Parotia berlepschi, Kleinschmidt, 1897, Kleinschmidt, 1897

Laman, Timothy G., 2017, Taxonomic status of Parotia berlepschi Kleinschmidt, 1897 based on analysis of external appearance, voice and behavior (Aves: Paradisaeidae), Zootaxa 4329 (6), pp. 560-573 : 561-569

Methods

Historical Review. We surveyed the literature, all known specimens from relevant ornithological collections, and the broader historical record in order to review the 121-year taxonomic history of the discovery of P. berlepschi , its long period of prolonged taxonomic uncertainty, and its discovery in the wild.

External Appearance. We examined specimens and/or photographs of specimens from the American Museum of Natural History in New York ( AMNH), Museum Zoologicum Bogoriense in Bogor, Indonesia ( MZB), Senckenberg Museum at Frankfurt-on-Main in Germany ( SMF) and Naturhistorisches Museum Wein in Vienna ( NHMW). ES examined and provided photographs of the three AMNH specimens. BB and Chris Milensky of the Smithsonian Institution (SI) examined and photographed the MZB specimens in Indonesia and in Washington, DC while on loan to SI. Curator Gerald Meyer provided photographs of three SMF specimens. Szabolcs Kókay examined and provided photographs of one specimen from NHMW. Photographs of free living and handheld P. berlepschi came from TL and BB. TL photographs of wild birds are archived in the Macaulay Library at the Cornell Lab of Ornithology and can be accessed online directly by appending desired catalog number to the URL: macaulaylibrary.org/image/catalog# (e.g., macaulaylibrary.org/image/48065581). TL provided photographs of wild P. carolae used for comparisons of living birds, which are similarly archived for reference. ES made all video recordings. BB and ES made sound recordings. Like photographs, video and sound recordings used for analysis are archived in the Macaulay Library at the Cornell Lab of Ornithology and can be accessed online at macaulaylibrary.org/video/catalog# for videos or macaulaylibrary.org/audio/catalog# for sounds.

Plumage and morphological comparisons were made between Parotia carolae and P. berlepschi . Because living individuals of nominate P. carolae from the Weyland Mountains of western New Guinea have not been documented, comparisons of living birds are made using images from within P. carolae complex, which (excluding P. berlepschi ) includes four named subspecies. Unless specifically noted, the characters used herein are not known to be variable within the P. carolae complex. Thus, when we compare P. berlepschi to P. carolae for the most part and unless explicitly stated, this comparison applies to the entire P. carolae complex since the characters that vary between P. berlepschi and P. carolae are not significantly variable within the P. carolae complex.

Field Observations. Observations, photographs, video and sound recordings of wild birds were collected as possible over the course of three expeditions to a field site known informally as “ Bog Camp ” in an upland forest clearing in the western half of the Foja Mountains (2° 34.363'S, 138° 43.032'E) . Expeditions took place in 2005, 2007, and 2008. The 2005 expedition was attended by BB and took place from 15 Nov–9 Dec. The 2007 expedition was attended by BB and TL and took place from 14–25 June. The final expedition to Bog Camp in 2008 was attended by ES and TL and took place from 27 Oct–26 Nov. During each of these expeditions, time spent observing P. berlepschi varied, but the most prolonged effort was during 2008. In all, two display courts were observed, one very near the Bog Camp clearing by BB in 2005 and one along the flank of a ridge to the south of Bog Camp by ES and TL in 2008.

Vocal and Behavioral Analysis. Behavioral observations, photographs and audio and video recordings were made by BB, TL, and ES. Videos of P. carolae in the wild were recorded by ES. Sound analysis was done using Raven Pro version 1.5 (Cornell Lab of Ornithology, Ithaca, NY, USA). Audio recordings examined are MLNS audio catalog numbers: 139542, 139631, 139538, 139536 and 163704. Video specimens examined are MLNS video catalog numbers: 457925 and 457926.

Historical Review

In December 1895, the private ornithological collection of Count Hans Karl Hermann Ludwig Graf von Berlepsch of Germany obtained two bird-of-paradise specimens from the genus Parotia ( Kleinschmidt 1897b) . Both were male, one juvenile and one adult, and they were thought to be a new form with close affinity to Parotia carolae ( Kleinschmidt 1897b) . The specimens were trade-skins of unknown provenance obtained from the Van Duivenbode family, Ternate-based traders and plume merchants ( Steinheimer 2005). Details are lacking, but the specimens likely came from the area of western New Guinea where the Van Duivenbodes conducted regular business with native hunters and traders ( Rothschild & Hartert 1903).

Otto Kleinschmidt described a new species from the two Berlepsch specimens, which he christened Parotia berlepschi in honor of Count von Berlepsch ( Kleinschmidt 1897a, b). Kleinschmidt argued that P. berlepschi was distinct from P. carolae in seven characters: (1) shorter supranarial tufts (sensu Scholes 2006), (2) loral tufts (sensu Scholes 2006) that lack white tips and do not curve inward, (3) a thicker and more curved bill, (4) black throat and cheeks, (5) smaller and darker eye-ring, (6) darker crown, and (7) a golden-brown (bronze) sheen on the hind neck ( Kleinschmidt 1897b).

In 1916, the two specimens examined by Kleinschmidt became incorporated into the Senckenberg Museum at Frankfurt-on-Main (SMF) when the Berlepsch collection was purchased from the Count’s widow during World War I ( Naumberg 1931). Although Frith and Beehler (1998) were unable to locate either of the P. berlepschi syntypes and suggested they may have been destroyed during World War II, they are in fact part of the SMF ornithological collections as noted by Steinheimer (2005) and confirmed by curator Gerald Mayr (pers. comm., Feb 2010). The juvenile specimen is SMF 64277 and the adult is SMF 64276.

Since Kleinschmidt did not fix a holotype and because no lectotype has been fixed subsequently, we hereby designate SMF 64276 as the lectotype according to Article 74 of the ICZN. In accordance with ICZN Article 74.1.3, SMF 64277 becomes a paralectotype.

On 16 December 1908 at a meeting of the British Ornithologists Club, Walter Rothschild exhibited a skin of an adult male P. berlepschi obtained for his collection at Tring ( Rothschild 1908). This specimen, along with two juvenile male specimens obtained previously by Rothschild (Rothschild & Hartert 1903) increased the worldwide specimen count at that time to five ( Rothschild 1908). One of the three Rothschild skins, the adult male, has a note on its original tag indicating it was purchased from Dunstall. This is believed to be the London feather merchant G. K. Dunstall from whom Rothschild was known to have acquired trade-skins (e.g., Tring Museum Correspondence TM 1/12/16 from 1895 documents correspondence between Rothschild and Dunstall). It is unclear if all three Rothschild specimens came via Dunstall or if the two juvenile males obtained sometime before 1903 came from another supplier like Van Duivenbode. Like the von Berlepsch trade-skins described by Kleinschmidt, all three of the Rothschild specimens were also of unknown provenance. In 1931 the three Rothschild specimens were sold, along with the entire Rothschild ornithological collection, to the American Museum of Natural History in New York and became AMNH 678170 (juvenile), 678171 (adult), and 678172 (juvenile).

A sixth specimen, an adult male, with the name Robert de Neufville and the date 1 Dec 1910 on the tag, resides in the ornithological collections of the SMF (Gerald Mayr, pers. comm., Feb 2010). The history and provenance of the “de Neufville specimen” is unknown. To our knowledge, Robert de Neufville was not a collector working in New Guinea, and the only information that we could find about him is that he was present at the von Berlepsch estate in 1916 to assist with the packing of specimens for relocation to SMF ( Naumberg 1931).

In 2013, while examining bird-of-paradise specimens in the natural history museum of Vienna (Naturhistorisches Museum Wein, NHMW), wildlife artist and illustrator Szabolcs Kókay uncovered another previously unrecognized P. berlepschi specimen (Kókay pers. comm., Jul 2013). It is an adult male dated 1894/95, with locality given only as “New Guinea.” The skin is registered as NHMW 12873 and is therefore the seventh historical P. berlepschi specimen in existence. It was likely acquired from a plume-trader (quite possibly the same one) at the time the von Berlepsch trade-skins were obtained.

In 1979 and 1981, Jared Diamond became the first ornithologist to explore the higher elevations of the uninhabited Foja Mountains ( Diamond 1982, 1985). The most celebrated finding from his time in the Foja was his discovery of the much sought-after bowerbird Amblyornis flavifrons Rothschild, 1895 , which was in a similar state of taxonomic uncertainty as P. berlepschi —i.e. known only from a few enigmatic specimens of unknown geographic origins obtained around the same time as the P. berlepschi skins ( Diamond 1982). Diamond encountered several female-plumaged birds-of-paradise from the genus Parotia , which he identified as belonging to the P. carolae group ( Diamond 1985). This observation, combined with the verified discovery of Amblyornis flavifrons , led Diamond to hypothesize that the Foja Mountains were also the geographic home of P. berlepschi ( Diamond 1985) .

Diamond’s hypothesis was consistent with previous speculations based on the time period when the A. flavifrons and P. berlepschi trade-skins were obtained and the same Dutch trade-merchant sources for both ( Mayr 1941). However, with only female-plumaged birds seen, conclusive evidence would await further observations.

A quarter century after Diamond, in 2005, a multi-national survey team conducted the first intensive biodiversity survey of the Foja region ( Beehler et al. 2007). This expedition yielded several major discoveries, including a new endemic species of wattled honeyeater from the genus Melipotes , and confirmation that the Foja Mountains are the home of P. berlepschi when several adult males were observed and two specimens, a male and female, were collected near an upland forest clearing ( Beehler et al. 2007). Follow-up surveys were conducted in 2007 and 2008, when the first detailed observations and audiovisual recordings of living P. berlepschi were made.

Male External Appearance

The external phenotype of adult male P. berlepschi is differentiated from P. carolae in the following six characters: (1) Frontal crest ( Fig. 1 , A). In P. berlepschi , the anterior-most loral tufts ( Fig.1, A1 ) of the frontal crest are entirely black and without white tips and do not curve strongly inward over the upper forehead and crown. As a result, the bulbous forehead feathering of P. berlepschi appears conspicuously velvet black at all times ( Fig. 2A–C and videos ML 457925 and 457926), especially when viewed head-on (e.g., photo ML 4813073). From the front, the stark black of the frontal crest contrasts with the coppery-bronze facial markings and nape (e.g., photo ML 48130731). Only the posterior-most loral feathers, those above the eyes, are tipped white (figs. 2C, 3C), which means that the only white visible on the head of P. berlepschi under most conditions in life (i.e. not in specimens) is a tiny patch on the crown above the eyes ( Fig. 2C ).

In contrast, P. carolae has a prominent white line that spans the entire forehead from culmen to fore-crown (figs. 2E–H; 3D, F). The line is created by the silver-white tips of the elongate loral tufts, which curve inwards from each side of the head to touch along the midline of the forehead ( Fig. 3F ).

Although described as such by Kleinschmidt, evidence of supranarial tufts (figs. 1, A2) being shorter than in P. carolae is inconclusive. The position of these feathers on the head make them susceptible to variation based on how a specimen is prepared (i.e. most specimens are prepared with the frontal crest “open”, e.g., figs. 3C, F), which makes it difficult to obtain consistent measurements. However, the proportion of white on the feather tips relative to the black base is smaller for P. berlepschi and, while difficult to quantify, the total area of the forehead covered by supranarial tufts seems to be less in P. berlepschi than P. carolae .

(2) Crown ( Fig. 1 , B). P. berlepschi , like P. carolae , has a highly reflective crown patch, which is made from the coppery-bronze forehead tufts ( Fig.1 , A 3 ), which lie folded along the top of the head and surrounding crown feathers with coppery-bronze tips ( Fig. 3C ). In life, the coppery-bronze crown patch of P. berlepschi is almost entirely concealed beneath the black crown and loral feathers ( Fig. 2C ) and is typically not visible unless purposely revealed when the frontal crest is spread (e.g., see video ML 457925 and photos ML 48066221 and ML48066211). In specimens, the loral feathers are relaxed and spread away from the crown and so that the patch appears more prominent in skins than it does in life ( Fig. 3C ).

In contrast, a sizable patch of the bronze-gold forehead tufts ( Fig. 1 , A 3 ) of P. carolae is prominently visible on the crown at all times (i.e. even without intentional feather movement) so that the top of the head appears to have a bronzy-gold patch on the crown ( Fig. 2G ). As with P. berlepschi , this patch appears more prominent in specimens because the loral feathers are relaxed and the frontal crest is “open” as in Fig. 3F .

(3) Facial markings. In P. carolae there is reflective bronzy-gold facial feathering that runs from the forecrown in front of the eyes onto the face to form a prominent bronzy-gold marking in front of each eye, which often appears crescent-shaped (figs. 2E, 3D). This marking is connected to an eye-ring of similar color, which can be difficult to discern under certain lighting conditions and viewing angles (traces of it are visible in figs. 2E, F, 3D).

In P. berlespschi , the there is no marking in front of the eye (figs. 2A, B, 3A). The stripe from the crown to eye is much broader and more conspicuous above the eye ( Fig. 2B, C ; videos ML 457925, 457926). This stripe forms a coppery-bronze “tiara” that spans the crown from eye to eye such that the silver-white crown patch appears as the centerpiece of the “tiara” ( Fig. 2C , but see also videos ML 457925, 4579256 and photo ML 48065351). The eyering in P. berlepschi is so dark as to be almost black and lacks the reflective quality of P. carolae . The stripe of P. berlepschi continues below the eye and onto the malar region so that the entire facial marking sometimes appears as a coppery-bronze eye-stripe running from crown to malar (e.g., figs. 2B, 3A). The malar stripe is also present in P. carolae (slightly visible in Fig. 2F ), but it is much less obvious (to the point of being indeterminate at times) because it lies adjacent to the less-contrasting golden-buff malar/throat, as opposed to the dark brown malar area in P. berlepschi (see below).

(4) Malar, cheeks, chin and throat. In P. berlepschi , the malar and cheek area is black, while the chin and throat, including the “whiskers” ( Fig. 1 , C) are dark-brown and the shorter feathers on the chin and throat are black ( Fig. 3A, B ). The gular and throat feathers of P. berlepschi are predominately rusty-brown with only the very slightest hint of white where the throat joins the upper breast ( Fig. 3B ).

Taxa in the P. carolae complex exhibit variation in the color of malar, cheek, chin, and throat feathering. However, in most members of the P. carolae complex, including P. carolae carolae , the malar, cheeks, and throat feathers, including the elongated “whiskers” of the chin and throat are straw colored with buff highlights ( Fig. 3D, E ). The shorter “non-whisker” feathers of the chin and upper throat are dark at the base. The feathers on the gular region and lower throat/upper breast are whitish and some have rusty-brown tips ( Fig. 3E ). It should be noted that within the P. carolae complex, P. carolae meeki Rothschild, 1910 has cheek, chin, throat/whisker feathers that are dark brown to black similar to P. berlepschi . However, unlike P. berlepschi , the malar of P. c. meeki is prominently straw/buff colored and the gular and upper breast are as in P. carolae .

(5) Iris color. Iris color of P. berlepschi is pale grey-blue with a dull orange outer ring ( Fig. 2A, B ). Under certain lighting conditions, iris color appears very blue and at other times (e.g., with flash photography at close range), it appears more pale grey. In contrast, the iris color of P. carolae is lemon-yellow with a subtle burnt-orange inner ring around the edge of the pupil and a pale-yellow outer ring ( Fig. 2E–G ). There is no known variation in eye color within the P. carolae complex ( Gilliard 1969; Frith & Beehler 1998).

(6) Nape and mantle ( Fig. 1 , D). The nape and upper part of the mantle “cape” of P. berlepschi has prominent coppery-bronze sheen of similar intensity to the coppery-bronze color on the head and face and is visible at all times ( Fig. 2A–D and videos ML 457925, 457926). In P. carolae , the nape and mantle has a very slight bronzy sheen visible only at close range and under certain light and viewing angles, but in most circumstances the mantle of P. carolae appears jet black ( Fig. 2H ).

Female External Appearance

The external phenotype of female-plumaged P. berlepschi is differentiated from female-plumaged P. carolae in head and facial markings and eye color ( Fig. 4 , and photo ML 48068341). P. berlepschi has more extensive white in the middle of the forehead, which extends almost to the fore-crown ( Fig. 4A ). The brown loral patch is also more extensive with less white near the base of the upper mandible. The supercilium extends further back onto the nape such that the nape of P. berlepschi is whiter than in P. carolae . As with males, eye color of female plumaged P. berlepschi is pale grey-blue ( Fig. 4A ), which is in contrast to the yellow eye color of female plumaged P. carolae ( Fig. 4B ).

Voice

The primary advertisement vocalizations of P. berlepschi are a shrill, ascending, two-note, “ whee-dee ” ( Fig. 5A ) reminiscent of a plastic squeak toy and a similarly shrill, four-note tremulous whistle or whinny, “we-e-e-et” ( Fig. 5B ) in which the first three are ascending (audio recordings ML 139538 and ML 139631). The “ whee-dee ” vocalization is occasionally truncated to just a single note, which sounds like “whee” or “ wheep.” At other times it is modified such that the two notes descend slightly with a more abrupt beginning and end to each note so that it sounds like “chee-deep” and the “ deep” note is a slightly lower frequency.

In comparison, the primary advertisements of P. carolae from Mt. Stolle are a powerfully whistled three-note “kwoi kwoi eeng” where each “kwoi” note is sharply ascending and the final “eeng” is a pure tone ( Fig. 5C ). P. carolae vocalizations from Crater Mountain are a single frantic quavering whistle-note that sounds like, “kwa-a-aa-ng”.

When excited, e.g., upon approach to the display court, P. berlepschi males were also heard to give a harsh, raspy white noise-like scold that ranges from a single to upwards of six ‘notes’ (e.g., the single scold note in video ML 457925 at 00:48 and also audio ML 139631) and several irregular chattery squeaks and chortles. Acoustically, the scold notes are similar to the advertisement vocalizations given by the “all-black” blue-eyed Parotia species: P. lawesii , P. sefilata , P. helenae , and P. wahnesi , but not known from the P. carolae complex.

Courtship Behavior

Due to the difficulty of locating display courts, total courtside observations were few and visitation rates infrequent. In all, five courtship behaviors were documented: two non-display, but courtship-related, and three actual courtship displays. Using the terminology of Scholes (2006), for P. carolae the behaviors identified for P. berlepschi are: 1) court clearing; 2) mat construction; 3) horizontal perch pivot display; 4) hop and shake display; and 5) leaf presentation. Each is briefly described below.

Court clearing behavior is evident from the presence of cleared terrestrial display courts and from bouts of documented court clearing/maintenance behavior. This behavior involves picking up leaves and other forest debris from the court floor and tossing them beyond the periphery of the court area. Court clearing behavior is vouchered in video ML 457925 (from time 00:00–00:31).

Mat construction behavior was observed when an adult male carried a large clump of dark brown rootlets onto its display court. This behavior was also indirectly observed by the presence of loose ‘carpet’ or mat of rootlets over the surface of a portion the display court floor. Creating a mat of rootlets on court floor is a behavior previously known only from P. carolae and not other Parotia species ( Scholes 2006).

The horizontal perch pivot display was observed on the display court and from tree branches above the court and nearby. This behavior involves the male adopting a rigid horizontal body posture, head and tail held up and flank plumes flared outward from the body so that the white portion forms two semi-circular ‘disks’ that emanate from either side of the body ( Fig. 6A ). In this posture, the body is pivoted side-to-side in a way that accentuates the white flank plumes. A low intensity version of this behavior is vouchered in video ML 457925 at time 00:48– 00:57.

A low intensity and truncated instance of the hop and shake display was documented on one occasion. While brief and incomplete, it nevertheless included enough of the behavior’s identifying postures and action patterns to be diagnosable. The observed elements were: standing upright on the court floor, fluttering the throat whiskers, “opening” the frontal crest, hopping in place, dipping the upper body and giving an exaggerated ruffling of the flank plumes and ritualized shake of the upper body. No distinct hop was observed, but otherwise the behavior pattern was very similar to the hop and shake of P. carolae . In P. carolae , there are two distinct phases, the puff shake and the skirt shake ( Scholes 2006), but only the puff shake was observed in P. berlepschi . This behavior is vouchered in video ML 457925 at time 00:30–00:33.

Leaf presentation is a modifier behavior that, in P. carolae , occurs as an accessory display along with several courtship displays. It involves holding a yellow or light colored leaf in the bill. We documented a male P. berlepschi holding a yellow leaf while on the horizontal perch of the display court ( Fig. 6B ).

Additional Information

As this is the first paper to examine the natural history of P. berlepschi in depth, we provide additional relevant species information below.

Subadult Male Plumage. As with P. carolae , subadult males range from primarily female-type plumage with partial adult male feathering appearing about the head (e.g., photo ML 48065341) to nearly full adult male plumage with only small patches of female-type plumage remaining (e.g., photos ML 48068171, ML 48068161).

Weights and Measurements. The following measurements were made from the individuals collected on the 2005 survey, which became the two specimens now housed in the Museum Zoologicum Bogoriense (MZB). Adult male (MZB 30.666) weight 162 g, testis 4x 9 mm, wing chord 152 mm, wing arc 153 mm. Female (MZB 30.667) weight 128 g, ova small, wing chord 136 mm, wing arc 139 mm, tail 91.5 mm, tarsus 45 mm, bill from base 28.5 mm, bill to edge of feathering 18 mm. An additional bird in female plumage that was netted, measured and released, had weight 139 mm, wing chord 138 mm, wing arc 142 mm, iris pale gray two-toned, mouth pale green.

Distribution and Habitat. Over the course of three expeditions to the Foja Mountains, which surveyed a broad range of elevations, (see Beehler et al. 2007, Beehler et al. 2012), P. berlepschi was found to be an uncommon species documented only from a tiny area of mid-montane forest between roughly 1,300–1,700 m.

Food and Feeding. No foraging observations were made, but presumably diet is similar to other Parotia species and includes a range of drupe and capsular fruits as well as the occasional arthropod.

Breeding. Little is known beyond the observation that males attend display courts in late November and early December in 2005 and 2008. In June of 2007, males were vocal, but were not observed attending courts although vocal activity was consistent with expectations for when courts are being actively maintained in the early part of breeding season.

Figures

FIGURE 1. Parotia-specific plumage ornaments referenced herein: (A) frontal crest, (A 1) loral tufts, (A 2) supranarial tufts and (A 3) forehead tufts, (B) crown patch, (C) throat “ whiskers ”, and (D) mantle cape. Modified from Scholes (2006), which was modified from Frith & Beehler (1998) and Schodde & McKean (1973).

FIGURE 2. Forehead, facial, crown and nape appearance of living P. berlepschi (A – D) as compared to living P. carolae (E – H). Notable characters that differentiate P. berlepschi from P. carolae include the all-black loral feathering on the frontal crest of P. berlepschi (A – C) versus the extensively white-tipped loral feathering of P. carolae (E – H); the eye-to-crown “ tiara ” of P. berlepschi (B, C) versus the more extensive coppery facial markings in front of and surrounding the eye of P. carolae; the darker malar, cheek and throat of P. berlepschi (A, B) versus the much lighter straw colored appearance of P. carolae (F); the almost nonexistent coppery-bronze hind crown patch of P. berlepschi (C) versus the extensive one visible at all times on the crown of P. carolae (G); the grey-blue eye color of P. berlepschi (A, B) versus the bright lemon yellow of P. carolae (E, F) and the rich bronzy sheen on the nape and upper mantle of P. berlepschi (D) compared to the black nape of P. carolae (H). Plates derived from photos ML 48065581 (A), 48065591 (B), 48065601 (C), 48065621 (D), 48068971 (E), 48068791 (F), 48068861 (G) and 48152301 (H).

FIGURE 3. Forehead, facial, and crown appearance of P. berlepschi specimen MZB 30.666 (A – C) compared to P. carolae specimen AMNH 302376. Notable characters differentiating P. berlepschi from P. carolae include the all black loral feathering on the frontal crest of P. berlepschi (A, C) versus the extensively white-tipped loral feathering of P. carolae (D, F); the narrow coppery-bronze stripe from above the eye to the crown of P. berlepschi (A) versus the more extensive, crescent shaped, coppery facial markings in front of the eye of P. carolae; the darker malar, cheek and throat, including the “ whiskers ” of P. berlepschi (B) compared to the straw-colored appearance of the same feathering in P. carolae (D, E). Note the appearance of the frontal crest in the “ open ” position (C, F).

FIGURE 4. Female facial markings and eye color of (A) P. berlepschi and (B) P. carolae. The forehead of P. berlepschi has more extensive white in the middle, which extends almost to the fore-crown; the brown loral patch is also more extensive with less white near the base of the upper mandible; and the supercilium extends further back onto the nape and is whiter than in P. carolae. Eye color of female-plumaged P. berlepschi is pale grey-blue (A) versus yellow in P. carolae (B). Note: the difference in tint between the two photos is largely due to differing white balance and flash settings, which make the plumage of the P. carolae image look more “ warm ” (orange) than the P. berlepschi image which is shifted slightly toward the “ cooler ” (blue) end. Image (A) is of a bird in the hand taken by BB. Image (B) is modified from photo ML 48068781.

FIGURE 5. Spectrograms of advertisement vocalizations of P. berlepschi (A, B) compared with those from two geographically varying populations within the P. carolae group (C, D). P. berlepschi has two primary advertisement vocalizations: (A) a shrill two-note form reminiscent of a squeaky toy and (B) a shrill four-note, slightly ascending whinny. In comparison, P. carolae from Mt. Stolle (C) gives a powerfully whistled three-note advertisement vocalization, and P. carolae from Crater Mountain (D) gives a single frantic quavering whistle-note. Spectrograms C and D are modified from Scholes (2006).

FIGURE 6. Two courtship behaviors documented in P. berlepschi: (A) horizontal perch pivot, which involves the male adopting a rigid horizontal body posture, head and tail held up and flank plumes flared outward from the body so that the white portions form two semi-circular ‘ disks’ that emanate from either side of the body while the body is pivoted side-to-side in a way that accentuates the white plumes; (B) leaf presentation, which is a modifier display that involves holding a yellow or light colored leaf in the bill while performing other displays. Derived from photos ML 48065351 and 48068531.