Relationship between developmental modes, flight styles, and wing morphology in birds

Figures & data

Table I. Basic statistic estimates of developmental mode – flight style groups: n = number of species in the group; mean = mean value; SD = standard deviation; CF = continuous flapping; FG = flapping and gliding; PT = passerine type; FS = flapping and soaring flight styles; hu = humerus; ul = ulna; mn = manus; fprim = primary feather; ta = total arm length.

			Developmental mode – fligh style group
			Altricial			Precocial	Intermediate
Row	Trait	Statistic	CF	FG	PT	CF	CF	FG	FS
1	Wingspan (m)	n	10	9	50	32	20	10	22
		mean	0.742	0.352	0.262	0.726	1.023	0.822	1.417
		SD	0.26	0.07	0.07	0.34	0.35	0.16	0.52
2	Wing area (m^2)	n	10	9	50	32	20	10	22
		mean	0.106	0.014	0.013	0.078	0.148	0.094	0.306
		SD	0.07	0.01	0.01	0.11	0.09	0.04	0.21
3	Body mass (g)	n	10	9	83	67	38	10	29
		mean	354.9	30.4	37.2	1045.5	726.8	326.2	1876.5
		SD	300.85	21.25	46.52	1697.8	771.16	215.08	2080.02
4	hu (mm)	n	6	4	61	53	31	4	20
		mean	49.3	14.0	21.0	87.9	92.8	60.5	145.9
		SD	13.41	1.55	8.52	52.98	41.24	18.11	67.94
5	ul (mm)	n	6	4	61	53	31	4	20
		mean	60.7	20.7	25.6	82.4	101.5	70.2	168.8
		SD	15.66	1.78	9.84	50.34	48.90	21.78	78.15
6	mn (mm)	n	6	4	61	53	31	4	20
		mean	48.6	21.2	20.5	69.7	66.6	65	104.1
		SD	18.50	5.13	9.76	42.34	28.63	18.67	37.62
7	fprim (mm)	n	6	4	61	53	31	4	20
		mean	181.0	112.0	77.4	201.0	224.6	211.0	322.6
		SD	61.83	23.05	30.92	104.06	80.99	57.38	126.69
8	hu/ul	n	6	4	61	53	31	4	20
		mean	0.810	0.673	0.818	1.068	0.941	0.864	0.865
		SD	0.033	0.022	0.057	0.120	0.156	0.020	0.047
9	hu/ta	n	6	4	61	53	31	4	20
		mean	0.313	0.251	0.314	0.363	0.356	0.309	0.345
		SD	0.013	0.031	0.021	0.038	0.037	0.003	0.026
10	ul/ta	n	6	4	61	53	31	4	20
		mean	0.387	0.372	0.385	0.341	0.382	0.358	0.399
		SD	0.017	0.043	0.022	0.025	0.036	0.006	0.021
11	mn/ta	n	6	4	61	53	31	4	20
		mean	0.300	0.377	0.301	0.296	0.262	0.333	0.257
		SD	0.027	0.074	0.036	0.051	0.050	0.005	0.044
12	fprim/ta	n	6	4	61	53	31	4	20
		mean	1.135	2.003	1.175	0.878	0.920	1.088	0.797
		SD	0.140	0.352	0.195	0.168	0.261	0.156	0.159

Table II. Association of evolutionary histories of flight style and developmental mode of birds: D = overall measure of evolutionary association; d_ij = measure of association for specific pairs of developmental mode and flight style; p = significance value of estimates; CF = continuous flapping; FG = flapping and gliding; FS = flapping and soaring; PT = passerine type flight styles.

Groups	Estimate	Observed estimate	p
General association	D	0.879	<0.001
Altricial – CF	dij	−0.145	0.001
Altricial – FG	dij	0.006	0.061
Altricial – FS	dij	−0.047	0.001
Altricial – PT	dij	0.185	<0.001
Precocial – CF	dij	0.150	<0.001
Precocial – FG	dij	−0.012	0.043
Precocial – FS	dij	−0.043	0.009
Precocial – PT	dij	−0.095	<0.001
Intermediate – CF	dij	−0.006	0.091
Intermediate – FG	dij	0.006	0.088
Intermediate – FS	dij	0.090	0.003
Intermediate – PT	dij	−0.090	0.001

Figure 1. Phylogenetic hypothesis of relationships amongst studied birds (Hackett et al. 2008) showing modes of development and flight styles: sp = number of the studied species; CF = continuous flapping; FG = flapping and gliding; FS = flapping and soaring; PT = passerine type flight styles.

Table III. The effect of body mass estimated with the analysis of best-fit PGLS models with the AICc differences (Δ) and multiple R²: dev.mode = developmental mode, flight = flight style; hu = humerus; ul = ulna; mn = manus; fprim = primary feather length. All models were significant (p_FDR < 0.03).

	Y
Regression formula	hu	ul	mn	fprim	wingspan	wing area
AICc differences	Δ	Δ	Δ	Δ	Δ	Δ
log(Y) ~ dev.mode	397.55	322.27	143.61	159.81	286.09	257.02
log(Y) ~ flight	399.59	323.44	144.00	156.16	280.09	251.42
log(Y) ~ log(mass) + dev.mode	0	0	0	2.78	0	0
log(Y) ~ log(mass) + flight	5.66	7.20	0.53	0	2.21	7.75
Multiple R^2	R^2	R^2	R^2	R^2	R^2	R^2
log(Y) ~ dev.mode	10.4	8.4	5.3	3.9	10.6	10.0
log(Y) ~ flight	10.4	8.9	6.2	6.9	15.3	14.5
log(Y) ~ log(mass) + dev.mode	90.4	85.0	58.0	60.5	87.1	84.2
log(Y) ~ log(mass) + flight	90.2	84.6	58.4	61.6	87.1	83.6

Figure 2. Phylogenetic discriminant analysis of developmental mode and flight style groups by wingspan (plots (a) and (c)) and wing elements (plots (b) and (d)) data sets. Convex hulls, which connect the most distant points of each group, are shown. Other group points were omitted from the plot. The length of arrows represents the contribution of the traits to discrimination. altr. = altricial; interm. = intermediate; prec. = precocial developmental modes; CF = continuous flapping; FG = flapping and gliding; FS = flapping and soaring; PT = passerine type flight styles; hu = humerus; ul = ulna; mn = manus; fprim = primary feather length.

Figure 3. Standard discriminant analysis of developmental mode and flight style groups by wingspan (plots (a) and (c)) and wing elements (plots (b) and (d)) data sets. Convex hulls, which connect the most distant points of each group, are shown. Other group points were omitted from the plot. The length of arrows represents the contribution of the traits to discrimination. altr. = altricial; interm. = intermediate; prec. = precocial developmental modes; CF = continuous flapping; FG = flapping and gliding; FS = flapping and soaring; PT = passerine type flight styles; hu = humerus; ul = ulna; mn = manus; fprim = primary feather length.

Figure 4. Cleveland’s dot chart of the relative length of wing elements. Each point represents a species. Developmental modes and flight styles are marked with different types of points. altr = altricial; interm = intermediate; prec= precocial developmental modes; CF = continuous flapping; FG = flapping and gliding; FS = flapping and soaring; PT = passerine type flight styles; hu = humerus; ul = ulna; mn = manus; fprim = primary feather; ta = total arm length.

Hackett SJ, Kimball RT, Reddy S, Bowie RCK, Braun EL, Braun MJ, Chojnowski JL, Cox WA, Han K-L, Harshman J, Huddleston CJ, Marks BD, Miglia KJ, Moore WS, Sheldon FH, Steadman DW, Witt CC, Yuri T. 2008. A phylogenomic study of birds reveals their evolutionary history. Science 320:1763–1768. DOI:10.1126/science.1157704.

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