Comparative wood anatomy of species of <i>Psychotria</i> L. (Rubiaceae) in Atlantic Rainforest remnants of Rio de Janeiro State, Brazil

Psychotria is the largest Rubiaceae genus and one of the most abundant in the Atlantic Rainforest of Rio de Janeiro State. The present work aims to characterize and compare the wood of nine of these species. ThePsychotria wood were characterized by: slightly distinct growth rings; diffuse porosity; solitary vessels or on radial multiples of 2-6 or clusters of 3-5 vessel elements, with terminal and lateral simple perforation plates and vestured and alternate intervessel and vessel-ray pits; septate fiber-tracheids; and rare axial parenchyma. Although, the wood anatomy of thePsychotria may be considered homogeneous, the statistical analyses, based on qualitative and quantitative features, allowed the segregation of the species and indicated the importance of habit (arboreous or shrubby) on the wood anatomy of the species. It is worth mentioning that the sampling was realized in only one studied site (Ilha Grande), which contributed to the conclusion that the wood characteristics are more related to the specificities of each studied taxon than to with environmental variations.

Keywords:
Ilha Grande; Psychotriae; Rubioideae; Tropical Forest; wood diagnostic characters

Authorship

Jonas B. C. Marques

Setor de Biologia Vegetal, Laboratório de Biologia Celular e Tecidual, Universidade Estadual do Norte Fluminense Darcy Ribeiro, 28013-602, Campos dos Goytacazes, RJ, Brazil Universidade Estadual do Norte FluminenseBrazilCampos dos Goytacazes, RJ, Brazil Setor de Biologia Vegetal, Laboratório de Biologia Celular e Tecidual, Universidade Estadual do Norte Fluminense Darcy Ribeiro, 28013-602, Campos dos Goytacazes, RJ, Brazil

Cátia H. Callado

Departamento de Biologia Vegetal, Instituto de Biologia Roberto Alcantara Gomes, Universidade do Estado do Rio de Janeiro, 20550-900 Rio de Janeiro, RJ, Brazil Universidade do Estado do Rio de JaneiroBrazilRio de Janeiro, RJ, Brazil Departamento de Biologia Vegetal, Instituto de Biologia Roberto Alcantara Gomes, Universidade do Estado do Rio de Janeiro, 20550-900 Rio de Janeiro, RJ, Brazil

Guilherme R. Rabelo

Sebastião J. da Silva Neto

Maura da Cunha ^** Corresponding author: maurauenf@gmail.com

* Corresponding author: maurauenf@gmail.com

SCIMAGO INSTITUTIONS RANKINGS

Figures | Tables

Figures (6)
Tables (2)

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Figure 1
Light microscopy micrographs of Psychotria woods sections. A: P. deflexa, distinct growth rings, diffuse porosity and vessel arrangement in a radial pattern (e = initial wood, l = later wood). B: P. vellosiana, diffuse porosity and vessel arrangement in a radial pattern. C: P. leiocarpa, vessels with circular to oval outline, libriform fibers thin- to thick-walled and prismatic crystals inside ray cells (arrowhead). D: P. nuda, multiseriate and aggregate rays with uniseriate portions (arrowhead). E: P. hoffmannseggiana, uniseriate and biseriate rays, and fused rays (arrowhead). F: P. vellosiana, multiseriate rays with sheath cells and septate fiber-tracheids (arrow). G: P. ruelliifolia, rays composed of procumbent, square and upright cells. H: P. hoffmannseggiana, rays composed of square and/or upright cells. I: P. carthagenensis, rays composed of procumbent, square and upright cells, and raphides crystals inside procumbent ray cells (arrowhead). Scale bar: A, B and D to I = 100 µm; C = 20 µm.

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Figure 2
Light microscopy micrographs. A: P. brasiliensis, fiber-tracheids. B: P. hoffmannseggiana, septate fiber-tracheids (arrowhead), simple fiber pits and fiber- wall thickness. C: P. deflexa, vessel elements (V). D: P. vellosiana, appendices and simple perforation plates on terminal and lateral wall of vessel (*). E: P. carthagenensis, intervessel vestured pits. F:P. vellosiana, ray-vessel vestured pits. G:P. carthagenensis, raphides crystals inside the upright ray cell (arrowhead). Scale bar: A and C = 100 µm; B, D, E, F and G = 20 µm.

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Figure 3
Scanning electron microscopy micrographs. A: P. brasiliensis, vestured intervessel pits viewed from outside the vessel. B: P. pubigera, vestured intervessel pits viewed from inside vessel. C: P. ruelliifolia, vessel with a simple perforation plate on terminal and lateral wall (*) and vessel appendices. D: P. deflexa, septate fiber-tracheids (arrowhead) with simple fiber pits (arrow). E: P. pubigera, ray composition of only square and upright cells. F: P. ruelliifolia, raphides crystals coming out from upright ray cells (arrowhead). Scale bar: A and B = 10 µm; C, D and F = 50 µm; E = 200 µm.

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Figure 4
Cluster similarity analysis of Psychotria andCoussarea qualitative characters.

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Figure 5
Principal Component Analysis of Psychotria andCoussarea qualitative characters.

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Figure 6
Principal Component Analysis of Psychotriaquantitative characters. A = Vessels/mm², B = vessel length, C = tangential diameter of vessel, D = radial diameter of vessel, E = vessel wall thickness, F = intervessel pits, G = vessel-ray pits, H = fiber diameter, I = fiber lumina, J = fiber length, L = fiber wall thickness, M = fiber pits, N = rays/mm', O = ray wide, P = ray length.

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Table 1
Characteristics of Psychotria species studied. N° Reg. = registration number in the UENF Wood collection, DBH = diameter at breast height in centimeters, H = height in meters, VL = vulnerability index, ME = mesomorphy index.

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Table 2
Quantitative wood anatomical features of the ninePsychotria studied. bar = P. hoffmannseggiana; bra = P. brasiliensis ; car = P. carthagenensis; rue = P. ruelliifolia; def = P. deflexa; lei = P. leiocarpa; nud = P. nuda; pub = P. pubigera; vell = P. vellosiana. Q25 = quartile 25, µ = medians, Q75 = quartile 75, KW = Kruskal-Wallis test between the medians.

Figure 1 Light microscopy micrographs of Psychotria woods sections. A: P. deflexa, distinct growth rings, diffuse porosity and vessel arrangement in a radial pattern (e = initial wood, l = later wood). B: P. vellosiana, diffuse porosity and vessel arrangement in a radial pattern. C: P. leiocarpa, vessels with circular to oval outline, libriform fibers thin- to thick-walled and prismatic crystals inside ray cells (arrowhead). D: P. nuda, multiseriate and aggregate rays with uniseriate portions (arrowhead). E: P. hoffmannseggiana, uniseriate and biseriate rays, and fused rays (arrowhead). F: P. vellosiana, multiseriate rays with sheath cells and septate fiber-tracheids (arrow). G: P. ruelliifolia, rays composed of procumbent, square and upright cells. H: P. hoffmannseggiana, rays composed of square and/or upright cells. I: P. carthagenensis, rays composed of procumbent, square and upright cells, and raphides crystals inside procumbent ray cells (arrowhead). Scale bar: A, B and D to I = 100 µm; C = 20 µm.

Figure 2 Light microscopy micrographs. A: P. brasiliensis, fiber-tracheids. B: P. hoffmannseggiana, septate fiber-tracheids (arrowhead), simple fiber pits and fiber- wall thickness. C: P. deflexa, vessel elements (V). D: P. vellosiana, appendices and simple perforation plates on terminal and lateral wall of vessel (). E: P. carthagenensis,* intervessel vestured pits. F:P. vellosiana, ray-vessel vestured pits. G:P. carthagenensis, raphides crystals inside the upright ray cell (arrowhead). Scale bar: A and C = 100 µm; B, D, E, F and G = 20 µm.

Figure 3 Scanning electron microscopy micrographs. A: P. brasiliensis, vestured intervessel pits viewed from outside the vessel. B: P. pubigera, vestured intervessel pits viewed from inside vessel. C: P. ruelliifolia, vessel with a simple perforation plate on terminal and lateral wall () and vessel appendices. D: P. deflexa, septate fiber-tracheids (arrowhead) with simple fiber pits (arrow). E: P. pubigera,* ray composition of only square and upright cells. F: P. ruelliifolia, raphides crystals coming out from upright ray cells (arrowhead). Scale bar: A and B = 10 µm; C, D and F = 50 µm; E = 200 µm.

Figure 4 Cluster similarity analysis of Psychotria andCoussarea qualitative characters.

Figure 5 Principal Component Analysis of Psychotria andCoussarea qualitative characters.

Figure 6 Principal Component Analysis of Psychotriaquantitative characters. A = Vessels/mm², B = vessel length, C = tangential diameter of vessel, D = radial diameter of vessel, E = vessel wall thickness, F = intervessel pits, G = vessel-ray pits, H = fiber diameter, I = fiber lumina, J = fiber length, L = fiber wall thickness, M = fiber pits, N = rays/mm', O = ray wide, P = ray length.

Table 1 Characteristics of Psychotria species studied. N° Reg. = registration number in the UENF Wood collection, DBH = diameter at breast height in centimeters, H = height in meters, VL = vulnerability index, ME = mesomorphy index.

Species	N° Reg.	Habit	DBH	H	VL	ME
P. hoffmannseggiana	112, 113, 155, 156 e 157	Shrubby	4.2 (±1.09)	1.8 (±0.44)	0.127	141.4
P. brasiliensis	134, 135, 136, 137 e 138	Arborecent	11.8 (±2.04)	6.4 (±1.51)	0.592	678.3
P. carthagenensis	144, 145, 146, 147 e 148	Arborecent	18.4 (±2.88)	12.8 (±2.16)	2.172	2115.9
P. ruelliifolia	107, 115, 152, 153 e 154	Shrubby	6.4 (±1.14)	2.3 (±0.27)	0.385	344.7
P. deflexa	110, 111, 159, 160 e 161	Shrubby	4.6 (±1.14)	2.3 (±0.57)	0.156	125.5
P. leiocarpa	106, 114, 158, 162 e 163	Shrubby	5.6 (±1.34)	1.9 (±0.41)	0.136	96.3
P. nuda	108, 130, 131, 132 e 133	Arborecent	9.8 (±1.48)	5.6 (±1.51)	0.148	132.9
P. pubigera	109, 116, 149, 150 e 151	Arborecent	7 (±1.87)	6 (±1.58)	0.315	307.6
P. vellosiana	139, 140, 141, 142, e 143	Arborecent	18.8 (±2.58)	10 (±2.12)	2.653	2318.7

Table 2 Quantitative wood anatomical features of the ninePsychotria studied. bar = P. hoffmannseggiana; bra = P. brasiliensis ; car = P. carthagenensis; rue = P. ruelliifolia; def = P. deflexa; lei = P. leiocarpa; nud = P. nuda; pub = P. pubigera; vell = P. vellosiana. Q25 = quartile 25, µ = medians, Q75 = quartile 75, KW = Kruskal-Wallis test between the medians.

Species		bar	bra	car	rue	def	lei	nud	pub	vel
Vessels
	Vessels/mm² (N 50)
		Q25	175	59	24	80	130	161	152	95	25
		µ	181	66	26	91	141	178	178	102	27
		Q75	189	75	28	98	152	193	190	114	30
	KW	a	be	d	b	c	a	ac	bc	de
	Length(µm) (N 125)
		Q25	931	1048	909	783	710	620	784	840	782
		µ	1120	1141	965	888	772	734	921	956	815
		Q75	1245	1277	1000	984	838	789	993	1068	887
	KW	a	a	d	be	c	c	bde	bd	e
	Tang. Diameter (µm) (N 125)
		Q25	19.4	32.2	48.1	28.4	20.4	21.4	22.3	28.5	60.5
		µ	28.0	36.8	53.6	33.2	26.2	24.0	24.9	32.9	67.8
		Q75	34.6	41.0	60.8	38.5	28.0	25.8	28.4	35.8	74.7
	KW	a	b	d	b	a	ac	c	b	d
	Rad. diameter (µm) (N 125)
		Q25	22.3	32.0	60.5	33.0	24.0	25.8	24.0	26.7	62.2
		µ	25.1	36.5	65.8	37.3	27.6	27.7	26.7	29.4	71.1
		Q75	28.5	43.2	71.1	41.8	30.2	30.4	30.2	32.9	79.2
	KW	a	b	d	b	ac	ac	ac	c	d
	Wall thickness (µm) (N 125)
		Q25	2.5	1.8	3.2	2.5	2.6	2.7	1.8	2.1	1.6
		µ	2.8	2.0	3.6	2.9	3.0	3.0	2.1	2.4	1.7
		Q75	3.2	2.5	4.0	3.2	3.4	3.4	2.5	2.7	2.0
	KW	a	b	c	a	a	a	b	b	d
	Intervessel pits (µm) (N 125)
		Q25	2.8	3.4	3.7	3.5	2.8	2.9	3.3	2.8	4.6
µ		3.1	3.9	4.1	4.2	3.1	3.2	3.6	3.0	4.9
Q75		3.5	4.3	4.6	4.8	3.5	3.4	3.9	3.3	5.3
KW	a	bd	b	b	ac	a	d	ac	e
Vessel-ray pits (µm) (N 125)
	Q25	2.9	3.3	3.8	3.6	2.9	2.8	2.9	2.8	4.5
	µ	3.1	3.8	4.2	4.1	3.2	3.1	3.3	3.0	4.8
	Q75	3.5	4.3	4.6	4.7	3.5	3.5	3.8	3.3	5.2
KW	ad	b	c	bc	ad	ad	a	d	e
Fiber
	Diameter (µm) (N 125)
		Q25	22.8	29.4	30.9	19.6	21.3	19.6	22.8	24.4	28.7
		µ	23.4	32.0	34.8	22.1	22.2	21.3	27.2	26.9	33.6
		Q75	31.1	37.3	37.1	25.2	28.2	22.4	29.5	28.6	38.4
	KW	a	d	d	b	c	b	ac	ac	d
	Lumen diam. (µm) (N 125)
		Q25	14.7	16.8	17.8	13.8	12.9	11.2	16.4	14.4	19.6
		µ	19.8	18.9	19.4	15.1	15.6	12.9	18.8	16.3	24.8
		Q75	23.1	25.1	22.3	17.5	19.4	14.3	21.4	17.7	30.5
	KW	a	a	ac	b	b	d	a	b	c
	Length (µm) (N 125)
		Q25	1388	1420	1459	1115	1046	1002	1278	1330	1199
		µ	1506	1543	1553	1249	1170	1063	1363	1411	1271
		Q75	1647	1614	1711	1342	1289	1150	1481	1514	1388
	KW	a	a	a	b	b	d	c	c	b
	Wall thickness (µm) (N 125)
		Q25	3.77	5.68	6.32	2.88	4.09	3.58	3.39	4.81	3.54
		µ	4.2	6.4	7.3	3.5	4.5	4.1	3.8	5.3	4.0
		Q75	4.6	7.2	7.9	4.2	5.0	4.8	4.2	5.6	4.4
KW	a	c	c	b	a	a	ab	d	ab
Pits (µm) (N 125)
	Q25	3.20	4.36	3.68	2.71	3.48	2.43	2.85	2.14	6.58
	µ	3.56	4.88	4.19	3.34	4.02	2.79	3.36	2.38	7.17
	Q75	3.9	5.9	4.7	3.9	4.7	3.2	3.9	2.9	8.3
KW	a	d	b	a	b	c	a	c	e
Rays
	Rays/mm' (N 100)
		Q25	8	4	2	3	3	5	5	3	4
		µ	8	5	3	4	3	5	5	4	4
		Q75	9	5	3	4	4	6	6	4	5
	KW	a	d	c	b	bc	d	d	bc	e
	Height (µm) (N 125)
		Q25	475	924	768	516	466	411	475	406	464
		µ	597	1187	987	584	553	510	548	503	689
		Q75	750	1351	1102	742	685	633	747	668	823
	KW	a	b	b	a	a	a	a	a	a
Width (µm) (N 125)
	Q25	23.2	37.3	65.6	37.4	23.1	23.2	33.8	24.0	58.7
	µ	28.4	44.5	69.5	42.8	26.7	26.7	39.2	26.7	69.3
Q75	34.0	52.8	76.5	50.1	30.2	30.3	48.9	29.3	78.2
KW	a	b	c	b	a	a	b	a	c

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Comparative wood anatomy of species of <i>Psychotria</i> L. (Rubiaceae) in Atlantic Rainforest remnants of Rio de Janeiro State, Brazil

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[1] * Corresponding author: maurauenf@gmail.com