Our analysis identified one high intensity display in both species, which we call the cape presentation display. Our analysis did not uncover substantial variation in the behaviors, postures and ornament forms among s. superba, s. addenda and s. latipennis, but did find minor variation in cadence of bouncing and snapping sounds (see below). As with the initial display activities, we use s. latipennis as the standard frame of reference for the superba complex and details on s. superba are provided in Fig. S1. As with the initial display activities above, our analysis of niedda found it to have numerous distinctive elements, which are described below.

Cape presentation display

The visual form and movements of this display comprise the quintessential features of courtship in the genus Lophorina (video vouchers: superba ML458003; niedda ML487557 and ML487562). When performed to a female, the cape presentation display is typically initiated from the posture of the pointing display—i.e., Fig. 3A superba and Figs. 3B and 4 niedda). However, if a female appears suddenly without stopping on a nearby perch on her approach, and during practice displays (no female present), the male will often transition directly into the cape presentation from any of initial display activities (e.g., in ML487557 the pointing display is omitted and the male appears to be caught off guard by the sudden arrival of the female).

The cape presentation display begins when the male perceives a female making her approach toward the log. In both superba and niedda, the cape-flapping of the pointing display (see above) increases in intensity. In niedda, the increase is greater as wings are lifted higher over the back and then quickly flicked open and shut in the exaggerated manner of superba during pointing (Figs. 5A and 5B). As in the pointing of superba, the cape feathers of niedda also flap higher such that the tips exceed the height the of the upward pointing bill (Fig. 5B). In conjunction with the exaggerated wing-flick/cape-flaps, the head is tilted back more sharply, the bill is also opened widely to reveal the bright yellow gape and the breast is thrust forward so that the feathers of breast shield ornament become fully expanded and conspicuous—its reflective surface angled upward toward the approaching female (Figs. 5A and 5B). The point at which the male presents his bright yellow gape is the best landmark for identifying of the start of the cape presentation display even though there is often a bout of the exaggerated wing-flick/cape-flaps preceding it.

Figure 5: Cape presentation display of superba and niedda. Lophorina. In both superba and niedda (A and B), cape-flapping increases in intensity. In niedda (B), the increase is greater because the wings are lifted higher over the back and then flicked open and shut in the exaggerated manner of superba. Simultaneously, the breast is thrust forward so that the breast shield becomes fully expanded and conspicuous with its reflective surface angled toward the approaching female. In superba (C), the cape is lifted into presentation position before to female arrives on the log. In L. niedda (D), wing-flick/cape-flaps continue for several more bouts after the female arrives and before the cape is put into presentation position. The appearance of the cape presentation phenotype differs dramatically between superba (E) and niedda (F). Image credit/source: (A, C and E) Edwin Scholes/ The visual form of this display is the quintessential feature of courtship in the genus. In bothand(A and B), cape-flapping increases in intensity. In(B), the increase is greater because the wings are lifted higher over the back and then flicked open and shut in the exaggerated manner of. Simultaneously, the breast is thrust forward so that the breast shield becomes fully expanded and conspicuous with its reflective surface angled toward the approaching female. In(C), the cape is lifted into presentation position before to female arrives on the log. In(D), wing-flick/cape-flaps continue for several more bouts after the female arrives and before the cape is put into presentation position. The appearance of the cape presentation phenotype differs dramatically between(E) and(F). Image credit/source: (A, C and E) Edwin Scholes/ ML458003 , (B) Tim Laman/ ML487538 (D and F) Tim Laman/ ML487557

The visual effect of the exaggerated wing-flick/cape-flaps with the breast shield fully revealed is more pronounced in niedda than superba because of the stark difference in the appearance of the breast shield during the pointing display (e.g., compare breast shield positions in Figs. 3A and 3B to those in Figs. 5A and 5B). In both species, the male will reorient himself as needed to track the position of the approaching female and keep his eye-spots and breast shield directed toward the female (e.g., video ML487530).

In niedda, bouts of wing-flick/cape-flapping coincide with a distinctive very rapid side-to-side wobbling of the head. As viewed from the front (i.e., the POV of the approaching female) the head wobbling has the effect of making the eye-spots flicker or shimmer. It is possible that the wobbling head movement may also be present in superba, but is not discernible from the available data.

Figure 6: Detailed comparison of the cape presentation form of superba and niedda. superba (A), the shape of the cape presentation form is oval—i.e., wider than it is high. The edge of the opened cape has a smooth fuzzy appearance, which is created by the distinct hair-like fringing along the terminal tips of the individual feathers that comprise the cape. In niedda (B), the shape of the cape presentation form is distinctly crescent shaped—i.e., it has a curved sickle shape that is broad in the center and tapers to a point at each end. The contour of the crescent is slightly ribbed and lacks the fuzzy appearance of superba. The shape and appearance of the breast shield also differs between the cape presentation phenotype of superba and neidda. In superba (A), the breast shield is wider with thicker terminal ends that make a relatively flat surface with a fairly straight edge along the top of the ornament. In neidda (B), the shorter and narrower terminal tips of the breast shield curve outward so that the surface of the ornament is slightly concave and the top edge is angled downward at the sides. This difference results in niedda looking like somewhat like “frowning face” whereas superba has more of a “smiling face” look. Also, the thinner profile of the narial tufts of niedda often makes the eye-spots look like they have thin blue “eyebrows” just above each eye-spots. Because the narial tufts fo superba are shorter and do not extend over the top margin of the eye-spot, “eyebrows” are possible. Image credit/source: (A) Edwin Scholes/ In(A), the shape of the cape presentation form is oval—i.e., wider than it is high. The edge of the opened cape has a smooth fuzzy appearance, which is created by the distinct hair-like fringing along the terminal tips of the individual feathers that comprise the cape. In(B), the shape of the cape presentation form is distinctly crescent shaped—i.e., it has a curved sickle shape that is broad in the center and tapers to a point at each end. The contour of the crescent is slightly ribbed and lacks the fuzzy appearance of. The shape and appearance of the breast shield also differs between the cape presentation phenotype ofand. In(A), the breast shield is wider with thicker terminal ends that make a relatively flat surface with a fairly straight edge along the top of the ornament. In(B), the shorter and narrower terminal tips of the breast shield curve outward so that the surface of the ornament is slightly concave and the top edge is angled downward at the sides. This difference results inlooking like somewhat like “frowning face” whereashas more of a “smiling face” look. Also, the thinner profile of the narial tufts ofoften makes the eye-spots look like they have thin blue “eyebrows” just above each eye-spots. Because the narial tufts foare shorter and do not extend over the top margin of the eye-spot, “eyebrows” are possible. Image credit/source: (A) Edwin Scholes/ ML458003 and (B) Tim Laman/ ML487563

Next comes the transition into the main part of cape presentation display—i.e., the presentation of the fully opened cape itself. In superba, the cape is lifted into its presentation position well before to female arrives on the log (Fig. 5C). In L. niedda, the wing-flick/cape-flaps with head wobbles continue for several more bouts after the female has arrived on the log and approaches the male become face-to-face with him (Fig. 5D). As the neidda female moves closer, the male will back up slightly and crouch lower before deploying his cape for full presentation (e.g., videos ML487530, ML487538, and ML487562).

Although the timing of cape deployment differs between superba and niedda, the cape of both is lifted into the full display position in largely the same way. First, it is spread wide, fan-like, over the back and then it is raised (anteriorly) and pushed over the head until it creates the super-black (McCoy et al., 2018), slightly concave, cape presentation display form that surrounds the entire head and upper body (Figs. 5E & 5F and S1E).

Once in position however, the form and appearance of the cape presentation phenotype differs dramatically between superba and niedda. In superba, the shape of the cape presentation form is oval—i.e., wider than it is high (Figs. 6A & S1F). In comparison, the shape of the cape presentation form of niedda is distinctly crescent shaped—i.e., it has a curved sickle shape that is broad in the center and tapers to a point at each end (Fig. 6B). In superba, the border edge of the fully opened cape has a smooth fuzzy appearance (Fig. 6A), which is created by the distinct hair-like fringing along the terminal tips of the individual feathers that comprise the cape. The border edge of the crescent in niedda is slightly ribbed and lacks the fuzzy appearance of superba (Fig. 6B). Correspondingly, the terminal tips of the niedda cape feathers have very minor hair-like fringing. The iconic oval shape of superba exhibits very little variation within the superba complex (e.g., Fig. 6A and Fig. S1F for a comparison of L. s. latipennis and L. s. superba and refer to the documentary films described in the Methods for comparative imagery of L. s. addenda).

The shape and appearance of the breast shield also differs between the cape presentation phenotype of superba and neidda. In superba, the breast shield is wider with thicker terminal ends that make a relatively flat surface with a fairly straight edge along the top of the ornament (Fig. 6A). In neidda, the shorter and narrower terminal tips of the breast shield curve outward so that the surface of the ornament is slightly concave and the top edge is angled downward at the sides (Fig. 6B). This difference results in niedda looking like somewhat like “frowning face” whereas superba has a “smiling face” look. The “smiling” vs “frowning” face appearance is further enhanced by the difference in how the eye-spot ornaments are created. In superba, the the narial tufts are thick and relatively short whereas in niedda they are narrow and longer (i.e., extend beyond the width of the head). This difference has the effect of making the eye-spots of niedda appear smaller with a straighter line along the top edge, which gives the “eyes” the appearance of having slightly droopy “eyelids” compared with the always larger and circular appearance of the “eyes” in superba (Figs. 6A and 6B). In addition, when alignment of the feathers is not perfect relative to the viewer, the thinner profile of the narial tufts often makes the eye-spots of niedda look like they have a thin blue “eyebrow” just above each eye-spots (Fig. partially visible in Fig. 5F). Because the narial tufts fo superba are shorter and do not extend over the top margin of the eye-spot, “eyebrows” are never visible nor even possible.

Throughout the cape presentation display, the transformed oval-shaped cape presentation form of superba is repeatedly lifted up and down, or bounced, with outstretched and bent legs (e.g., ML458003). The repeated up and down movements coincide with a series of alternating shuffle–hop steps to the side, which create the distinct bouncing cadence: up–down shuffle–hop, first to one side and then the other. When a female is present, the male’s bouncing shuffle–hop movements revolve around the female in either a semi-circular or full circle path.

In niedda, the overall appearance of the rotational movement during the cape presentation display is very different from that of superba. Instead of the characteristic bouncing of superba, the feet of niedda move in a faster, more pronounced, side-step motion without bouncing and with no visible up and down movement of the body. This creates the effect of niedda appearing to move in a quick but steady glide, rather than bounce, around the female in semi- or full circle (e.g., ML487557 and ML487562).

Figure 7: Wing-flick snaps of superba and niedda. superba (A), distinct snapping sounds are a characteristic acoustic component of the cape presentation display and occur rhythmically with the up–down bouncing movement on the down beat of the bounces, but not on every bounce. In superba latipennis, the cadence of snaps and bouncing is down-snap-up, down-snap-up, down-up such that snaps occur in doublets with a distinct pause in between, which sounds like “snap-snap (pause) snap-snap”. In niedda (B), where the rotational movement lacks the bouncing of superba, the overall pattern and effect of the wing/tail flicks and snaps is very different because they occur at a regular interval that is not timed with any particular aspect of the rotational movement. The snaps of niedda are given at uniform beat of approximately one snap per second for a steady “snap-snap-snap” sound. Audio credit/source: (A) Edwin Scholes/ In(A)distinct snapping sounds are a characteristic acoustic component of the cape presentation display and occur rhythmically with the up–down bouncing movement on the down beat of the bounces, but not on every bounce. In, the cadence of snaps and bouncing is down-snap-up, down-snap-up, down-up such that snaps occur in doublets with a distinct pause in between, which sounds like “(pause). In(B)where the rotational movement lacks the bouncing ofthe overall pattern and effect of the wing/tail flicks and snaps is very different because they occur at a regular interval that is not timed with any particular aspect of the rotational movement. The snaps ofare given at uniform beat of approximately one snap per second for a steady “sound. Audio credit/source: (A) Edwin Scholes/ ML458003 and (B) Edwin Scholes/ ML8576351

Along with the bouncing (superba) or smooth quick (niedda) rotational movement of the cape presentation form along the display log, the tail of both species is lifted to about 45 degrees above the plane of the back and rectrices are fanned. The secondary feathers of the wings are spread such that they become partially intertwined with the rectrices (see Fig. S1E). In one explosive motion, the wings are flicked open and shut to the sides, which causes a violent interaction among the secondaries and rectrices. In superba, the explosive wing-flicks occur rhythmically with the up–down bouncing movement and coincides with the distinct snapping sounds that are a characteristic acoustic component of the display. In superba, the wing-flicks happen on the “down beat” of the bounces, but not on every bounce. In s. latipennis, the cadence of snaps and bouncing is down-snap-up, down-snap-up, down-up such that snaps occur in doublets with a distinct pause in between, which sounds like “snap-snap (pause) snap-snap” (Fig. 7A). In niedda, where the rotational movement lacks the bouncing of superba, the overall pattern and effect of the wing/tail flicks and snaps is very different because they occur at a regular interval that is not timed with any particular aspect of the rotational movement. The snaps of niedda are given at uniform beat of approximately one snap per second for a steady “snap-snap-snap” sound (Fig. 8B).

In s. superba, the cadence of the snapping is more frequent with less obvious choreography among the snaps and bouncing movements. In s. addenda, to the extent discernible from the documentary video sources, both the cadence of the bouncing and snapping seems to be more frequent with wing-flicks (and presumably the snaps) occurring with each downward bouncing stroke, “down-snap-up, down-snap-up, down-snap-up”. In both niedda and superba, the exact relationship between the wing movement and snapping sound production is unclear and warrants further investigation using high-speed videography.

In both superba and niedda, the cape presentation display ends when the female moves quickly away from the male such that he is unable to continue rotating around her, or when the female solicits copulation. At times the female moves along the log ahead of the male and at other times she will fly away from the log entirely. When females depart, males of both superba and niedda maintain the full cape presentation posture directed toward her and slowly lower it down over the back to readopt the pointing posture. In niedda, as the female departs, the male opens his bill to reveal the bright yellow gape once again. This end of display reveal of the gape was not observed in superba.

Figure 8: Advertisement vocalizations of superba and niedda. niedda and superba are phrases of 2–8 notes, with five notes common. While the number of notes is similar in both species, superba (A) gives a characteristic harsh raspy screech. The acoustic quality of niedda (B) vocalizations are very different. The basic note of niedda is a high-pitched nasal “yiap” sound, which has a piercing whistle-like quality that ascends and descends slightly in frequency. Audio credit/source: (A) Edwin Scholes/ Advertisement vocalizations of bothandare phrases of 2–8 notes, with five notes common. While the number of notes is similar in both species,(A) gives a characteristic harsh raspy screech. The acoustic quality of(B) vocalizations are very different. The basic note ofis a high-pitched nasal “sound, which has a piercing whistle-like quality that ascends and descends slightly in frequency. Audio credit/source: (A) Edwin Scholes/ ML458003 and (B) Edwin Scholes/ ML85763571

If a female solicits copulation (e.g., ML487568 from 0:07–0:17), she beings fluttering her wings as the male rotates around her and then will tip forward with breast low and tail high. At this point, the male circles closely and mounts the female while maintaining the fully open cape posture. The cape is held in the open position during copulation and wing-snaps are given as part of, or just after, the wing flapping that occurs with copulation. After copulation, the male dismounts from the back of the female and resumes the cape presentation display directed at the female (e.g., ML487562). If she departs the log, the male holds the opened cape posture for a few seconds and the slowly lowers it and often returns to the pointing display and sometimes even reverts backwards through the entire sequence from pointing, to horizontal to advertisement displays.