y temporal de cada lek. Los cantos de despliegue de los machos fueron registrados para obtener el repertorio vocal. Nuestros resultados mostraron que los machos de C. thalassinus forman leks en ambientes espacialmente variables y con requerimientos específicos de las características del hábitat. Los leks fueron establecidos en zonas de vegetación secundaria, la cual generalmente estuvo bordeada por bosque de pino y campos de cultivo. Los machos frecuentemente usan pinos para establecer sus perchas de canto y las distancias entre estos territorios vocales individuales (8ñ15 m) define a estos leks como “explosivos”. La información registrada durante los tres años del estudio, indican que algunos de estos leks fueron establecidos en sitios previamente ocupados, incluyendo el uso de las mismas perchas de canto entre años. Finalmente, presentamos la primer descripción del repertorio vocal desplegado en los leks, incluyendo el número de notas emitidas. Son necesarios más estudios enfocados en la descripción de los requerimientos ambientales para el establecimiento de los leks. Abstract. – In spite of the fact that hummingbirds are the second most diverse group of birds, knowledge of lekking behavior in this family is limited. The characteristics of 11 established leks of Green Violet-ear (Colibri thalassinus) were analyzed in a temperate forest of central Mexico from 2009–2011. In each lek, we counted the number of males attending the lek area, type of perch used for the vocal displays, and measured the distance between adjacent perches. In addition, we verified the spatial and temporal per- manence of each lek, and the male display songs were recorded to obtain the vocal repertoire. Our results show that males of C. thalassinus form leks in variable spatial environments with specific require- ments regarding habitat characteristics. The leks were established in secondary vegetation zones, which generally bordered pine forest and cornfields. The males frequently used pine trees to establish their singing perches and the distance between these individual singing territories (8–15 m apart) define these leks as “explosive.” Recorded information during the 3-year study period indicate that some leks were established in previously occupied sites including the use of the same singing perches in consecutive years. Finally, we provide the first description of the vocal repertoire of static vocal displays at the leks including the number of notes emitted. Further studies are needed focusing on the description of the environmental requirements for lek establishment. Accepted 13 August 2013. 183 MARTÍNEZ-GARCÍA ET AL. Key words: Colibri thalassinus, Trochilidae, habitat, lek behavior, vegetation composition, vocal reper- toire, Mexico. INTRODUCTION Violet-ears are highly flexible to the environ- mental conditions they use to form their A lek is a gathering of males occupying and aggregations and to perch during singing dis- defending the same small territories on a mat- plays. Nevertheless, specific data about the ing display arena (Payne 1984, Höglund & characteristics of the vegetation and possible Alatalo 1995, MacDougall-Shackleton & Har- variation in location and time the leks sites are bison 1998, Almeida & Macedo 2001). These established require further exploration. Fur- display territories do not function to defend thermore, although there is information on resources required by females but are the song of this species (Gaunt et al. 1994, defended by males for display to females Barrantes et al. 2008), the vocal repertoire dis- (Pizo & Silva 2001). Birds are one of the most played by males forming leks has not been studied groups in the formation of lek arenas. described. For instance, aggregations of male display ter- The objective of our study is to describe ritories have been reported in over 130 spe- the characteristics of leks formed by Green cies from six families (Cotingidae, Paradi- Violet-ears in a temperate forest in central saeidae, Pipridae, Ptilonorhynchidae, Pycno- Mexico based on measurements and observa- notidae, and Tyrannidae). In spite of the fact tions over a 3-year study. In addition, our that hummingbirds (Trochilidae) are the study presents the first description of the rep- second most diverse group of birds in the ertoire of vocal displays for this species dur- New World, knowledge of lekking behavior in ing lek formation. this family is limited (Bleiweiss 1997, 1998; Hayes 2002). In reviewing the existing litera- METHODS ture, we found reports of 49 different species of hummingbirds that suggest the formation Study site. The fieldwork was carried out of leks (Table 1). Four of these reports during three reproductive seasons of provide a brief description of leks, five Green Violet-ears, from June to October include a spatial and temporal description of 2009–2011 in the “La Malinche” National the mating system, and the rest only mention Park (MNP), Tlaxcala, Mexico (Fig. 1). The that leks are formed during the mating season MNP forms part of the Trans-Mexican without providing more details. Also, the Volcanic Belt. It has an area of 46,093 ha, vocal repertoire displayed by males in these being located at 19°14’N, 98°14’W, with an arenas is rarely described. Clearly further altitude ranging from 2300 to 4461 m a.s.l. studies are necessary to generate more basic (López-Domínguez & Acosta Pérez 2005). information about leks in hummingbirds. The vegetation is mainly a mosaic of pine Since the first description of its natural forest (dominated by Pinus montezuma, P. history by Wagner (1945) in Mexico, the pseudostrobus, P. leiophylla, and P. hartwegii) Green Violet-ear (Colibri thalassinus) has been and second growth vegetation (Villers et al. considered a lekking species. Later studies 2006). performed in Costa Rica confirmed these observations (Feinsinger 1977, Gaunt et al. Study species. Colibri thalassinus (Swainson 1827) 1994, Barrantes et al. 2008). The information is a medium-sized, sexually monochromatic gathered from these studies reveal that Green species. Both sexes are iridescent green in 184 LEKS AND SONG OF GREEN VIOLET-EAR HUMMINGBIRDS TABLE 1. Hummingbird species with evidence in forming leks. Some species have shown a description of leks, and details of the spatial and temporal variation of thereof, but most only documents the formation of leks without offering details. References referring to indices: 1Wagner 1945, 2Skutch 1958, 3Wiley 1971, 4Barash 1972, 5Snow 1974, 6Feinsinger 1977, 7Stiles & Wolf 1979, 8Payne 1984, 9Armstrong 1987, 10Tamm et al. 1989, 11Atwood et al. 1991, 12Bleiweiss 1997, 13Dastagir et al. 1997, 14Johnsgard 1997, 15Bleiweiss 1998, 16MacDougall & Harbison 1998, 17Pizo & Silva 2001, 18Ornelas et al. 2002, 19Boughman & Moss 2003, 20González & Ornelas 2009, 21del Hoyo et al. 1999, 21Ramjohn et al. 2003, 22Pizo 2012, 23Hayes 2002. Species Description Reference Band-tailed Barbthroat Lek formation (3–5 individuals) during the breeding season. There 7, 11, 12, Threnetes ruckeri is no description of the system. 15 Pale-tailed Barbthroat Lek formation during the breeding season. There is no description 7, 12 Threnetes leucurus (niger) of the system. Green Hermit Lek formation during peak activity (12–15 individuals). In the lek, 6, 7, 11, 12, Phaethornis g. guy the members are separated by less than 10 m without visual con- 14, 16, 20 tact. Males consistently sang from one perch within these vocal territories. Green Hermit Lek formation during the breeding season. There is no description 12 Phaethornis guy coruscans of the system. Long-tailed Hermit Formation of “explosive” leks (20–25 individuals), with an area of 7, 11, 12, Phaethornis superciliosus approximately 1 km2. Distance between individuals ranging from 13, 14, 17, 20 to 500 m. If environmental conditions are favorable the leks are 20 formed throughout the year. Little Hermit Leks are established in areas with tall trees of 27–30 m. The 3, 5, 7, 11, Phaethornis longuemareus height of the song perches was 22 to 45 cm, and these were 12, 14, 20 located in the deep shade near the forest floor. In the lek, the members are separated by less than 10 meters and without visual contact. Straight-billed Hermit Lek formation during the breeding season. There is no description 12 Phaethornis bourcieri of the system. Reddish Hermit Lek formation during the breeding season. There is no description 7, 8, 12, 14 Phaethornis ruber of the system. Tawny-bellied Hermit Lek formation during the breeding season. There is no description 12 Phaethornis syrmatophorus of the system. Grey-chinned Hermit Lek formation during the breeding season. There is no description 12 Phaethornis griseogularis of the system. White-whiskered Hermit Lek formation during the breeding season. There is no description 7, 12 Phaethornis yaruqui of the system. White-bearded Hermit Lek formation during the breeding season.
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