- Order: Galliformes
- Family: Cracidae
Penelopina is a monotypic genus. Highland Guan is pheasant-sized and sexually dimorphic. Females are on average larger than males. The adult male is glossy black with red legs, bill, and wattle. The female is brown barred black, with gray bill and dull reddish legs. The male’s display call and wing rattling are unmistakable. Warning calls are distinctive.
The altitudinal range of Highland Guan extends from 700 to 3300 m, where Crested Guan (Penelope purpurascens) is sympatric in the area below 2000 m, Plain Chachalaca (Ortalis vetula) is sympatric on the Atlantic slope up to 2000 m, and White-bellied Chachalaca (Ortalis leucogastra) on the Pacific slope up to 900 m. West Mexican Chachalaca (Ortalis poliocephala) is sympatric in the westernmost part of Highland Guan’s range. The range of Highland Guan overlaps also with Horned Guan (Oreophasis derbianus) in areas above 2000 m, and with Great Curassow (Crax rubra) below 1600 m. Highland Guan is rather shy and often only seen running away, therefore it can be confused with all of the above mentioned cracids, with Crested Guan and Plain Chachalaca being the most similar species. Howell and Webb (1995) pointed out that based on a poor illustration of Crested Guan in Peterson and Chalif (1973), this species was frequently misidentified as Highland Guan in Mexico. The ground-dwelling sympatric Thicket Tinamou (Crypturellus cinnamomeus), Slaty-breasted Tinamou (C. boucardi), and Buffy-crowned Wood-Partridge (Dendrortyx leucophrys) have red legs similar to Highland Guan, but are shorter tailed. Under good observation conditions, however, Highland Guan is unmistakable: the male is the only all black cracid in its range, further distinguished by its bright red tarsi, and the uniformly barred plumage of the female also is unique among sympatric cracids. Flushing birds usually utter a distinctive warning call.
For the following account on Highland Guan vocalizations, recordings were analyzed using software Raven Pro. Notes were marked in sonograms with the curser and following data were measured: peak frequency (frequency with the highest power), delta time, and center time. All means are given with standard deviation.
Male whistle: The most obvious sounds of Highland Guan are produced by the male. Mainly during the first part of the breeding season (January-April), males utter single, loud and far-carrying whistles which can be heard over more than a kilometer. This call is usually given while perched in the upper forest canopy, midstory, or sometimes on ground (Pullen 1983, González-García et al. 2001; K. Eisermann, personal observations), but it has also been reported during flight (Rowley 1984). A male’s whistle often provokes other males to vocalize (K. Eisermann, personal observations). Often the whistle precedes a wing-rattling which is produced during glides between perches. The whistle has been described as “long whistle, steady in volume but rising in pitch” (Land 1970). Based on recordings and sonograms of 5 individuals in the Guatemalan highlands, whistles ranged in length 2.3-3.1 s (mean: 2.6 ± 0.3, n = 6 calls of 5 individuals; K. Eisermann, personal observations). Peak frequency ranged 2062-2906 Hz (mean: 2469 ± 278, n = 6 calls of 5 individuals; K. Eisermann, personal observations). Sonogram 1 shows a typical whistle. The function of the whistle and combined wing-rattling is thought to be a territorial advertisement and a signal for sexual selection (Pullen 1983, del Hoyo and Motis 2004). Seasons of call activity may change between years. At Montaña Caquipec, Alta Verapaz, Guatemala, there was no call activity from September 1997 to February 1998, but in 1998 guans were calling until October (Eisermann 1999).
Alarm calls: In sight of potential danger, such as humans, male and female Highland Guans utter alarm calls which have been described as “sharp yelp” (Griscom 1932), “pia and noise resembling the neighing of a horse” (Pullen 1978), and “see-uh, may be run into twittering” (Howell and Webb 1995). The short see-uh notes are the low arousal alarm call. Once in a while a see-uh note runs into a twitter. Twitters are given in short intervals in a stage of high arousal. A females with chicks uttered a see-uh note each 0.5-3.0 s (mean interval between center times of see-uh notes: 0.9 ± 0.3 s, n = 266 notes) with twitterings given each 0.9-36.6 s (mean interval between center times of twitters: 15.0 ± 10.9 s, n = 15 twitters; K. Eisermann, personal observations). The length of see-uh notes ranged 0.1-0.4 s (0.22 ± 0.02 s, n = 472 notes of 2 individuals) and peak frequency ranged 375-3750 Hz (1807 ± 772 Hz, n = 472 notes of 2 individuals). The length of twitters, including the introductory see-uh note, ranged 0.5-1.1 s (0.8 ± 0.2 s, n = 21 notes of 3 individuals), and peak frequency ranged 1312-2625 Hz (2158 ± 304 Hz, n = 21 notes of 3 individuals; K. Eisermann, personal observations). Sonogram 2 shows typical see-uh and twitter calls.
Juvenile calls: About two weeks old chicks gave series of soft short whistles wi-wi-wi, while an adult female and an adult male were uttering warning calls (K. Eisermann, personal observations, 13 June 2011).
In the first part of the breeding season, males produce a sound during glides between perches, which has been pictorially described by several authors: “habit of ‘drumming’ ... emitted a sort of crushing or rushing noise, like that produced by a falling tree” (Salvin and Godman 1897-1904), “curious, whirring rattle ... sounds very much like a policeman’s rattle” (Dickey and van Rossem 1938), “loud crack, followed by a cascade of descending notes” (Land 1970), “ripping or drumming sound ... similar to the sound of a piece of cloth being forcefully torn” (Pullen 1983), “bizarre ‘crashing tree’ sound ... initially 1-2 raps, then a creaking, falling, rattling crash, ke’k’a,arrrrrrr” (Howell and Webb 1995).
The sound characters of the wing-drumming of five males in the Guatemalan highlands were as follows (K. Eisermann, personal observations). Each wing-drumming consisted of two types of introductory cracks, followed by a rattling (Sonogram 3).
The first kind of crack was a series of 1-4 notes, ranging in length 0.025-0.04 s (mean 0.035 ± 0.008 s, n = 11 notes of 5 individuals). Peak frequency of these notes ranged 422-985 Hz (mean 482 ± 170 Hz, n = 11 notes of 5 individuals). The interval between the center time of each crack ranged 0.10-0.27 s (mean 0.18 ± 0.05 s , n = 4 series of notes of 4 individuals). The interval between the center time of the last note of these cracks and the center time of the first note of the second kind of cracks was 0.3 s.
The second kind of crack was a series 2-3 notes, ranging in length 0.06-0.09 s (mean 0.08 ± 0.01 s, n = 11 notes of 5 individuals). The interval between the center time of each crack ranged 0.13-0.18 s (mean 0.16 ± 0.02 s , n = 5 series of notes of 5 individuals). Peak frequency ranged 468-2531 Hz (mean 1611 ± 600 Hz, n = 11 notes of 5 individuals). After the last note, 0.03-0.10 s passed until the beginning of the rattling.
The rattling lasted 1.2-1.7 s (mean 1.5 ± 0.2 s, n = 5 rattlings of 5 individuals). The peak frequency ranged 1172-3844 Hz (mean 1913 ± 1140 Hz, n = 5 rattlings of 5 individuals). In the field the number of introductory notes can hardly be distinguished by ear. The entire wing-drumming, from the first introductory note to the end of the rattling, lasted 2.24-2.44 s (2.33 ± 0.07 s, n = 5 wing-drummings of 5 individuals). Sonogram 3 shows a typical series wing-drumming.
The wing-drumming was thought to be of vocal nature, produced by “rapid expulsion of air from the throat (Rowley 1984), until Pullen (1983) observed the feather vibration during the rattling. The introductory notes are assumed to be generated by wing-flaps during take-off from the perch. An obvious wing-flap noise is produced during wing beats after take off and before landing (sonogram 4).
Detailed Description (appearance)
Adult male: Based on specimens from throughout the Highland Guan’s range, Vaurie (1968) described the plumage of the adult male as “pure black, with a dark blue or greenish gloss which varies individually and is highly developed on the upper parts with the exception of the rump, the lower abdomen being sooty, rather than pure black, and not glossy”.
Adult female: Based on specimens from throughout the guan’s range, Vaurie (1968) described the plumage of the adult female as “the ground color of the feathers of the upper parts varies from pale brown to dark fuscous brown on the head, and from buffy brown to ochraceous cinnamon, or pale russet on the back and wings, the ground color of the tail being more rufous than the color of the back. The feathers of the head are dark brown at the center, the dark center contrasting with the pale edges; the feathers of the back and tail are very heavily vermiculated, or barred with dark fuscous brown, the pale bars being narrower than the dark interspaces, except on the tail where the dark and pale bars are of about the same width. The bars of the tail are not invariably regular, as the dark bars are in some cases more or less wavy and regular. The under parts are paler than the upper parts and are chiefly grayish buffy brown, less rufous than the back, and are less heavily vermiculated and barred, the dark bars being paler and more irregular, especially on the upper breast which in some is more mottled than barred”.
Immature male: Vaurie (1968) described the plumage of the immature male as “blacker than that of the adult female, but is not uniformly black as in the adult male. The head is dull black, but not perfectly uniform, as the feathers are more or less edged with reddish brown. The feathers of the back and tail are chiefly black, but are interrupted by bars or heavy vermiculation which varies from dull pale chestnut to rufous cinnamon, the pale bars being more regular and broader on the upper tail coverts, and the tail being vermiculated rather than barred. The under parts are grayish buffy brown, but darker than in the adult female, and more or less regularly barred with very dark buff.”
Immature female: The plumage of the immature female “resembles that of the adult female but is much less regularly and distinctly patterned” (Vaurie 1968).
Juveniles: Chicks were described for the first time by Andrle (1967), summarized by Delacour and Amadon (1973) as follows: “A broad, black band down top of the head and hindneck, bordered by buffy stripes. The black line becomes more extensive on the back, but paler, mixed with brown. Wing coverts, quills and tail quills somewhat browner, darkest on breast, palest on throat.”
Iris: reddish brown (Blake 1977; K. Eisermann, personal observations)
Eye ring: “brownish red”, “reddish brown” (Vaurie 1968); dull reddish brown (Andrle 1967)
Bill: bright red-orange or coral red (Vaurie 1968)
Wattle: bright red-orange or coral red (Vaurie 1968)
Tarsi and toes: bright red-orange or coral red (Vaurie 1968)
Iris: reddish brown (Blake 1977; K. Eisermann, personal observations)
Eye ring: dusky (van Rossem 1934)
Bill: brown (Vaurie 1968)
Tarsi and toes: brownish red (Vaurie 1968)
Juveniles (about 1 week old)
Iris: dark, bluish gray (Andrle 1967)
Bill: black, with white tip and pale pinkish-white cere (Andrle 1967)
Tarsi and toes: purplish vinaceous (Andrle 1967)
Highland Guan is sexually dimorphic with females on average larger than males. Total length ranges 590-650 mm (Blake 1977). The following table gives data of the largest series of published measurements (Blake 1977). Previous smaller series of measurements (van Rossem 1934, Ridgway and Friedmann 1946) did not show the differences and are not included here. Mass was reported 866-914 g (del Hoyo 1994, Dunning 2008).
Males (n = 29)
Females (n = 15)
|wing (flat)||220-250 (236.2)||236-270 (247.1)|
|tail||231-290 (263.7)||253-290 (270.5)|
|tarsus||60-77 (68.8)||60-74 (67.2)|
|exposed culmen||20-28 (24.2)||23-26 (24.8)|
There is no detailed information available on molt of Highland Guan. Juvenile cracids are precocial. Remiges develop already inside the egg, which enables chicks to short flights few days after hatching (Wagner 1953). Because juvenile remiges are soon too small to carry the adult weight, they are molted for the first time in the first weeks of life, described on Plain Chachalaca (Ortalis vetula; Stresemann and Stresemann 1966, Marion 1977, Pyle 2008). Young resemble adults when still much smaller than adults (del Hoyo 1994). Young male Highland Guans resemble females, and acquire the black plumage in the second year (Wagner 1953, Vaurie 1967, del Hoyo 1994).
Two subspecies distinct from the nominate form have been described based on limited material (van Rossem 1934, Hellmayr and Conover 1942, Ridgway and Friedman 1946), resulting in three subspecies: nominate Penelopina nigra nigra in Mexico, Guatemala, and southwestern El Salvador; P. n. dickeyi in El Salvador and Honduras; and P. n. rufescens in Nicaragua. Examination of larger series revealed no geographic variation and no constancy of the subtle morphological subspecies characters (color of gloss, color of eye ring) used by van Rossem (Vaurie 1967). Therefore all subspecies now are considered synonyms of nominate Penelopina nigra (Vaurie 1967, Dickinson 2003, Clements et al. 2011).
Highland Guan belongs to the family Cracidae (curassows, guans, and chachalacas), which is placed in the order Galliformes together with the families of megapodes (Megapodiidae), turkeys (Melaegrididae), grouse (Tetraonidae), New World quails (Odontophoridae), pheasants and partridges (Phasianidae), and guineafowl (Numididae) (del Hoyo 1994). Highland Guan was originally described as Penelope niger (Fraser 1850). The monotypic genus Penelopina was established by Reichenbach (1862). Penelopina differs in some unique characters from other members of the subfamily Penelopinae, such as a distinct juvenile male plumage and females being larger than males (Vaurie 1968).
Traditionally, the family Cracidae is divided into two subfamilies, Cracinae and Penelopinae, based on morphological differences of the pelvic bone (Huxley 1868, Ridgway and Friedmann 1946, del Hoyo 1994, Dickinson 2003). Penelopina has been grouped together with the genera Ortalis, Penelope, Pipile, Aburria, Chamaepetes, and Oreophasis in the latter subfamily, and the genera Crax, Nothocrax, Mitu, and Pauxi in the Cracinae.
This division, however, has been controversial. Based on morphological characters, Sclater and Salvin (1870) divided the Cracidae in three subfamilies by establishing the additional subfamily Oreophasinae for Oreophasis. Vaurie (1968) agreed on the special position of Oreophasis and suggested three tribes Penelopini, Oreophasini, and Cracini. Vuilleumier (1965) and AOU (1998) did not recognize subfamilies.
Based on mitochondrial segments and nuclear genes, Pereira et al. (2002) identified four clades in the family Cracidae: curassows (Mitu, Pauxi, Nothocrax, and Crax), Horned Guan (Oreophasis), chachalacas (Ortalis), and guans (Aburria, Pipile, Penelope, Chamaepetes, and Penelopina). In this analysis, Penelopina was closely related to Chamaepetes. A cladistic study combining molecular (mitochondrial genes), morphological, and behavioral characters (Frank-Hoeflich et al. 2007) resulted in Penelopina as a sister group of Penelope, Pipile, and Aburria. These four genera are a sister group to Chamaepetes. These five genera are a sister group to Ortalis, and all of these genera form the subfamily Penelopinae. Oreophasis resulted as a sister lineage to the other genera of Cracinae (Nothocrax, Pauxi, Mitu, and Crax).
Eisermann, Knut. 2012. Highland Guan (Penelopina nigra), Neotropical Birds Online (T. S. Schulenberg, Editor). Ithaca: Cornell Lab of Ornithology; retrieved from Neotropical Birds Online: http://neotropical.birds.cornell.edu/portal/species/overview?p_p_spp=80391