Studies in Avian Biology No. 37:53–68
HABITAT USE OF WINTERING BIRD COMMUNITIES IN SONORA, MEXICO: THE IMPORTANCE OF RIPARIAN HABITATS
JOSÉ FERNANDO VILLASEÑOR-GÓMEZ
Abstract. Riparian systems are dynamic and diverse despite their limited areal extent. They are espe- cially important for breeding bird communities in southwestern US and are highly used as migra- tory corridors; however, their importance for wintering birds has not been assessed systematically. Information from 1,816 standard 10-min point counts was gathered at 85 locations in the State of Sonora, Mexico from sea level to 2,175 m during January and February 2004–2006. I detected 253 bird species across 14 vegetation types, including nine categories of riparian vegetation. Eighty percent of total species were detected in riparian habitats, and 72% were detected in non-riparian habitats. The mean number of species and individuals detected per count were signifi cantly higher in ripar- ian habitats than in non-riparian habitats for migratory species, but not for residents. A hierarchical classifi cation analysis showed that riparian bird communities are different from those in non-riparian communities, and they contribute 22% of the species that comprise the regional avifauna, which is more than any other habitat type.
Key Words: habitat use, Mexico, neotropical migrant birds, riparian, Sonora, wintering.
USO DE HABITAT DE COMUNIDADES DE AVES INVERNANTES EN SONORA, MEXICO: IMPORTANCIA DE LOS HABITATS RIPARIOS Resumen. Los sistemas riparios son dinámicos y diversos a pesar de su limitada cobertura espacial. Son especialmente importantes para las comunidades de aves que se reproducen en el Suroeste de los Estados Unidos y son usados extensamente como corredores migratorios; sin embargo, su importancia para las aves invernantes no se ha determinado de forma sistemática. Información de 1,816 conteos estándar de 10-minutos se obtuvo en 85 localidades en el Estado de Sonora, México, desde el nivel del mar hasta 2,175m durante Enero-Febrero de 2004–2006. Detecté 243 especies en 14 tipos de vegetación, incluyendo nueve categorías de vegetación riparia. Ochenta por ciento de las especies fueron registradas en hábitats riparios y 72% en hábitats no-riparios. El número promedio de especies e individuos detectados por conteo fue signifi cativamente mayor en hábitats riparios que en no-riparios para las especies migratorias, pero no para las residentes. Un análisis de clasifi cación jerárquica mostró que las comunidades de aves riparias son diferentes a las comunidades de sitios no-riparios, y contribuyen con 22% de las especies que conforman la avifauna regional, la mayor contribución entre los hábitats estudiados.
Riparian habitats are dynamic and generally light of the very small percentage (generally more biologically diverse than surrounding <1%) of land area composed of riparian habitat uplands, especially in arid regions (Hunt 1985). (Mosconi and Hutto 1982, Knopf 1985, Schmitt This faunal diversity is due to the presence of 1976, Hubbard 1971). water, high productivity, and abundance of The importance of riparian systems during habitat edge, which is maximized by the linear periods of migratory passage and as wintering shape of riparian habitats (Gregory et al. 1991). habitats has begun to receive more attention, In the arid Southwest, riparian systems support and we know these periods may be equally at least 80% of all wildlife species (Hunt 1985, or more important than the breeding season USDI Bureau of Land Management 1998). They in terms of population regulation (Fretwell support the highest densities of small mammals 1972, Sherry and Holmes 1995, Hutto 1998). (Andersen 1994) and richest communities of During the energetically demanding migra- butterfl ies (Fleishman et al. 1999, Nelson and tion period, landbirds have to make important Anderson 1999). Riparian habitats support choices about which stopover habitats will some of the highest densities and most species- provide enough food, cover, and water to rich avian communities in the US (Knopf et al. enable a rapid and safe replenishment of fuels 1988) and western Mexico (Hutto 1995), and (Moore and Simons 1992). Mortality rates may infl uence the ecological dynamics of adjacent be considerable (especially for young birds) as upland habitats (Strong and Bock 1990, Farley migrants compete for resources while avoid- et al. 1994a, Skagen et al. 1998, Powell and Steidl ing potential predators in unfamiliar locations. 2000). In addition, many riparian bird species We currently know little about the specifi c are riparian obligates, which is signifi cant in habitat needs of migrants. Studies have shown,
53 54 STUDIES IN AVIAN BIOLOGY NO. 37 however, that landbirds use riparian habitats of wintering birds across the complete array disproportionately more frequently during of extant vegetation types in the state, and migration. This is especially the case during determined the value of each vegetation type spring migration, when the productivity of in terms of its contribution to regional avifaunal riparian habitats is higher than that of the sur- diversity. rounding uplands (Skagen et al. 1998, Finch and Yong 2000, Skagen et al. 2005, Kelly and Hutto METHODS 2005). Understanding migration ecology is now considered key to the conservation of migrant STUDY AREA landbird populations (Moore et al. 1995, Hutto 2000, Heglund and Skagen 2005). Sonora is the second largest state in Mexico, Information on the importance of ripar- covering 179,156 km2. It is located at the north- ian systems compared to other habitat types western corner of mainland Mexico between for wintering birds is also scant and is based 26°18’ and 32°29’N, and 108°25’ and 115°03’W. largely on occasional species- and site-specifi c The geographical features of the state make it records (Russell and Lamm 1978, Terrill 1981, rich and biologically diverse. Sonora is located Rosenberg et al. 1991, USDI Bureau of Land at the latitude where the tropics meet the south- Management 1996), Christmas Bird Counts, ern limit of the temperate region, and includes and studies that happen to include the complete elements from both regions. It is composed of a annual cycle of riparian birds (Anderson and complex mix of landscapes with elevations from Ohmart 1977, Wells et al. 1979, Strong and Bock sea level to 2,630 m. The lowland plains are veg- 1990, Farley et al. 1994a). In the only published etated primarily by desert and xeric shrubs. studies focused on wintering bird distribution At middle elevations the northernmost exten- among a variety of vegetation types in western sions of tropical deciduous forest are found in Mexico, the abundance of Neotropical migrants the south and southeastern part of the state. in riparian habitats and gallery forests are In the highlands a diversity of oak woodlands among the highest recorded (Hutto 1980, 1995), and mixed coniferous forest along the eastern thus suggesting that riparian corridors are section border the state of Chihuahua. Riparian important for wintering birds. During winter, communities composed mainly of associations riparian habitats may be especially important in of cottonwood (Populus spp.) and willows the Sonoran Desert, which lies at the northern (Salix spp.) are present discontinuously along edge of the wintering range of many western the river courses, and mangroves (Avicennia, North American migratory bird species, and Conocarpus, Laguncularia, and Rhizophora) are which represents the primary wintering area distributed in isolated patches along the coast for short-distant migrant species from the cen- of the Sea of Cortés (INEGI 2000). Because of tral and western US. In lowland Sonora, the these characteristics and its location, Sonora only other habitats for wintering birds are drier, supports a diverse community of breeding, hotter, and structurally less diverse. migrating, and wintering landbirds of western Riparian zones are known to be habitats North America (Kelly and Hutto 2005) of critical conservation concern worldwide, Rivers and subsurface water continue to including the southwestern US and northwest- play an important role in the economy of the ern Mexico. These habitats, which have always state. Most rivers originate in the Sierra Madre constituted a small component of arid land- Occidental and run to the coastal plains and into scapes, have been reduced drastically and are the Sea of Cortés. The most important perma- now fragmented and modifi ed by desiccation, nent fl ows are from north to south and east to dam construction, water diversion, invasion of west, and include the Colorado, Sonoyta, Altar, exotic species, overgrazing, and other factors. Magdalena, San Miguel, Sonora, Moctezuma, In this study I sought to determine how Bavispe, Mátape, Sahuaripa, Yaqui, Cedros, signifi cant riparian areas are for wintering bird and Mayo Rivers (Bojórquez-Tapia et al. 1985). communities in the state of Sonora, Mexico. Several large dams create important impound- Specifi cally, I addressed two main questions: ments to supply water for the irrigation of (1) are wintering bird communities associated extensive agricultural fi elds in the lowlands. with riparian habitats signifi cantly different Sonora is also one of Mexico’s main produc- from those associated with upland habitat types ers of high quality beef cattle; fi fteen million in the State of Sonora, and; (2) are the riparian hectares, including pasturelands, woodlands, habitats important in terms of their contribu- shrublands, and prairies with buffelgrass tion to regional diversity? In order to answer (Pennisetum ciliare) are used for raising and these questions, I examined the community breeding beef cattle, having potential negative composition and species abundance patterns impacts, especially on the composition and HABITAT USE BY WINTERING BIRDS IN SONORA—Villaseñor-Gómez 55 structure of riparian habitats due to overgraz- Russell and Monson 1998), and personal experi- ing (Saab et al. 1995). ence (Appendix 1). The total number of species and individu- SURVEY PROTOCOLS als detected, the mean number of species and individuals per count, and the percentage of During January and February of 2004, 2005, resident, partial migrants, and migrant species and 2006, avian surveys were conducted at were computed for each vegetation type and 85 locations in 14 vegetation types following riparian association. ANOVA was performed to INEGI (2000) and nine riparian associations compare the mean number of species and indi- throughout Sonora, ranging from sea level to viduals detected in point counts (25-m radius) elevations over 2,000 m (Table 1). Non-repeated by residency status, as well as the number of and randomly selected standard 10-min point individuals detected for each species to deter- counts with unlimited radius were conducted mine the ones showing signifi cant differences between 0700 and 1100 H. Most aquatic spe- between non-riparian and riparian habitats. cies and birds fl ying over were recorded but A chi-square test was performed to examine not used in any of the analysis. Raptors, swal- differences in the mean percentage of species lows, and other aerial species were recorded recorded belonging to the residency status only if they were perched on the vegetation or groups between riparian and non-riparian habi- the ground within the point count area (Hutto tats. I also used a contingency analysis of species et al. 1986, Ralph et al. 1993). Unidentifi ed spe- richness to look for differences in the expected cies, such as hummingbirds and Empidonax proportions of species with preferences for non- species, were grouped and included on the list riparian and riparian habitats, as well as those as unknown hummingbird and Empidonax sp., with no preferences for either habitat. respectively. The location and distance from the To determine if wintering bird communi- observer to each detected bird were mapped; ties associated with riparian habitats were sig- the information gathered was entered into nifi cantly different from those associated with Excel and was managed for the analysis with other habitat types, I used two classifi cation SPSS 11.5.1 software package (SPSS 2002). Only methods. A hierarchical classifi cation method the detections within a radius of 25 m from the (cluster analysis; Manly 2004) was used to observer were used in the analyses. Although compare similarities among bird communities much information was unused by restricting the in each habitat type, producing a dendrogram data in this way, and rarer and shy or cryptic (based on presence-absence data and using species may have been missed as a result, it was complete linkage and the Ochiai measure). done to decrease the potential error caused by Secondly, a two-way indicator species analy- the inclusion of individuals detected in an adja- sis (TWINSPAN; McCune and Mefford 1997) cent but different habitat type and to decrease was used, based on the concept that samples bias due to inherent differences in detectability which constitute a group will have a corre- of birds among habitats. sponding set of species that characterize that group (indicator species). TWINSPAN fi nds ANALYSES the relationships between species and samples through correspondence analysis ordination. It Species were assigned to one of three resi- initially classifi es the samples into two groups dency status categories—residents, migrants, and then refi nes the classifi cation through and partial migrants. Residents are those spe- detrended correspondence analysis (DCA), cies that remain in the same area year-round. fi nding the indicator and associated species for Migrants are those species that move far from the resulting groups, and based on those spe- their breeding areas and occupy a completely cies, regroups iteratively within the groups into different geographical region to the south dur- smaller clusters until a limit is met. An indicator ing the winter, with no overlapping popula- species is the species (or the group of species) tions (all long-distance migrant species). Partial present in all of the clustered vegetation types; migrants are those species that move seasonally an associated species is present primarily in a but not for distances of such magnitude. As a given group although it could also be present consequence, in the southern portions of some in other vegetation types or associations. With species’ distributions, populations of resident, TWINSPAN I defi ned the species that charac- transient, and wintering individuals could terize the general vegetation groups identifi ed overlap during migration and winter. The spe- by the dendrogram produced in the clustering cies were assigned to one of these residency technique. In order to perform these analyses, status categories based on published informa- I used the software PC-ORD for Windows, ver- tion (van Rossem 1945, Howell and Webb 1995, sion 3.17 (McCune and Mefford 1997). 56 STUDIES IN AVIAN BIOLOGY NO. 37 2006). AND (2004, 2005, EBRUARY F ) 111 28°00’50” 111°02’57” 21 AND
Ficus ANUARY J and DURING , EXICO Washingtonia , M ONORA S IN
SAMPLED
TYPES
VEGETATION
AND
OCATIONS 1. L ABLE T cottonwoods, cottonwood-willow 588 29°48’14” 110°13’37” 72 76 110°13’37” 30 109°10’03” 29°48’14” 69 112°13’25” 28°51’45” 588 110°10’29” 30°47’35” 22 516 324 29°43’27” cottonwood-willow 110°07’37” willow-mesquite 30°56’34” scrubland 554 Location 1,372 Aconchi—Río Sonora cottonwoods, Aconchi, Río Sonora, Agua Caliente Springs sycamores Aribabi—Sierra Alta Arroyo La Poza Bámori—Río Sahuaripa Baviácora—Río Sonora cottonwood-willow Cerro La Pintada—Tetabejo Caborca Sarcocrassicaulescent Cajón de Onapa, Riparian cottonwood-mesquite, Río Sahuaripa Camino a San Lázaro-San Antonio Camino a San Marcial Riparian mesquite Cañón de Evans, road Vegetation sampled Highland oaklands Cananea—Bacoachi Riparian Riparian cottonwood-willow, willow, Cañón de Nacapule, San Carlos, Nuevo Guaymas Riparian mesquite Riparian cottonwoods, Sarcocaulescent scrubland Coteco Cucurpe—Río San Miguel Thornscrub Desv. Playa San Nicolás Elevation (m) Ejido Ganadero Puente El Tigre, Guaymas El Carrizo, Rancho San Darío 606 Latitude N El Chiculi, Hornos, Río Yaqui, Sonora 1,490 El Cochito—km. 179, road Riparian mesquite, willow 188 Sarcocaulescent scrubland Grasslands Agua Prieta-Moctezuma Longitude W El Llano—Moctezuma Willow-mesquite Oasis ( 29°50’26” N Counts El Resbalón—Sahuaripa 30°03’25” 188 El Sahuaral, San José de Guaymas 28°34’41” Riparian willow-mesquite-chino Estero del Soldado, San Carlos, Nuevo Guaymas Riparian cottonwoods Estero Paraíso—Punta Checa 110°16’33” Mangroves 1,189 Estero Santa Cruz—Kino 109°06’10” 28°52’04” Grasslands, subtropical scrub Coastal sarcocaulescent scrubland Estero Santa Rosa—Punta Chueca 82 111°00’43” Granados—Río Bavispe 57 Huásabas—Río Bavispe 1 31°03’00” 30 Jécori—Río Moctezuma 110°57’43” Coastal sarcocaulescent scrubland 5 20 28°06’22” La Aduana Microphyllous scrubland Coastal sarcocaulescent scrubland 730 587 La Majada, between Moctezuma-Mazocahui 27°46’26” 849 326 Thornscrub La Mesa del Campanero Tropical deciduous forest 110°38’32” 15 28°49’43” Mangroves Mangroves 5 110°59’27” 22 Lowland oaklands 30°02’59” 28°44’09” 109°53’52” 30°19’20” 28°34’53” Mangroves 1,164 5 111°48’19” 27°59’15” 22 26°34’35” 0 111°12’19” 109°07’59” Riparian cottonwood-willow 20 654 110°42’35” 110°15’18” Riparian cottonwood-willow 31°10’52” Riparian cottonwoods 37 110°50’30” 109°18’11” 27°57’23” 61 3 29°07’57” 14 20 1,011 Highland coniferous forest 109°33’53” 540 638 22 554 28 110°58’53” Tropical deciduous forest 109°16’51” 0 0 29°42’57” 30 29°51’46” 29°43’25” 716 29°55’13” 2,139 17 1 33 29°02’21” 28°58’37” 109°50’45” 109°18’01” 109°39’04” 109°17’31” 29°58’01” 28°22’12” 28°47’49” 602 112°09’59” 112°09’48” 31 20 9 10 109°45’39” 109°01’50” 111°52’34” 27°01’53” 16 6 67 19 16 109°00’34” 15 HABITAT USE BY WINTERING BIRDS IN SONORA—Villaseñor-Gómez 57 . ONTINUED 1. C ABLE T Riparian sycamores 1,358 30°33’09” 109°39’21” 35 109°39’21” 30°33’09” 28 110°57’37” 1,358 30°36’18” 19 sycamores 784 scrubland 110°38’33” Location Los Alisos Riparian Los Torreones Magdalena, road Magdalena-Imuris Sarcocrassicaulescent Mazatán, ca. 10 km W Mesa del Toro-Ejido Ignacio Zaragoza, Cananea Sierra Pinacate (between Elegante and Tecolote) 31°09’03” Sierra Pinacate Crater Vegetation of sandy deserts Cerro Colorado Puerto Peñasco Rancho Monte Alto-road to Puerto Libertad Rancho El 44, between Cobachi and Road to Yécora Subtropical scrub Rancho El Perú Rancho La Cuesta, Coastal sarcocaulescent scrubland 219 km 94 road to Yécora Vegetation sampled Highland oaklands, Grasslands Rancho La Noria, N Subtropical scrub Subtropical scrub Rancho Los Cuervos, N El Carrizo) 131 Rancho Piedras Negras 31°51’21” (Carretera Minera Nyco) Rancho San Darío ca. Carrizo Rancho San Fermín, ca. Cobachi 29°18’38” Elevation (m) Reserva de la Biosfera Pinacate 113°21’28” 431 Río Altar, Tubutama Vegetation of sandy deserts Latitude N Río Cuchujaqui Subtropical scrub 1,284 Vegetation of sandy deserts Río Cocóspera—Rancho 111°51’26” 10 Aribabi 28°47’22” 517 857 Longitude W Río Cuchujaqui at El Mentidero Subtropical scrub N Counts 1,556 Subtropical scrub Río Cuchujaqui at La Isleta Río Matape—San José 31 Sarcocaulescent scrubland Subtropical scrub Subtropical scrub 185 110°21’28” de Pimas 29°00’00” Sarcocaulescent scrubland 29°17’23” Vegetation of sandy deserts Río Mayo—Presa Mocúzari 16 31°03’38” Río Mayo—Tetapeche 31°55’08” 110°13’45” Río San Miguel, 715 109°54’12” 1 Rancho San Esteban Riparian baldcypress-willow 31°21’36” 352 110°03’53” Riparian cottonwoods Río Santa Cruz, San Lázaro Riparian cottonwood-willow, 167 352 cottonwood-willow 113°18’05” 391 30°06’17” 709 Río Yaqui, Ónavas Riparian baldcypress-willow 1 5 Riparian baldcypress-willow 113°27’41” Río Yaqui, Sitio Nochebuena 29°03’01” 30 549 724 31°48’16” 29°16’16” 227 Riparian mesquite, willow, Riparian willow-mesquite-chino, 10 28°35’44” 111°12’37” 30°20’38” 20 110°37’56” 28°55’14” 30°04’18” 224 Cottonwood-mesquite, cottonwoods 634 113°17’45” 111°05’18” 239 26°54’44” Riparian willow-mesquite-chino 110°11’00” 111°04’39” 2 Riparian cottonwood-mesquite, 974 10 110°07’55” 111°14’27” willow-mesquite 26°54’38” 30°53’09” 30 Riparian mesquite 18 26°56’29” 108°54’46” Riparian willow-mesquite 10 1 49 30°51’15” 1 108°54’51” 1 111°27’39” Riparian mesquite 108°53’05” 9 tropical deciduous forest 27°12’23” 110°39’51” 8 203 21 9 109°21’35” 350 40 285 28°47’45” 98 19 155 28°43’02” 29°17’13” 109°38’41” 27°12’57” 110°52’02” 110°20’50” 28°27’45” 1 109°06’47” 30 35 109°32’01” 20 1 58 STUDIES IN AVIAN BIOLOGY NO. 37 . ONTINUED 1. C ABLE T Tónichi Riparian willow-mesquite 186 28°35’42” 109°34’05” 31 19 88 15 109°34’05” 111°21’11” 109°39’04” 111°27’21” 28°35’42” 31°09’25” 186 29°40’41” 30°53’17” Location mesquite, Río Yaqui, San Antonio de la Huerta Río Yaqui, Tónichi Río Yaqui, S El Novillo Río Yaqui, Soyopa 786 San Carlos—ca. willow-mesquite Cañón de Nacapule San Ignacio—Terrenate Riparian willow-mesquite San Javier San José de Pimas San Lázaro, S San Miguel, road to Punta Chueca Santa Cruz 565 187 Sáric—Río Altar 684 Riparian willow-mesquite Riparian willow-mesquite cottonwood-willow Sierra La Elenita, Cananea Vegetation sampled 28°37’44” Riparian willow-mesquite, mesquite Sierra Ladrilleros-El Pinacate Riparian cottonwoods Sarcocaulescent scrubland Sierra Mazatán Sierra Mazatán, NE Presa Teópari 188 Terapa—Río Moctezuma 109°36’27” 214 cottonwood-willow, 166 Thornscrub cottonwoods Elevation (m) Tónichi Riparian 28°45’02” Tubutama - Río Altar Tropical deciduous forest 28°49’43” 1 47 Lowland oaklands Latitude N cottonwood-willow 28°35’37” 784 Unámichi—Río Sonora Sarcocaulescent scrubland Riparian mesquite Sarcocaulescent scrubland Highland coniferous forest 109°37’32” Yécora Riparian cottonwood-mesquite, Longitude W Yecora—La Palmita 109°37’46” 27°58’58” Grasslands, highland oaklands N Counts 109°34’08” 30°42’51” 813 4 Riparian cottonwood-mesquite, 1,920 3 111°02’24” 1,417 177 Lowland oaklands 154 2 110°55’27” 1,279 384 28°34’58” cottonwoods, mesquite, willow, 663 31°00’30” 31 Riparian cottonwoods, 31°19’56” 28°57’51” 31°44’28” 19 31°05’49” Riparian cottonwood-willow 28°43’54” 109°44’51” 110°23’06” 29°13’15” 110°36’30” 112°03’29” 113°18’23” Highland oaklands 110°39’00” 1,357 110°21’49” 32 1,048 30 110°04’44” 62 30 6 Highland coniferous forest 6 29°05’38” 21 30°39’39” 22 110°12’54” 1,462 109°58’58” 1,538 9 28°22’19” 20 28°22’54” 109°03’59” 108°54’39” 21 11 HABITAT USE BY WINTERING BIRDS IN SONORA—Villaseñor-Gómez 59
RESULTS The mean numbers of species and individuals (all groups combined) detected per count in Survey data were gathered on 1,816 stan- riparian habitats were signifi cantly higher than dard 10-min point counts (944 in non-riparian in non-riparian habitats (Table 3). The same and 872 in riparian sites, respectively), at 85 pattern held for mean number of species and locations from sea level to 2,175 m (Table 1). A individuals per count within the migrant and total of 32,570 individuals of 253 bird species partial migrant groups, which also had sig- was recorded across all vegetation types. Eighty nifi cantly higher numbers in riparian habitats. percent of total species (203) were recorded in In contrast, I found no signifi cant differences riparian associations, and 72% (183) in all other in mean number of species or individuals per non-riparian vegetation types. count between riparian and non-riparian habi- The number of bird species in riparian veg- tats for resident species. etation was greater than in any other vegetation I found no statistically signifi cant differ- type. A tally of species richness by vegetation ences in the proportions of residents, migrants, types shows that species richness in all non- and partial migrants in each count between riparian vegetation types was less than 65 spe- riparian and non-riparian vegetation types cies per habitat type, with all but four having (χ2 = 4.105, df = 2, P = 0.128; Fig. 1a). However, less than 50 species per category. In contrast, the general pattern suggests that the propor- riparian habitats for the most part were richer, tion of total resident species was higher in with all but two riparian vegetation types hav- non-riparian habitats, while the proportions ing 75 or more species per type, and all but of total partial migrants and migrant species three having more than 95 species per category were higher in riparian vegetation types. When (Table 2). partial migrants and migrants were combined By limiting data analyses to observations into a single migrant group and compared with within a 25-m radius, a total of 8,237 individuals residents, the difference became signifi cant (χ2 = of 168 species was detected (82 residents, 33 par- 4.083, df = 1, P = 0.03; Fig. 1b), with a greater tial migrants, and 53 migrants [see Appendix 1 proportion of resident species in non-riparian for the residency status and number of individ- habitats and a greater percentage of migrant uals recorded for the most common species]). species in riparian habitats. The contribution
TABLE 2. VEGETATION TYPES, NUMBER OF POINTS SAMPLED, AND NUMBER OF SPECIES RECORDED IN SONORA, MEXICO, DURING JANUARY AND FEBRUARY (2004, 2005, AND 2006).
Vegetation Type Counts/species Elevation range (m) Non-riparian vegetation Mangroves 54/53 Sea level (0) Coastal sarcocaulescent scrubland 138/58 (0–234) Vegetation of sandy deserts 70/17 (3–231) Microphyllous scrubland 30/17 (1,159–1,217) Tropical deciduous forest 79/61 (102–983) Sarcocrassicaulescent scrubland 58/20 (307–824) Thornscrub 45/37 (184–1,189) Subtropical scrub 25/43 (391–878) Sarcocaulescent scrubland 152/47 (13–551) Grasslands 83/14 (316–1,592) Low oaklands 54/41 (970–1,250) High oaklands 75/61 (1,400–2,010) Highland coniferous forest 60/37 (1,525–2,175) Oases 21/37 (50–233) Non-riparian vegetation (general) 944/183 (Sea level–2,175) Riparian vegetation Willow–mesquite-chino 57/96 (36–102) Mesquite desert riparian 89/108 (117–823) Willow-mesquite 91/96 (166–823) Willow 104/96 (348–840) Willow-baldcypress (Salix-Taxodium) 31/75 (222–277) Cottonwood-willow 230/125 (527–1,282) Cottonwood-mesquite 23/45 (555–1,305 Cottonwood 214/125 (505-1,288) Sycamores (Platanus) 33/40 (1,322–1,402) Riparian sites (general) 872/203 (33–1,402) 60 STUDIES IN AVIAN BIOLOGY NO. 37
TABLE 3. ANOVA RESULTS COMPARING THE MEAN NUMBER OF SPECIES AND INDIVIDUALS DETECTED IN POINT COUNTS (25-M RADIUS) BY RESIDENCY STATUS.
General Non-riparian Riparian mean habitats habitats Residency status (N = 1,816) (N = 944) (N = 872) F P Species All species 2.55 1.93 3.24 164.40 <0.01 Residents 1.32 1.36 1.29 1.15 0.28 Partial migrants 0.59 0.32 0.88 234.99 <0.01 Migrants 0.64 0.25 1.06 424.78 <0.01 Individuals All species 4.29 3.34 5.33 58.43 <0.01 Residents 2.06 2.15 1.94 1.92 0.17 Partial migrants 1.12 0.65 1.63 46.14 <0.01 Migrants 1.11 0.53 1.74 106.41 <0.01
FIGURE 1. Mean percentage of bird species by residency status in riparian and non-riparian habitat groups in Sonora (mean percentage of species recorded for resident, partial migrant, and migrant species per habitat type); (a) considering the three residency status groups; (b) considering partial migrants and migrants together as a single group. of migrant and partial migrant species and Considering the community as a whole, the individuals to the avifauna in riparian sites is contingency analysis (chi-square test) shows particularly important, as is the contribution of that the residents are overrepresented in the resident species and individual to non-riparian non-riparian habitats and the frequency of the habitats (Fig. 2 a, b). Numerically speaking, migrants is signifi cantly higher in the riparian migrants are an important element of the win- environments (χ2 = 13.72, df = 4, P = 0.008; Table tering avifauna of riparian environments in the 4). If the analysis is limited to those species state of Sonora. with at least 20 detections (66 species) to avoid However, such generalizations could be mis- the infl uence of those species with low sample leading because species respond ecologically sizes, the differences are even more evident and behaviorally in different ways, and they (χ2 = 18.35, df = 4, P = 0.001; Table 4). have to be assessed individually. Of the 168 spe- The dendrogram resulting from the cluster cies noted above, 59 (35.1%) showed signifi cant analysis (Fig. 3) separates the habitats into differences in their abundances between the three main groups. The fi rst, near the bottom riparian and non-riparian habitats; 18 (10.7%) of the dendrogram, represents highland habi- were primarily associated with non-riparian tats, the second clusters desertscrub habitats habitats, and 41 (24.4%) with riparian habitats. and mangroves, and the last, at the top of the HABITAT USE BY WINTERING BIRDS IN SONORA—Villaseñor-Gómez 61
FIGURE 2. Proportional contribution of residents, partial migrants, and migrants to (a) species richness, and (b) total individuals for each riparian and non-riparian habitat (habitat types: NR = non-riparian: 1 = mangroves, 2 = microphyllous scrubland, 3 = oasis, 4= coastal sarcocaulescent scrubland, 5 = subtropical scrub, 6 = tropical deciduous forest, 7 = thornscrub, 8 = sarcocrassicaulescent scrubland, 9 = sarcocaulescent scrubland, 10 = veg- etation of sandy deserts, 11 = grasslands, 12 = low elevation oaklands, 13 = high elevation oaklands, 14 = high- land coniferous forest; R= riparian: 1= willow-mesquite-chino, 2 = willow-mesquite, 3 = mesquite, 4 = willow, 5 = baldcypress-willow, 6 = cottonwood, 7 = cottonwood-willow, 8 = cottonwood-mesquite, 9 = sycamores.) 62 STUDIES IN AVIAN BIOLOGY NO. 37
dendrogram, groups riparian associations (except for the sycamores [Platanus sp.] which are grouped within the highland habitats) with tropical deciduous forest and oases. The micro- phyllous scrub habitat stands by itself and inde- pendent of the other groups as a result of its poor avifauna (only six species recorded). The contribution of these grouped habitat types to regional diversity was assessed by determining the species that were found exclu- sively in a particular habitat or group of habi- tats and would not occur if that habitat were not present in the region. Based on these criteria, riparian habitats contributed the most species unique to the regional avifauna, followed in importance by highlands, desertscrub habitats, tropical deciduous forest, oases, and mangroves
. (Fig. 3). The two-way indicator species analysis (TWINSPAN) results identifi ed those indicator HABITATS
and associated species that characterize each one of the vegetative clusters produced by the hierarchical analysis. It defi ned the existence of RIPARIAN
six groups of bird species: (1) highland vegeta- cantly Non-riparian Riparian Total
fi tion species, (2) microphyllous scrubland, (3) AND desert scrubby vegetations and grasslands, (4) mangroves, (5) lower elevation riparian asso- ciations, and (6) tropical deciduous forest and RIPARIAN - oases (Table 5). NON DISCUSSION
BETWEEN Riparian environments have been identifi ed as a key habitat component of ecological sys- tems through maintenance of dynamic ecologi- cal processes along a gradient of landscapes, which links wildlife, vegetation, and soils DIFFERENCES : within terrestrial systems. They are very pro- ductive systems and represent the most valu- able habitat for wildlife in general, especially in RICHNESS the xeric regions of the world. The results of this study support this concept.
SPECIES Cottonwood riparian woodlands and their
OF associations have been identifi ed as the most
important habitat for birds in the southwest- ern US (Hubbard 1971, Stamp 1978, Strong
ANALYSIS and Bock 1990; Farley et al. 1994a, b; Skagen et al. 1998), and in every case they support the highest number of species and/or densi- cantly Non-riparian Riparian Total signi N
fi ties among the studied habitats. Based on the above-cited papers that included a list of CONTINGENCY the species recorded in riparian habitats, the OF percentage of those riparian species in the US that were also found wintering in Sonoran
ESULTS cottonwood riparian associations in this study ranged from 63–86%. The differences are due 4. R primarily to resident species with distributional : Expected frequencies shown in parenthesis.
ABLE ranges restricted to the United States or to spe- N signi N All species (N = 168) Species with at least 20 detections (N = 66) (53.2) (8.8) (20.0) (15.0) (7.5) (5.5) (15.0) (7.5) (6.1) (5.9) (3.0) (20.0) 15 2 12 (12.9) (8.8) 38 Status 53 1 82 9 2 19 53 15 14 Residents (5.7) different species (53.2) species migrants Partial 24 18 11 Migrants 32 (34.4) Total (7.1) 168 (3.5) (8.1) (21.4) 18 41 109 1 different species species 66 (2.6) (5.3) 26 13 27 8 33 7 0 6 13 T Note cies whose distribution does not include the HABITAT USE BY WINTERING BIRDS IN SONORA—Villaseñor-Gómez 63
FIGURE 3. Grouping of the vegetation types resulting from a cluster analysis based on the wintering avifauna in Sonora, Mexico, based on presence-absence data. (The number of species and percentages represent the number of species unique to that habitat or cluster of habitats).
US. Although we do not know the details of extensions of sycamores adjacent to oak wood- connectivity among the populations involved lands, which explains the composition of their here, the high proportion of riparian-breeding bird community and the affi nity with this group species that also use riparian associations (and of habitats. Tropical deciduous forest and the mostly lowland riparian areas) in the winter in oases clustered near the riparian associations; northwestern Mexico indicates the importance they share a good number of species, maybe of these habitat types for wintering migrants. as a result of the extension of tropical dry for- Knowing that riparian environments are impor- est tree species farther north along the riparian tant for wintering birds, an additional question corridors, as well as the effect of the elevational might be asked: how unique are riparian bird gradient previously mentioned. communities during the winter in comparison My approach to assessing the contribution of with communities in other habitat types? Are riparian habitats to regional diversity is similar they important in terms of their contribution to to an approach advocated by Hylander (2006). regional diversity? My results indicate that riparian habitats con- Cluster analysis results show an interesting tribute substantially to regional diversity (22% pattern in the composition of the avian com- of species) in Sonora, Mexico, and are consistent munities—an elevational gradient separates in general with Sabo and Soykan’s (2006) fi nd- the highland habitats from the low elevation ings which state that on average, the percent- ones, and a gradient of humidity separates age of unique riparian species is 24%, and that dry habitat communities from ones associated riparian zones increase regional richness by with more humid riparian areas. This pattern 38%. My results indicate that wintering bird is in accordance with the TWINSPAN ordina- communities in riparian habitats are richer in tion, which in the highlands grouped a set of 19 comparison to adjacent uplands, and of the 134 coniferous forest bird species, a set of 17 species species recorded in our counts, 37 were exclu- in drier areas such as grasslands and scrubby sive to riparian habitats. These differences are vegetations, dry forest and oases shared 12 due mostly to the increased number of migrant species, and the lower elevation riparian asso- and partial migrant species and individuals ciations were typifi ed by a group of 24 species. detected in riparian habitats. Mangroves had a combination of four species Several important factors may explain the that defi ned its community. importance of riparian habitats to winter avian It is important to note that the riparian vege- diversity in Sonora. Riparian habitats provide tation in the highlands is represented by limited structural complexity, which is important for 64 STUDIES IN AVIAN BIOLOGY NO. 37
TABLE 5. RESULTS OF A TWO-WAY INDICATOR SPECIES ANALYSIS (TWINSPAN) TO IDENTIFY INDICATOR AND ASSOCIATED SPECIES CHARACTERIZING THE HABITAT CLUSTERS PRODUCED BY THE HIERARCHICAL ANALYSIS.
HIGHLAND VEGETATION CLUSTER MANGROVES Indicator species: Indicator species: Acorn Woodpecker (Melanerpes formicivorus) American Redstart (Setophaga ruticilla) Associated species: Associated species: Sharp-shinned Hawk (Accipiter striatus) Yellow Warbler (Dendroica petechia) Band-tailed Pigeon (Patagioenas fasciata)a Northern Waterthrush (Seiurus noveboracensis) White-eared Hummingbird (Hylocharis leucotis) Lincoln’s Sparrow (Melospiza lincolnii) Hairy Woodpecker (Picoides villosus) Williamson’s Sapsucker (Sphyrapicus thyroideus) b LOWER ELEVATION RIPARIAN Hutton’s Vireo (Vireo huttoni) ASSOCIATIONS Mexican Chickadee (Poecile sclateri) Indicator species: Eastern Bluebird (Sialia sialis) Violet-crowned Hummingbird (Amazilia American Robin (Turdus migratorius) violiceps) Brown Creeper (Certhia americana) Green Kingfi sher (Chloroceryle americana) Olive Warbler (Peucedramus taeniatus) Sinaloa Wren (Thryothorus sinaloa) Crescent-chested Warbler (Parula superciliosa) Associated species: Townsend’s Warbler (Dendroica townsendi) Elegant Quail (Callipepla douglasii) Hermit Warbler (Dendroica occidentalis) a Cooper’s Hawk (Accipiter cooperii) Grasshopper Sparrowb (Ammodramus Wilson Snipe (Gallinago delicata) savannarum) Plain-capped Starthroat (Heliomaster constantii) Yellow-eyed Junco (Junco phaeonotus) Belted Kingfi sher (Ceryle alcyon) Pine Siskin (Carduelis pinus) Pacifi c-slope Flycatcher (Empidonax diffi cilis) Scott’s Oriole (Icterus parisorum) Nutting’s Flycatcher (Myiarchus nuttingi) Cassin’s Vireo (Vireo cassinii) MICROPHYLLOUS SCRUBLAND Plumbeous Vireo (Vireo plumbeus) Indicator species: Warbling Vireo (Vireo gilvus) Rock Wren (Salpinctes obsoletus) Happy Wren (Thryothorus felix) Associated species: none American Pipit (Anthus rubescens) Lucy’s Warbler (Vermivora luciae) a DESERT SCRUBBY VEGETATION AND MacGillivray’s Warbler (Oporornis tolmiei) GRASSLANDS Black-and-white Warbler (Mniotilta varia) Indicator species: Painted Redstart (Myioborus pictus) Green-tailed Towhee (Pipilo chlorurus) Hepatic Tanager (Piranga fl ava) Associated species: Lazuli Bunting (Passerina amoena) Gambel’s Quail (Callipepla gambelii) Varied Bunting (Passerina versicolor) a c American Kestrel (Falco sparverius) Red-winged Blackbird (Agelaius phoeniceus) Common Ground-Dove (Columbina passerina) Lawrence’s Goldfi nch (Carduelis lawrencei) a b Broad-billed Hummingbird (Cynanthus latirostris) c TROPICAL DECIDUOUS FOREST AND OASES Costa’s Hummingbird (Calypte costae) a Indicator species: none Say’s Phoebe (Sayornis saya) Associated species: Brown-crested Flycatcher (Myiarchus tyrannulus) White-tipped Dove (Leptotila verreauxi) Vermilion Flycatcher (Pyrocephalus rubinus) Northern Beardless-Tyrannulet (Camptostoma Loggerhead Shrike (Lanius ludovicianus) b c imberbe) c Horned Lark (Eremophila alpestris) Nutting’s Flycatcher (Myiarchus nuttingi) Black-capped Gnatcatcher (Polioptila nigriceps) a Ash-throated Flycatcher (Myiarchus cinerascens) Bendire’s Thrasher (Toxostoma bendirei) a b c Canyon Wren (Catherpes mexicanus) Phainopepla (Phainoepla nitens) Five-striped Sparrow (Aimophila quinquestriata) Vesper Sparrow (Pooecetes gramineus) Rock Wren (Salpinctes obsoletus) White-crowned Sparrow (Zonotrichia leucophrys) Black-and-white Warbler (Mniotilta varia) Pyrrhuloxia (Cardinalis sinuatus) Wilson’s Warbler (Wilsonia pusilla) Rufous-capped Warbler (Basileuterus rufi frons) Rufous-crowned Sparrow (Aimophila rufi ceps) Streak-backed Oriole (Icterus pustuatus) a PIF Watch List. b USFWS birds of conservation concern national level. c USFWS birds of conservation concern regional level [Southwestern Region] USDI Fish and Wildlife Service 2002, Rich et al. 2004). HABITAT USE BY WINTERING BIRDS IN SONORA—Villaseñor-Gómez 65 breeding and wintering birds (MacArthur 1964, (2006) explored the ties between food resources Anderson et al. 1983, Farley et al. 1994a, Sanders and predation. They suggest that the avail- and Edge 1998, McComb et al. 2005). Wintering ability of food resources while feeding young migrants were found to be more abundant in affects the risk of nest predation; where fewer Acacia sp. patches with relatively high tree den- food resources exist parents have to increase sity and understory height in managed pasture- the number of feeding trips, attracting preda- lands in eastern Chiapas, Mexico (Greenberg et tors and reducing nest guarding time, therefore al. 1997). Compared with surrounding uplands, increasing the likelihood of nest predation. riparian habitats have a more complex vertical To my knowledge, in western Mexico no and horizontal structure, higher plant diversity, studies have addressed these possible expla- and more woody vegetation, especially in arid nations (diet composition and seasonality, landscapes like Sonora. predation, and habitat disturbance), and it Another possible factor is that vegetative was beyond the scope of this study. Because of productivity, which is higher in riparian eco- their importance to resident and migrant bird systems in general due to higher humidity species, and because they experience impacts and available water, and translates into a more on and variation in the ecological factors of abundant and diverse array of food items avail- interest, riparian corridors in Sonora provide a able to birds during all times of year. If this good opportunity to assess these concepts and were the case, one might expect resident species hypotheses. However, caution should be used to be equally or more abundant than migrants in designing such studies. Avian assemblages in riparian environments. However, according on riparian tracts and adjoining uplands are to my results this is not so; therefore, how can not independent (Strong and Bock 1990, Knopf we explain the higher numbers of migrant spe- and Samson 1994, Saab 1999, Martin et al. 2006), cies and individuals detected during winter in and it is complicated to defi ne the effects these riparian environments? habitats exert on each other. Birds can move The integration of migrant and resident spe- along the riparian corridor, as well as back and cies in the tropics and related wintering grounds forth between adjacent vegetation types. In this can be described as a paradox; ecosystem study, I made the practical assumption that productivity (and especially the abundance of by limiting the analyses to detections within a arthropods) is low when bird abundances reach 25-m radius I avoided most of these effects. their annual high during winter. Numerous During this study I did not detect any ter- hypotheses have been proposed to explain this ritorial behavior in insectivorous migrants, paradox. Greenberg (1995) proposed that a sea- but did fi nd that individuals of several species sonal abundance of large, protein-rich insects stayed in the same areas during the winter and supports the breeding productivity (feeding showed a certain degree of site fi delity (indi- of young) of resident populations, while small viduals banded in November were recaptured arthropods are available year-round in suffi cient in the same sites, and even in the same nets in biomass to support self-maintenance of adults, February of the next year, and even after two both resident and migrant. In Jamaica, Johnson consecutive years). Additional, research would et al. (2005) found some supporting evidence be valuable to increase our understanding of for one of Greenberg’s predictions. They found the importance of food abundance, water, and more wintering migrants using habitats that structural complexity of riparian areas in com- provided less breeding season food resources parison with non-riparian habitats, and the way for residents, and total abundance of birds was numerous birds fl eeing the cold, harsh northern correlated with total arthropod biomass in win- winters make use of these environments, as ter. Johnson et al. (2005), however, suggest that well as deeper insights on riparian winter site other ecological factors act in synergy with food fi delity and the extent to which wintering birds availability to affect migrant-to-resident ratios. move about and make use of adjoining vegeta- One of these other ecological factors could tion. be predation. Hutto (1980) suggested that dis- turbed habitats supported higher abundances MANAGEMENT IMPLICATIONS of wintering migrants because these habitats were underutilized by residents looking for The Sonoran habitats included in this study nesting sites safe from predation, leaving them fall with the Southwest Avifaunal Biome available for wintering species. This pattern has described in Rich et al.(2004). More particularly, been found in habitats with edges and those they are located within two bird conserva- that form habitat corridors (Kricher and Davis tion regions (BCR): the Sonoran and Mojave 1992, Hutto 1995, Villaseñor-Gómez and Hutto Desert BCR and the Sierra Madre Occidental 1995, Warkentin et al. 1995). Johnson et al. BCR. The southwest avifaunal biome includes 66 STUDIES IN AVIAN BIOLOGY NO. 37 more than half of the landbird Species of con- densities of spring migrating birds (Kelly and tinental importance identifi ed in this plan and Hutto 2005). For these reasons, riparian areas in many of the species have small population northwestern Mexico and southwestern US are sizes, restricted ranges, high threats, and/or unique and essential habitats for the wintering declining population trends. Of the primary and migrating bird species of western North habitats within this biome, riparian woodlands America as well as resident species, many of support the highest diversity of landbird spe- them of conservation concern. cies (Rich et al. 2004); according to my results, among the species associated to riparian envi- ACKNOWLEDGEMENTS ronments, three are included in the Partners in Flight Watch List: Lucy´s Warbler (Vermivora This research was part of the Western luciae), Varied Bunting (Passerina versicolor), and North American Land Bird Project: Wintering Lawrence´s Goldfi nch (Carduelis lawrencei); the Habitats in Sonora, a project funded by the Varied Bunting is also included by the USDI Neotropical Migratory Bird Conservation Act Fish and Wildlife Service as a species of con- (USDI Fish and Wildlife Service Agreement servation concern for the Southwestern Region No. 201814J856), the Sonoran Joint Venture, the (Region 2), and the Lawrence´s Goldfi nch as a University of Montana, and Instituto de Medio species of conservation concern at the national Ambiente y Desarrollo Sustentable de Sonora level (USDI Fish and Wildlife Service 2002). (IMADES). I appreciate the support of Robert Human settlements and activities are often Mesta, Mike Scott, Dick Hutto, Rob Hazlewood, closely associated with and dependent on ripar- and Carol Beardmore. I was also supported by ian ecosystems, imposing ecological pressures PROMEP through the Universidad Michoacana on riparian environments. Rich et al. (2004) also de San Nicolás de Hidalgo. Field activities were identifi es agricultural and suburban develop- possible thanks to the hard work and great spirit ment, grazing management, and habitat frag- of Rita Guadalupe Dávila Vindiola, Eduardo mentation as conservation issues or threats to Gómez Limón, Juan José Rivera Díaz, Leonardo this biome. Villaseñor Gómez, Fátima Méndez, Philip J. Due to the restricted areal extent of riparian Micheal, Denise Avila Jiménez, Sahira Aracely habitats in comparison to desertscrub and high- Leyva Briseño, and Rosalío León Carrasco; I land forests habitats in Sonora and elsewhere am deeply indebted to them. I express my deep in the Southwest, lowland riparian habitats gratitude to Creagh Breuner, Lisa Eby, Erick contribute signifi cantly to regional species Greene, Chris Guglielmo, Don Jenni, and espe- richness, supporting 22% of the total avifauna cially Dick Hutto, for their guidance, support in the state of Sonora. In addition to the impor- and important suggestions in all phases of this tance of Sonoran riparian areas as habitat for research. I also thank David Krueper, Janet M. wintering birds, as documented in this study, Ruth, and two anonymous reviewers who pro- they also act as corridors that permit the north- vided an impressive review; their suggestions ward expansion of tropical species and faunal improved this manuscript enormously. mixture on a broader scale, and support high HABITAT USE BY WINTERING BIRDS IN SONORA—Villaseñor-Gómez 67
d RIPARIAN
AND
RIPARIAN - preference c NON
IN
DETECTIONS ) P SE >15 WITH
SPECIES
) Mean ( ) Mean SE COMMON
MOST
THE
) Mean ( ) Mean FOR SE ) RADIUS
M (25- Mean ( Mean COUNT b
PER
N a INDIVIDUALS
OF
) R 16 0.01 (0.002) 0.001 (0.001) 0.02 (0.004) 13.69 (<0.01) Rip ) R 107 0.06 (0.01) 0.07 (0.01) 0.04 (0.01) 5.40 (0.02) Nrip ) R 18 0.01 (0.003) 0.01 (0.002) 0.01 (0.01) 3.53 (0.06) ) R 83 0.05 (0.01) 0.08 (0.01) 0.01 (0.004) 27.70 (<0.01) Nrip ) R 153 0.08 (0.01) 0.14 (0.02) 0.02 (0.01) 48.81 (<0.01) Nrip ) R 64 0.03 (0.01) 0.03 (0.01) 0.04 (0.01) 2.32 (0.13) ) R 24 0.013 (0.01) 0.001 (0.001) 0.03 (0.01) 3.65 (0.06) ) R 19 0.01 (0.004) 0.02 (0.01) 0.003 (0.003) 3.48 (0.06) ) PM 39 0.021 (0.01) 0.04 (0.02) 0.00 (0.00) 3.04 (0.08) ) M 17 0.01 (0.002) 0.004 (0.002) 0.01 (0.004) 5.63 (0.02) Rip ) R 50 0.03 (0.004) 0.001 (0.001) 0.06 (0.01) 38.52 (<0.01) Rip NUMBER
2004–2006). ) R 56 0.03 (0.004) 0.00 (0.00) 0.06 (0.01) 56.44 (<0.01) Rip ) R 215 0.12 (0.01) 0.10 (0.01) 0.13 (0.01) 1.80 (0.18) ) R 51 0.03 (0.01) 0.03 (0.01) 0.02 (0.01) 0.63 (0.43) THE ) PM 24 0.01 (0.003) 0.02 (0.004) 0.01 (0.003) 2.06 (0.15)
) M 49 0.03 (0.004) 0.001 (0.001) 0.05 (0.01) 47.85 (<0.01) Rip ) R 2 0.03 (0.004) 0.04 (0.01) 0.02 (0.01) 3.16 (0.08) ) R 16 0.01 (0.004) 0.01 (0.01) 0.01 (0.004) 1.08 (0.30) ) M 29 0.02 (0.003) 0.01 (0.003) 0.02 (0.01) 7.12 (<0.01) Rip ) R 43 0.02 (0.004) 0.02 (0.01) 0.03 (0.01) 0.45 (0.50) ) R 113 0.06 (0.01) 0.04 (0.01) 0.09 (0.03) 2.57 (0.11) sp.) M 148 0.08 (0.01) 0.01 (0.004) 0.16 (0.01) 116.42 (<0.01) Rip ) R 173 0.09 (0.02) 0.11 (0.04) 0.08 (0.02) 0.39 (0.53) ) R 79 0.04 (0.01) 0.06 (0.02) 0.02 (0.01) 3.46 (0.06) ) R 25 0.01 (0.003) 0.03 (0.01) 0.00 (0.00) 15.79 (<0.01) Nrip ) R 19 0.01 (0.003) 0.004 (0.003) 0.02 (0.01) 5.64 (0.02) Rip ) R 21 0.01 (0.003) 0.02 (0.004) 0.01 (0.003) 3.18 (0.08) EBRUARY Amazilia violiceps ) R 222 0.12 (0.01) 0.004 (0.002) 0.25 (0.02) 177.40 (<0.01) Rip PM 73 0.04 (0.01) 0.01 (0.003) 0.07 (0.01) 37.60 (<0.01) Rip ) R 70 0.04 (0.01) 0.06 (0.03) 0.01 (0.01) 3.05 (0.08) ) R 16 0.01 (0.002) 0.02 (0.004) 0.001 (0.001) 10.0 (<0.01) Nrip R 26 0.01 (0.004) 0.02 (0.01) 0.01 (0.003) 3.11 (0.08) ) R 31 0.02 (0.001) 0.03 (0.01) 0.01 (0.003) 4.50 (0.03) Nrip –F Picoides scalaris Cynanthus latirostris ) R 25 0.01 (0.004) 0.004 (0.002) 0.02 (0.01) 5.40 (0.02) Rip ) R 17 0.01 (0.01) 0.002 (0.002) 0.02 (0.01) 1.20 (0.27) PM 29 0.02 (0.003) 0.01 (0.003) 0.02 (0.01) 5.56 (0.02) Rip COMPARING
Myiarchus tuberulifer ) R 405 0.22 (0.01) 0.31 (0.02) 0.12 (0.01) 47.32 (<0.01) Nrip ) R 34 0.02 (0.01) 0.01 (0.01) 0.02 (0.01) 0.37 (0.54) Myiarchus cinerascens Columbina passerina Empidonax Calypte costae ANUARY Sphyrapicus nuchalis Pyrocephalus rubinus Tachycineta thalassina Zenida asiatica (J Melanerpes formicivorus RESULTS Actitis macularius Lanius ludovicianus ( aviceps Chloroceryle americana fl Melanerpes uropygials Baeolophus wollweberi Corvus corax) Colaptes auratus Zenaida macroura Empidonax wrightii Cathartes aura Callipepla gambelii Thryomanes bewickii Colaptes chrysoides Coragyps atratus ONORA Thryothorus sinaloa Sayornis nigricans Sayornis saya) Campylorhynchus brunneicapillus Troglodytes aedon) Aphelocoma ultramarina S Salpinctes obsoletus sher ( Columbina inca IN fi
haradrius vociferus Auriparus 1. ANOVA Psaltriparus minimus PPENDIX Turkey Vulture ( Inca Dove ( Common Ground-Dove ( Broad-billed Hummingbird ( Costa’s Hummingbird ( Red-naped Sapsucker ( Ladder-backed Woodpecker ( Violet-crowned Hummingbird ( Unknown hummingbird Green King Gila Woodpecker ( Northern Flicker ( Gilded Flicker ( PM Dusky-capped Flycatcher ( 17 0.01 (0.003) 0.002 (0.001) 0.02 (0.01) 9.08 (<0.01) Rip Gray Flycatcher ( Acorn Woodpecker House Wren ( Killdeer (C White-winged Dove ( Bewick’s Wren ( Black Vulture ( Spotted Sandpiper ( A VEGETATIONS Gambel’s Quail ( Mourning Dove ( Bushtit ( Bridled Titmouse ( Verdin ( Black Phoebe ( Vermilion Flycatcher ( Unknown Empidonax ( Say’s Phoebe ( Ash-throated Flycatcher ( Loggerhead Shrike ( Overall Non-riparian Riparian Non-riparian Overall Species Status (N = 1,816) (N = 944) (N = 872) Habitat Rock Wren ( Sinaloa Wren ( Cactus Wren ( Violet-green Swallow ( Mexican Jay ( Common Raven ( 68 STUDIES IN AVIAN BIOLOGY NO. 37 d preference c ) P SE ) Mean ( ) Mean SE ) Mean ( ) Mean SE Mean ( Mean b N a ) M 36 0.02 (0.004) 0.01 (0.003) 0.03 (0.01) 10.77 (<0.01) Rip ) M 172 0.09 (0.02) 0.04 (0.02) 0.15 (0.04) 8.40 (<0.01) Rip ) PM 26 0.01 (0.01) 0.02 (0.01) 0.00 (0.00) 3.62 (0.06) ) R 84 0.05 (0.01) 0.02 (0.01) 0.07 (0.01) 14.73 (<0.01) Rip ) R 46 0.02 (0.01) 0.04 (0.01) 0.01 (0.01) 3.41 (0.07) ) R 50 0.03 (0.01) 0.04 (0.01) 0.01 (0.01) 2.55 (0.11) ) R 85 0.05 (0.01) 0.07 (0.01) 0.02 (0.01) 6.79 (<0.01) Nrip ) R 217 0.12 (0.0) 0.16 (0.02) 0.07 (0.01) 17.59 (<0.01) Nrip ) M 436 0.24 (0.02) 0.06 (0.02) 0.43 (0.03) 116.18 (<0.01) Rip ) R 74 0.04 (0.01) 0.05 (0.01) 0.03 (0.02) 1.59 (0.21) ) M 145 0.08 (0.01) 0.06 (0.01) 0.11 (0.01) 9.17 (<0.01) Rip ) PM 32 0.02 (0.01) 0.00 (0.00) 0.04 (0.02) 2.57 (0.11) ) PM 180 0.10 (0.03) 0.15 (0.07) 0.05 (0.01) 2.06 (0.15) ) R 21 0.01 (0.01) 0.01 (0.001) 0.01 (0.01) 0.09 (0.76) ) M 506 0.28 (0.02) 0.06 (0.01) 0.51 (0.030) 209.68 (<0.01) Rip ) PM 170 0.09 (0.01) 0.031 (0.01) 0.16 (0.01) 66.33 (<0.01) Rip ) PM 440 0.24 (0.01) 0.13 (0.02) 0.36 (0.02) 65.25 (<0.01) Rip ceps ) PM 21 0.01 (0.01) 0.02 (0.02) 0.001 (0.001) 0.82 (0.37) fi ) R 134 0.07 (0.01) 0.05 (0.01) 0.10 (0.01) 9.78 (<0.01) Rip ) R 34 0.02 (0.003) 0.001 (0.001) 0.04 (0.01) 28.87 (<0.01) Rip ) M 39 0.02 (0.01) 0.04 (0.02) 0.00 (0.00) 3.88 (<0.05) Nrip ) M 174 0.10 (0.01) 0.03 (0.01) 0.16 (0.02) 53.29 (<0.01) Rip ) M 47 0.03 (0.01) 0.004 (0.003) 0.05 (0.01) 16.67 (<0.01) Rip PM 22 0.01 (0.01) 0.02 (0.02) 0.00 (0.00) 1.35 (0.25) ) M 255 0.14 (0.03) 0.07 (0.02) 0.21 (0.07) 4.33 (0.04) Rip ) PM 515 0.28 (0.053) 0.14 (0.04) 0.44 (0.10) 8.32 (<0.01) Rip ) R 21 0.01 (0.01) 0.02 (0.01) 0.00 (0.00) 5.58 (0.02) Nrip ) R 222 0.12 (0.01) 0.13 (0.02) 0.11 (0.02) 0.76 (0.38) ) R 124 0.07 (0.01) 0.02 (0.01) 0.12 (0.03) 15.48 (<0.01) Rip ) M 127 0.07 (0.02) 0.09 (0.03) 0.05 (0.03) 0.67 (0.41) ) M 68 0.04 (0.01) 0.001 (0.001) 0.08 (0.01) 55.95 (<0.01) Rip ) PM 17 0.01 (0.003) 0.01 (0.01) 0.01 (0.002) 2.56 (0.11) ) PM 351 0.19 (0.01) 0.001 (0.001) 0.401 (0.03) 247.84 (<0.01) Rip ) M 60 0.03 (0.01) 0.06 (0.02) 0.001 (0.001) 7.65 (<0.01) Nrip ) R 41 0.02 (0.01) 0.01 (0.004) 0.03 (0.01) 5.48 (0.02) Rip ) R 47 0.03 (0.01) 0.04 (0.01) 0.01 (0.003) 12.79 (<0.01) ) R 56 0.03 (0.01) 0.04 (0.01) 0.01 (0.01) 9.57 (<0.01) Nrip Dendroica nigrescens Aimophila ru Polioptila nigriceps Vermivora celata Dendroica coronata Polioptila melanura Zonotrichia leucophrys Aimophila carpalis Amphispiza bilineata Geothlypis trichas Mimus polyglottos Regulus calendula Polioptila caerulea Toxostoma curvirostre Spizella pallida Agelaius phoeniceus Sturnella neglecta Pipilo chlorurus Icterus pustuatus Quiscalus mexicanus . Passerculus sandwichensis Junco phaeonotus Cardinalis cardinalis Spizella passerina Melospiza lincolnii Spizella breweri Wilsonia pusilla Pipilo fuscus Turdus migratorius) Carduelis psaltria Pipilo maculatus Junco hyemalis Melospiza melodia Chondestes grammacus Carpodacus mexicanus Phainoepla nitens Cardinalis sinuatus ONTINUED nch ( nch fi 1. C PPENDIX Rip = preference for riparian and Nrip non-riparian habitats. Number of individuals detected. Status: R = resident, PM partial migrant, M migrant. ANOVA (P) analysis. Northern Cardinal ( Western Meadowlark ( Dark-eyed Junco ( Yellow-eyed Junco ( Pyrrhuloxia ( Red-winged Blackbird ( White-crowned Sparrow ( Clay-colored Sparrow ( Black-throated Sparrow ( Savannah Sparrow ( Lincoln’s Sparrow ( Song Sparrow ( Blue-gray Gnatcatcher ( Black-tailed Gnatcatcher ( Rufous-crowned Sparrow ( Overall Non-riparian Riparian Non-riparian Overall Species Status (N = 1,816) (N = 944) (N = 872) Habitat A Ruby-crowned Kinglet ( Black-capped Gnatcatcher ( American Robin ( Black-throat. Gray Warbler ( Spotted Towhee ( Canyon Towhee ( Rufous-winged Sparrow ( Chipping Sparrow ( Brewer’s Sparrow ( Great-tailed Grackle ( a b c d Green-tailed Towhee ( Lark Sparrow ( Northern Mockingbird ( Curve-billed Thrasher ( Phainopepla ( Orange-crowned Warbler ( Yellow-rumped Warbler ( Common Yellowthroat ( Wilson’s Warbler ( Streak-backed Oriole ( House Finch ( Lesser Gold Studies in Avian Biology No. 37:146–165
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