Pipridae - Manakins

Manakins are small frugivores that can occupy all levels of the interior forest and tend to tolerate edges and fragmentation because of its positive effects on fruiting trees, especially Melastomataceae, which are favored by many manakins. The classic manakins are sexually dichromatic; males are brightly colored while females are mostly green. Some are sexually monochromatic. These are small frugivores that are variably yellow-green and flycatcher-like. Most manakins have atypical social structures in which males form leks and parental care is done solely by the female. This social structure causes differences in movements and capture rates among age and sex classes. In general, adult males are most sedentary and are least frequent in nets. Females are slightly more mobile as they wander between leks and raise young without the help of males. Juveniles and subadults wander widely in search of fruits because they are not yet tied to a breeding location; they also probably also develop and test their social skills during this time and investigate potential future leks in which to participate as breeding adults. The juvenal apparently is held extremely briefly. In the most frequently captured species at the BDFFP, the preformative molt is partial resulting in molt limits among the wing coverts. In most species, the replaced greater coverts are a few millimeters longer and brighter green than the juvenal greater coverts. The second prebasic molt is complete, but depending on the species, it may or may not be the definitive plumage. Not until the third prebasic molt in some species is it possible to separate males from females. Thus a green bird without molt limits may either be a male or female in Basic II or a female in Basic III or later. Some green have a few male-like feathers among the green feathers, but these individuals could be either advanced immature males or old females. The extent and location of these feathers may provide clues of the sex and age, but may not be 100% reliable and are species- specific. The skull ossifies in most (all?) species, but probably does not add much insight to aging birds because the preformative molt is often partial such that molt limit criteria are available as the skull ossifies. Molt-breeding overlap is rare probably because manakins have highly predictable molting and breeding seasons. Only females incubate and develop BPs. Breeding peaks during the middle to end of the dry season and molt occurs afterward and peaks during the early wet season.

# species at BDFFP: 9 # species netted: 9 # primaries: 10 # secondaries: 9 # rectrices: 12 chloris tshudii(?) – Wing-barred Piprites Band size: Unknown

# Individuals Captured: 2 Similar Species: Combination of bold yellow wingbar, yellow underparts, yellow eyering, and yellow crown stripe are unique. Measurements: Wing 64.5 – 65.0 (64.8 ± 0.4, n2) Tail 43.0 (n2) Mass 15.5-17.0 (16.3 ± 1.1, n2) Skull: Probably does not ossify in adults (LSU specimens). Brood Patch: No data available. Molt: Unknown. Age/Sex:

BASIC I: ♀=♂ (?) BASIC II: ♀=♂ Undescribed. Rect shape may be Both sexes have yellow crown stripe. useful, being distinctly pointed (LSU Rects are rounded. Other plumage specimen, n1). Otherwise very similar criteria for aging are not known. to adult.

Note: Taxonomic status is not well understood and is currently treated as incertae sedis in the SACC checklist (2009). chrysocephalum (no subsubspecies) – Saffron-crested Tyrant-Manakin Band size: Unknown

# Individuals Captured: 1 Similar Species: Drab smallish flycatcher-like bird with grayish crown, bright yellow semi- concealed crown patch, yellowish belly, and a creamy-white iris. Otherwise it is like other flycatchers with nondescript olive upperparts and dirty grayish-olive throat and chest. Measurements: Wing 67.0 (n1) Tail 48.5 (n1) Mass 12.3 (n1) Skull: Unknown. Brood Patch: No data available. Molt: Unknown. Probably Complex Basic Strategy. PF partial or incomplete?, PBs complete, PA probably absent. Age/Sex:

BASIC I: : ♀=♂ (?) BASIC II: ♀=♂ Undescribed. Both sexes have large bright yellow concealed crown patch. Not known FORMATIVE I ♀=♂ how to separate from B1 and F1. The only bird captured at the BDFFP had juvenile-like growth bar alignment on remiges and rectrices, but was otherwise adult-like, suggesting a bird in F1 and that the PF1 is partial- incomplete. Wing coverts should be examined for molt limits to understand the extent of PF1 molt.

Note: Typically a species of campina (white sand woodland), this species is rarely encountered in terra firme forest. The few observations at the BDFFP may have been of dispersing subadult birds. Captured birds should be closely examined for juvenile-like features. virescens (no subspecies) – Tiny Tyrant-Manakin Band size: C

# Individuals Captured: 11 Similar Species: Flycatcher-like, but note lack of rictal bristles. Otherwise, combination of gray iris, semi-concealed yellow crown patch, very small size, and relatively short tail is unique. Measurements: Wing 42.0-51.0 (46.7 ± 3.0, n10) 100 Tail 12.0-25.0 (21.1 ± 4.5, n9) Mass 6.0-9.0 (7.5 ± 1.1, n9) 80 Bill 6.4 (n1) 60 Tarsus 13.4 (n1)

Skull: Unknown. 40 Brood Patch: No data available. 20 Molt: Complex Basic Strategy. PF1 partial, PBs complete. PF1 appears to 0 % birds with(solidmolt circles)bp or (open circles) include the replacement of body feathers, Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec less covs, med covs, and inner gr covs, but Molt; n =202010021201 not outer gr covs, pp covs, or remiges. Bp; n =001000000100 Age/Sex: BASIC I: : ♀=♂ (?) Unknown.

BASIC II: ♀=♂ (?) The iris is brownish-gray. The yellow crown patch is partially concealed or absent. The inner gr covs are brighter olive-green and longer than the retained outer gr covs, which are dull olive-green and relatively short. Tyranneutes virescens (no subspecies) – Tiny Tyrant-Manakin

BASIC II: ♀=♂ (?) The iris is gray. Large yellow crown patch is partially concealed. ♂♂ are shorter- tailed and average larger yellow crown patches than ♀♀, but we do not yet have reliable quantitative data to separate ♂♂ from ♀♀. Most birds in the database have been aged as ♂ presumably because of the presence of a large crown patch, but ♀♀ also have this crown patch. Almost certainly some of these are incorrectly sexed based on tail measurements. gutturalis (no subspecies) – White-throated Manakin Band size: C

# Individuals Captured: 208 Similar Species: Definitive ♂unmistakable. ♀ and juvenile like other small green manakins, but note small size and pale bill. 60 Measurements:

Wing ♂ 51.5-59.0 (55.0 ± 1.2, n80) 50 ♀ 51.0-56.5 (53.1 ± 1.2, n80) 40 Tail ♂ 23.0-31.0 (26.6 ± 1.3, n78) ♀ 21.0-30.0 (26.2 ± 1.6, n77) 30

Mass ♂ 5.8-10.0 (7.8 ± 0.7, n86) 20 ♀ 6.0-10.9 (8.4 ± 0.7, n87) Bill ♂ 4.1-5.6 (4.8 ± 0.4, n9) 10

♀ 4.5-5.4 (5.1 ± 0.4, n6) line) (dotted bp or line) (solid molt with birds % 0 Tarsus ♂ 16.4-17.5 (17.1 ± 0.4, n9) Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec ♀ 16.4-17.5 (17.0 ± 0.5, n6) Molt, n = 16 6 7 5 20 12 33 11 23 51 32 11 Bp, n = 0 0 0 1 1 6 24 2 7 19 15 3 Skull: Completes during F1. Brood Patch: Only ♀♀ develop BPs, which have been observed between Jul and Nov, peaking from Aug to Nov, but few data are available from Dec to Jun. Molt-breeding overlap is 0.0% (n9). Molt: Complex Basic Strategy. PF1 partial, PBs complete. PF1 includes all body feathers, rects, less and med covs, and 1-5 (usually 3) gr covs. The timing of molt remains elusive as only 5 of 227 captures were in molt. All molting birds were seen between Feb and Jun and this is probably the molting period, which is consistent with other manakins and BP data. Age/Sex: BASIC I: ♀=♂ Pale green upperparts. This plumage is probably only briefly held as some F1 feathers begin to emerge while some juvenile feathers are still in pin.

FORMATIVE I: ♀≠♂ Molt limits are evident among the wing covs. The retained gr covs and pp covs are duller green and shorter than the fresher, brighter green, longer replaced inner gr covs. Corapipo gutturalis (no subspecies) – White-throated Manakin FORMATIVE I: ♀≠♂ ♀ - Green upperparts with molt limits among the gr covs. Throat is grayish-white with slightly grayish-olive chest, grayish-white underparts, and no gray or black in the auriculars.

♂ -Like F1 ♀. Some (all?) ♂♂ can be distinguished by having a whiter throat (rather than pale grayish-white) and some (most?) have gray or black in the face. Because it is possible that not all ♂♂ are distinguishable, F1 birds without ♂- like features should be sexed as “unknown.”

BASIC II: ♀≠♂ ♀ - The definitive plumage. Green upperparts without molt limits among the gr covs. Throat is grayish-white with slightly grayish-olive chest and grayish-white underparts. More study is needed to confirm that all green birds without molt limits are ♀, because some Basic II ♂♂ may lack black body feathers. P10 is broader than in ♂ B2 and may be reliable to make sex determinations in the absence of plumage characteristics.

♂ - Green upperparts without molt limits among the gr covs. May always (but see BASIC II ♀) have some black feathers in the face. The throat is bright white. The width of p10 may be diagnostically narrow, even if ♂-like plumage characteristics are not present. Corapipo gutturalis (no subspecies) – White-throated Manakin

BASIC III ♂ - The definitive plumage. Mostly glossy blue-black with a white throat and chest. Pp1-4 are marked white, but occasionally p3 and/or p4 lack white on the outer webbing, which may be related to age as it often is seen when definitive ♂♂ have a few black body feathers that are tipped green (are these always B3?). serena (no subspecies) – White-fronted Manakin Band size: C

# Individuals Captured: 573 Similar Species: Definitive ♂ unmistakable. ♀♀ and subadults like other small green manakins, but note yellow wash on belly and short wing. 35 Measurements: Wing ♂ 48.0-58.0 (52.3 ± 1.7, n150) 30

♀ 47.0-59.0 (53.7 ± 1.5, n232) 25

Tail ♂ 19.0-31.0 (25.5 ± 1.9, n146) 20 ♀ 21.0-33.0 (27.1 ± 1.6, n227) 15 Mass ♂ 8.0-12.0 (10.2 ± 0.7, n185) ♀ 8.0-13.0 (10.8 ± 0.9, n275) 10 Bill ♂ 5.1-6.4 (5.8 ± 0.3, n22) 5

♀ 5.6-6.6 (6.1 ± 0.3, n18) molt% birds with line) (solid or bp (dotted line) 0 Tarsus ♂ 16.9-19.2 (17.9 ± 0.7, n20) Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec ♀ 16.6-18.8 (17.9 ± 0.7, n18) Molt, n = 32 25 49 57 83 61 117 62 78 87 100 56 Bp, n = 0 0 8 10 16 19 56 24 38 35 36 23 Skull: Completes during F1. Brood Patch: Only ♀♀ develop BPs, which have been observed between Aug and Dec(-Mar?), peaking (>35%) in Oct. Molt-breeding overlap is 0.0% (n13). Molt: Complex Basic Strategy. PF1 partial, PBs complete. PF1 includes all body feathers, rects, less and med covs, and 1-5 (usually 3) inner gr covs. The timing of molt is well defined, peaking (>30%) in Apr and is least frequent (<5%) between Jul and Nov. Age/Sex: BASIC I: ♀=♂ Generally green overall, with faint dull yellow in belly.

FORMATIVE I: ♀=♂ (usually) Green upperparts and yellow underparts with molt limits among the gr covs. The retained gr covs are duller green and shorter than the replaced brighter green and longer inner gr covs. Lepidothrix serena (no subspecies) – White-fronted Manakin

FORMATIVE I: ♀=♂ (usually) ♂ - At least some ♂♂ will have a trace to lots of definitive ♂-like plumage in the body feathers, especially in the forehead and back. Because some ♂♂ probably do not have definitive ♂-like feathers, all “green and yellow” birds with molt limits should be sexed as unknown.

BASIC II: ♀≠♂ ♀ - The definitive plumage. Green overall with a yellow belly and without molt limits among the gr covs.

♂ - The definitive plumage. Mostly black with a yellow belly, yellow spot in chest, white patch above bill, and blue rump. Occasionally a few black feathers will have green tips, which may be birds in their 1st definitive plumage (i.e. B2). manacus manacus – White-bearded Manakin Band size: D-E

# Individuals Captured: 20 Similar Species: ♂ unmistakable. ♀ like other green manakins, but with bright orange legs. Measurements: Wing ♂ 51.0-52.0 (51.5 ± 0.6, n4) 30

♀ 50.0-55.0 (52.3 ± 1.5, n9) 25 Tail ♂ 31.0-33.0 (32.3 ± 1.0, n4) 20 ♀ 28.0-33.0 (31.2 ± 1.6, n9)

Mass ♂ 15.5-17.0 (16.3 ± 0.9, n4) 15 ♀ 12.4-17.0 (15.3 ± 1.3, n9) 10 Skull: Probably completes during F1. 5 Brood Patch: Only in ♀♀, but has not

0

been observed in the 3 definitive ♀♀ circles) (open bp or circles) (filled molt with birds % examined (in Jun, Sep, and Nov) so Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec the timing remains unknown, but Molt, n =000403412060 should be expected to be similar to Bp, n =000003312030 , Corapipo, and Lepidothrix. Molt: Complex Basic Strategy. PF1 partial, PBs complete. PF1 includes all body feathers, rects, less and med covs, and 1-5 (usually 3) inner gr covs. The timing of molt is well defined, peaking (>30%) in Apr and is least frequent (<5%) between Jul and Nov. Age/Sex: BASIC I: ♀=♂ Unknown and probably only briefly held as some F1 feathers begin to emerge while juvenile feathers are still in pin (as in Pipra pipra). Leg color is probably duller than in adults.

FORMATIVE I: ♀=♂ (usually) Green overall and definitive ♀-like, but with molt limits among the gr covs. The retained gr covs are duller green than the brighter green, replaced inner gr covs. The legs are bright orange. Manacus manacus manacus – White-bearded Manakin

FORMATIVE I: ♀=♂ (usually) ♂ - At least some ♂♂ will have a trace of definitive ♂-like plumage in the body feathers, especially in the forehead and back. Because some ♂♂ probably do not have definitive ♂-like feathers, all “green” birds with molt limits should be sexed as unknown.

BASIC II: ♀≠♂ ♀ - The definitive plumage. Green overall without molt limits. Legs are bright orange.

♂ - The definitive plumage. Upperparts are black, the throat and cheek are white and the breast and belly are gray. Legs are bright orange. Pipra pipra pipra – White-crowned Manakin Band size: D

# Individuals Captured: 2149 Similar Species: ♂ distinct, but ♀ and juvenile like other green manakins. From Lepidothrix serena by lack of yellow on underparts. Measurements are useful to separate it from the shorter- tailed P. erythrocephala. Manacus manacus has orange (not dull grayish-black) legs. Measurements: 50 Wing ♂ 48.0-58.0 (52.3 ± 1.7, n150) 40 ♀ 47.0-59.0 (53.7 ± 1.5, n232)

Tail ♂ 19.0-31.0 (25.5 ± 1.9, n146) 30 ♀ 21.0-33.0 (27.1 ± 1.6, n227)

Mass ♂ 8.0-12.0 (10.2 ± 0.7, n185) 20 ♀ 8.0-13.0 (10.8 ± 0.9, n275) Bill ♂ 5.1-6.4 (5.8 ± 0.3, n22) 10 ♀ 5.6-6.6 (6.1 ± 0.3, n18) % birds with molt (solid line) or bp (dotted line) or bp (dotted line) (solid molt with % birds 0 Tarsus ♂ 16.9-19.2 (17.9 ± 0.7, n20) Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

♀ 16.6-18.8 (17.9 ± 0.7, n18) Molt, n = 93 85 156 151 254 384 424 282 217 240 230 115 Skull: Completes during F1, usually Bp, n = 0 0 22 13 37 107 177 89 76 106 86 27 before Jun. Brood Patch: Only develops in ♀♀. Very rarely F1 birds can have a BP, but breeding probably does not usually occur until B2. Breeding strictly occurs between Aug and Dec(-Jan?), peaking in Oct and Nov. Molt-breeding overlap is 1.4% (n73). Molt: Complex Basic Strategy. PF1 partial, PBs complete. PF1 includes all body feathers, rects, less and med covs, and 1-5 (usually 3) gr covs. Rarely all gr covs are replaced. Molt primarily occurs between Jan and May, peaking (>25%) in Mar and Apr. Age/Sex:

BASIC I: ♀=♂ Dull green with dull green wing covs, remiges, and rects. Iris is brown. Plumage replaced very soon after leaving the nest. Pipra pipra – White-crowned Manakin

FORMATIVE I: ♀=♂ Bright green upperparts and definitive ♀-like, but with molt limits often among gr covs (see Molt). Retained outer gr covs are duller green and about 3mm shorter than the brighter replaced inner covs or occasionally all gr covs are replaced, contrasting with dull green pp covs. The iris is typically brown upon completion of the PF1, but becomes light orange, orange, or dark orange by Aug, before PB2 begins.

BASIC II: ♀=♂ (not always) ♀ - The definitive plumage. Green upperparts without molt limits. The head may be mostly green or with some gray, but probably averages less gray than in B2 ♂♂. The iris is orange, dark orange, or occasionally red(?). Green birds without (and with) molt limits (See FORMATIVE I) should be sexed as unknown unless there is evidence of a brood patch. Some old ♀♀ can have some ♂-like white crown feathers or black body feathers and may not be separable from B2 ♂♂ - these should be sexed as unknown. Pipra pipra – White-crowned Manakin

BASIC II: ♀=♂ (not always) ♂ - Can be identical to B2 ♀♀ so green birds without (and with: see Formative I) molt limits should be sexed as unknown, but may average grayer-headed than definitive ♀♀.B2 ♂♂ can show varying amounts of definitive ♂-like feathers throughout the body from none (these are definitive ♀-like) to many. Iris may be dark orange to red to dark red. Some B2 ♂♂ have few to many definitive feathers, but beware of confusion with old ♀♀that also have several ♂-like feathers. Birds with the most ♂-like feathers and/or dark red irises are almost certainly ♂ but there are many intermediately-plumaged birds that should be sexed as unknown.

BASIC III: ♀≠♂ ♂ - The definitive plumage. All black with a white crown and dark red iris. Some show a trace of green on the body and/or wings; these may be birds in B3 (always?), but more study is needed. Pipra erythrocephala erythrocephala – Golden-headed Manakin Band size: D

# Individuals Captured: 302 Similar Species: ♀♀ and juveniles very much like other dull green manakins (especially P. pipra). The short tail and short tarsus are the best quantitative features. Also note its pale bill and brown (not orange-red) iris, but beware of very young P. pipra with brown eyes.

Measurements: 60 Wing ♂ 51.0-59.0 (54.5 ± 1.7, n65) 50 ♀ 52.0-59.0 (55.5 ± 1.3, n142) Tail ♂ 15.0-23.0 (19.1 ± 1.6, n65) 40

♀ 15.0-24.0 (20.5 ± 1.6, n140) 30 Mass ♂ 9.0-14.3 (11.3 ± 1.1, n72) ♀ 10.0-15.5 (12.0 ± 1.0, n154) 20

Bill ♂ 5.3-5.7 (5.5 ± 0.2, n3) 10

♀ 5.8-6.9 (6.4 ± 0.3, n10) % birds with molt (solid line) or line) bp (dotted 0 Tarsus ♂ 14.6-15.6 (15.2 ± 0.6, n3) Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec ♀ 14.5-16.0 (15.4 ± 0.4, n10) Molt, n =8 5 20133742422019323117 Skull: Completes during F1. Bp, n = 0 0 5 7 18 20 27 13 12 23 15 10 Brood Patch: Only develops in ♀♀. Breeding occurs between Jul and Dec(-Feb?), peaking in Dec. Molt-breeding overlap is 0.0% (n19). Molt: Complex Basic Strategy. PF1 partial, PBs complete. PF1 includes all body feathers, rects, less and med covs, and 1-5 (usually 3) gr covs. Peaks in Mar and not observed between Aug and Dec. Age/Sex:

BASIC I: ♀=♂ Probably held for a very short time as in P. pipra and juvenile characteristics will be evident (e.g. weak plumage and gape). The iris is brown.

FORMATIVE I: ♀=♂ (not always) Green with molt limits among the gr covs. The inner replaced gr covs are brighter green than the retained outer gr covs and pp covs. Pipra erythrocephala erythrocephala – Golden-headed Manakin

FORMATIVE I: ♀=♂ (not always) ♂ - Some develop white splotches in the brown iris or occasionally the iris will be entirely white. Additionally, some ♂♂ can be identified by having traces of definitive ♂-like yellow feathers in the crown or black feathers on the body. Birds with molt limits and without definitive ♂-like features described above should be sexed as unknown.

BASIC II: ♀≠♂ ♀ - The definitive plumage. Dull olive-green without molt limits. The iris is brown, but older ♀♀ often develop minute white specs in iris (not splotches or blocks as in F1 ♂♂).

♂ - The definitive plumage. Black with a yellow head narrowly bordered red with black body and red “leggings.” The iris is white.

Note: The tarsus is too short to place both a metal and color band on the same leg, but two color bands will fit well if trimmed.