Integrating voice and phenotype in a revision of the brush cuckoo Cacomantis variolosus (Aves: Cuculidae) complex Author Wu, Meng Yue 0000-0002-8562-7667 National University of Singapore, Department of Biological Sciences, 14 Science Drive 4, Singapore. & dbswmy @ nus. edu. sg; https: // orcid. org / 0000 - 0002 - 8562 - 7667 dbswmy@nus.edu.sg Author Schodde, Richard 0000-0002-8255-4773 Australian National Wildlife Collection, National Research Collections Australia, CSIRO, Australian Capital Territory 2601, Australia. rschodde @ grapevine. com. au; https: // orcid. org / 0000 - 0002 - 8255 - 4773 rschodde@grapevine.com.au Author Rheindt, Frank E. 0000-0001-8946-7085 National University of Singapore, Department of Biological Sciences, 14 Science Drive 4, Singapore. & dbsrfe @ nus. edu. sg; https: // orcid. org / 0000 - 0001 - 8946 - 7085 dbsrfe@nus.edu.sg text Zootaxa 2022 2022-01-12 5091 1 69 106 journal article 2633 10.11646/zootaxa.5091.1.3 22e66303-82d5-4c5d-b688-a4d511f551de 1175-5326 5840523 964647F3-9828-4E34-A495-67E03BAFC2EF

TABLE 2. Measurements and proportions of regional forms and populations of the Cacomantis variolosus complex. Bracketed measurements are of extreme ranges; those in between cover 75% of variation. Records of numbers of morphs per population are not exhaustive because morphs were not recorded for all museum collections. Abbreviations: M—males, F—females.

Taxon (distribution)	Sex and	Wing (mm)	Tail (mm)	Tail/wing ratio	Exposed cul-	Width of max-	Width of	Morph
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	sample size				men (mm)	illa (mm)	maxilla/culmen
							length ratio
Sunda Brush Cuckoo (sepulcralis) —Greater Sundas to Philippines and west Nusa Tenggara
Greater Sundas incl. Malay	35 M	(110–)113–	(110–)115–	(0.97–)1.02–	(14.0–)14.8–	(5.6–)6.0–6.8(–	(0.34–)0.37–	20 basic, 1grey
Peninsula east to central	16 F	120(–123)	129(–137)	1.10(–1.14)	17.4(–18.1)	7.2)	0.45(–0.48)	3 basic, 2 barred, 2
Nusa Tenggara (Flores, Alor,		(104–)110–	(107–)111–	(0.97–)1.00–	(14.9–)16.3–17.5	6.0–7.2	0.36–0.42(–0.46)	muted
Sumba)—(sepulcralis)		117(–120)	121(–125)	1.05(–1.07)
Philippines—(sepulcralis)	10 M	112–116(–120)	122–128(–	1.06–1.12	(14.7–)15.4–17.8	(5.6–)6.2–6.7	0.35–0.42(–0.44)	7 basic, 1 grey
	9 F	107–118	130)	(0.94–)0.99–1.05	14.8–17.1	(5.6–)5.9–6.8	0.38–0.42	3 basic, 5 barred
			108–118
Sulu Archipelago—(everetti)	1 M (juv.)	119	123	1.03	14.4	6.3	0.44	‒
	2 F	117–118	126–133	1.08–1.13	15.3–15.9	5.9–6.2	0.39	2 basic
Sulawesi Brush Cuckoo (virescens)—Sulawesi and Sula Archipelago
Sulawesi and satellite islands—	34 M	(104–)108–	(115–)125–	(1.10–)1.12–1.23	(13.7–)14.4–	(5.4–)5.8–6.7	(0.35–)0.38–	30 basic, 1 grey
(virescens)	13 F	116(–120)	137(–141)	(–1.28)	16.1(–16.8)	(–6.9)	0.43(–0.46)	11 basic
		(106–)110–	(122–)124–	1.10–1.22(–1.25)	(13.5–)14.2–	(4.9–)5.4–6.4	(0.36–)0.38–
		114(–117)	134(–142)		15.4(–16.1)	(–6.8)	0.43(–0.45)
Sula Archipelago—(virescens)	1 M	113	137	1.21	14.5	6.2	0.42	1 basic
	1 F	108	121	1.12	14.6	6.5	0.46	1 barred
	3 unsexed	110–114	124–136	1.09–1.21	15.1–15.3	5.6–6.2	0.37–0.41	3 basic
Moluccan Brush Cuckoo (aeruginosus)—Moluccas
North Moluccas (Obi, Bacan,	7 M	111–121	114–131	(1.02–)1.05–1.14	(14.9–)15.7–16.8	(5.7–)6.4–6.9	(0.34–)0.41–0.43	7 basic
Halmahera)—(heinrichi)	5 F	110–117	112–125	1.02–1.07	13.7–16.2	5.2–7.1	0.35–0.44	1 basic, 4 barred
South Moluccas (Buru)—(aeru-	10 M	(113–)115–	123–132	1.04–1.13(–1.17)	(13.6–)14.0–15.8	5.7–6.9	(0.40–)0.41–0.46	10 basic
ginosus)	6 F	120(–122)	(–135)	(1.00–)1.06–	(13.7–)15.1–16.7	6.4–7.4	0.42–0.46	2 basic, 2 barred
	1 unsexed	(112–)114–	(119–)122–133	1.12(–1.14)	–	–	–	1 basic
		119(–120)	137	1.15
		119
	sample size				men (mm)	illa (mm)	maxilla/culmen
							length ratio
South Moluccas (Ambon,	4 M	114–120(–124)	125-133	1.09–1.11	15.6–16.9	6.4–7.5	0.39–0.44	4 basic
Seram)—(aeruginosus)	4 unsexed	115–122(–124)	118-135	1.03–1.11	16.1–17.3 (n	5.9–6.5 (n = 2)	0.34–0.40 (n = 2)	4 basic
= 2)
Australian Brush Cuckoo (major)—North Moluccas
Obi, Bacan, Halmahera, Ter-	10 M	(126–)128–133	(105–)109–	(0.83–)0.85–	15.7–17.3	6.2–7.1	0.38–0.43	8 basic, 2 grey
nate—(pale plumaged major)	6 F	125–131	120	0.89(–0.90)	16.3–17.3	6.1–6.4	0.35–0.39	3 basic, 3 barred
			102–111	0.82–0.89
Bacan, Halmahera, Ternate—	19 M	(119–)121–	(105–)108–	(0.85–)0.87–	(15.6–)16.3–	5.9–7.4	0.36–0.42(–0.44)	14 basic, 3 grey
(putative major X infaustus)	10 F	128(–130)	115(–122)	0.92(–0.94)	17.5(–18.3)	(5.3–)6.5–7.5	(0.31–)0.37–0.43	5 basic, 4 barred, 1
		118–125(–128)	102–113	0.83–0.90(–0.96)	16.6–17.6			muted
Bacan, Halmahera, Ternate,	4 M	118–123	107–112	0.87–0.94	16.5–17.4	6.1–6.5	0.37–0.41	3 basic, 1 grey
Morotai—(dark-plumaged	2 F	116–123	105–115	0.91–0.94	16.3–16.6	6.2–6.6	0.38–0.40	2 muted
infaustus phenotype)
Australian Brush Cuckoo (tymbonomus) ‒ Timor and satellite islands
Timor, Rote,?Wetar—(tym-	3 M	123–130	119–130	0.97–1.02	16.2–16.9	6.7	‒	2 basic
bonomus)	3 F	123–126	114–120	0.90–0.96	‒	‒	‒	1 basic, 2 barred
	4 unsexed	121–128	111–124	0.92–1.00	14.7–16.3	6.2–7.0	0.42–0.44	4 basic
Australian Brush Cuckoo (variolosus)—Australian variolosus spring and summer (mid-September to mid-March) sampling
Kimberley Division east to	14 M	125–132	(101–)104–	(0.80–)0.83–0.88	(15.6–)16.0–	(5.7–)6.3–7.2	(0.34–)0.37–0.43	14 basic
Victoria River, 14°–19°S	4 F	(122–)126–132	113(–118)	0.81–0.85	17.4(–17.6)	6.2–6.9	0.39–0.43	3 barred, 1 muted
			(99–)103–110		15.7–16.6
Arnhem Land between Victo-	30 M	(119–)121–130	(98–)103–	(0.80–)0.82–	(15.3–)15.9–	(5.8–)6.2–7.1(–	(0.35–)0.37–	27 basic, 1 grey
ria (Northern Territory) and	9 F	(–135)	113(–120)	0.88(–0.91)	17.2(–17.7)	7.7)	0.42(–0.46)	4 basic, 3 barred, 2
Leichhardt (Gulf of Carpen-		120–128	97–107	0.80–0.86	15.0–16.3(–17.8)	(5.1–)6.2–7.4	0.38–0.45(–0.48)	muted
taria) Rivers, 12°–18°S
South Torres Strait / Cape York	15 M	122–128(–131)	101–110(–113)	0.81–0.88	16.0–18.0(–18.4)	6.4–7.3	0.37–0.43	14 basic
Peninsula, 10°–15°S (south	4 F	120–123	100–105	0.83–0.86	16.6–17.8	7.0–7.2	0.40–0.43	3 basic, 1 muted
to middle Normanby–lower
Mitchell Rivers)
	sample size				men (mm)	illa (mm)	maxilla/culmen
							length ratio
South Torres Strait / Cape York	10 M	(118–)122–	100–115	(0.80–)0.82–0.89	16.2–18.9(–19.2)	6.3–7.3	(0.35–)0.38–0.42	10 basic
Peninsula, 10°–16°S (south to	6 F	135(–138)	(94–)96–109	(0.77–)0.80–0.86	15.8–17.3	5.9–6.8	0.37–0.41	1 basic, 4 barred, 1
Annan–Palmer–lower Mitchell		121–128						muted
Rivers)
	5 M	128–138	102–111	0.79–0.82	15.9–17.4	6.7–6.9	0.39–0.41	5 basic
Northeast Queensland, 16°–20°	7 F	119–127(–136)	(96–)99–112	(0.78–)0.80–0.86	16.1–18.2(–18.8)	6.0–7.5	0.37–0.40	4 basic, 1 barred, 2
S (Annan–Palmer Rivers to								muted
north Flinders–Burdekin River
drainages)
	1 M	138	121	0.88	18.7	6.5	0.35	1 basic
Central east Queensland,	2 F	124–127	102–107	0.82–0.84	17.0–17.5	7.1–7.2	0.41–0.42	2 barred
20°–27°S (south Flinders–Bur-
dekin River drainages to upper
Brisbane– Balonne River
drainages)
	nil	‒	‒	‒	‒	‒	‒	‒
Southeast Queensland to
central east New South Wales,
27°–33°S (Brisbane–Balonne
River drainages to Hunter
River)
Southeast Australia, 33°–39°S	1 M	135	112	0.83	‒	6.9	‒	1 basic
(Hunter River to all east Vic-	2 F	132–133	114	0.86	17.1–17.6	7.5–7.6	0.43–0.44	1 basic, 1 muted
toria)
Australian Brush Cuckoo (variolosus)—New Guinea region
	sample size				men (mm)	illa (mm)	maxilla/culmen
							length ratio
small islands southeast of	9 M	(113–)114–118	(99–)101–108	0.86–0.92	16.5–17.0(–17.9)	6.3–7.2	0.37–0.41(0.44)	7 basic, 2 grey
south Moluccas (Seram Laut,	4F	114–120	96–103	0.81–0.88	15.2–17.2	6.5–7.3	0.38–0.44	2 basic, 2 barred
Watubela & Kai Archipela-
gos)—(infaustus)
West Papuan Islands, Aru	30 M	(113–)116–	(100–)103–	(0.85–)0.86–	(15.0–)15.6–	(5.1–)5.5–7.2(–	(0.34–)0.36–	22 basic, 5 grey
Islands, Vogelkop, its neck	17 F	123(–125)	113(–116)	0.93(–0.96)	17.5(–18.1)	7.5)	0.42(–0.45)	4 basic, 8 barred, 4
and north New Guinea east to		(110–)113–	(90–)93–104(–	(0.80–)0.82–	(15.1–)15.8–	(6.2–)6.5–7.3(–	(0.36–)0.38–	muted
Astrolabe Bay—(infaustus)		118(–121)	110)	0.89(–0.92)	17.8(–19.0)	7.5)	0.43(–0.47)
Numfor Island—(obscuratus)	6 M	123–130	111–117	0.86–0.94	17.7–19.3	6.2–7.1	0.34–0.39	4 basic, 2 grey
	1 F	124	‒	‒	15.5	6.3	0.41	1 barred
Schouten Islands—(infaustus)	4 M	113–118	106–108	0.90–0.95	16.0–18.1	6.7–7.4	0.41–0.44	4 basic
	1 F	115	107	0.93	17.2	7.4	0.43	1 basic
Manam and Karkar Islands—	8 M	117–126	(102–)110–117	0.88–0.98	(16.8–)17.1–18.8	(6.5–)7.2–7.9(–	(0.35–)0.41–0.45	8 basic
(infaustus)	8 F	118–123	100–113	0.84–0.89(–0.93)	17.5–19.8	8.1)	0.38–0.42	1 basic, 5 barred, 2
						7.2–7.9		muted
D’Entrecasteaux Archipela-	7 M	124–131(–138)	105–117(–123)	0.87–0.94	17.8–19.0	6.8–7.7	0.36–0.41	6 basic, 1 grey
go—(fortior)	1 F	127	112	0.88	19.0	7.3	0.38	1 barred
Huon and Papuan Peninsulas	53 M	(112–)114–	(98–)102–	(0.83–)0.86–	(14.9–)15.5–	(5.9–)6.2–7.3(–	(0.34–)0.37–	46 basic, 5 grey
and all south New Guinea	13 F	123(–126)	110(–112)	0.91(–0.93)	18.0(–18.7)	7.9)	0.44(–0.46)	6 basic, 7 barred
(excl. Trans–Fly region)—(or-		(113–)115–	(99–)102–112	(0.84–)0.86–0.89	15.2–17.6(–18.4)	6.4–7.4	(0.36–)0.39–0.46
eophilus)		122(–124)
	sample size				men (mm)	illa (mm)	maxilla/culmen
							length ratio
Trans–Fly region, south New	15 M	118–123(–125)	(101–)104–	(0.84–)0.87–0.94	(15.9–)16.6–	(5.5–)6.0–7.0(–	0.34–0.39(–0.42)	14 basic, 1 grey
Guinea—(oreophilus)	11 F	115–121(–124)	113(–115)	(0.81–)0.85–0.89	18.1(–18.4)	7.3)	0.36–0.43	7 basic, 3 barred, 1
			98–106(–109)		(15.7–)16.4–17.6	6.4–6.9(–7.3)		muted
Australian Brush Cuckoo (variolosus) group—North Melanesian Islands
Bismarck Archipelago (New	21 M	(119–)121–	(118–)122–	(0.98–)1.01–	(15.2–)15.8–	(7.0–)7.2–7.9(–	(0.38–)0.41–	17 basic, 4 grey
Britain and satellite islands)—	10 F	128(–130)	134(–138)	1.06(–1.10)	17.9(–18.8)	8.5)	0.47(–0.49)	3 basic, 6 barred, 1
(macrocercus)		(117–)120–127	(115–)118–	(0.94–)0.96–	(15.9–)16.4–	6.7–7.8(–8.1)	(0.36–)0.40–	muted
			126(–131)	1.02(–1.07)	18.2(–18.8)		0.47(–0.50)
Bismarck Archipelago (New	17 M	117–125(–127)	(118–)121–	(0.98–)1.00–1.05	(15.8–)16.5–	(6.9–)7.2–7.8	(0.39–)0.41–	13 basic, 4 grey
Ireland, central and south	4 F	116–121	128(–132)	0.99–1.04(–1.09)	17.8(–19.2)	7.0–7.7	0.45(–0.47)	2 basic, 2 barred
Hibernian Islands)—(mac-			116–127		16.1–17.2		0.43–0.45
rocercus)
Bismarck Archipelago (North	2 M	126–132	132–136	1.03–1.05	20.0–20.1	8.1–8.3	0.41	2 grey
Hibernian islands)—(tabaren-
sis)
Bismarck Archipelago (New	7 M	116–123	120–126		(16.6–)17.2–19.1	7.3–8.0	0.40–0.45(–0.46)	7 basic (all grey)
Hanover)—(websteri)	3 unsexed	115–121	114–123	1.02–1.06	15.9–17.6	7.0–7.3	0.41–0.44	3 basic (all grey)
0.99–1.05
Manus Brush Cuckoo (blandus)—Admiralty Islands
	sample size				men (mm)	illa (mm)	maxilla/culmen
							length ratio
Admiralty Islands (Manus)—	10 M	(104–)106–	(102–)107–	(0.96–)0.98–	(15.5–)16.0–	(6.9–)7.3–7.8	0.42–0.48	10 basic
(blandus)	7 F	115(–117)	114(–118)	1.02(–1.04)	17.8(–18.2)	7.0–7.5	0.42–0.45(–0.47)	5 basic, 2 barred
		109–113	107–116	0.96–1.02	(15.4–)16.4–17.2
Solomons Brush Cuckoo (addendus)—Solomon Islands
Solomon Islands—(addendus)	28 M	(115–)117–	(124–)128–	(1.07–)1.09–	(16.2–)16.7–	(6.1–)6.5–7.6(–	(0.37–)0.39–	27 basic; 1 grey
	13 F	123(–126)	140(–144)	1.14(–1.19)	18.2(–18.5)	8.1)	0.45(–0.47)	basic 10, 2 barred,
		(111–)113–	(122–)126–	(1.05–)1.09–	(15.5–)16.0–	(6.1–)6.7–7.5(–	(0.39–)0.41–0.46	1 muted
		120(–125)	136(–142)	1.15(–1.19)	17.2(–17.4)	7.8)