Flowering and fruiting phenology of a subtropical rain forest, Fush

an, Taiwan

張楊家豪 1 、呂佳陵 1 、孫義方 2 、謝長富 1

1 臺灣大學生態學與演化生物學研究所2 東海大學熱帶生態學與生物多樣性研究中心

Introduction

Phenology is the study of the recurrent biological events, the causes of the timing with regard biotic and abiotic forces, and the interralation among phases of the same species or different species. (Lieth 1974)

Plant reproductive patterns have a major impact on the temporal structure of a community. (Bawa 1983; Herrera 1986)

The activity of pollinators, frugivores, and flower and fruit predators

The timing of seed dispersal, seed germination The breeding periods of many animals

Introduction

In seasonal environments the timing of fruiting and flowering is usually correlated with climatic conditions. Photoperiod (Heinrich 1976)

Temperature (Arroyo et al 1981; Smith-Ramirez and Armeto 1994)

Precipitation (van Schaik et al 1993)

Irradiance (van Schaik et al 1993; Hamann 2004; Zimmermann 2007)

Few records exist for Taiwan and no attempts have been made to evaluate reproductive phenology at the community level over an extended period of time.

Introduction

The climate in Fushan exhibits a clear seasonality in temperature and irradiance but not in precipitation. average temperature: 18.2 ℃ annual precipitation: 4271 mm

The objectives of this research: Understand the flowering and fruiting patterns of the subtropical

rain forest in Fushan Evaluate the relationship between reproductive activity and clima

tic factors (irradiance and temperature)

Data Collection

87 seed traps in the Fushan Forest Dynamics Plot Established in Aug. 2002

Surface area: 0.5 m2 Trap mesh: 1.6 mm

All reproductive parts (flowers, fruit, seeds) were sorted, counted, and identified to species each week

Sept. 2002 – Aug. 2007 (260 weeks)

The locations of 106 seed traps

Quantitative phenological records

Flower production No. of traps with flower collected in each month for each species

Seed production No. of seeds collected in each month for each species

All analyses were restricted to species with ≥ 10 records and encountered in ≥ 5 traps.

Data analysis

Flowering and fruiting phenology Circular vector algebra (Batschelet 1981; Wright and Calderon 1995)

Relationship between reproductive activity and climatic factors Time series cross correlation (Zar 1999; Zimmerman et al 2007)

Correspondence analysis

Flowering and fruiting phenology

Mean flowering and fruiting dates Vector algebra (Batschelet 1981; Wright and Calderon 1995)

0

1

2

3

4

J F M A M J J A S O N D

Per

cent

of t

raps

mean vector

Periodicity of flowering (fruiting) phenology Autocorrelation analyses (Zimmerman et al, 2007)

lag = 0r = 1

lag = 1r = 0.618

lag = 6r = -0.346

lag = 12r = 0.712

Periodicity of flowering (fruiting) phenology Autocorrelation analyses (Zimmerman et al, 2007)

annual biannual supra-annual continuous

Regularity of flowering (fruiting) phenology

vector length = 0.99

Second mean vector (Hamann 2004)

Annual and supra-annual Irregular intervals: vector length < 0.7

Mean flowering dates n = 44 species

Mean flowering dates n = 44 species

annual: 41 species annual but irregular: 1species biannual: 2 species

Mean fruiting dates n = 28 species

Mean fruiting dates n = 28 species

annual: 19 species continuous: 1species supra-annual & regular : 4 species irregular: 4

species

Relationship between the reproductive activity and climatic variables

Cross correlation analyses (Zar 1999; Zimmerman et al 2007)

Community-level reproductive patterns No. of species with peak reproductive activity Fewest months comprised ≥ 75% of total records

Climatic variables (1992 – 2006) Temperature

Growing Degree Index (GDI) Tavg – Tbase

1, 3, 6-month GDI0

20

40

60

80

100

120

J F M A M J J A S O N D

Month

Gro

win

g D

egre

e In

dex(

°C)

GDI1

GDI3

GDI6

Community-wide peak months of reproductive activity

0

5

10

15

20

J F M A M J J A S O N D

Month

No.

of s

peci

es p

eak

activ

ity

0

5

10

15

20

25

J F M A M J J A S O N D

Month

No.

of s

peci

es p

eak

activ

ityFlowering

Fruiting

Cross-correlation: Flowering vs 1-month GDI

0

5

10

15

20

J F M A M J J A S O N D

Month

No.

of s

peci

es p

eak

activ

ity

0

5

10

15

20

25

J F M A M J J A S O N D

Month

Gro

win

g D

egre

e (°

C) GD1

Cross-correlation: Flowering vs Temperature change

0

5

10

15

20

J F M A M J J A S O N D

Month

No.

of s

peci

es p

eak

activ

ity

-4

-3

-2

-1

0

1

2

3

4

J F M A M J J A S O N D

Month

Tem

pera

ture

(°C

)

temperature change

Cross-correlation: Fruiting vs 1-month GDI

0

5

10

15

20

25

J F M A M J J A S O N D

Month

No.

of s

peci

es p

eak

activ

ity

0

5

10

15

20

25

J F M A M J J A S O N D

Month

Gro

win

g D

egre

e (°

C) GD1

Cross-correlation: Fruiting vs 3-month GDI

0

5

10

15

20

25

J F M A M J J A S O N D

Month

No.

of s

peci

es p

eak

activ

ity

0

10

20

30

40

50

60

J F M A M J J A S O N D

Month

Gro

win

g D

egre

e (°

C) GD3

Cross-correlation: Fruiting vs 6-month GDI

0

5

10

15

20

25

J F M A M J J A S O N D

Month

No.

of s

peci

es p

eak

activ

ity

0

20

40

60

80

100

120

J F M A M J J A S O N D

Month

Gro

win

g D

egre

e (°

C) GD6

0

5

10

15

20

25

30

2002

-09

2002

-12

2003

-03

2003

-06

2003

-09

2003

-12

2004

-03

2004

-06

2004

-09

2004

-12

2005

-03

2005

-06

2005

-09

2005

-12

2006

-03

2006

-06

2006

-09

2006

-12

2007

-03

2007

-06

flow

er p

rodu

ctio

n(f

requ

ency

)

0

50

100

150

200

250

300

350

frui

t pro

duct

ion

(no.

of s

eeds

)

Fruiting Flowering

02468

1012141618

2002

-09

2002

-12

2003

-03

2003

-06

2003

-09

2003

-12

2004

-03

2004

-06

2004

-09

2004

-12

2005

-03

2005

-06

2005

-09

2005

-12

2006

-03

2006

-06

2006

-09

2006

-12

2007

-03

2007

-06

flow

er p

rodu

ctio

n(f

requ

ency

)

0

200

400

600

800

1000

1200

1400

frui

t pro

duct

ion

(no.

of s

eeds

)

Fruiting FloweringGlochidion acuminatum (Euphorbiaceae)

Machilus thunbergii (Lauraceae)

0123456789

10

2002

-09

2002

-12

2003

-03

2003

-06

2003

-09

2003

-12

2004

-03

2004

-06

2004

-09

2004

-12

2005

-03

2005

-06

2005

-09

2005

-12

2006

-03

2006

-06

2006

-09

2006

-12

2007

-03

2007

-06

flow

er p

rodu

ctio

n(f

requ

ency

)

020

406080

100120140

160180

frui

t pro

duct

ion

(no.

of s

eeds

)

Fruiting Flowering

0

2

4

6

8

10

12

14

16

2002

-09

2002

-12

2003

-03

2003

-06

2003

-09

2003

-12

2004

-03

2004

-06

2004

-09

2004

-12

2005

-03

2005

-06

2005

-09

2005

-12

2006

-03

2006

-06

2006

-09

2006

-12

2007

-03

2007

-06

flow

er p

rodu

ctio

n(f

requ

ency

)

0

50

100

150

200

250

300

350

400

frui

t pro

duct

ion

(no.

of s

eeds

)

Fruiting FloweringPyrenaria shinkoensis (Theaceae)

Myrsine seguinii (Myrsinaceae)

AMPECA ARALDE ARDIQU CLEMCR CLEYJA CRYPCH DIOSMO ELAEJA ENGERO GLOCAC HELIFO

2003 157 0 6 20 58 1 1 0 82 1807 3

2004 194 232 9 6 322 1 51 96 649 2896 8

2005 135 0 1 15 12 0 1 0 267 506 4

2006 129 0 20 18 25 23 6 0 121 138 16

HOYACA LAGESU LASIAN LITSAC MACHTH MACHZU MELISQ MISCFL MYRSSE PERIFO PIPESI

2003 12 752 31 578 370 225 0 91 17 15 3

2004 23 575 116 52 411 283 73 47 319 15 28

2005 11 735 22 28 2 0 0 28 16 1 0

2006 32 5235 29 17 23 114 0 2 375 4 0

PRUNPH PSYCSE PYRESH RUBUPY SABISW SCHEOC SMILAX SYMPTH SYZYBU

2003 31 20 147 16 2 2266 94 10 0

2004 181 7 132 20 1 1588 4 14 35

2005 53 11 578 2 0 0 0 0 0

2006 2 16 19 11 22 1308 1 3 1

Correspondence Analysis

Correspondence Analysis

CA1 CA2 CA3

lambda 0.41 0.19 0.11

accounted 0.58 0.84 1.00

Summary

The flowering and fruiting phenology exhibit a clear seasonality.

Most species are annual flowering and fruiting.

No. of species peak flowering are positively correlated with monthly temperature changes.

No. of species peak fruiting are positively correlated with 6-month growing degree index.

Frost occurring in 2005 affected the fruit production of many species.

Acknowledgements 2004– 2005 林試所林業科技委辦計畫 福山亞熱帶雨林永久樣區設置與幼苗更新狀態之研究 2006 林務局委託研究計畫 台灣北部福山地區亞熱帶雨林種子雨之研究 2006 林務局委託研究計畫 台灣北部福山地區亞熱帶雨林小苗短期動態之研究 2007 林務局委託研究計畫 台灣北部福山地區亞熱帶雨林種子雨與喬木小苗動態之研究 林試所 蘇聲欣 先生 林試所福山研究中心 福山樣區調查團隊 臺灣大學 黃思博 黃雅怡 林均雅 族繁不及備載的調查義工們

Mean flowering and fruiting dates

Vector algebra (Batschelet 1981)

mean vector angle mean vector length

Cross correlation: Flowering vs Irradiance

0

5

10

15

20

J F M A M J J A S O N D

Month

No.

of s

peci

es p

eak

activ

ity

0

100

200

300

400

500

600

J F M A M J J A S O N D

Month

Tot

al r

adia

tion

(MJ

m2) irradiance

0

100

200

300

400

500

600

D J F M A M J J A S O N

Month

Tot

al r

adia

tion

(MJ

m2) irradiance lag=1

0

100

200

300

400

500

600

N D J F M A M J J A S O

Month

Tot

al r

adia

tion

(MJ

m2) irradiance lag=2

Cross correlation: Flowering vs Irradiance

Temperature vs Irradiance