Phenological dynamics of <i>Croton heliotropiifolius</i> populations in a savanna/caatinga gradient, Chapada Diamantina, Brazil

Costa, Thayse Moreira; Santos, Marília Grazielly Mendes dos; Neves, Sâmia Paula Santos; Miranda, Lia d`Afonsêca Pedreira de; Funch, Ligia Silveira

doi:10.1590/2175-7860202172130

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Original Paper • Rodriguésia 72 • 2021 • https://doi.org/10.1590/2175-7860202172130 copy

Phenological dynamics of Croton heliotropiifolius populations in a savanna/caatinga gradient, Chapada Diamantina, Brazil

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Abstract

The relationship between phenology and environmental factors is critical to understanding population dynamics in environmental gradients. We evaluated phenological variations in Croton heliotropiifolius Kunth in sites with contrasting water resource availability in caatinga, cerrado/caatinga transition, and cerrado in the Chapada Diamantina, Brazil. The phenophases of 81 individuals (n = 27 individuals/area) were recorded monthly for 12 months. Multiple regression, Spearman correlation, circular statistics, Shannon-Wiener diversity, and Morisita-Horn indices were used to test relationships between phenophases and abiotic factors, phenological seasonality, diversity, and similarities between the three populations. The vegetative behaviors in the study sites were distinct in terms of their intensity, seasonality, and synchrony; but reproductive phenophases maintained similar characteristics. Phenological events were positively related to rainfall and soil water availability. C. heliotropiifolius populations exhibited high levels of vegetative phenological diversity, except in the caatinga during the dry season. Reproductive phenological diversity varied along the studied period in the three sites, with higher reproductive than vegetative similarities among populations. Differences in soil types and rainfall volumes in the dry season, even at small distances, therefore make the savanna/caatinga gradient a suitable model for investigating phenological responses related to plant eco-hydrological strategies in seasonally tropical dry ecosystems.

Key words
phenological diversity; rainfall; seasonally dry tropical ecosystems; soil water availability; synchrony

Areas	Phenophases	Abiotic factors
		Rainfall	Temperature	Photoperiod	Soil moisture
		p	p	p	p
caatinga	Leaf flushing	0.003	-	-	0.001
	Young leaf	0.007	-	-	0.005
	Leaf fall	0.048	-	-	0.025

savanna-caatinga transition	Leaf flushing	0.018	-	-	0.009
	Young leaf	0.013	-	0.013	0.014
	Leaf fall	0.024	0.044	0.010	0.016

savanna	Leaf flushing	0.045	-	-	0.013
	Young leaf	-	-	-	-
	Leaf fall	-	-	0.012	-

Areas	Variables	Reproductive phenophases
Areas	Variables	Male bud	Female bud	Male flower	Female flower	Immature fruit	Mature fruit
caatinga	N° of observations	351	182	283	145	393	367
	LMV (r)	0.447	0.462	0.352	0.448	0.599	0.638
	Circular deviation	72.725°	71.188°	82.786°	72.646°	57.993°	54.364°
	Mean data	07 feb.	02 jan.	25 feb.	11 feb.	04 feb	22 feb.
	Mean Vector (µ)	37.197°	1.761°	55.048°	41.441°	34.724°	51.824°
	Rayleigh Test (Z)	70.085	38.873	35.084	29.053	141.078	149.169
	Rayleigh Test (p)	< 1E-12	< 1E-12	< 1E-12	< 1E-12	< 1E-12	< 1E-12

savanna-caatinga transition	N° of observations	271	156	86	40	277	263
	LMV (r)	0.582	0.552	0.570	0.519	0.796	0.693
	Circular deviation	59.612°	62.433°	60.754°	65.582°	38.652°	49.051°
	Mean data	04 jan.	10 dec.	28 feb.	26 feb.	24 feb.	04 mar.
	Mean Vector (µ)	3.926°	345.341°	58.792°	57.834°	55.15°	64.291°
	Rayleigh Test (Z)	91.803	47.583	27.938	10.791	175.725	126.372
	Rayleigh Test (p)	< 1E-12	< 1E-12	< 1E-12	1.02E-05	< 1E-12	< 1E-12

savanna	N° of observations	302	170	175	52	371	344
	LMV (r)	0.364	0.324	0.630	0.549	0.386	0.382
	Circular deviation	81.457°	85.972°	55.059°	62.774°	79.035°	79.516°
	Mean data	17 jan.	02 dec.	24 mar.	08 apr.	13 mar.	04 mar.
	Mean Vector (µ)	16.07°	337.045°	82.922°	97.873°	72.571°	63.427°
	Rayleigh Test (Z)	40.014	17.891	69.500	15.657	55.334	50.129
	Rayleigh Test (p)	< 1E-12	0.000000017	< 1E-12	1.59E-07	< 1E-12	< 1E-12

Areas	Phenophases	Abiotic factors
		Rainfall		Photoperiod		Soil moisture
		p	r	p	r	p	r
caatinga	Male bud	0.0018	0.7997	-	-	0.0022	0.7913
	Female bud	0.0004	0.8552	-	-	0.0004	0.8578
	Male flower	0.0002	0.8708	-	-	0.0013	0.8136
	Female flower	<0.0001	0.9176	-	-	0.0004	0.8527
	Immature fruit	<0.0001	0.8970	-	-	0.0002	0.8808
	Mature fruit	0.0033	0.7711	-	-	0.0051	0.7488

savanna-caatinga transition	Male bud	0.0114	0.6989	0.0224	0.6490	0.0073	0.7274
	Female bud	0.0119	0.6961	0.0355	0.6091	0.0052	0.7469
	Male flower	0.0007	0.8377	-	-	-	-
	Female flower	0.0004	0.8557	-	-	0.0166	0.6721
	Immature fruit	-	-	-	-	0.0255	0.6382
	Mature fruit	-	-	-	-	-	-

savanna	Male bud	<0.0001	0.8952	-	-	0.0046	0.7535
	Female bud	0.0001	0.8831	-	-	0.0127	0.6918
	Male flower	0.0015	0.8062	-	-	-	-
	Female flower	0.0104	0.7051	-	-	-	-
	Immature fruit	0.0133	0.6885	-	-	0.0275	0.6316
	Mature fruit	-	-	-	-	-	-

Years	Month	Similarity (Morisita-Horn)
Years	Month	Vegetative	Reproductive
2017	J	0.043	0.001
	A	0.027	0.623
	S	0.000	0.512
	O	0.000	0.634
	N	0.034	0.005
	D	0.115	0.022
2018	J	0.057	0.349
	F	0.106	0.011
	M	0.031	0.002
	A	0.120	0.016
	M	0.039	0.244
	J	0.037	0.569
Mean ± standard deviation		0.061 ± 0.001	0.249 ± 0.074

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[1] 8Corresponding author: ligiafunch@yahoo.com