FRONTISPIECE. Male (top) and female (other three individuals) berlepschi, Esmeraldas Wood- star, an endangered endemic to western visiting flowers of spicata (Gesneriaceae) and Cornutia pyramidata (Lamiaceae). The original description of female C. berlepschi was incorrect because it came from mis-labeled specimens of juvenile males. Original painting by Paul J. Greenfield. Published by the Wilson Ornithological Society

VOL. 121, NO. 2June 2009 PAGES 227–456

The Wilson Journal of Ornithology 121(2):227–239, 2009

DISTRIBUTION, PLUMAGE, AND OF THE ENDEMIC (CHAETOCERCUS BERLEPSCHI) OF WESTERN ECUADOR

J. BERTON C. HARRIS,1,4,6 ANA E. A´ GREDA,2 MERY E. JUIN˜ A,1 AND BERND P. FREYMANN3,5

ABSTRACT.—The Esmeraldas Woodstar (Chaetocercus berlepschi) is a poorly known and endangered hum- mingbird endemic to lowland and foothill moist forest in coastal western Ecuador. We encountered 11 new localities, observed two copulations, and found 26 nests of the from October 2007 to April 2008. We observed the generally accepted descriptions of the female must have come from mis-labeled specimens of juvenile males and were incorrect. We collected the first three confirmed females of the species and describe their characteristics. The correct identification of female C. berlepschi and recognition of the species’ breeding habitat should facilitate more effective conservation of the species. Received 17 June 2008. Accepted 24 November 2008.

Chaetocercus (Trochilidae) is presently tle Woodstar; C. heliodor, ; considered to comprise six species (C. mul- C. astreans, ; C. berlep- sant, White-bellied Woodstar; C. bombus, Lit- schi, Esmeraldas Woodstar; and C. jourdanii, Rufous-shafted Woodstar) which inhabit semi- 1 Fundacio´n de Conservacio´n Jocotoco, Avenida arid to humid forest and woodland in northern Los Shyris N37-146 y El Comercio, Quito, Ecuador. and western South America from 0 to 4,000 2 Aves & Conservacio´n (Corporacio´n Ornitolo´gica m above sea level (Graves 1986, Schuchmann del Ecuador), Pasaje Joaquı´n Tinajero E3-05 y Jorge 1999). Four of the six species are generally Drom, Casilla 17-17-906, Quito, Ecuador. rare to uncommon, three (C. bombus, C. ber- 3 Alexander Koenig Research Institute and Museum of lepschi, and C. astreans) have restricted rang- Zoology (ZFMK), Leibniz Institute for Terrestrial Biodi- versity, Research Group: Biology and Phylogeny of Trop- es, and two are globally threatened (C. bom- ical , Adenauerallee 160, 53113 Bonn, Germany. bus, Vulnerable; C. berlepschi, Endangered) 4 Current address: Research Institute for Climate (Schuchmann 1999; BirdLife International Change and Sustainability, School of Earth and Envi- 2000, 2008a, b). Chaetocercus woodstars are ronmental Sciences, University of Adelaide, Australia. difficult to detect in the field because of their 5 Current address: Community and Conservation small body size and inconspicuous behavior Ecology Group, Centre for Ecological and Evolution- ary Studies, University of Groningen, P. O. Box 14, outside of the breeding season. Their small 9750 AA Haren, The Netherlands. size makes collection and specimen prepara- 6 Corresponding author; e-mail: tion challenging and, coupled with a reduction [email protected] in collecting effort, has resulted in a dearth of 227 228 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 121, No. 2, June 2009 specimens for some species (Freymann and C. berlepschi is sympatric with C. bombus, Schuchmann 2005). The great majority of and accurate identification of either species specimens that exist were collected before depends on careful consideration of the other. 1960 (Freymann and Schuchmann 2005). C. berlepschi males are characterized by green Female Chaetocercus woodstars are similar upperparts with a faint bluish sheen and white to each other in plumage and are, at times, underparts with a narrow green chest band. considered to be inseparable in the field (Hilty They have a purple gorget (Purple 1; Smithe and Brown 1986, Hilty 2003). These factors 1975), and a forked tail with a short and have combined to make the genus poorly rounded rectrix 1 (R1), a longer and narrower known as evidenced by few recent records of pointed R2, and long, distinctive R3-5 re- several species and a paucity of detailed duced nearly to shafts (Ridgely and Greenfield breeding biology information for any 2001; Frontispiece). Males of C. bombus have (Schuchmann 1999). All species except C. a ruby-pink gorget, more extensive green on jourdanii were formerly placed in the genus the underparts, a cinnamon-buff pectoral col- Acestrura but most recent authors and check- lar, and usually display bronzier green upper- lists follow Schuchmann (1999) in merging parts (Schuchmann 1999, Ridgely and Green- these species into Chaetocercus because there field 2001, Gurney 2006, Mata et al. 2006, was insufficient morphological justification Restall et al. 2006, Schulenberg et al. 2007). for their separation (e.g., Dickinson 2003, Re- Published works illustrate C. berlepschi fe- stall et al. 2006, Remsen et al. 2008). males as white below with a white postocular One of the least-known Chaetocercus is the stripe, and a buff-tinged throat (Meyer de localized Ecuadorian endemic Esmeraldas Schauensee 1970, Schuchmann 1999, Ridgely Woodstar (Simon 1889). This hummingbird and Greenfield 2001, Mata et al. 2006, Restall inhabits semi-deciduous to evergreen moist et al. 2006). The central rectrices of the female forest (ϳ1,500 mm annual rainfall) along the are green and R2-5 are cinnamon with a black Pacific coast of western Ecuador from near subterminal band and white tips. Depictions sea level to 750 m elevation (Becker et al. of female C. bombus vary but all suggest that 2000, Ridgely and Greenfield 2001, A´ greda its cinnamon-buff underparts and postocular 2007) (Fig. 1). The vast majority of records stripe are the main identifying features come from the rainy season, mid October to (Schuchmann 1999, Ridgely and Greenfield late May. The species is found along a gra- 2001, Gurney 2006, Mata et al. 2006, Restall dient from low elevation (0–250 m), partially et al. 2006, Schulenberg et al. 2007). The tail disturbed areas (Becker et al. 2000, Ridgely is shown as cinnamon with a black subter- and Greenfield 2001) to more intact, higher minal band (the tips are also cinnamon) and elevation (250–750 m), misty garu´a forest in green is at times shown on the central rectri- the hills of the Cordillera Chongo´n-Colonche ces. The short and narrow cheekstripe (exten- (A´ greda 2007). Contrary to Mata et al. (2006), sion of the auriculars) is also a diagnostic fea- C. berlepschi is not known from subtropical ture of female C. bombus (Gurney 2006). or temperate forests. The species seems to Much of the lowland humid forest in west- breed in lower elevation, disturbed areas along ern Ecuador has been cleared, threatening nu- the central Ecuadorian coast and move to merous species including C. berlepschi (Dod- northwestern Ecuador for the non-breeding son and Gentry 1991, Best and Kessler 1995), season (MEJ, pers. obs.; C. D. Becker, pers. and the species is considered rare to uncom- comm.). Data are still limited because move- mon (Becker et al. 2000, Ridgely and Green- ments and breeding biology of the species re- field 2001). The combination of these factors main poorly known. Before this study, C. ber- has led to the perception that the species is lepschi had been recorded from 11 localities Endangered with an estimated population size but was only regularly observed at two: Rı´o of 250–999 individuals (Collar et al. 1992; Ayampe (01Њ 41Ј S, 80Њ 48Ј W; Manabı´ Prov- BirdLife International 2000, 2008a) and ince), and Reserva Ecolo´gica Loma Alta should perhaps even be treated as Critically (Loma Alta; 1Њ 50Ј S, 80Њ 39Ј W; Santa Elena Endangered (Ridgely and Greenfield 2001). Province) (Collar et al. 1992, Ridgely and The Esmeraldas Woodstar has received lit- Greenfield 2001, A´ greda 2007). tle research attention despite its restricted Harris et al. • ESMERALDAS WOODSTAR DISTRIBUTION AND PLUMAGE 229

FIG. 1. Western Ecuador showing confirmed records of Chaetocercus berlepschi (Esmeraldas Woodstar) as well as larger cities and landmarks. range and Endangered status, and little was and Santa Elena provinces, western Ecuador. known of its reproductive biology. The objec- We searched for the species at Ayampe in Oc- tives of our study were to: (1) gather funda- tober and November and documented the ar- mental natural history and distributional in- rival date of male and female C. berlepschi. formation on the species, (2) assess its con- We continued our searches from December servation status, and (3) recommend possible 2007 to April 2008 along the coast from San conservation actions. Jose´(1Њ 45Ј S, 80Њ 45Ј W; 30 m; Santa Elena Province) north to Cabo Pasado (0Њ 24Ј S, 80Њ METHODS 29Ј W; 25 m; Manabı´ Province; Figs. 1,2). We We searched for and observed C. berlepschi surveyed areas Ͻ20 km from the coast from from October 2007 to April 2008 in Manabı´ 0 to 700 m elevation that varied from arid to 230 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 121, No. 2, June 2009 humid. We were based in Ayampe and data Vuilleumier 1998, 2000; Freymann and were collected from that site across all months Schuchmann 2005; Cuervo et al. 2006). of the study. We also documented the location We described the plumage colors of our and elevation of each C. berlepschi nest to specimens with Munsell (1994) and Smithe obtain information on which areas are impor- (1975) color standards. Lower case color tant for reproduction. We complemented names refer to the Munsell (1994) classifica- breeding season observations with intensive tion; capitalized color names refer to Smithe searches in August and September 2008 that (1975). attempted to locate non-breeding localities. We compared our specimens to the nine C. We searched along the coast from Ayampe to berlepschi specimens (AMNH 37925, San Lorenzo (1Њ 4Ј S, 80Њ 53Ј W; Manabı´ 118529–34, 119766, 119768) and specimens Province) to Bilsa Biological Station (0Њ 22Ј of the other five species of Chaetocercus (C. N, 79Њ 45Ј W; Esmeraldas Province) from 3 to astreans, C. bombus, C. jourdanii, C. helio- 20 August 2008. We also surveyed the north- dor, and C. mulsant) at the American Museum ern and central Cordillera Chongo´n-Colonche of Natural History (AMNH). We examined all in Manabı´ Province (0 to 700 m elevation) of the specimens of each species with partic- from 22 August to 9 September 2008. We ular attention to the plumage of the tail, un- Њ Ј Њ Ј searched from Agua Blanca (1 32 S, 80 44 derparts, postocular stripe, cheek stripe, and Њ Ј Њ Ј W) to Pedro Pablo Go´mez (1 37 S, 80 33 chest band. We drew detailed figures of the Њ Ј Њ Ј W) to Puerto Cayo (1 21 S, 80 44 W) to tail patterns of the plumages of greatest im- Њ Ј Њ Ј above Gala´n (1 20 S, 80 40 W), and back portance to this study (female C. berlepschi, to Ayampe. female C. bombus, and juvenile male C. ber- We recorded detailed field notes on plum- lepschi; Fig. 3). We used a caliper accurate to age and took photographs with a 10 megapixel 0.02 mm to measure wing chord, length of R1 Canon EOS 40D digital camera with a 300 and R5, and bill length from the anterior edge mm telephoto lens for future plumage analysis of the nostril to the bill tip to compare mor- of each female that was courted by a male C. phology of the different species. We measured berlepschi (n Ͼ 40) or that was in attendance a representative specimen-series of all con- at a nest (n ϭ 21). generics (Table 1). We also analyzed mea- We realized that female C. berlepschi were surements and photographs (when available) incorrectly described and collected three fe- of the following specimens: an adult male C. males on 4–6 March 2008 at Ayampe. The berlepschi from the Academy of Natural Sci- specimens (AS 1374, AS 1375, and AS 1377; MECN uncatalogued) were deposited at the ences, Philadelphia (ANSP 183118), a juve- Museo Ecuatoriano de Ciencias Naturales, nile male from the Museum and Institute of Quito, Ecuador (MECN). We also saved tissue Zoology, Polish Academy of Sciences, War- samples from all three specimens and blood saw, Poland (MIZPAN 44432), a male from from the two for which it was possible (AS Museu de Cie`ncies Naturals, Zoologia, Bar- 1375 and 1377) to enable future genetic anal- celona, Spain (MCNC-MZB 85-0542), and an yses. adult male C. bombus from Zoologisches For- Collection of endangered species must be schungsinstitut und Museum A. Koenig, Ger- carefully considered and we collected the many (ZFMK 9972). We also reviewed de- minimum number of individuals required to scriptions and measurements of the different describe the female. Conservation measures species in the literature (Simon 1889, Hilty for the species necessitate an accurate descrip- and Brown 1986, Schuchmann 1999, Cle- tion of the female, and our judgment was the ments and Shany 2001, Ridgely and Green- need to eliminate any possible uncertainties field 2001, Hilty 2003, Gurney 2006, Mata et justified taking this small sample. Collecting, al. 2006, Restall et al. 2006, Schulenberg et especially of under-represented genera such as al. 2007). Chaetocercus, is considered to be necessary We used General Linear Models (GLMs) to and is amply supported in the literature test whether there were significant differences (Winker et al. 1991; Remsen 1995, 1997; between means of the four measured morpho- Winker 1996, 2005a, b; Peterson et al. 1998; metric parameters between female C. berlep- Harris et al. • ESMERALDAS WOODSTAR DISTRIBUTION AND PLUMAGE 231

FIG. 2. Focal area of study (inset in Fig. 1) showing confirmed records of Chaetocercus berlepschi (Es- meraldas Woodstar) as well as towns and landmarks. schi and C. bombus. Analyses were conducted 49Ј W; 40 m; Manabı´ Province), Puerto Rico in Statistica 6.1 (StatSoft Inc. 2003). (1Њ 37Ј S, 80Њ 49Ј W; 50–100 m; Manabı´ Prov- ince), Las Tunas (1Њ 39Ј S, 80Њ 48Ј W; 15 m; RESULTS Manabı´ Province), Ayampe/Atamari/Cinco We encountered Chaetocercus berlepschi at Cerros (30–140 m), Rı´o San Jacinto (1Њ 40Ј 12 sites separated from each other by at least S, 80Њ 46Ј W; 40 m; Manabı´ Province), and 2 km (Figs. 1,2): San Lorenzo (200 m), Tro- San Jose´ (30 m). C. berlepschi had been re- pieso (1Њ 9Ј S, 80Њ 51Ј W; 140 m; Manabı´ corded in only one of these areas, Ayampe/ Province), above Gala´n (350 m), San Sebas- Atamari/Cinco Cerros, before this study. tia´n (1Њ 36Ј S, 80Њ 42Ј W; 675 m; Manabı´ We began to search the Ayampe area inten- Province), Rı´o Blanco (1Њ 36Ј S, 80Њ 45Ј W; sively on 11 October 2007 but did not en- 175–240 m; Manabı´ Province), Salango (1Њ counter female C. berlepschi until 4 Novem- 35Ј S, 80Њ 50Ј W; 50 m; Manabı´ Province), ber. We did not monitor the area between 14 Rı´o Chico/Hacienda Tocuyo (1Њ 36Ј S, 80Њ November and 6 December but, when we re- 232 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 121, No. 2, June 2009 0.67 (11.3–13.3) 0.45 (11.5–14.1) 0.34 (12.0–15.0) 0.23 (12.0–13.0) 0.32 (10.4–14.0) 0.29 (11.7–13.8) 0.30 (13.0–15.6) 0.16 (17.2–21.6) 0.45 (13.6–15.0) 0.23 (10.9–13.5) Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ 0.15 (13.2–13.7)0.24 (12.0–13.4)0.18 (10.9–11.5) 12.63 0.25 (13.6–16.1) 12.88 14.5 0.19 (13.4–14.0) 13.98 0.17 (8.9–11.1) 12.63 12.59 0.4 (11.4–14.5)0.14 (8.1–9.7)0.22 (14.6–17.3) 12.55 0.16 (14.9–16.8) 14.43 12.8 19.28 Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ C. j. rosae. 4 for ϭ . Bill length was measured from the anterior edge of the nostril 0.43 (28.1–29.4)0.26 (27.0–29.0) 13.48 0.22 (29.2–31.8) 12.68 0.43 (35.9–37.8) 14.64 0.19 (32.8–35.0) 13.63 10.09 0.13 (27.4–28.6)0.68 (32.7–41.6)0.27 (38.0–40.9) 8.81 16.01 15.85 0.36 (25.4–26.6) 11.23 0.21 (31.2–32.9) 13.29 Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ C. h. cleavesi. C. j. andinus, n 2 for ϭ 7for Chaetocercus ϭ C. h. heliodor, n C. j. jourdanii, n

FIG. 3. Tail patterns of (A) adult female Chaeto- 9 for ϭ cercus berlepschi (Esmeraldas Woodstar), (B) adult fe- 2 for n ϭ male C. bombus (), and (C) juvenile 0.41 (12.02–12.84)0.1 (14.3–15.0) 28.55 0.17 (14.2–17.0) 28.08 30.46 0.17 (15.0–16.7)0.44 (16.3–21.7) 27.89 39.09 n 0.3 (14.7–15.3)0.36 (14.0–15.7)0.22 25.9 (12.9–15.3) 36.85 0.22 (15.7–17.1) 34.05 0.2 (18.1–20.7) 32.3 39.44 male C. berlepschi. Ϯ Ϯ Ϯ Ϯ Ϯ Males: Ϯ Ϯ Ϯ Ϯ Ϯ Males: SE, range in parentheses) of 12.43 14.63 15.0

turned on 7 December, we observed the first Ϯ male C. berlepschi of the season. C. berlep- ¯ C. h. cleavesi. C. j. rosae. c schi was seasonally (mid Nov–Apr) fairly d 4for a,b 2 for bill. common in partially disturbed forest and 2for ϭ ϭ ϭ n

woodland along the Rı´o Ayampe. We ob- 2 for bill. served several individual C. berlepschi on ϭ n Female (1)Male (1) 16.2 16.7 33.9 30.3 14.1 9.4 14.1 13.3 Juv male (8) Ad male (3) Female (15)Male (1)Male (13) 15.35 14.4Male (11) 14.4 Female (13)Male (13) 15.86 20.21 19.6 25.8 9.3 13.7 Female (3) nearly every visit to the field at Ayampe dur- Female (4)Female (8) 14.9 16.6

ing that time period. We recorded the last 6 for R1 and R5. C. j. andinus, n C. h. heliodor, n male C. berlepschi on 4 May and the last fe- ϭ n 2for 4for 7; male on 24 May 2008. We did not encounter e f ϭ ϭ ϭ n n

C. berlepschi during our non-breeding season n searches. Taxon Gender Bill Wing chord Rectrix 1 Rectrix 5 Bill destroyed on AS 1374, Bill destroyed on AMNH 118532, Sample size in parentheses. For bill Females: Females:

The most important floral resource for C. TABLE 1. Measurements (mm) (x a b c d e f C. heliodor C. astreans C. jourdanii C. bombus C. mulsant C. berlepschi berlepschi at Ayampe was Kohleria spicata to the tip of the bill. Harris et al. • ESMERALDAS WOODSTAR DISTRIBUTION AND PLUMAGE 233

(Kunth) Oerst (Gesneriaceae, Frontispiece). C. these displays at six localities (San Lorenzo, berlepschi abundance was highest at Ayampe Tropieso, Gala´n, Rı´o Chico, Puerto Rico, and during the peak of the Kohleria spicata flow- Ayampe). All displays involved females with ering event from January to March. At all oth- pale buffy underparts; none involved females er locations, Cornutia pyramidata (Lamiaceae) with white underparts. We observed two cop- was the plant most commonly-visited. We did ulations on 9–10 February 2008 between male not encounter flowers of Psychotria hazennii C. berlepschi and females of the above de- Standl. (Rubiaceae) or Razisea cf. ericae scription. We encountered 26 Chaetocercus Mildbraed ex Wassh. (Acanthaceae), both of nests, of which 21 were active, at nine sites. which are important resources for the species All active nests were attended by females with in high elevation garu´a forests. We observed pale buffy underparts, white postocular C. berlepschi at several sites (e.g., Puerto stripes, and the same characteristic tail pattern. Rico, Rı´o Chico) along wooded riparian cor- Female nestlings (n ϭ 10) were buffy below ridors in a matrix of deforested land. These with green central rectrices tipped cinnamon riparian corridors connect lower elevation, de- in a pattern similar to the females we collect- forested areas to large forest patches of the ed, and unlike adult female C. bombus. A beg- upland Cordillera Chongo´n-Colonche. ging male fledgling observed on 11 March All C. berlepschi nests were Ͻ14 km from 2008 was distinguishable from female nest- the coast from 30 to 350 m elevation and most lings. Its tail and underparts were similar to were in areas disturbed by cattle ranching and that of other juvenile males, but it lacked any agriculture, but adjacent to large forest patch- obvious purple gorget feathers. es. Nests at a few sites (e.g., Rı´o Blanco) were We observed birds that generally matched in mature moist forest. Female C. berlepschi the illustrations of ‘‘female C. berlepschi’’ on usually placed their nests near the top of small several occasions. Our data suggest these dead trees (ϳ5 m tall) near creeks or roads. birds were juvenile male C. berlepschi rather The nest, eggs, nestlings, nest placement, and than females. These birds have white under- parental care will later be described in detail parts similar to that of adult males and have (MEJ, in prep.). a buff-tinged throat with several gorget feath- Male C. berlepschi and female Chaetocer- ers that are the same color purple as adult cus woodstars with pale buffy underparts, males. Their tails are distinctive: R1 is rela- white postocular stripes, and distinctive tail tively short, rounded and green, R2 is long, patterns (Frontispiece, Fig. 3A) were season- narrow and dusky, and R3-5 have a rufous- ally fairly common at all of our observation cinnamon base, black subterminal band, and sites, while male and female C. bombus were tips that are pale cinnamon to whitish with a rare or absent. C. bombus has a different tail pale pinkish buff tinge (Fig. 3C; the tips are pattern (no green on the outer webs of the shown as white without any buff tinge in the central rectrices, a narrower black subterminal sources that illustrate the ‘‘female’’ of the spe- band, and less black on R5; Fig. 3B), a buffier cies). We usually observed birds of this de- postocular stripe, and richer buff underparts scription when they challenged adult males compared to the females we observed. We ini- for display perches and we did not observe tially assumed these females were variable in- them copulate. dividuals of C. bombus because their buffy Museum specimens provide strong support underparts differed strikingly from published for our conclusion. AMNH specimen 37925 descriptions of female C. berlepschi. and MIZPAN 44432, labeled as juvenile Male C. berlepschi defend perches at the males, display the same white underparts and top of small (ϳ10 m tall) dead trees. When a tail pattern of the juvenile males we observed female arrives at a perch, the male flies ver- in the field. AMNH specimens 118533, tically above the perch until no longer visible 119766, and 119768, and MCNC-MZB 85- to the naked eye, and then quickly drops to 0542, which are labeled as males but were not the perch while making a mechanical noise. classified to age, also show these characteris- The mechanical noise is likely produced by tics and are likely juvenile males. Two AMNH the distinctive narrow rectrices (R. S. Ridgely, specimens labeled as females (118529 and pers. comm.). We observed males perform 118534) seem to be the basis for the original 234 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 121, No. 2, June 2009 description of the female of the species (R. S. tips of R3-5 in C. bombus are at times shaped Ridgely, pers. comm.). These two skins show into distinctive diamonds by the subterminal the characteristics of juvenile male C. berlep- band, but this is variable. C. bombus’s post- schi, most notably their tail pattern (Fig. 3C). ocular stripe tends to be rich cinnamon while The only difference between the juvenile the postocular of C. berlepschi is white and males and one of these ‘‘females’’ is the lack only occasionally includes some buff feathers. of purple gorget feathers in specimen 118529, C. berlepschi shows a long and wide cheek an absence that is probably related to age. The stripe that is often mostly green compared to third ‘‘female’’ (118531) which, like AMNH C. bombus’s usually short, narrow, and dusky 118529–530, 118532–534, 119766, and cheek stripe. In addition, the underparts of C. 119768, was collected by W. B. Richardson, bombus are usually rich cinnamon-buff com- is also mis-labeled. The skin is actually an pared to the pale buff underparts of C. ber- adult male with a fully-grown adult male tail lepschi. The most posterior feather tracts of (measurements of each rectrix are similar to the uppertail coverts in C. bombus are at times the morphometric measurements of the other buffy, while the uppertail coverts of C. ber- 2 adult males). The specimen had lost most of lepschi are green. its gorget feathers (only skin remained) but C. bombus is significantly larger than C. the remnant gorget feathers were all purple berlepschi. Females of C. bombus have sig- ϭ and it is likely that if the other feathers had nificantly longer bills (GLM: F1,12 40.91, P Ͻ ϭ not been lost, the specimen would have a full 0.001), longer wings (GLM: F2,14 13.36, ϭ ϭ adult gorget. This obvious error, along with P 0.002), and longer R1 (GLM: F2,9 5.96, the more subtle errors, suggests the gonads P ϭ 0.032), whereas the length of R5 is not were not carefully examined on some of the significantly different between females of the ϭ ϭ AMNH specimens collected by W. B. Rich- two species (GLM: F2,8 4.87, P 0.052; ardson. It is not especially surprising that mis- Table 1). takes were made because gonads of Chaeto- Among the other four Chaetocercus, C. cercus are extremely small. We conclude there berlepschi females are most similar to C. mul- actually are no specimens of female C. ber- sant in tail pattern. C. mulsant usually has lepschi currently in any museum collection green central rectrices with a black subtermi- and describe the gender. nal band and extensive cinnamon-rufous tips. The tail pattern of C. berlepschi is subtly dif- Description of Female ferent; the cinnamon-rufous tips are small in C. berlepschi females are characterized pri- extent and unevenly bi-lobed in shape (Fig. marily by their green central rectrices with 3A). The central rectrices of C. mulsant are unique small, unevenly bi-lobed cinnamon-ru- variable and, at times, are different from the fous tips and their white postocular stripe pattern in C. berlepschi. C. mulsant can dis- (Frontispiece; Fig. 3A). C. berlepschi is subtly play a small area of rufous-brown in the outer different from all other Chaetocercus females, web of the basal part of the central rectrix. but the species shows several characteristics The central rectrices at times can be almost similar to other members of the genus such as entirely dusky with a greenish sheen and a buffy underparts, green central rectrices, cin- small cinnamon-rufous tip. C. mulsant is no- namon R2-4 with a black subterminal band, ticeably larger than C. berlepschi (Table 1) and green patches on the sides of the chest and the species are not known to be sympatric. connected by a faint dusky band. C. berlep- C. jourdanii and C. astreans differ from C. schi is best distinguished by a combination of berlepschi in their entirely green central rec- plumage characteristics, morphometrics, and trices and R2 differs in pattern from R3-5. C. distribution. The only sympatric Chaetocercus heliodor is distinguished by its rich cinnamon- is C. bombus which is similar in size (Table rufous rump and underparts. C. heliodor also 1), but shows a different tail pattern with has the least amount of green on the central green present only as a sheen on parts of the rectrices of any Chaetocercus (just a fleck of inner webs of the central rectrices, a narrower green proximal to the subterminal band). black subterminal band on R2-5, and much The following description is based on the more brown on R5 (Fig. 3B). The cinnamon sexually mature female collected 5 March Harris et al. • ESMERALDAS WOODSTAR DISTRIBUTION AND PLUMAGE 235

TABLE 2. Characteristics of three female Chaetocercus berlepschi (Esmeraldas Woodstar) specimens col- lected at Rı´o Ayampe (Manabı´ Province, Ecuador) in March 2008.

Specimen numbera AS 1374 AS 1375 AS 1377 Collection date 4 Mar 2008 5 Mar 2008 6 Mar 2008 Wing chord (mm) 28.2 29.4 28.1 Anterior edge of nostril to tip of bill (mm) NAb 12.8 12.0 Rectrix 1 length (mm) 13.4 13.4 13.7 Rectrix 2 length (mm) 14.6 14.5 15.1 Rectrix 3 length (mm) 14.4 14.5 15.4 Rectrix 4 length (mm) 13.4 13.4 14.5 Rectrix 5 length (mm) 11.3 13.3 13.3 Body mass (g) 2.0 2.1 1.9 Molt strong general none light general Ovary dimensions (mm) 3.0 ϫ 2.0 3.0 ϫ 2.0 1.6 ϫ 1.0 Number ovarian follicles none-ovary smooth nine none-ovary very smooth Dimensions of largest follicle (mm) NA 1.0 ϫ 1.0 NA

a Specimens are deposited in Museo Ecuatoriano de Ciencias Naturales, Quito, Ecuador (MECN; uncatalogued). b Bill destroyed on AS 1374.

2008 by Aldo Sornoza M. (AS 1375), in Rı´o life the iris was dark brown, and the tarsi and Ayampe, Manabı´ Province, Ecuador, (01Њ 41Ј bill were black. The stomach contained insect S, 80Њ 48Ј W; 40 m). Our small sample did not remains. The other two specimens, a sexually permit us to distinguish between First Basic immature female (AS 1374) and a juvenile fe- and Definitive Basic plumages in female C. male (AS 1377) were similar to AS 1374 (Ta- berlepschi. Upper tail coverts, rump, scapu- ble 2) but differed in the following ways. lars, crown, and wing coverts are Parrot Green AS 1374 has upperparts Parrot Green but (260) with a weak bronzy sheen. Green of up- with a stronger bronzy sheen, except for central perparts is darkest and dullest on the crown. rectrices and uppertail coverts. In contrast to AS Central rectrices are mostly Parrot Green but 1375, the crown is not duller than back. The fade into dusky and are finally tipped with a cheek stripe is dusky and the green patch on small, unevenly bi-lobed area of pale pinkish sides of chest is much reduced. One throat buff (7.5 YR 7/6). Tips of other rectrices are feather is Purple (1) as for male. The postocular the same pale pinkish buff. Rectrices 2–5 have stripe is white with a few pallid pinkish buff cinnamon-rufous (2.5 YR 5/8) base and black feathers (7.5 YR 8/3 to 8/4). The dusky chest (GLEY, diagram 1, 2.5/N) subterminal band. band is seemingly absent but difficult to identify Shafts of rectrices match color of surrounding because of specimen condition. The longest up- feather barbs except for proximal shaft of R1 pertail covert feathers are tipped dusky. which is black surrounded by green barbs. AS 1377 is a juvenile female with a yellow The remiges are dusky and the postocular gape. The entire upperparts (back, crown, up- stripe is white. The long and wide cheek stripe per tail coverts) appear scaly because the (auriculars and below) is Parrot Green with feathers have thin blackish subterminal bars some dusky feathers. The throat, upper breast, and cinnamon-rufous (2.5 YR 5/8) tips. The and central belly are pale pinkish buff (7.5 YR belly is uniform vinaceous pink (5 YR 5/6) 7/5) (slightly grayer than 7/6). Sides of breast without the areas of lighter buff shown by AS have patches of Parrot Green that are con- 1375. The throat is pallid pinkish buff (7.5 YR nected by a vague dusky chest band. The up- 8/3 to 8/4), paler than AS 1375. The cheek per breast is similar in color to the throat but stripe is dusky with cinnamon-rufous (2.5 YR slightly more rufescent above the chest band. 5/8) tips. The tail is similar to other two spec- Sides and lower edge of green patch on sides imens but both proximal and distal edges of of the breast are vinaceous pink (5 YR 5/6). the subterminal band are more regular; black Undertail coverts are dull cinnamon-rufous (5 is not as extensive down shaft or along sides YR 5/8). Flank feathers are white and extend of feather. The green patch on sides of breast onto sides of rump. Leg puffs are white. In and dusky chest band are seemingly absent. 236 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 121, No. 2, June 2009

The soft parts and stomach contents of AS southwest of Su´a (ϳ0Њ 50Ј N, 79Њ 53Ј W) 1374 and 1377 were similar to AS 1375. All (Ridgely and Greenfield 2001). Recent sites in specimens had little fat. Manabı´ Province are Agua Blanca (50 m; Becker et al. 2000) (Fig. 2), Rı´o Ayampe (40 DISCUSSION m; Ridgely and Greenfield 2001), and nearby We present the first description of female Cinco Cerros (1Њ 41Ј S, 80Њ 46Ј W; 151 m; C. Chaetocercus berlepschi to facilitate the cor- D. Becker, pers. comm.). Reports further south rect identification of the gender which will be in Santa Elena Province are from Loma Alta important in future studies of the species. Our (400–750 m; Becker and Lo´pez Lanu´s 1997, new distributional records significantly aug- A´ greda 2007) and 7 km northeast of Loma ment the number of known localities for C. Alta, at Cerro La Culebra above Dos Mangas berlepschi and suggest the species may not be (1Њ 47Ј S, 80Њ 37Ј W; 500–600 m; A´ greda as threatened as previously thought. We iden- 2007). tify an important breeding zone which ad- We regard the report of the species from vances our understanding of the breeding bi- Isla de la Plata (1Њ 16Ј S, 81Њ 04Ј W; 50 m; ology of C. berlepschi and should enable Manabı´ Province; Becker et al. 2000) as un- more effective conservation of the species. certain. The island is more arid than any other C. berlepschi was known from 10 published site from which the species is known; the and two previously unpublished specimens from was viewed from 25 m and might have been western Ecuador before this study, only 10 of confused with Short-tailed Woodstar (Myrmia which have certain locality data (Collar et al. micrura), a common species on the island. 1992, Ridgely and Greenfield 2001). Three Four weeks of avian sampling (Cisneros-He- males and three ‘‘females’’ (AMNH 118529-34) redia 2005) and frequent visits by scientists were collected in October to December 1912 at and birdwatchers have not produced any fur- Esmeraldas (ϳ0Њ 58Ј N, 79Њ 39Ј W; Esmeraldas ther records for this area. Province) (Fig. 1). Two males (AMNH 119766 The distributional records we present con- and 119768) were taken in December 1912 at firm the importance of the southern Manabı´ Chone (ϳ0Њ 42Ј S, 80Њ 05Ј W; Manabı´ Prov- Province coastal zone for C. berlepschi. Our ince). A previously unpublished specimen of a data, along with previous observations (Ridgely male (MCNC-MZB 85-0542) was collected in and Greenfield 2001; D. M. Brinkhuizen, pers. September 1934 at San Mateo (ϳ0Њ 54Ј N, 79Њ comm.), indicate that C. berlepschi is present 37Ј W; Esmeraldas Province). Among the spec- in the Ayampe area from mid-October to late- imens collected at uncertain localities are a ju- May. Future research should seek to identify venile male (AMNH 37925), probably collected which areas and habitats are used by C. ber- in the late 19th century and supposedly from lepschi during the non-breeding season from ‘‘Rı´o Napo, eastern Ecuador’’, and a previously June to September. The lack of records from unpublished juvenile male (MIZPAN 44432) our non-breeding season searches in the north- collected in ‘‘Ecuador’’. One adult male speci- ern Cordillera Chongo´n-Colonche, combined men mentioned by Collar et al. (1992), labeled with the dearth of sightings from Ayampe as C. berlepschi, is at the Muse´um d’Histoire (Ridgely and Greenfield 2001), Loma Alta (C. Naturelle, Paris, France (MNHN CG2003-17), D. Becker, pers. comm.), and San Sebastia´n but BPF identified the skin to be C. jourdanii (R. S. Ridgely, pers. comm.) suggest that C. in 2002. After the MCNC-MZB specimen, the berlepschi leaves the Cordillera during this species went unrecorded for nearly 60 years un- season. We hypothesize the species migrates til an adult male was collected (ANSP 183118) to Esmeraldas Province for the non-breeding in January 1991 in the hills south of the lower period. The only two non-breeding records of Rı´o Ayampe in what is now northwest Santa the species come from Esmeraldas: a juvenile Elena Province (ϳ1Њ 43Ј S, 80Њ 46Ј W; ϳ75 m male (MCNC-MZB 85-0542) from San Mateo elevation). in September 1934, and a male seen at Finca C. berlepschi has been reported since 1991 Paraı´so de Papagayos from 15 to 23 Septem- on single occasions from two sites in Esmer- ber 1995. We searched Esmeraldas for only 3 aldas Province: Finca Paraı´so de Papagayos, days in August, and it is likely that future north of Quininde´(ϳ0Њ 20Ј N, 79Њ 28Ј W) and sampling in the province during the non- Harris et al. • ESMERALDAS WOODSTAR DISTRIBUTION AND PLUMAGE 237 breeding season will record C. berlepschi. In- needed to identify the habitat requirements of tensive searches combined with stable isotope C. berlepschi. (Hobson et al. 2003, Fraser et al. 2008) and Two of our observation sites, San Sebastia´n mark-recapture techniques should produce the and Rı´o Blanco, and one previously published most data on movements of C. berlepschi. Ef- site, Agua Blanca (Becker et al. 2000), are fective long term management of this species within the boundaries of Machalilla National will depend upon knowledge of its non-breed- Park (Fig. 2). The park provides nominal pro- ing distribution. tection for the species but problems with reg- ulation enforcement in the park have been not- CONSERVATION IMPLICATIONS ed in the past (BirdLife International 2000). C. berlepschi was considered rare to un- We observed adverse human impacts such as common (Becker et al. 2000, BirdLife Inter- logging, hunting, and farming inside the park national 2000) and Ridgely and Greenfield during our study, and several small human set- (2001) proposed that Critically Endangered tlements are within the boundaries of the park. may be the appropriate conservation status for Significant changes in the management of the species (IUCN 2001). The large number Machalilla National Park should be imple- of observations we made suggest the species mented to help ensure the survival of C. ber- may not be as seriously threatened as previ- lepschi and numerous other range-restricted ously thought. C. berlepschi seems to be sea- species that occur there. These problems are sonally more numerous in disturbed, lower el- not limited to Machalilla National Park; a ma- evation areas where we made our observations jor priority for Ecuadorian bird conservation than in high-elevation garu´a forest at Loma is to strengthen management of the country’s Alta where it appears to be genuinely uncom- protected areas (Freile and Rodas 2008). mon (C. D. Becker, pers. comm.). Our data Currently, Machalilla National Park does suggest the global population size of the spe- not provide reliable protection for C. berlep- cies is likely to be ϳ1,000 to 3,000 individ- schi. The only other protected area where the uals, although more work is needed to confirm species is known to occur is Reserva Ecolo´- this estimate. The species is localized in a gica Loma Alta, a 3,000 ha community-owned small range that has been mostly deforested reserve that protects higher elevation moist (Dodson and Gentry 1991) and, in our view, forest and drier forest at lower elevations should still be considered Endangered. Our (Becker 1999, Becker et al. 2005). C. berlep- data suggest that C. berlepschi meets criterion schi is regularly recorded in the reserve but A4 (suspected population reduction of Ն50% no evidence of breeding has been observed. over a 10-year period) and possibly C1 (pop- The establishment of a new protected area in ulation size Ͻ2,500 mature individuals with the breeding habitat of C. berlepschi would an estimated continuing decline) (IUCN help ensure the long-term persistence of the 2001). C. berlepschi is a high priority species species. This protected area should ideally be that should receive conservation action as placed along the coast somewhere between soon as possible. San Jose´ and Salango, and should include as Our sampling area was limited to within 20 much intact forest and as many riparian cor- km of the coast; the large number of nests we ridors as possible. encountered suggests this coastal zone from 0 The description error corrected by this to 700 m elevation from San Jose´ (Santa study emphasizes there is still much to be Elena Province) north to San Lorenzo (Man- learned about the avifauna of northern South abı´ Province) is an important breeding area for America. Our findings underscore the need for C. berlepschi. Future conservation actions for more fundamental natural history research on this species should prioritize habitat protection the lesser known birds of this extremely di- in this coastal area. That C. berlepschi indi- verse continent. viduals and nests were found at least adjacent to large patches of forest suggests the species ACKNOWLEDGMENTS may require large areas of intact forest, even We are grateful to the American Bird Conservancy though the species is a seasonal resident in (William Belton conservation grant) and the World partially disturbed areas. More research is Land Trust for funding. We thank the executive direc- 238 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 121, No. 2, June 2009 tor and board of Fundacio´n Jocotoco for placing con- versity and conservation in Tumbesian Ecuador tinued emphasis on this project, and for providing and Peru. BirdLife International, Cambridge, funding and logistical support throughout. We are in- United Kingdom. debted to Aldo Sornoza M. for his efforts in collecting BIRDLIFE INTERNATIONAL. 2000. Threatened birds of the specimens. AEA´ thanks the Ministerio del Am- the world. Lynx Edicions and BirdLife Interna- biente for issuing national investigation and collection tional, Barcelona, Spain and Cambridge, United permits. Specimen collection was supported by sources Kingdom. separate from the American Bird Conservancy and BIRDLIFE INTERNATIONAL. 2008a. Species factsheet: World Land Trust funds. This project depended on the Chaetocercus berlepschi. BirdLife International, hard work of our field assistants: Matt Brady, B. R. Cambridge, United Kingdom. www.birdlife.org/ Hickman, J. D. Ritterson, and R. S. Terrill, J. G. Loor, datazone/ (accessed 9 October 2008). P. A. Parraga, Nildo Parrales, and Francisco Sornoza BIRDLIFE INTERNATIONAL. 2008b. Species factsheet: M. also provided valuable assistance in the field. We Chaetocercus bombus. BirdLife International, thank Matt Brady, B. E. Lyon, and R. S. Terrill for Cambridge, United Kingdom. www.birdlife.org/ discussions on the Ayampe woodstar situation. We are datazone/ (accessed 9 October 2008). grateful to P. J. Greenfield for work on the magnificent CISNEROS-HEREDIA, D. F. 2005. La avifauna de la isla frontispiece, and Scott Panciera for work on figure 3. de la Plata, Parque Nacional Machalilla, con notas Niels Krabbe provided valuable assistance with the sobre nuevos registros. Cotinga 24:22–27. color description of the specimens. We are grateful to CLEMENTS,J.F.AND N. SHANY. 2001. A field guide to the following individuals for their generous help at the the birds of Peru. Ibis Publishing Company, Te- museums: Peter Capainolo, M. E. Hart and P. R. Sweet mecula, California, USA. at AMNH, N. H. Rice at ANSP, Albert Burgas R. and COLLAR, N. J., L. P. GONZAGA,N.KRABBE,A.MAD- E. G. Franquesa at MCNC-MZB, Dominika Mierzwa RON˜ O NIETO,L.G.NARANJO,T.A.PARKER III, AND at MIZPAN, and P.-Y. Henry and Eric Pasquet at D. C. WEGE. 1992. Threatened birds of the Amer- MNHN. David Agro, A. A. Chenault, G. R. Graves, P. icas: the ICBP/IUCN Red Data Book. Internation- J. Greenfield, J. M. Harris, Salvador Herrando-P., J. L. al Council for Bird Preservation, Cambridge, Reid, R. S. Ridgely, B. R. Scheffers, K.-L. Schuch- United Kingdom. mann, N. S. Sodhi, L. W. Traill, and T. C. Wanger CUERVO, A. M., C. D. CADENA, AND J. L. PARRA. 2006. provided helpful comments on the manuscript. We also Seguir colectando aves en Colombia es imprescin- thank C. D. Becker, D. M. Brinkhuizen, D. F.Cisneros- dible: un llamado a fortalecer las colecciones or- H., M. P. L. Fogden, Johan Ingels, R. L. Mumme, Peter nitolo´gicas. Ornitologı´a Colombiana 4:51–58. Pyle, H. V. M. Stein, S. M. Wethington, and the Fun- DICKINSON, E. C. (Editor). 2003. The Howard and dacio´n Jocotoco office staff for assistance. AEA´ is Moore complete checklist of the birds of the grateful to the Royal Society for the Protection of world. Third Edition. Princeton University Press, Birds and the Percy Sladen Fund for financial support, Princeton, New Jersey, USA. and Aves & Conservacio´n for institutional and logis- DODSON,C.H.AND A. H. GENTRY. 1991. Biological tical support. BPF thanks the German National Merit extinction in western Ecuador. Annals of the Mis- Foundation for financial support. souri Botanical Garden 78:273–295. FRASER, K. C., T. K. KYSER, AND L. M. RATCLIFFE. LITERATURE CITED 2008. Detecting altitudinal migration events in A´ GREDA, A. E. 2007. Feeding ecology and conserva- neotropical birds using stable isotopes. Biotropica tion of Esmeraldas Woodstar Chaetocercus ber- 40:269–272. lepschi in the Chongo´n-Colonche Hills, western FREILE,J.AND F. R ODAS. 2008. Conservacio´n de aves Ecuador. Cotinga 27:38–41. en Ecuador: ¿co´mo estamos y que´ necesitamos BECKER, C. D. 1999. Protecting a garu´a forest in Ec- hacer? Cotinga 29:48–55. uador: the role of institutions and ecosystem val- FREYMANN,B.AND K.-L. SCHUCHMANN. 2005. Collect- uation. Ambio 28:156–161. ing history of the hummingbird genera Chaeto- BECKER,C.D.AND B. LO´ PEZ LANU´ S. 1997. Conser- cercus Gray 1855 and Lophornis Lesson 1829. vation value of a Garua forest in the dry season: Journal of Ornithology 146:61–64. a bird survey in Reserva Ecologica de Loma Alta, GRAVES, G. R. 1986. Systematics of the gorgeted Ecuador. Cotinga 8:66–73. woodstars (Aves: Trochilidae: Acestrura). Pro- BECKER, D., A. A´ GREDA,A.RICHTER, AND O. RODRI- ceedings of the Biological Society of Washington GUEZ. 2000. Interesting bird records from the Co- 99:218–224. lonche Hills, western Ecuador. Cotinga 13:55–58. GURNEY, M. 2006. Identification of Little Woodstar BECKER, C. D., A. A´ GREDA,E.ASTUDILLO,M.CON- Chaetocercus bombus in Ecuador. Neotropical STANTINO, AND P. T ORRES. 2005. Community-based Birding 1:38–41. monitoring of fog capture and biodiversity at HILTY, S. L. 2003. Birds of Venezuela. Princeton Uni- Loma Alta, Ecuador enhance social capital and versity Press, Princeton, New Jersey, USA. institutional cooperation. Biological Conservation HILTY,S.L.AND W. L. BROWN. 1986. A guide to the 14:2695–2707. birds of Colombia. Princeton University Press, BEST,B.J.AND M. KESSLER (Editors). 1995. Biodi- Princeton, New Jersey, USA. Harris et al. • ESMERALDAS WOODSTAR DISTRIBUTION AND PLUMAGE 239

HOBSON, K. A., L. I. WASSENAAR,B.MILA´ ,I.LOVETTE, RESTALL, R., C. RODNER, AND M. LENTINO. 2006. Birds C. DINGLE, AND T. B. SMITH. 2003. Stable isotopes of northern South America: an identification as indicators of altitudinal distributions and move- guide. Yale University Press, New Haven, Con- ments in an Ecuadorean hummingbird communi- necticut, USA. ty. Oecologia 136:302–308. RIDGELY,R.S.AND P. J. GREENFIELD. 2001. The birds INTERNATIONAL UNION FOR CONSERVATION OF NATURE of Ecuador. Cornell University Press, Ithaca, New (IUCN). 2001. IUCN Red List categories and cri- York, USA. teria. Version 3.1. IUCN Species Survival Com- SCHUCHMANN, K.-L. 1999. Family Trochilidae (hum- mission. IUCN, Gland, Switzerland and Cam- mingbirds). Pages 468–680 in Handbook of the bridge, United Kingdom. www.iucnredlist.org (ac- birds of the world. Volume 5 (Barn Owls to hum- cessed 8 November 2008). mingbirds) (J. del Hoyo, A. Elliot, and J. Sargatal, MATA,J.R.R.,F.ERIZE, AND M. RUMBOLL. 2006. Birds Editors). Lynx Edicions, Barcelona, Spain. of South America, nonpasserines: rheas to wood- SCHULENBERG, T. S., D. S. STOTZ,D.F.LANE,J.P. peckers. Princeton University Press, Princeton, O’NEILL, AND T. A . P ARKER III. 2007. Birds of New Jersey, USA. Peru. Princeton University Press, Princeton, New MEYER DE SCHAUENSEE, R. 1970. A guide to the birds Jersey, USA. of South America. Academy of Natural Sciences, SIMON, E. 1889. Description d’une espe`ce nouvelle du Philadelphia, Pennsylvania, USA. genre Chætocercus.Me´moires de la Socie´te´ Zool- MUNSELL COLOR. 1994. Munsell soil color charts. Mac- ogique de France 2:230–231. beth Division of Kollmorgan Instruments Corpo- SMITHE, F. B. 1975. Naturalist’s color guide. American ration, New Windsor, New York, USA. Museum of Natural History, New York, USA. PETERSON,A.T.,A.G.NAVARRO-SIGU¨ ENZA, AND H. STATSOFT INC. 2003. STATISTICA. Version 6.1. BENI´TEZ-DI´AZ. 1998. The need for continued sci- StatSoft Inc., Tulsa, Oklahoma, USA. entific collecting; a geographic analysis of Mexi- VUILLEUMIER, F. 1998. The need to collect birds in the can bird specimens. Ibis 140:288–294. Neotropics. Ornitologı´a Neotropical 9:201–203. REMSEN JR., J. V. 1995. The importance of continued VUILLEUMIER, F. 2000. Response: further collecting of collecting of bird specimens to ornithology and birds in the Neotropics is still needed. Ornitologı´a bird conservation. Bird Conservation International Neotropical 11:269–274. 5:145–180. WINKER, K. 1996. The crumbling infrastructure of bio- REMSEN JR., J. V. 1997. Museum specimens: science, diversity: the avian example. Conservation Biol- conservation and morality. Bird Conservation In- ogy 10:703–707. ternational 7:363–366. WINKER, K. 2005a. Bird collections: development and REMSEN JR.,J.V.,C.D.CADENA,A.JARAMILLO,M. use of a scientific resource. Auk 122:966–971. NORES,J.F.PACHECO,M.B.ROBBINS,T.S.SCHU- WINKER, K. 2005b. Natural history museums in a post- LENBERG,F.G.STILES,D.F.STOTZ, AND K. J. ZIM- biodiversity era. BioScience 54:455–459. MER. 2008. A classification of the bird species of WINKER, K., B. A. FALL,J.T.KLICKA,D.F.PARMELEE, South America. American Ornithologists’ Union. AND H. B. TORDOFF. 1991. The importance of avi- www.museum.lsu.edu/ϳRemsen/SACCBaseline. an collections and the need for continued collect- html (accessed 1 April 2008). ing. The Loon 63:238–246.