363 SEED STORAGE BEHAVIOUR OF AMAZONIAN TREES Classification of seed storage behaviour of 67...

363

SEED STORAGE BEHAVIOUR OF AMAZONIAN TREES

* Author for correspondence

Lima, M.de Jr., Hong, T.D., Arruda, Y.M.B.C., Mendes, A.M.S. and Ellis, R.H.(2014), Seed Sci. & Technol., 42, 363-392, http://doi.org/10.15258/sst.2014.42.3.06

Classification of seed storage behaviour of 67 Amazonian tree species

M. DE JR. LIMA1, T.D. HONG2, Y.M.B.C. ARRUDA1, 3, A.M.S. MENDES1 AND R.H. ELLIS2*

1 Amazonas Native Seed Center, Faculdade de Ciências Agraria, University Federal of Amazonas, Av. General Rodrigo Otávio Jordão Ramos, 3000, Coroado, Manaus- AM, Brazil

2 School of Agriculture, Policy and Development, The University of Reading, Earley Gate, P.O. Box 237, Reading RG6 6AR, UK (E-mail: [email protected])

3 Institute of Biology, University Federal of Amazonas, Av. General Rodrigo Otávio Jordão Ramos, 3000, Coroado, Manaus- AM, Brazil

(Accepted August 2014. Handling editor: A.A. Powell)

Summary

Information was collated on the seed storage behaviour of 67 tree species native to the Amazon rainforest of Brazil; 38 appeared to show orthodox, 23 recalcitrant and six intermediate seed storage behaviour. A previously developed, double-criteria key based on thousand-seed weight and seed moisture content at shedding to estimate likely seed storage behaviour, showed good agreement with the above classifications. The key can aid seed storage behaviour identification considerably.

Introduction

The fi ve most forest-rich countries of the world, of which Brazil is one, account for more than half of the world’s total forest area of some four billion hectares (FAO, 2010). The Amazon rainforest of South America covers about 550 million hectares, of which 60% is in Brazil, is the largest tropical rainforest in the world. Brazil’s bio-diverse and genetic-resources-rich forests have been severely depleted, however. At a regional level, South America suffered the largest net loss of forests between 2000 and 2010 – about 4 million hectares per year (FAO, 2010). The Amazon region is reported to have lost about 580 million hectares through deforestation since 1980 (Butler, 2012). Around one-third of the world’s threatened plant species are in Latin America and the Caribbean (Royal Botanic Gardens Kew, 2010).

Reforestation with native species to support biodiversity conservation and environmental protection requires an adequate supply of high quality tree seeds for seedling production. The science and technology of tropical tree seed collection, storage, dormancy and germination is under-researched in relation to the magnitude of the task. In addition to in-situ conservation by maintaining or replanting native species, ex-situ conservation using seed banks for wild species is valuable to biodiversity conservation (Pritchard et al., 2004b). Many countries

364

M. DE JR. LIMA, T.D. HONG, Y.M.B.C. ARRUDA, A.M.S. MENDES AND R.H. ELLIS

have therefore established national seed banks. Brazil, for example, has seven seed banks (León-Lobos et al., 2012). International cooperation in wild species conservation in seed banks has been effective: for example, the Millennium Seed Bank Partnership now covers about 80 countries, including Brazil, and has already banked 13% of the world's wild plant species, including many woody species (www.kew.org/science-conservation/millennium-seed-bank-partnership/about-millennium-seed-bank-partnership; accessed 23 July 2014).

Identifi cation of the seed storage behaviour of a species is an important component of developing strategies for plant biodiversity conservation, because long-term seed storage is not possible for all species. Three categories of seed storage behaviour have been identifi ed. When mature, orthodox seeds can be dried, without incurring damage, to low moisture contents and in the air-dry range their longevity increases with decrease in seed storage moisture content and/or temperature (Roberts, 1973) in a quantifi able and predictable way (Ellis and Roberts, 1980). Long-term seed storage is possible for such species in cool, dry environments (e.g. Pérez-García et al., 2009). Recalcitrant seeds, in contrast, do not survive more than mild desiccation (Roberts, 1973) and have to be stored at high moisture contents close to fully imbibed, which is not possible for more than short periods. Finally, between the above two extremes, a third category of intermediate seed storage behaviour has been recognised where the seeds are able to tolerate desiccation to moisture contents in equilibrium with about 40-50% relative humidity (r.h.), i.e. about 7-10% moisture content depending upon species, but further drying may result in more rapid loss in viability of stored seeds and sometimes immediate damage (Ellis et al., 1990; Hong and Ellis, 1996). Dry intermediate seeds of species of tropical origin may also show more rapid loss in viability at cooler temperatures, often below about 10°C (Ellis et al., 1990; Ellis et al., 1991; Hong and Ellis, 1992). Successful medium-term conservation is thus feasible for intermediate seeds of tropical lowland species at about 10°C with seed moisture contents in equilibrium with about 40-50% r.h. (Hong and Ellis, 2002).

Seed storage behaviour was identified for 6,919 plant species by Hong et al. (1996, 1998a, b). That information is now replicated and supplemented as new understanding becomes available through the Royal Botanic Garden Kew’s Seed Information Database (data.kew.org/sid/sidsearch.html; accessed on 21 July 2014). Given that such sources of information cover less than 5% of the world’s flowering plant species, rapid approaches to estimating likely seed storage behaviour, if only limited information is available, are helpful to conservation activities, especially on seed collection missions.

Relevant indirect information can include plant ecology, plant taxonomy, fruit/seed characters, seed size, seed shape and seed moisture content at shedding (Hong et al., 1996, 1998a). Information on only one of the above is far from diagnostic, but the use of multiple-criteria keys combining seed weight and moisture content at shedding (or maturity or natural dispersal, depending upon data available; the term shedding is used herein for these similar, but not identical, seed developmental stages) was shown to provide a good indication of likely seed storage behaviour (Hong and Ellis, 1996, 1997, 2002; Ellis et al., 2007). Similar approaches have been developed to estimate the likelihood of seed desiccation tolerance from information on seed mass and rainfall in the month of seed dispersal (Pritchard et al., 2004a) or seed mass and seed coat ratio (Daws et al., 2006).

365


In this paper we collate, and in some cases interpret, information on the seed storage survival and behaviour of several major Amazonian tree species. We then use this independent information to test the validity of a multiple-criteria key developed previously (Hong and Ellis, 1996, 1997, 2002; Ellis et al., 2007).

Materials and methods

Information was collated on seed storage survival and behaviour and the experimental drying and storage conditions used for 67 major trees of the Amazon region (table 1). For each species, information on natural habitat, fruit or seed dispersal period, thousand-seed weight (TSW), seed shape, and seed moisture content at shedding (MCS) was also collated (table 2). The majority of this information was provided by members of the Amazon Seed Network, Brazil, in particular the Instituto Nacional de Pesquisas da Amazônia (INPA) and Universidade Federal do Amazonas (UFAM).

To classify seed storage behaviour from the diverse information available we used the criteria defined by Hong and Ellis (1996), namely:

(i) seeds which tolerate desiccation to approximately 5% moisture content or to equilibrium with approximately 10-13% r.h. at 20°C are likely to show orthodox seed storage behaviour;

(ii) if the majority of seeds tolerate desiccation to approximately 10-12.5% moisture content (i.e. in equilibrium with 40-50% r.h. at 20°C), but further desiccation damages viability and/or reduces subsequent survival then intermediate seed storage behaviour is probable;

(iii) if all seeds fail to survive by desiccation to 15-20% moisture content (i.e. values in equilibrium with > 70% r.h. at 20°C) then recalcitrant seed storage behaviour is probable.

In some cases information on desiccation tolerance was not available. In these cases, if all dry seeds survived hermetic storage at sub-zero temperatures (-18°C) or in liquid nitrogen (-196°C) then orthodox seed storage behaviour was assumed, whilst in other cases, where seeds survived desiccation to 10-12.5% moisture content but not subsequent hermetic storage at -18°C, they were classified as intermediate (Hong and Ellis, 1996).

Results

The 67 tree species were from 23 different families, of which 38 species appeared to show orthodox (56.7%), 23 recalcitrant (34.3%) and six intermediate seed storage behaviour (9%) (table 3). Some 38 of these species are not yet represented in the Seed Information Database.

Bixa orellana and Swietenia macrophilla were previously classified as intermediate in their seed storage behaviour (Hong et al., 1996, 1998b; Hong and Ellis, 1998; Lima and Galvão, 2005). This had been assumed because viability declined on desiccation to about5% moisture content, with a further reduction in subsequent hermetic storage at -20°C (Goldbach, 1979; Hong and Ellis, 1996). However, we now classify both species as orthodox.

366


The amended classification is because later research showed that hardseededness in Bixa orellana (Poornima and Ambika, 2012) and dormancy in Swietenia macrophylla were imposed upon desiccation to < 10% moisture content (Ellis et al., 2007). The failure to germinate was overcome by scarification for Bixa orellana or providing a more suitable alternating temperature germination regime (35/20°C for 16 hours / 8 hours) for Swietenia macrophylla, and viability was shown to be maintained on desiccation to low moisture contents and in hermetic storage at sub-zero temperatures (Ellis et al., 2007; Poornima and Ambika, 2012).

Desiccation sensitivity was reported in Calophyllum brasiliense: loss in viability on desiccation to 20.9% (Carlota, 2006) or to 10.7% moisture content (Carvajal et al., 2013) when drying from initial values of 60 or 46% moisture content, respectively. The species was therefore classified as recalcitrant (Hong et al, 1996; Carlota, 2006; Carvajal et al., 2013). However, elsewhere the seeds have been reported to show limited germination (4%) after desiccation from 40 to 4.8% moisture content (Vásquez-Carballo et al., 2005), with best seed storage survival at 45% r.h. and 8°C (Carlota, 2006). This range of responses has several potential explanations. Earlier reports of considerable desiccation sensitivity may have been based on investigation with immature seeds; at the other extreme, the survival of a few seeds after considerable desiccation might have resulted from variation in drying rate with those surviving being the wettest in the seed population. Given this uncertainty, we have provisionally classified Calophyllum brasiliense as “Recalcitrant?” (table 3).

Virola surinamensis has been classified as recalcitrant (Cunha et al., 1992; Cesarino, 2006; Limas et al., 2007). However, while viability was lost on high-temperature drying at 50°C to 18% moisture content, drying at 15% r.h. and 24°C to 11.7 and 5.4% moisture content provided 23 and 4% germination, respectively (Cunha et al., 1992). This species has therefore been classified as intermediate (Hong et al., 1996, 1998b).

Amongst the 23 families, there is evidence of contrasting seed storage behaviour within families. For Fabaceae 18 species are listed, 14 with orthodox (Adenanthera pavonina, Cassia ferruginea, Cenostigma tocantinum, Dinizia excelsa, Enterolobium schomburgkii, Hymenaea courbaril, Ormosia excelsa, Parkia discolour, P. multijuga, P. nitida, P. pendula, Schizolobium amazonicum, Stryphnodendron guianense, S. microstachyum), two recalcitrant (Cynometra bauhiniifolia, Inga cinnamomea) and two with intermediate (Copaifera multijuga, Dipteryx odorata) seed storage behaviour. Of the nine species listed for Arecaceae, eight show recalcitrant (Bactris gasipaes, Euterpe oleracea, E. precatoria, Mauritia flexuosa, Oenocarpus bacaba, O. bataua, O. mapora) and one intermediate (Astrocaryum aculeatum) seed storage behaviour (table 3). Within Lecythidaceae, six show orthodox (Cariniana micrantha, C. atrovinosa, C. guianensis, C. longipedicellata, C. stellata, Lecythis pisonis) and one intermediate (Bertholletia excelsa) seed storage behaviour.

367

SEED STORAGE BEHAVIOUR OF AMAZONIAN TREEST

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Tabl

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con’

t

368

M. DE JR. LIMA, T.D. HONG, Y.M.B.C. ARRUDA, A.M.S. MENDES AND R.H. ELLIST

able

1.

Con

tinu

ed

Spec

ies

Fam

ilyM

etho

d of

dry

ing

Met

hod

of s

tora

geR

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ence

10C

arap

a gu

iane

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A

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t m

c =

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olye

thyl

ene

bags

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eds

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7% m

c.F

erra

z (2

004)

12C

arin

iana

mic

rant

ha

Duc

keL

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hida

ceae

Ove

r si

lica

gel

, to

6%

mc.

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met

ical

ly i

n cl

osed

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ss b

ottl

eat

5 ±

3°C

, fo

r th

ree

year

s.

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argo

and

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(200

7)

13C

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ria

sylv

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Air

con

diti

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m,

low

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mc

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t 5°

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(200

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(S

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pera

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5°C

, 60

-80%

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(20

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clos

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aine

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d 15

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n ov

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2b)

Tabl

e 1

con’

t

369


able

1.

Con

tinu

ed

Spec

ies

Fam

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d of

dry

ing

Met

hod

of s

tora

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efer

ence

22C

ouep

ia l

ongi

pend

ula

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ysob

alan

acea

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mai

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ned

at 1

5°C

, dr

y to

39.

5% m

c.N

ot d

escr

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ilva

et

al.

(201

0)

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(M

art.)

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pocy

nace

aeO

ver

sili

ca g

el,

to 3

% m

c.H

erm

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sto

rage

at

3-5%

mc

and

-18°

C.

Sil

va e

t al

. (2

010)

24C

oura

tari

atr

ovin

osa

Lec

ythi

dace

aeO

ver

sili

ca g

el,

seve

n da

ys,

to 3

.8%

mc.

Her

met

ic s

tora

ge a

t 5°

C.

Oli

veir

a (2

010)

25C

oura

tari

gui

anen

sis

Lec

ythi

dace

aeO

ver

sili

ca g

el,

seve

n da

ys,

to 5

.3%

mc.

H

erm

etic

sto

rage

at

5°C

.O

live

ira

(201

0)

26C

oura

tari

lo

ngip

edic

ella

taL

ecyt

hida

ceae

Ove

r si

lica

gel

, se

ven

days

, to

4.8

% m

c.

Her

met

ic s

tora

ge a

t 5°

C.

Oli

veir

a (2

010)

27C

oura

tari

ste

llat

a

A.C

. S

m.

Lec

ythi

dace

aeO

ver

sili

ca g

el,

seve

n da

ys,

to 7

.9%

mc.

Her

met

ic s

tora

ge a

t 5°

C.

Oli

veir

a (2

010)

28C

ynom

etra

bau

hini

ifol

ia

Ben

th.

Fab

acea

eD

ry t

o 28

.2%

mc.

Moi

st s

tora

ge.

Naz

ário

et

al.

(200

8)

29D

iniz

ia e

xcel

sa D

ucke

Fab

acea

eO

ver

sili

ca g

el,

to 5

% m

c.H

erm

etic

sto

rage

at

5°C

, -1

8°C

w

ith

5, 1

0% m

c.M

esqu

ita

et a

l. (2

009)

30D

ipte

ryx

odor

ata

(A

ubl.)

Wil

ld.

Fab

acea

eD

esic

cate

to

12%

mc

and

belo

w.

Not

des

crib

ed.

BC

SN

A (

2013

),

Car

valh

o (2

009b

)

31E

nter

olob

ium

sc

hom

burg

kii

(B

enth

.) B

enth

.F

abac

eae

Roo

m t

empe

ratu

re

(25°

C,

60-8

0% r

.h.)

.H

erm

etic

sto

rage

, 11

.9%

mc

and

5°C

.B

CS

NA

(20

13),

H

orin

g et

al.

(201

2)

32E

ugen

ia s

tipi

tata

M

cVau

ghM

yrta

ceae

Des

icca

te t

o 15

% m

c.

Moi

st s

tora

ge f

or 5

0 da

ys.

Del

gado

and

Bar

bedo

(2

007)

, G

enti

l an

d F

erre

ira

(199

9)

33E

uter

pe o

lera

cea

Mar

t.A

reca

ceae

Des

icca

te t

o 26

% m

c.M

oist

sto

rage

in

poly

ethy

lene

bag

s,

37-4

3% m

c an

d 20

°C.

Nas

cim

ento

(20

08)

Tabl

e 1

con’

t

370


able

1.

Con

tinu

ed

Spec

ies

Fam

ilyM

etho

d of

dry

ing

Met

hod

of s

tora

geR

efer

ence

34E

uter

pe p

reca

tori

a

Mar

t.A

reca

ceae

Des

icca

te t

o 17

% m

c.N

ot d

escr

ibed

.B

CS

NA

(20

13)

35G

enip

a am

eric

ana

L.

Rub

iace

aeD

esic

cate

to

5% m

c.H

erm

etic

sto

rage

at

10%

mc

and

0°C

.M

agis

tral

i et

al.

(201

3)

36G

oupi

a gl

abra

Aub

l.C

elas

trac

eae

Roo

m t

empe

ratu

re,

to 6

% m

c.H

erm

etic

sto

rage

at

sub-

zero

te

mpe

ratu

res.

Sch

wen

gber

and

S

mid

erle

(20

05)

37H

andr

oant

hus

se

rrat

ifol

ius

(Vah

l)

S.O

. G

rose

[T

abeb

uia

se

rrat

ifol

ia (

Vah

l)

G.

Nic

hols

on]

Big

noni

acea

eD

ryin

g to

equ

ilib

rium

wit

h 30

% r

.h.

Lab

orat

ory

at 2

5°C

, th

ree

days

, to

5.

4% m

c.S

tora

ge i

n 30

% r

.h.;

in l

iqui

d N

2.F

erre

ira

et a

l. (2

004)

, S

alom

ão (

2002

)

38H

elic

osty

lis

tom

ento

sa

(Poe

pp.

& E

ndl.)

R

usby

Mor

acea

eD

esic

cate

to

16%

mc.

Moi

st s

tora

ge i

n sa

nd a

t 20

°C f

or

12 m

onth

s.A

rrud

a an

d F

erra

z (2

008)

39H

evea

bra

sili

ensi

s

(Wil

ld.

ex A

. Ju

ss.)

M

üll.

Arg

.E

upho

rbia

ceae

Des

icca

te t

o 15

% m

c.

Moi

st s

tora

ge (

moi

st c

harc

oal

and

saw

dust

) in

pol

yeth

ylen

e ba

g, a

t 7-

10°C

for

thr

ee m

onth

s; m

oist

sto

rage

at

10°

C,

20°C

for

sev

en m

onth

s.

Bon

ome

(200

6)

40H

ymen

aea

cour

bari

l L

.F

abac

eae

Roo

m t

empe

ratu

re,

for

35 d

ays,

de

sicc

ate

to 1

0.2%

mc.

Her

met

ic s

tora

ge a

t 10

.2%

mc

and

-18°

C,

for

90 d

ays.

Car

valh

o et

al.

(200

6)

41In

ga c

inna

mom

ea

Spr

uce

ex B

enth

.F

abac

eae

Not

des

crib

ed.

Moi

st s

tora

ge,

none

sur

vive

d (2

0 da

ys).

Car

valh

o et

al.

(200

1)

42L

ecyt

his

piso

nis

Cam

b.L

ecyt

hida

ceae

Ove

r si

lica

gel

, to

6.3

% m

c.H

erm

etic

sto

rage

at

-18°

C a

nd 6

.3%

m

c, f

or 9

0 da

ys.

Car

valh

o et

al.

(200

6)

43M

auri

tia

flex

uosa

L.f

.A

reca

ceae

Ove

r si

lica

gel

, re

duce

d m

c fr

om 4

5.6

to 1

3.6%

.N

ot d

escr

ibed

.S

ousa

et

al.

(200

4)

44M

inqu

arti

a gu

iane

nsis

A

ubl.

Mor

acea

eN

ot d

escr

ibed

.M

oist

sto

rage

at

45-5

0% m

c an

d 15

°C

in p

olye

thyl

ene

bags

, ni

ne m

onth

s.C

amar

go a

nd F

erra

z (2

005)

Tabl

e 1

con’

t

371


able

1.

Con

tinu

ed

Spec

ies

Fam

ilyM

etho

d of

dry

ing

Met

hod

of s

tora

geR

efer

ence

45M

yrci

aria

dub

ia (

Kun

th)

M

cVau

gh

Myr

tace

aeN

ot d

escr

ibed

.M

oist

sto

rage

at

46%

mc

and

15°C

, fi

ve m

onth

s.

Fer

reir

a an

d G

enti

l (2

003)

, G

enti

l et

al.

(200

4)

46O

chro

ma

pyra

mid

ale

(C

av.

ex L

am.)

Urb

.M

alva

ceae

Dry

ing

cham

ber,

to

10%

mc

or b

elow

.

Ope

n st

orag

e in

dry

ing

cham

ber,

or

labo

rato

ry a

t 13

± 2

% m

c fo

r th

ree

year

s, i

n he

rbar

ium

for

44

year

s.L

eão

et a

l. (2

008)

47O

enoc

arpu

s ba

caba

Mar

t.A

reca

ceae

Des

icca

te a

t 15

°C,

from

33.

9 to

26

.5%

mc.

Moi

st s

tora

ge.

José

et

al.

(201

2)

48O

enoc

arpu

s ba

taua

Mar

t.A

reca

ceae

Des

icca

te t

o 24

.3%

mc.

Moi

st s

tora

ge.

Naz

ário

and

Fer

reir

a (2

012)

49O

enoc

arpu

s m

apor

a

H.

Kar

st.

Are

cace

aeN

ot d

escr

ibed

.N

ot d

escr

ibed

.C

arva

lho

et a

l. (2

001)

50O

enoc

arpu

s m

inor

Mar

t.A

reca

ceae

Des

icca

te a

t 15

°C t

o 30

% m

c.M

oist

sto

rage

at

15°C

for

tw

o m

onth

s.B

CS

NA

(20

13)

51O

rmos

ia e

xcel

sa B

enth

.F

abac

eae

Roo

m t

empe

ratu

re

(25°

C,

60-8

0% r

.h.)

.H

erm

etic

sto

rage

, 11

% m

c an

d 5°

C.

BC

SN

A (

2013

)

52P

arki

a di

scol

or S

pruc

e

ex B

enth

.F

abac

eae

Roo

m t

empe

ratu

re

(25°

C,

60-8

0% r

.h.)

.H

erm

etic

sto

rage

, 11

% m

c an

d 5°

CB

CS

NA

(20

13)

53P

arki

a m

ulti

juga

Ben

th.

Fab

acea

eL

abor

ator

y te

mpe

ratu

re,

four

mon

ths.

Not

des

crib

ed.

Car

valh

o (2

009a

)

54P

arki

a ni

tida

Miq

.F

abac

eae

Roo

m t

empe

ratu

re (

25°C

, 60

-80%

r.h

.).

Her

met

ic s

tora

ge,

11%

mc

and

5°C

.B

CS

NA

(20

13)

55P

arki

a pe

ndul

a B

enth

.F

abac

eae

Des

icca

te t

o 5.

9% m

c.5.

9% m

c an

d 5°

C f

or s

ix m

onth

s.R

osse

to (

2006

)

56P

seud

obom

bax

mun

guba

(M

art.

& Z

ucc.

)

Dug

and

Mal

vace

aeL

abor

ator

y at

25°

C,

thre

e da

ys,

to 7

% m

c.L

iqui

d N

2.S

alom

ão (

2002

)

57P

sidi

um a

cuta

ngul

um D

C.

Myr

tace

aeN

ot d

escr

ibed

.H

erm

etic

sto

rage

, 5%

mc

and

-18°

C.

Car

valh

o et

al.

(200

1)

Tabl

e 1

con’

t

372


able

1.

Con

tinu

ed

Spec

ies

Fam

ilyM

etho

d of

dry

ing

Met

hod

of s

tora

geR

efer

ence

58Sc

heff

lera

mor

otot

oni

(A

ubl.)

Mag

uire

,

Ste

yerm

. &

Fro

din

Ara

liac

eae

Not

des

crib

ed.

Pap

er b

ag,

10%

mc

and

12°C

and

30

% r

.h.

for

11 m

onth

s.O

hash

i an

d L

eão

(200

5)

59Sc

hizo

lobi

um a

maz

onic

um

Hub

er e

x D

ucke

Fab

acea

eN

ot d

escr

ibed

.S

tora

ge a

t 0-

5°C

and

40%

r.h

. fo

r th

ree

year

s.S

ouza

et

al.

(200

5)

60Sp

ondi

as m

ombi

n L

.A

naca

rdia

ceae

Not

des

crib

ed.

Her

met

ic s

tora

ge,

at 5

% m

c an

d -1

8°C

. In

liq

uid

N2.

Car

valh

o et

al.

(200

1),

Sal

omão

(20

02)

61St

ryph

node

ndro

n

guia

nens

e (A

ubl.)

B

enth

.F

abac

eae

Roo

m t

empe

ratu

re

(25°

C,

60-8

0% r

.h.)

.H

erm

etic

sto

rage

, 10

% m

c an

d 5°

C.

BC

SN

A (

2013

)

62St

ryph

node

ndro

n

mic

rost

achy

um P

oepp

.

& E

ndl.

Fab

acea

eR

oom

tem

pera

ture

(2

5°C

, 60

-80%

r.h

.).

Her

met

ic s

tora

ge,

10%

mc

and

5°C

.B

CS

NA

(20

13)

63Sw

iete

nia

mac

roph

ylla

K

ing

Mel

iace

aeO

ver

sili

ca g

el,

to 4

% m

c.P

olye

thyl

ene

bags

, 4-

5% m

c at

2-5

°C

for

eigh

t ye

ars.

Lim

a an

d G

alvã

o (2

005)

64T

apir

ira

guia

nens

is A

ubl.

Ana

card

iace

aeO

ven

at 4

0°C

, fo

r 0,

3,

6, 9

, …

to 2

4 ho

urs,

red

uced

mc

from

39

to

8.3%

.N

ot d

escr

ibed

.C

arva

lho

et a

l. (2

006)

, S

anto

s-M

oura

et

al.

(201

2)

65T

etra

gast

ris

pana

men

sis

(E

ngl.)

Kun

tze

Bur

sera

ceae

Ove

r si

lica

gel

, to

5%

mc.

Her

met

ic s

tora

ge,

10%

mc

at 5

°C,

or 5

% m

c at

-5°

C.

BC

SN

A (

2013

)

66T

heob

rom

a gr

andi

flor

um

(Wil

ld.

ex S

pren

g.)

K

. S

chum

.M

alva

ceae

For

ced

air

oven

, at

24-

32°C

and

43

- 98

% r

.h.,

redu

ced

mc

from

57

.1 t

o 15

.6%

.M

oist

sto

rage

.C

ruz

and

Cic

ero

(200

8)

67V

irol

a su

rina

men

sis

(R

ol.)

War

b.M

yris

tica

ceae

Dry

ing

at 2

7 ±

3°C

and

75

± 5

% r

.h.

to 1

4.6%

mc;

ove

n at

50°

C f

or 0

, 15

, 60

, 90

, 12

0 an

d 30

0 m

inut

es t

o 14

.8%

m

c; d

ryin

g ch

ambe

r at

15%

r.h

. an

d 24

°C t

o 5.

4% m

c.

Her

met

ic s

tora

ge a

t 5 a

nd -

20°C

.C

esar

ino

(200

6),

Cun

ha e

t al

. (1

992)

, L

imas

et

al.

(200

7)

373


Tabl

e 2

con’

t

Ade

nant

hera

pav

onin

a30

4.00

9.0

Ovo

idL

owla

nd f

ores

tD

ry s

easo

nO

Ali

bert

ia e

duli

s 18

.00

25.5

Fla

tL

owla

nd f

ores

t, fl

oode

d fo

rest

Rai

ny s

easo

nO

Ani

ba r

osae

odor

a 35

00.0

048

.5O

void

Low

land

for

est

Dry

sea

son

R

Ast

roca

ryum

acu

leat

um

1495

0.00

28.9

Sph

eroi

dU

plan

d fo

rest

Rai

ny s

easo

n/D

ry s

easo

nI

Bac

tris

gas

ipae

s 22

30.0

044

.1O

blon

g, O

void

Upl

and

fore

stR

ainy

sea

son

R

Ber

thol

leti

a ex

cels

a 65

00.0

035

.0T

rian

gula

rU

plan

d fo

rest

Rai

ny s

easo

nI

Bix

a or

ella

na31

.12

31.4

Ovo

idU

plan

d fo

rest

, lo

wla

nd f

ores

t R

ainy

/Dry

sea

son

O

Cal

ophy

llum

bra

sili

ense

27

00.0

060

.0O

blon

gU

plan

d fo

rest

, lo

wla

nd f

ores

tD

ry s

easo

nR

?

Cal

ycop

hyll

um s

pruc

eanu

m

0.13

5.5

Fla

tL

owla

nd f

ores

t, fl

oode

d fo

rest

Dry

sea

son

O

Car

apa

guia

nens

is

1938

.00

47.0

Cun

eifo

rmL

owla

nd f

ores

t, fl

oode

d fo

rest

Dry

sea

son

R

Car

apa

proc

era

1500

.00

52.0

Cun

eifo

rmL

owla

nd f

ores

t, fl

oode

d fo

rest

Rai

ny s

easo

n/D

ry s

easo

nR

Car

inia

na m

icra

ntha

15

7.00

12.0

Fla

tL

owla

nd f

ores

tD

ry s

easo

nO

Cas

eari

a sy

lves

tris

1.

9622

.9O

blon

gL

owla

nd f

ores

tD

ry s

easo

nO

Cas

sia

ferr

ugin

ea

60.5

011

.3F

lat

Upl

and

fore

stD

ry s

easo

nO

Ced

rela

fis

sili

s 22

.60

22.4

Fla

tU

plan

d fo

rest

, m

ount

ain

fore

stD

ry s

easo

nO

Ced

rela

odo

rata

48

.00

13.8

Fla

tL

owla

nd,

upla

nd f

ores

tD

ry s

easo

nO

Cei

ba p

enta

ndra

49

.00

20.0

Sph

eroi

dL

owla

nd f

ores

tD

ry s

easo

nO

Cen

osti

gma

toca

ntin

um

487.

8012

.3F

lat

Low

land

for

est

Dry

sea

son

O

Cla

risi

a ra

cem

osa

1000

.00

52.2

Obl

ong

Upl

and

fore

st,

low

land

for

est

Rai

ny s

easo

nR

Cop

aife

ra m

ulti

juga

14

16.0

034

.0O

blon

gU

plan

d fo

rest

, lo

wla

nd f

ores

t, F

lood

ed f

ores

tD

ry s

easo

nI

Cor

dia

goel

dian

a 3

5.00

18.0

Obl

ong

Low

land

for

est

Dry

sea

son

O

Cou

epia

lon

gipe

ndul

a31

034.

0079

.0O

blon

gL

owla

nd f

ores

tR

ainy

sea

son

R

Cou

ma

util

is

330.

0026

.6F

lat

Low

land

for

est

Rai

ny s

easo

nO

Tab

le 2

. In

form

atio

n on

tho

usan

d-se

ed w

eigh

t (T

SW

), s

eed

moi

stur

e co

nten

t at

she

ddin

g (M

CS

), s

eed

shap

e, e

colo

gy a

nd h

abit

at a

nd p

erio

d of

fru

it o

r se

ed

disp

ersa

l in

rel

atio

n to

ort

hodo

x (O

), r

ecal

citr

ant

(R)

or i

nter

med

iate

(I)

see

d st

orag

e be

havi

our

(SS

B).

Spec

ies

TSW

(g

)

M

CS

(%

, w.b

.)Se

ed s

hape

Hab

itat

Fru

it/s

eed

disp

ersa

l pe

riod

SSB

374

M. DE JR. LIMA, T.D. HONG, Y.M.B.C. ARRUDA, A.M.S. MENDES AND R.H. ELLISTa

ble

2. C

onti

nued

Spec

ies

TSW

(g

)

M

CS

(%

, w.b

.)Se

ed s

hape

Hab

itat

Fru

it/s

eed

disp

ersa

l pe

riod

SSB

Cou

rata

ri a

trov

inos

a13

0.00

26.7

Fla

tL

owla

nd f

ores

t D

ry s

easo

nO

Cou

rata

ri g

uian

ensi

s11

0.00

32.6

Fla

tL

owla

nd f

ores

tR

ainy

sea

son

O

Cou

rata

ri l

ongi

pedi

cell

ata

110.

0034

.4F

lat

Low

land

for

est

Dry

sea

son

O

Cou

rata

ri s

tell

ata

170.

0026

.1F

lat

Low

land

for

est

Dry

sea

son

O

Cyn

omet

ra b

auhi

niif

olia

14

14.0

051

.3

Obl

ong

Low

land

for

est,

floo

ded

fore

stD

ry s

easo

nR

Din

izia

exc

elsa

27

1.70

20.0

Fla

tL

owla

nd f

ores

tD

ry s

easo

nO

Dip

tery

x od

orat

a 22

88.0

032

.0E

llip

soid

Low

land

for

est,

floo

ded

fore

stD

ry s

easo

nI

Ent

erol

obiu

m s

chom

burg

kii

47.8

011

.9F

lat

Low

land

for

est,

floo

ded

fore

stD

ry s

easo

nO

Eug

enia

sti

pita

ta

2834

.00

51.4

Obl

ong

Low

land

, up

land

for

est

Dry

sea

son/

Rai

ny s

easo

nR

Eut

erpe

ole

race

a 11

29.0

038

.5

Sph

eroi

dU

plan

d fo

rest

, lo

wla

nd f

ores

tD

ry s

easo

nR

Eut

erpe

pre

cato

ria

837.

1038

.6S

pher

oid

Upl

and

fore

st,

low

land

for

est

Dry

sea

son

R

Gen

ipa

amer

ican

a 7.

5040

.6F

lat

Upl

and

fore

st,

low

land

for

est

Rai

ny s

easo

nI

Gou

pia

glab

ra

1.70

6T

rian

gula

rL

owla

nd f

ores

t, fl

oode

d fo

rest

Dry

sea

son/

Rai

ny s

easo

nO

Han

droa

nthu

s se

rrat

ifol

ius

125.

006.

0F

lat

Low

land

for

est,

upla

nd f

ores

tD

ry s

easo

nO

Hel

icos

tyli

s to

men

tosa

25

0.00

33.0

Obl

ong

Upl

and

fore

st,

low

land

for

est

Rai

ny s

easo

nR

Hev

ea b

rasi

lien

sis

4232

.00

26.9

Sph

eroi

dL

owla

nd f

ores

t, fl

oode

d fo

rest

Rai

ny s

easo

nR

Hym

enae

a co

urba

ril

4097

.00

20.0

Ovo

idL

owla

nd f

ores

t, fl

oode

d fo

rest

Dry

sea

son

O

Inga

cin

nam

omea

65

05.0

050

.7O

blon

gU

plan

d fo

rest

, lo

wla

nd f

ores

tD

ry s

easo

n/R

ainy

sea

son

R

Lec

ythi

s pi

soni

s 87

87.0

020

.8O

blon

gL

owla

nd f

ores

t, fl

oode

d fo

rest

Dry

sea

son

O

Mau

riti

a fl

exuo

sa

1900

0.00

45.6

Ovo

idL

owla

nd f

ores

t, fl

oode

d fo

rest

Dry

sea

son/

Rai

ny s

easo

nR

Min

quar

tia

guia

nens

is

700.

0040

.0O

blon

gL

owla

nd f

ores

t, fl

oode

d fo

rest

Dry

sea

son

R

Myr

ciar

ia d

ubia

43

4.00

56.8

Bea

n sh

ape

Upl

and

fore

st,

low

land

for

est

Dry

sea

son

R

Och

rom

a py

ram

idal

e 6.

306.

6P

irif

orm

Upl

and

fore

st,

low

land

for

est

Dry

sea

son

O

Oen

ocar

pus

baca

ba

1440

1.00

53.6

Ovo

idU

plan

d fo

rest

, lo

wla

nd f

ores

tD

ry s

easo

n/R

ainy

sea

son

R

Oen

ocar

pus

bata

ua

9100

.00

41.7

Ovo

idL

owla

nd f

ores

t, fl

oode

d fo

rest

Dry

sea

son/

Rai

ny s

easo

nR

Tabl

e 2

con’

t

375

SEED STORAGE BEHAVIOUR OF AMAZONIAN TREESTa

ble

2. C

onti

nued

Spec

ies

TSW

(g

)

M

CS

(%

, w.b

.)Se

ed s

hape

Hab

itat

Fru

it/s

eed

disp

ersa

l pe

riod

SSB

Oen

ocar

pus

map

ora

2487

.00

31.8

Ovo

idL

owla

nd f

ores

t, fl

oode

d fo

rest

Dry

sea

son

R

Oen

ocar

pus

min

or

2564

.00

46.5

Ovo

idL

owla

nd f

ores

tD

ry s

easo

n/R

ainy

sea

son

R

Orm

osia

exc

elsa

13

58.0

018

.0O

void

Flo

oded

for

est

Dry

sea

son

O

Par

kia

disc

olor

36

1.00

9.8

Obl

ong

Low

land

for

est

Dry

sea

son

O

Par

kia

mul

tiju

ga

4000

.00

10.0

Obl

ong

Low

land

for

est

Dry

sea

son

O

Par

kia

niti

da

989.

5010

.3O

blon

gL

owla

nd f

ores

t, fl

oode

d fo

rest

Dry

sea

son

O

Par

kia

pend

ula

101.

507.

3O

blon

gL

owla

nd f

ores

t, fl

oode

d fo

rest

Dry

sea

son

O

Pse

udob

omba

x m

ungu

ba

22.1

015

.2S

pher

oid

Flo

oded

for

est

Dry

sea

son

O

Psi

dium

acu

tang

ulum

11

5.00

32.6

Sph

eroi

dU

plan

d fo

rest

, lo

wla

nd f

ores

tD

ry s

easo

n/R

ainy

sea

son

O

Sche

ffle

ra m

orot

oton

i 13

.50

35.0

Fla

tU

plan

d fo

rest

, lo

wla

nd f

ores

t,F

lood

ed f

ores

tD

ry s

easo

nO

Schi

zolo

bium

am

azon

icum

90

0.00

12.5

Fla

tL

owla

nd f

ores

t, fl

oode

d fo

rest

Dry

sea

son

O

Spon

dias

mom

bin

2143

.00

*4.1

Obl

ong

Low

land

for

est,

floo

ded

fore

stR

ainy

sea

son

O

Stry

phno

dend

ron

guia

nens

e 10

2.00

23.1

Obl

ong

Upl

and

fore

st,

low

land

for

est

Dry

sea

son

O

Stry

phno

dend

ron

mic

rost

achy

um

100.

0020

.0O

blon

gU

plan

d fo

rest

, lo

wla

nd f

ores

tD

ry s

easo

nO

Swie

teni

a m

acro

phyl

la

2000

.00

35.0

Fla

tU

plan

d fo

rest

, lo

wla

nd f

ores

tD

ry s

easo

nO

Tap

irir

a gu

iane

nsis

5

2.0

36.0

Ovo

idL

owla

nd f

ores

t. fl

oode

d fo

rest

Rai

ny s

easo

nR

Tet

raga

stri

s pa

nam

ensi

s 45

9.20

17.0

Ovo

idU

plan

d fo

rest

, lo

wla

nd f

ores

tD

ry s

easo

nO

The

obro

ma

gran

difl

orum

48

78.0

057

.6B

ean

shap

eL

owla

nd f

ores

t, fl

oode

d fo

rest

Rai

ny s

easo

nR

Vir

ola

suri

nam

ensi

s10

0.00

24.9

Ovo

idL

owla

nd f

ores

t, fl

oode

d fo

rest

Rai

ny s

easo

nI

* N

o dr

ying

trea

tmen

t app

lied

afte

r co

llect

ion

(Sal

omão

, 200

2).

376


able

3.

Sum

mar

y of

see

d dr

ying

and

sto

rage

dat

a w

ith

prob

able

see

d st

orag

e be

havi

our

of 6

7 tr

ee s

peci

es f

rom

Am

azon

ia:

O (

orth

odox

), I

(in

term

edia

te),

R (

reca

lcit

rant

) (m

c =

moi

stur

e co

nten

t).

Spe

cies

Germination (%) at shedding/harvest

Lowest safe mc (wb)Critical mc (wb)

Moisture content (%) at which germination

reduced to 0%

Germination (%)on desiccation to

10% mc


5% mc

Viability in subsequent hermetic

air-dry storage (5-10% mc) -18°C

Seed storagebehaviour

R

efer

ence

C

omm

ent

Ade

nant

hera

pa

voni

na91

NR

O§

NR

O§

91-

-O

Hon

g et

al.

(199

6),

Oli

veir

a et

al.

(201

2)

No

loss

in

viab

ilit

y on

des

icca

tion

to

5% m

c, a

nd

herm

etic

sto

rage

at

-18°

C (

Ell

is e

t al

., 20

07).

Ali

bert

ia

edul

is91

NR

O§

NR

O§

--

-O

BC

SN

A (

2013

)

No

loss

in

viab

ilit

y in

sto

rage

wit

h 5-

10%

mc

and

5°C

(B

CS

NA

(20

13);

cla

ssif

ied

as O

? (H

ong

et a

l., 1

996)

; de

sicc

atio

n to

lera

nt (

Daw

s et

al.,

20

06).

Ani

ba

rosa

eodo

ra76

4020

0-

-R

Sam

paio

et

al.

(200

3)

Cla

ssif

ied

as R

(H

ong

et a

l., 1

996)

.

Ast

roca

ryum

ac

ulea

tum

589.

85.

7-6.

940

0-

IR

ebou

ças

(201

0)N

ot r

epre

sent

ed i

n S

ID**

.

Bac

tris

ga

sipa

es87

4012

-16

--

-R

Bov

i et

al.

(200

4),

Fer

reir

a (2

005)

D

esic

cati

on s

ensi

tive

(D

aws

et a

l., 2

005,

200

6).

Ber

thol

leti

a

exce

lsa

6412

< 4

.2-

2-

I

Fig

ueir

edo

et a

l. (1

990)

, H

ong

et a

l. 19

96),

San

tos

et a

l. (2

006)

Cla

ssif

ied

as I

(H

ong

et a

l., 1

996)

.

Bix

a or

ella

na86

NR

O§

NR

O§

8662

39-4

0O

Gol

dbac

h (1

979)

Bas

ed o

n da

ta g

iven

by

Gol

dbac

h (1

979)

, H

ong

et a

l.(19

96)

clas

sifi

ed a

s I,

but

rec

lass

ifie

d no

w

as O

, si

nce

viab

ilit

y m

aint

aine

d fo

r tw

o ye

ars

in

herm

etic

sto

rage

at

-20°

C w

ith

< 7

% m

c (E

ira

and

Mel

lo,

1997

); n

o lo

ss a

t 5.

2% m

c an

d 0°

C

for

20 m

onth

s (P

oorn

ima

and

Am

bika

, 20

12);

no

loss

in

viab

ilit

y af

ter

nine

mon

ths

at 7

% m

c an

d -2

0°C

(C

orle

tt e

t al

., 20

07).

Tab

le 3

con

’t

§ N

RO

, co

lum

n no

t re

leva

nt f

or o

rtho

dox

seed

sto

rage

beh

avio

ur;

-, n

ot d

eter

min

ed;

* D

T,

desi

ccat

ion

tole

rant

;**

SID

dat

a.ke

w.o

rg/s

id/s

idse

arch

.htm

l; a

cces

sed

on 2

1 Ju

ly 2

014.

377


Spe

cies




reduced to 0%


10% mc


5% mc




R

efer

ence

C

omm

ent

Tab

le 3

. C

onti

nued

Cal

ophy

llum

br

asil

iens

e96

28.2

20.9

--

-R

?

Sil

va (

2005

),

Car

lota

(20

06),

Jo

ker

and

Sal

azar

(2

000)

, C

arva

jal

et

al.

(201

3)

Sil

va (

2005

), C

arlo

ta (

2006

), J

oker

and

Sal

azar

(2

000)

, C

arva

jal

et a

l. (2

013)

cla

ssif

ied

as R

, bu

t 4%

of

seed

s ge

rmin

ated

on

desi

ccat

ion

to 4

% m

c (C

arlo

ta,

2006

) su

gges

ting

it

may

not

be

R.

Cal

ycop

hyll

um

spru

cean

um70

NR

O§

NR

O§

-D

T*

at

2.9%

m

cN

o lo

ssO

Alm

eida

(20

04),

C

onse

rva

et a

l. (2

013)

Not

rep

rese

nted

in

SID

**.

Car

apa

gu

iane

nsis

90-

180

--

RF

erra

z (2

003)

, F

erra

z an

d S

ampa

io

(199

6)

Cla

ssif

ied

as R

in

SID

**.

Car

apa

proc

era

90-

200

--

RF

erra

z (2

004)

, H

ong

et a

l. (1

996)

Cla

ssif

ied

as R

in

SID

**.

Car

inia

na

mic

rant

ha73

NR

O§

NR

O§

-D

T*

at

6% m

c-

OC

amar

go a

nd

Fer

raz

(200

7)S

imil

arly

, C

. es

trel

lens

is s

how

ed O

(H

ong

et a

l.,

1996

). N

ot r

epre

sent

ed i

n S

ID**

.C

asea

ria

sy

lves

tris

29.6

NR

O§

NR

O§

-D

T*

at

7% m

c-

OM

atom

i et

al.

(200

9)N

o lo

ss i

n vi

abil

ity

at 7

% m

c w

ith

ambi

ent

tem

pera

ture

aft

er o

ne y

ear

(Mat

omi

et a

l., 2

009)

.

Cas

sia

ferr

ugin

ea67

NR

O§

NR

O§

DT

*N

o lo

ss-

OB

CS

NA

(20

13)

No

loss

in

viab

ilit

y in

her

met

ic s

tora

ge a

t 11

%

mc

and

5°C

(B

CS

NA

, 20

13).

Ced

rela

fis

sili

s86

.5N

RO

§N

RO

§N

o lo

ssN

o lo

ssN

o lo

ssO

Cor

vell

o et

al.

(199

9),

Mei

deir

os

and

Eir

a (2

006)

Sur

viva

l in

LN

2 (S

alom

ão,

2002

).

Ced

rela

odo

rata

92N

RO

§N

RO

§96

9292

OL

ima

and

Ell

is

(200

5)

Cei

ba p

enta

ndra

88N

RO

§N

RO

§90

8580

OL

ima

and

Ell

is

(200

5),

Lim

a et

al.

(200

0)

Tab

le 3

con

’ t

§ N

RO

, co

lum

n no

t re

leva

nt f

or o

rtho

dox

seed

sto

rage

beh

avio

ur;

-, n

ot d

eter

min

ed;

* D

T,

desi

ccat

ion

tole

rant

;**

SID

dat

a.ke

w.o

rg/s

id/s

idse

arch

.htm

l; a

cces

sed

on 2

1 Ju

ly 2

014.

378


Spe

cies




reduced to 0%


10% mc


5% mc




R

efer

ence

C

omm

ent

Tab

le 3

. C

onti

nued

Cen

osti

gma

to

cant

inum

94N

RO

§N

RO

§94

92.5

-O

BC

SN

A (

2013

);

Gar

cia

et a

l. (2

008)

N

ot r

epre

sent

ed i

n S

ID**

.

Cla

risi

a

race

mos

a89

5015

.90

--

RS

anto

s et

al.

(200

8)N

ot r

epre

sent

ed i

n S

ID**

.

Cop

aife

ra

mul

tiju

ga94

-10

-12

--

-I

Bru

m e

t al

. (2

009)

But

C.

lang

dorf

gii

and

othe

r C

opai

fera

spp

, or

thod

ox (

SID

**).

Cor

dia

go

eldi

ana

54N

RO

§N

RO

§-

--

OH

ong

et a

l. (1

996)

, V

iann

a (1

982b

)

See

ds s

urvi

ved

in s

tora

ge a

t 30

% r

.h.

for

seve

n m

onth

s (V

iann

a, 1

982b

). O

ther

Cor

dia

spec

ies

orth

odox

(S

ID**

).

Cou

epia

lo

ngip

endu

la87

.5-

9.8

--

-R

Sil

va e

t al

. (2

010)

Des

icca

tion

sen

siti

ve (

Sil

va e

t al

., 20

10).

Not

re

pres

ente

d in

SID

**.

Cou

ma

ut

ilis

95.8

19.2

NR

O§

NR

O§

-D

T*

23.5

Fea

sibl

eO

Hon

g et

al.

(199

6),

Sil

va e

t al

. (2

010)

T

oler

ance

of

dryi

ng a

nd f

reez

ing,

lon

g-te

rm

cons

erva

tion

in

gene

bank

s is

fea

sibl

e (E

copo

rt).

Cou

rata

ri

atro

vino

sa10

0N

RO

§N

RO

§-

98

-O

Oli

veir

a (2

010)

Not

rep

rese

nted

in

SID

**.

Cou

rata

ri

guia

nens

is40

N

RO

§N

RO

§-

88

-O

Oli

veir

a (2

010)

Not

rep

rese

nted

in

SID

**.

Cou

rata

ri

long

iped

icel

lata

52

NR

O§

NR

O§

-77

-

OO

live

ira

(201

0)N

ot r

epre

sent

ed i

n S

ID**

.

Cou

rata

ri

stel

lata

53

NR

O§

NR

O§

-25

-

OO

live

ira

(201

0)N

ot r

epre

sent

ed i

n S

ID**

.

Cyn

omet

ra

bauh

inii

foli

a94

41.4

-46

.6<

28.

6 >

23

.90

--

RB

CS

NA

(20

13),

N

azár

io e

t al

. (2

008)

Not

rep

rese

nted

in

SID

**.

Tab

le 3

con

’t

§ N

RO

, co

lum

n no

t re

leva

nt f

or o

rtho

dox

seed

sto

rage

beh

avio

ur;

-, n

ot d

eter

min

ed;

* D

T,

desi

ccat

ion

tole

rant

;**

SID

dat

a.ke

w.o

rg/s

id/s

idse

arch

.htm

l; a

cces

sed

on 2

1 Ju

ly 2

014.

379


Spe

cies




reduced to 0%


10% mc


5% mc




R

efer

ence

C

omm

ent

Tab

le 3

. C

onti

nued

Din

izia

ex

cels

a90

NR

O§

NR

O§

DT

*D

T*

Fea

sibl

eO

Mes

quit

a et

al.

(200

9)N

ot r

epre

sent

ed i

n S

ID**

.

Dip

tery

x

odor

ata

97-

< 1

213

-16

00

0I

BC

SN

A (

2013

),

Car

valh

o (2

009b

),

Hon

g et

al.

(199

6)

Des

icca

tion

sen

siti

ve (

Car

valh

o, 2

009b

), b

ut

Dip

tery

x al

ata

Ort

hodo

x (H

ong

et a

l., 1

996)

.

Ent

erol

obiu

m

scho

mbu

rgki

i99

NR

O§

NR

O§

DT

* at

10

%

mc

DT

* at

5%

mc

-O

BC

SN

A (

2013

),

Hor

ing

et a

l. (2

012)

Via

bili

ty m

aint

aine

d in

her

met

ic s

tora

ge a

t 11

.9%

m

c an

d 5

°C (

BC

SN

A,

2013

). N

ot r

epre

sent

ed

in S

ID**

.

Eug

enia

stip

itat

a80

-25

.815

-20

--

-R

Del

gado

and

B

arbe

do (

2007

),

Lop

es (

2010

),

Gen

til

and

Fer

reir

a (1

999)

Not

rep

rese

nted

in

SID

**.

Eut

erpe

ol

erac

ea90

40 37.4

20-3

021

0 0-

-R

BC

SN

A (

2013

),

Car

valh

o et

al.

(200

1),

Hon

g et

al.

(199

6),

Nas

cim

ento

(2

006,

200

8)

Not

rep

rese

nted

in

SID

**.

Eut

erpe

pr

ecat

oria

84.8

-17

0-

-R

BC

SN

A (

2013

),

Hon

g et

al.

(199

6)

Gen

ipa

am

eric

ana

98 96c.

10

4.2

91 DT

*

45L

ittl

e lo

ss

0 0I

Mag

istr

ali

et a

l. (2

013)

SID

** c

lass

ifie

d as

O?

Gou

pia

glab

ra95

NR

O§

NR

O§

No

loss

No

loss

Fea

sibl

eO

Sch

wen

gber

and

S

mid

erle

(20

05)

Not

rep

rese

nted

in

SID

**.

Tab

le 3

con

’t

§ N

RO

, co

lum

n no

t re

leva

nt f

or o

rtho

dox

seed

sto

rage

beh

avio

ur;

-, n

ot d

eter

min

ed;

* D

T,

desi

ccat

ion

tole

rant

;**

SID

dat

a.ke

w.o

rg/s

id/s

idse

arch

.htm

l; a

cces

sed

on 2

1 Ju

ly 2

014.

380


Spe

cies




reduced to 0%


10% mc


5% mc




R

efer

ence

C

omm

ent

Tab

le 3

. C

onti

nued

Han

droa

nthu

s

serr

atif

oliu

s10

098

NR

O§

NR

O§

-98

-O

Fer

reir

a et

al.

(200

4),

Sal

omão

(20

02)

No

loss

in

LN

2 (S

alom

ão,

2002

).

Hel

icos

tyli

s

tom

ento

sa73

3816

0-

-R

Arr

uda

and

Fer

raz

(200

8)N

ot r

epre

sent

ed i

n S

ID**

.

Hev

ea

bras

ilie

nsis

7336 20

12 150 -

- -0 0

RB

CS

NA

(20

13),

H

ong

et a

l. (1

996)

, B

onom

e (2

006)

Des

icca

tion

sen

siti

ve (

Daw

s et

al.,

200

5, 2

006)

.

Hym

enae

a

cour

bari

l87 21

NR

O§

NR

O§

87 19- -

85 20O

BC

SN

A (

2013

),

Car

valh

o et

al.

(200

6),

Hon

g et

al.

(199

6)

No

loss

in

viab

ilit

y in

her

met

ic s

tora

ge a

t 10

.2%

m

c an

d -1

8°C

, fo

r th

ree

mon

ths

(Car

valh

o et

al.,

20

06).

Inga

ci

nnam

omea

97.5

--

--

-R

Car

valh

o et

al.

(200

1),

Med

eiro

s an

d E

ira

(200

6)N

ot r

epre

sent

ed i

n S

ID**

.

Lec

ythi

s pi

soni

s51

NR

O§

NR

O§

-58

60

O C

arva

lho

et a

l. (2

006)

Not

rep

rese

nted

in

SID

**.

Mau

riti

a

flex

uosa

82-

> 1

3.6

--

-R

Sou

sa e

t al

. (2

004)

Cla

ssif

ied

as R

(S

ID**

).

Min

quar

tia

gu

iane

nsis

7540

27-

--

RC

amar

go a

nd

Fer

raz

(200

5)N

ot r

epre

sent

ed i

n S

ID**

.

Myr

ciar

ia d

ubia

93-

190

--

RF

erre

ira

and

Gen

til

(200

3),

Gen

til

et a

l. (2

004)

Not

rep

rese

nted

in

SID

**.

Och

rom

a

pyra

mid

ale

75N

RO

§N

RO

§N

o lo

ssN

o lo

ss-

OB

CS

NA

(20

13),

H

ong

et a

l. (1

996)

, P

into

et

al.

(200

4)D

esic

cati

on t

oler

ance

(D

aws

et a

l, 20

05,

2006

).

Tab

le 3

con

’t

§ N

RO

, co

lum

n no

t re

leva

nt f

or o

rtho

dox

seed

sto

rage

beh

avio

ur;

-, n

ot d

eter

min

ed;

* D

T,

desi

ccat

ion

tole

rant

;**

SID

dat

a.ke

w.o

rg/s

id/s

idse

arch

.htm

l; a

cces

sed

on 2

1 Ju

ly 2

014.

381


Spe

cies




reduced to 0%


10% mc


5% mc




R

efer

ence

C

omm

ent

Tab

le 3

. C

onti

nued

Oen

ocar

pus

ba

caba

85-

26.5

19.1

0-

-R

BC

SN

A (

2013

),

José

et

al.

(201

2),

Sil

va e

t al

. (2

010)

Des

icca

tion

sen

siti

ve (

Daw

s et

al.,

200

5).

Not

re

pres

ente

d in

SID

**.

Oen

ocar

pus

ba

taua

93-

24.3

0-

-R

BC

SN

A (

2013

),

Naz

ário

and

F

erre

ira

(201

2)N

ot r

epre

sent

ed i

n S

ID**

.

Oen

ocar

pus

m

apor

a86

.3-

--

--

RC

arva

lho

et a

l. (2

001)

Not

rep

rese

nted

in

SID

**.

Oen

ocar

pus

m

inor

98-

--

--

RC

arva

lho

et a

l. (2

001)

Not

rep

rese

nted

in

SID

**.

Orm

osia

ex

cels

a77

NR

O§

NR

O§

No

loss

--

OB

CS

NA

(20

13)

No

loss

in

viab

ilit

y in

her

met

ic s

tora

ge a

t 11

%

mc

and

5°C

(B

CS

NA

(20

13).

O.

fast

igia

ta

surv

ived

in

LN

2 (S

alom

ão,

2002

). N

ot

repr

esen

ted

in S

ID**

.

Par

kia

disc

olou

r99

NR

O§

NR

O§

No

loss

--

OB

CS

NA

(20

13)

Via

bili

ty m

aint

aine

d in

her

met

ic s

tora

ge a

t 5°

C

(BC

SN

A (

2013

). N

ot r

epre

sent

ed i

n S

ID**

.

Par

kia

mul

tiju

ga90

NR

O§

NR

O§

No

loss

--

OC

arva

lho

(200

9a),

S

anto

s (2

011)

Not

rep

rese

nted

in

SID

**.

Par

kia

niti

da80

NR

O§

NR

O§

No

loss

--

OC

ruz

et a

l. (2

001)

, F

erra

z et

al.

(200

4)N

ot r

epre

sent

ed i

n S

ID**

.

Par

kia

pend

ula

63N

RO

§N

RO

§99

99-

OR

osse

to (

2006

)V

iabi

lity

mai

ntai

ned

in h

erm

etic

sto

rage

at

5.9%

m

c an

d 5°

C (

Ros

seto

, 20

06).

Not

rep

rese

nted

in

SID

**.

Pse

udob

omba

x

mun

guba

78N

RO

§N

RO

§N

o lo

ss-

-O

BC

SN

A (

2013

)S

urvi

val

in L

N2

(Sal

omão

, 20

02).

Not

re

pres

ente

d in

SID

**.

Tab

le 3

con

’t

§ N

RO

, co

lum

n no

t re

leva

nt f

or o

rtho

dox

seed

sto

rage

beh

avio

ur;

-, n

ot d

eter

min

ed;

* D

T,

desi

ccat

ion

tole

rant

;**

SID

dat

a.ke

w.o

rg/s

id/s

idse

arch

.htm

l; a

cces

sed

on 2

1 Ju

ly 2

014.

382


Spe

cies




reduced to 0%


10% mc


5% mc




R

efer

ence

C

omm

ent

Tab

le 3

. C

onti

nued

Psi

dium

ac

utan

gulu

m69

.8N

RO

§N

RO

§N

o lo

ssN

o lo

ssN

o lo

ssO

Car

valh

o et

al.

(200

1)

Via

bili

ty m

aint

aine

d in

her

met

ic s

tora

ge a

t 5%

mc

and

-18°

C (

Car

valh

o et

al.,

200

1).

Not

re

pres

ente

d in

SID

**.

Sche

ffle

ra

mor

otot

oni

70N

RO

§N

RO

§D

T*

--

OO

hash

i an

d L

eão

(200

5)V

iabi

lity

mai

ntai

ned

in h

erm

etic

sto

rage

at

10%

m

c at

12°

C (

Oha

shi

and

Leã

o, 2

005)

.

Schi

zolo

bium

am

azon

icum

100

NR

O§

NR

O§

No

loss

--

OH

ong

et a

l. (1

996)

, S

ouza

et

al.

(200

5)V

iabi

lity

mai

ntai

ned

in c

old

room

(0-

5°C

) w

ith

40%

r.h

. (S

ouza

et

al.,

2005

).

Spon

dias

mom

bin

76.3

NR

O§

NR

O§

-D

T*

No

loss

O

Hon

g et

al.

(199

6),

Car

valh

o et

al.

(200

1),

Wor

ld

Agr

ofor

estr

y

50%

ger

min

atio

n af

ter

27 m

onth

s st

ored

air

-dry

at

2-5

°C (

Wor

ld A

grof

ores

try)

; lo

ng-t

erm

sto

rage

at

5%

mc

and

-18°

C (

Car

valh

o et

al.,

200

1);

surv

ival

in

LN

2 (S

alom

ão,

2002

).

Stry

phno

dend

ron

gu

iane

nse

97N

RO

§N

RO

§N

o lo

ss-

-O

BC

SN

A (

2013

)S

urvi

ved

in s

tora

ge a

t 10

% m

c an

d 5°

C (

BC

SN

A

(201

3).

Not

rep

rese

nted

in

SID

**.

Stry

phno

dend

ron

m

icro

stac

hyum

89N

RO

§N

RO

§-

70-

OL

opes

(20

10)

Sur

vive

d in

sto

rage

at

10%

mc

and

5°C

(B

CS

NA

(2

013)

. N

ot r

epre

sent

ed i

n S

ID**

.

Swie

teni

a

mac

roph

ylla

100

NR

O§

NR

O§

100

98N

o lo

ssO

Ell

is e

t al

. (2

007)

For

mer

ly c

lass

ifie

d as

int

erm

edia

te (

Hon

g et

al.

1996

; H

ong

and

Ell

is,

1998

b; L

ima

and

Gal

vão,

20

05),

but

now

rec

lass

ifie

d as

ort

hodo

x (E

llis

et

al.,

2007

).

Tap

irir

a

guia

nens

is97

- -16

< 1

5.5

- -0 -

0 -R

Car

valh

o et

al.

(200

6),

San

tos-

Mou

ra e

t al

. (2

012)

Not

rep

rese

nted

in

SID

**.

Tet

raga

stri

s

pana

men

sis

87N

RO

§N

RO

§91

9278

O

BC

SN

A (

2013

)N

o lo

ss i

n vi

abil

ity

in h

erm

etic

sto

rage

at

5%

mc

and

-5°C

(B

CS

NA

(20

13).

Not

rep

rese

nted

in

SID

**.

Tab

le 3

con

’ t

§ N

RO

, co

lum

n no

t re

leva

nt f

or o

rtho

dox

seed

sto

rage

beh

avio

ur;

-, n

ot d

eter

min

ed;

* D

T,

desi

ccat

ion

tole

rant

;**

SID

dat

a.ke

w.o

rg/s

id/s

idse

arch

.htm

l; a

cces

sed

on 2

1 Ju

ly 2

014.

383


Spe

cies




reduced to 0%


10% mc


5% mc




R

efer

ence

C

omm

ent

Tab

le 3

. C

onti

nued

The

obro

ma

gr

andi

flor

um93

.841

.414

.50

--

RC

ruz

and

Cic

ero

(200

8)N

ot r

epre

sent

ed i

n S

ID**

.

Vir

ola

su

rina

men

sis

4119

-23

40

IH

ong

et a

l. (1

996)

, S

ID**

Alt

houg

h cl

assi

fied

as

reca

lcit

rant

by

mos

t au

thor

s (C

unha

et

al.,

1992

; C

esar

ino,

200

6;

Daw

s et

al.,

200

5, 2

006;

Lim

as e

t al

. 20

07),

bu

t 23

% a

nd 4

% g

erm

inat

ed o

n de

sicc

atio

n (a

t 15

% r

.h.

and

24°C

) to

11.

7% a

nd 5

.4%

mc,

re

spec

tive

ly,

and

no s

eeds

at

5.4-

26.5

% m

c su

rviv

ed t

hree

day

s in

sub

sequ

ent

stor

age

at 5

°C

or -

20°C

(C

unha

et

al.,

1992

), h

ence

rec

lass

ifie

d as

int

erm

edia

te (

Hon

g et

al.,

199

6);

SID

**.

§ N

RO

, co

lum

n no

t re

leva

nt f

or o

rtho

dox

seed

sto

rage

beh

avio

ur;

-, n

ot d

eter

min

ed;

* D

T,

desi

ccat

ion

tole

rant

;**

SID

dat

a.ke

w.o

rg/s

id/s

idse

arch

.htm

l; a

cces

sed

on 2

1 Ju

ly 2

014.

384


Discussion

The Amazon basin is a vast area with different microclimates, but with the typical attributes of tropical rainforests. Temperatures average 27.9°C during the dry season and 25.8°C during the rainy season with relative humidity averaging 88% in the rainy season and 77% in the dry season. Rainfall follows a seasonal pattern, with considerable spatial and temporal variation. For example, Iquitos in Peru receives all-year round rainfall averaging 2,616 mm per year, while Manaus in Brazil receives 1,771 mm and experiences a dry season (figure 1).

0

50

100

150

200

250

300

350

Rai

nfa

ll (m

m)

0

5

10

15

20

25

Ave

rag

e ra

iny

day

s(≥

0.1

mm

)

Figure 1. Rainfall (mm) and number of rainy days (> 0.1 mm) in Manaus, Brazil. Data from http://www.weather-and-climate.com/; accessed 21 July 2014.

The ecology of the Amazon rainforest is compatible with the survival of recalcitrant seeds, but ecology cannot be used as a diagnostic tool here to identify seed storage behaviour since a large number of indigenous species show orthodox or intermediate seed storage behaviour also (table 3). Nevertheless, the Amazon rainforest in Brazil contains a higher proportion of species with recalcitrant seed storage behaviour (34%, table 3), than does Panama (27.9%, Daws et al., 2005), Vietnam (21.7%, Ellis et al., 2007), African drylands (11%, Pritchard et al., 2004a), or worldwide (7.4%, Hong et al., 1996). This agrees with the observation of Tweddle et al. (2003) that desiccation-sensitive seeds are most common in tropical rainforests, where they suggest they constitute some 47% of higher plants.

385


Any putative association between seed storage behaviour and either natural habitat or seasonal period of seed dispersal is weak within this environment. The tree species tabulated here occur in one or more of the following habitats: upland or mountain forest at elevations > 600 m above sea level (a.s.l.); lowland forest from sea level to 600 m a.s.l.; and flooded (lowland) forest, where flooding occurs periodically for about four months each year. All three categories of seed storage behaviour can be found in each of upland, lowland or flooded forests with a higher proportion of orthodox species in the former (61 vs. 28% recalcitrant) than the latter (55 vs. 38% recalcitrant) (table 2).

A majority of species with seeds maturing and dispersing in the dry season showed orthodox seed storage behaviour (65%), whilst those species with seeds maturing and dispersing in the wet season showed the greatest frequency of recalcitrant seed storage behaviour (59%), but all three categories were present in each season (table 2). These findings tally with those of Daws et al. (2005, 2006).

Neither was seed shape a diagnostic tool, since species with ovoid, oblong or spheroid seeds provided all three categories of seed storage behaviour. However, flat seeds showed a degree of desiccation tolerance: of the 19 species with flat seeds 18 showed orthodox (95%) and one intermediate seed storage behaviour (5%) with no recalcitrant species with flat seeds identified (table 2). Similarly, flat seeds in Meliaceae showed either orthodox (80%) or intermediate seed storage behaviour (20%) (Hong and Ellis, 1998).

Seed moisture content at shedding (MCS) was not a useful single predictor of seed storage behaviour, although some discrimination was possible despite the overlapping values: MCS for species with orthodox seed storage behaviour varied between 5 and 35%, intermediate between 25 and 40%, and recalcitrant between 27 and 79% moisture content (table 2). Some discrimination, but with considerable overlaps, was also evident for seed size (weight) alone: orthodox seeds ranged from 0.00013 g / seed for Calycophyllum spruceanum to 8.79 g / seed for Lecythis pisonis; intermediate from 0.0075 for Genipa americana to 14.95 g / seed for Astrocaryum aculeatum; and recalcitrant from 0.052 g / seed for Tapirira guianensis to 31.03 g/seed for Couepia longipendula (table 2).

To test the utility of multiple-criteria keys to estimate seed storage behaviour, the exact schema developed previously by Hong et al. (1996, 1998a), Hong and Ellis (1996, 1997, 2002) and validated by Ellis et al. (2007) combining information on thousand-seed weight (TSW, g) and seed moisture content at shedding (MCS) and seed storage behaviour was applied to the data for 67 tree species from the Amazon (figure 2). In this presentation, all 26 species (39%) in zone I (left of solid line) show orthodox seed storage behaviour, whereas all 12 species (18%) in zone II (right of broken line) show recalcitrant seed storage behaviour. Hence the species in zones I or II matched precisely the expectation of orthodox or recalcitrant classification, respectively.

Zone III (located centrally between the two lines) is, as expected (Hong and Ellis, 1997), more difficult to discern since of these 29 species (43% of the total investigated), 12 show orthodox, six intermediate and 11 recalcitrant seed storage behaviour (figure 2). Within zone III, the additional (third) criterion of seed shape is most helpful: since they have flat seeds, we can estimate (correctly) that Aliberta edulis, Cedrela fissilis, Couma utilis, Couratari attrovinosa, C. guianensis, C. longipedicellata, C. stellata, Genipa americana and Schefflera morototoni are either orthodox or intermediate and not recalcitrant (table 2).

386


0.01

0.1

1

10

100

1000

10000

0 10 20 30 40 50 60 70 80

TS

W (

g, l

og

sca

le)

Seed moisture content (% w.b.) at shedding

I

III

II

Figure 2. Probable seed storage behaviour of 67 tree species from the Amazon forest in relation to seed moisture content (%, w.b.) at shedding (MCS) and thousand-seed fresh weight (TSW, g, logarithmic scale). Zone I, to the left of the vertical solid line at 22% moisture content, contains 26 species with orthodox seed storage behaviour ( ): Adenanthera pavonina, Bixa orellana, Calycophyllum spruceanum, Cariniana micrantha, Cassia ferruginea, Cedrela odorata, Ceiba pentandra, Cenostigma tocantinum, Cordia goeldiana, Dinizia excelsa, Enterolobium schomburgkii, Goupia glabra, Handroanthus serratifolius, Hymenaea courbaril, Lecythis pisonis, Ochroma pyramidale, Ormosia excelsa, Parkia discolour, P. multijuga, P. nitida, P. pendula, Pseudobombax munguba, Schizolobium amazonicum, Spondias mombin, Stryphnodendron microstachyum, Tetragastris panamensis. Zone II, to the right of the broken line (x1 = 40, y1 = 45000; and x2 = 65, y2 = 5) contains 12 species with recalcitrant seed storage behaviour ( ): Aniba rosaeodora, Calophyllum brasiliense, Carapa procera, Clarisia racemosa, Couepia longipendula, Cynometra bauhiniifolia, Eugenia stipitata, Inga cinnamomea, Mauritia flexuosa, Myrciaria dubia, Oenocarpus bacaba, Theobroma grandiflorum. Zone III, between the solid and broken lines, contains all three categories of seed storage behaviour: six intermediate ( ); Astrocaryum aculeatum, Bertholletia excelsa, Copaifera multijuga, Dipteryx odorata, Genipa americana, Virola surinamensis; 12 orthodox ( ); Alibertia edulis, Casearia sylvestris, Cedrela fissilis, Couma utilis, Couratari atrovinosa, C. guianensis, C. longipedicellata, C. stellata, Swietenia macrophylla, Psidium acutangulum, Schefflera morototoni, Stryphnodendron guianense; and 11 recalcitrant seed storage behaviour ( ): Bactris gasipaes, Carapa guianensis, Euterpe oleracea, E. precatoria, Helicostylis tomentosa, Hevea brasiliensis, Minquartia guianensis, Oenocarpus bataua, O. mapora, O. minor, Tapirira guianensis. Detailed information of TSW and MCS for each species is provided in table 2. The boundaries of the three zones are taken from figure 10 of Hong et al. (1996).

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Another potential explanatory variable to consider within zone III (i.e. in concert with seed weight and moisture content at shedding) is the season of shedding (the equivalent variable to rainfall in the month of shedding in Pritchard et al. [2004a]). This variable provided no discrimination therein. Of the 12 orthodox species in zone III, eight were shed in the dry season, three in the rainy and one in both; of the six intermediate species, two were shed in the dry season, three in the rainy and one in both; of the 11 recalcitrant species, five were shed in the dry season, four in the rainy season, and two in both. Moreover, within each of these nine subgroups in zone III there were no discernible trends with respect to seed weight or moisture content at shedding. The failure of season of shedding to discriminate within zone III for seed storage behaviour matches conclusions from Daws et al. (2006) who reported no significant effect of rainfall in the month of shedding within their probability of desiccation model.

The boundaries between zones I and III and between zones II and III are exactly thesame as the original introduction of this approach (Hong et al., 1996) and its subsequent validation with Vietnamese species (Ellis et al., 2007). Together these three studies demonstrate the consistency in the patterns and further confirm the position of the boundary values developed originally. Uncertain seed storage behaviour (zone III) ranged from 37% (Ellis et al., 2007) to 55% (Hong and Ellis, 1997) of species examined. This implies that about 45 to 63% of the workload required to investigate seed storage behaviour of wild species, where this is not known, can be avoided using the multiple criteria keys proposed previously (Hong and Ellis, 1997).

There are similarities between this approach to estimate likely seed storage behaviour from information on seed weight and moisture content at shedding and those to predict seed desiccation tolerance using seed weight and either rainfall in the month of shedding (Pritchard et al., 2004a) or the seed coat ratio (Daws et al., 2006). Both approaches are, similarly, also imperfect: zone III of the former is uncertain seed storage behaviour whilst, for example, a 50% probability of desiccation tolerance in the latter is equally unhelpful (albeit both outcomes are indirectly useful). The most obvious difference between the approaches is that the former attempts to classify seed storage behaviour into one of three categories whilst the latter provides a probability of desiccation tolerance. While neither provides a perfect diagnostic tool, the relative successes of the different approaches illustrate the considerable utility in combining several criteria to estimate how to conserve seeds of species where desiccation tolerance and/or seed storage behaviour are unproven; and that seed weight, shape (if flat), moisture content at shedding, rainfall in the month of shedding (perhaps), and the seed coat ratio are the pertinent variables where several are considered in conjunction. The approach to use before or during a seed collecting mission will depend on what information can be obtained. For example, detailed long-term meteorological data is now readily accessible for many locations, whereas information on the seed coat ratio may not be available.

In conclusion, 38 of the 67 tree species from Amazonian Brazil appeared to show orthodox (57%), 23 recalcitrant (34%) and six intermediate (9%) seed storage behaviour. The protocol and multiple criteria keys introduced and tested previously (Hong and Ellis, 1996, 1997; Hong et al., 1996; Ellis et al., 2007) have utility in reducing the workload in wild plant species biodiversity conservation, with the criteria seed weight, moisture content at shedding, and seed shape being most helpful in this respect when considered in concert.

388


Acknowledgements

We thank an anonymous reviewer for several helpful comments, the many institutes and authors whose publications cited here provided data for this paper, and the Conselho de Pesquisa Apoio ao Ensino Superior (CAPES), Brazil for providing M.J.V. Lima with a post-doctoral fellowship to study at the Seed Conservation Department, Royal Botanic Gardens, Kew, and at the University of Reading.

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363 SEED STORAGE BEHAVIOUR OF AMAZONIAN TREES Classification of seed storage behaviour of 67...

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