Morphoanatomical leaf traits of herbaceous species raised in a cronosequence of naturally regenerated areas of tropical deciduous forest

Figures & data

Table 1. Average soil moisture, vapor pressure deficits, and photosynthetic photon flux densities in sites with different regeneration periods (22 years, R22; 44 years, R44; and more than 60 years, R > 60) after land abandonment, in a northeastern Brazilian seasonally dry tropical forest (Santa Teresinha, Paraíba state, Brazil).

	R22	R44	R >60
Soil moisture (%)	6.66	9.33	9.00
Vapor pressure deficits (kPa)	2.96	1.15	1.52
Photosynthetic photon flux densities (µmol m^−2 s^−1)	1600	1300	1500

Figure 1. Morphological characteristics of the leaves of Mesosphaerum suaveolens (A), Stachytarpheta sanguinea (B) and Jacquemontia evolvuloides (C) harvested from individuals growing in sites with different regeneration periods after land abandonment in a northeastern Brazilian seasonally dry tropical forest (Santa Teresinha, Paraíba state, Brazil).

Table 2. Morphological characteristics in leaves of three herbaceous species (one tall, one short, and one climbing species) growing in forest sites with different regeneration periods after land abandonment in a northeastern Brazilian seasonally dry tropical forest (Santa Teresinha, Paraíba state, Brazil).

Species	Regeneration periods
	R22	R44	R >60
Leaf area (cm2)
Mesosphaerum suaveolens	11.9 ± 3.8^a	12.6 ± 5.6^a	11.9 ± 3.3^a
Stachytarpheta sanguinea	9.45 ± 3.24^a	6.97 ± 1.77^b	5.9 ± 1.3^b
Jacquemontia evolvuloides	6.83 ± 2.18^a	7.38 ± 1.73^a	7.97 ± 2.58^a
Specific leaf area (cm^2 g^−1)
Mesosphaerum suaveolens	196.3 ± 20.3^a	164.02 ± 27.7^a	156.2 ± 27.2^a
Stachytarpheta sanguinea	304.1 ± 41.7^b	383.2 ± 54.4^a	231.27 ± 28.9^c
Jacquemontia evolvuloides	87.1 ± 10.8^c	191.6 ± 18.01^b	240.61 ± 26.2^a
Leaf blade length (cm)
Mesosphaerum suaveolens	4.49 ± 0.86^b	5.38 ± 0.99^a	4.98 ± 0.70^ab
Stachytarpheta sanguinea	5.38 ± 1.04^a	4.65 ± 0.64^b	4.250 ± 0.47^b
Jacquemontia evolvuloides	3.55 ± 0.62^a	3.66 ± 0.44^a	3.38 ± 0.51^a
Leaf blade width (cm)
Mesosphaerum suaveolens	3.41 ± 0.62^a	3.67 ± 0.73^a	3.59 ± 0.55^a
Stachytarpheta sanguinea	2.38 ± 0.49^a	2.03 ± 0.33^b	1.87 ± 0.26^b
Jacquemontia evolvuloides	2.68 ± 0.48^a	2.752 ± 0.38^a	2.82 ± 0.54^a

Averages in the column followed by the same letter are not significantly different by the Tukey test at 5% probability level (n = 50 ± standard deviation). R22 = site under regeneration for 22 years; R44 = site under regeneration for 44 years; R > 60 = site under regeneration for over 60 years.

Figure 2. Mesosphaerum suaveolens harvested from individuals growing in sites with different regeneration periods after land abandonment in a northeastern Brazilian seasonally dry tropical forest (Santa Teresinha, Paraíba state, Brazil). Diacytic stomata; tector and glandular trichomes in frontal view (A-C); cross-section of leaf blade (D-E) and midrib (F). (D) GT = trichomes; (E) PP = palisade parenchyma; LP = lacunar parenchyma; ep aba = abaxial epidermis; ep ada = adaxial epidermis; (F) VB = vascular bundle; ep = epidermis; FP = fundamental parenchyma; Phl= phloem; xy = xylem; col = collenchyma;; sto = stomata.

Figure 3. Stachytarpheta sanguinea harvested from individuals growing in sites with different regeneration periods after land abandonment in a northeastern Brazilian seasonally dry tropical forest (Santa Teresinha, Paraíba state, Brazil). Diacytic stomata on the adaxial leaf surface; hook-shaped tector trichomes; multicellular tector trichomes; tector trichomes at high density in the vein (A-D) in frontal view; cross-section of leaf blade (E) and midrib (F). (E) PP = palisade parenchyma; LP = lacunar parenchyma; TT = tector trichomes; sto = stomata; ep aba = abaxial epidermis; ep ada = adaxial epidermis; (F) ep = epidermis; FP = fundamental parenchyma; phl = phloem; xy = xylem; col = collenchyma; TT = tector trichomes.

Figure 4. Jacquemontia evolvuloides leaves harvested from individuals growing in sites with different regeneration periods after land abandonment in a northeastern Brazilian seasonally dry tropical forest (Santa Teresinha, Paraíba state, Brazil). Paracytic stomata and star-shaped tector trichomes (A-B) in frontal view; cross-section of leaf blade (C) and midrib (D). (C) PP = palisade parenchyma; LP = lacunar parenchyma; T = trichome; ep aba = abaxial epidermis; ep ada = adaxial epidermis; (D) ep = epidermis; FP = fundamental parenchyma; phl = phloem; xy = xylem.

Table 3. Morphometric characteristics in leaves of three herbaceous species (one tall, one short, and one climbing species) growing in sites with different regeneration periods after land abandonment in a northeastern Brazilian seasonally dry tropical forest (Santa Teresinha, Paraíba state, Brazil).

Species	Leaf Surface	Regeneration periods
		R22	R44	R >60
Epidermis thickness (μm)
Mesosphaerum suaveolens	Abaxial	6.5 ± 1.4^Aa	7.2 ± 2.35^Aa	6.3 ± 1.73^Ab
	Adaxial	6.9 ± 1.7^Aa	7.75 ± 2.14^Aa	7.7 ± 2.53^Aa
Stachytarpheta sanguinea	Abaxial	6.72 ± 1.4^Ba	7.0 ± 1.8^Ba	8.2 ± 1.3^Aa
	Adaxial	6.92 ± 1.44^Ba	7.3 ± 1.6^Ba	8.13 ± 2.08^Aa
Jacquemontia evolvuloides	Abaxial	10.1 ± 2.86^Aa	10.7 ± 2.1^Aa	11.64 ± 2.4^Aa
	Adaxial	10.9 ± 2.48^Aa	11.2 ± 2.6^Aa	11.28 ± 1.7^Aa
Cuticle thickness (μm)
Mesosphaerum suaveolens	Abaxial	1.3 ± 0.4^Aa	1.2 ± 0.5^Aa	1.1 ± 0.3^Aa
	Adaxial	1.3 ± 0.4^Aa	1.3 ± 0.4^Aa	1.2 ± 0.4^Aa
Stachytarpheta sanguinea	Abaxial	1.1 ± 0.3^Aa	1.1 ± 0.3^Aa	1.0 ± 0.0^Aa
	Adaxial	1.3 ± 0.3^Aa	1.2 ± 0.4^Aa	1.03 ± 0.1^Ba
Jacquemontia evolvuloides	Abaxial	1.2 ± 0.4^Aa	1.1 ± 0.3^Aa	1.1 ± 0.3^Aa
	Adaxial	1.3 ± 0.5^Aa	1.0 ± 0.0^Aa	1.3 ± 0.4^Aa
Stomatal density (n mm−2)
Mesosphaerum suaveolens	Abaxial	275.1 ± 73.1^ABa	247.8 ± 41.0^Ba	301.8 ± 54.1^Aa
	Adaxial	183.6 ± 25.0^Ab	135.7 ± 15.5^Bb	183.0 ± 40.4^Ab
Stachytarpheta sanguinea	Abaxial	175.7 ± 35.0^Aa	196.9 ± 32.3^Aa	195.1 ± 34.9^Aa
	Adaxial	154.5 ± 33.1^Ba	155.7 ± 24.6^Bb	192.7 ± 57.9^Aa
Jacquemontia evolvuloides	Abaxial	105.4 ± 18.9^Aa	123.0 ± 23.0^Aa	124.84 ± 32.4^Aa
	Adaxial	100.0 ± 20.7^Aa	76.3 ± 13.1^Ab	56.96 ± 12.4^Bb
Trichome density (n mm−2)
Mesosphaerum suaveolens	Abaxial	5.0 ± 7.9^Aa	0.3 ± 0.2^Ba	1.1 ± 0.4^Ba
	Adaxial	1.14 ± 0.6^Ab	1.1 ± 0.3^Aa	1.7 ± 0.5^Aa
Stachytarpheta sanguinea	Abaxial	12.1 ± 8.3^Aa	14.28 ± 15.1^Aa	15.1 ± 3.48^Aa
	Adaxial	0.8 ± 0.4^Bb	1.6 ± 0.5^Bb	7.3 ± 3.3^Ab
Jacquemontia evolvuloides	Abaxial	3.0 ± 0.7^Ba	3.8 ± 1.2^Ba	7.4 ± 1.6^Aa
	Adaxial	2.7 ± 1.0^Aa	2.5 ± 0.6^Ab	3.4 ± 0.5^Ab
Trichome size (μm)
Mesosphaerum suaveolens	Abaxial	58.8 ± 10.8^Aa	43.4 ± 5.02^Ba	44.4 ± 6.5^Ba
	Adaxial	48.0 ± 7.38^Aa	56.2 ± 6.2^Aa	52.4 ± 7.9^Aa
Stachytarpheta sanguinea	Abaxial	31.9 ± 23.6^Aa	28.4 ± 18.0^Aa	26.7 ± 19.8^Ab
	Adaxial	35.0 ± 26.4^Aa	33.4 ± 25.2^Aa	45.3 ± 34.0^Aa
Jacquemontia evolvuloides	Abaxial	22.6 ± 2.07^Aa	23.4 ± 2.7^Aa	16.8 ± 4.7^Ab
	Adaxial	27.2 ± 3.7^Aa	33.6 ± 7.09^Aa	33.8 ± 8.1^Aa
Mesophyll (µm)
Mesosphaerum suaveolens		78.1 ± 17.6^B	89.6 ± 9.3^A	93.8 ± 5.5^A
Stachytarpheta sanguinea		79.8 ± 8.7^B	69.9 ± 12.4^C	89.9 ± 16.15^A
Jacquemontia evolvuloides		74.2 ± 8.1^A	65.2 ± 11.8^B	81.3 ± 7.7^A
Palisade parenchyma (μm)
Mesosphaerum suaveolens		31.4 ± 10.3^A	32.5 ± 4.1^A	35.6 ± 5.1^A
Stachytarpheta sanguinea		31.16 ± 4.57^A	26.8 ± 7.07^B	30.5 ± 7.02^AB
Jacquemontia evolvuloides	Abaxial	12.1 ± 2.0^Ab	11.2 ± 3.6^Bb	14.2 ± 3.3^Ab
	Adaxial	23.5 ± 3.9^Aa	19.1 ± 5.5^Ba	27.0 ± 2.6^Aa
Lacunar parenchyma (μm)
Mesosphaerum suaveolens		33.2 ± 8.8^B	43.8 ± 5.7^A	43.6 ± 4.9^A
Stachytarpheta sanguinea		35.5 ± 6.4^B	28.8 ± 5.7^C	42.9 ± 11.8^A
Jacquemontia evolvuloides		13.6 ± 3.1^AB	12.6 ± 2.8^B	16.0 ± 3.4^A

Averages (± standard deviation) in the column followed by the same capital letter for species comparisons and by the same lower case letter for surface comparisons are not significantly different by the Tukey test at 5% probability level (n = 90). R22 = site under regeneration for 22 years; R44 = site under regeneration for 44 years; R > 60 = site under regeneration for over 60 years.