Evolutionary Biology of Acacia caven (Leguminosae, Mimosoideae): Infraspecific Variation in Fruit and Seed Characters

11 Pages·1992·7.3 MB·English

by  James Aronson

960
Annals the
of
Garden
Missouri Botanical
-Icm
Figure Typical 29343.— Variety
1. fruits of the six varieties of Acacia caven.—a. Variety caven Burkart h.
&
stenocarpa Krnpovickas 983.— Ka-
Variety microcarpa 7697.— Tressans
c. Rojas sphaerocarpa
Variety
d.
3539.—
dovancich drawings
Variety mnrrocarpa 17577.—
e. Burkart Burkart 15730. All
Variety dehiscens
{.
size.
life
incompatible, both an
in Argentinian
population
^^^^ DiSPERSAL
near Cordoba Anton,
(A. comm.)
pers. and
a Chil-
ean
pod
K'^
Pollin
fashion.
at least seeds are presumably dispersed in this
central Chile ^ prim^ ^a^^Br^^^y^mJ
in
beetles
in activ
seasonally
Chaco was near
habitat within the
(C
J^^
bees
today
waterways, where seen
often
it is
mo^
group common
pollinator the most
is for the genus same time,
Toledo, comm.)- At the
pers.
(Arroyo, 1981).
Ion/''
caven seem adapted for
varieties of A.
Number
4
Volume Aronson
79, 961
1992 Evolutionary Biology of Acacia caven
FicuRE
Each
2. Continental distribution of Acacia caven in southern South America. point represents a single
field
collection.
&
tance 48 Aronson, unpublished
dispersal by mega-vertebrates. In prehistoric with grains (Cuinet
numerous
data).
camelids,
stegomastodons, eden-
notoungulates,
Methods
tates, and giant sloths in the Chaco (Bucher, 1987). Materials and
At
present,
domestic livestock are undoubtedly the
FRUITS AND SEEDS
umportant
Armesto, pod
1981). Medium-sized such as
birds
((
charata" f^
{Ortalis and "corzuela covering
canicollis)^
{Mazama more 750
than her-
americana, M. Acacia caven. In addition,
as well as tapir
rufa),
caven were
{Tapirus specimens of A.
guanaco {Lama guanicoe), barium fruit-bearing
terreslris),
«nd and vegetative
sury (Rhea compared carpological traits.
americana) are probable dis- for
all
appeared
and seed characters
Parsers (pod)
of seed (C. Saravia Toledo, pers. comm.). Several fruit
and
three
populations,
constant within llip
relatively
earh
from populations of of
samples
largest fruit
Cytology
and
Palynology
analy
ted for
pod h of five or
ibe
^n
each
recorded 2n 26 1945) and more individuals was sufficient to represent
as (Castronovo,
= pods
52 since the addition of further failed
2^1 26, (Atchison, 1948). SimUarly v.a«riable population,
numbers
^__.eralized
occur A.farnesiana, „._
in the closely related
^hich = & Ashton, 1978). Only ripe pods free
is said to have 2n 52, 104 (Atchison,
Voucher specimens
damage were used.
1948).
Pollen shed large polyads
in relatively O
is
herbaria
are deposited at in
<^on£ibtingof 32 (Caeca van, 1970; Heus- julation
ca. grains
Kew and Missouri (M<
SCO), (K), in
*^r, 1971; Exceptions are at
Peralla
et in press).
al.,
pods per
grams
meapurwl
were
SCO) population
.
962
Annals
of the
Missouri Garden
Botanical
Table Leaf and peduncle
1. characters in six varieties of Acacia caven. Based on examination
of
fully developed
750
leaves of herbarium specimens.
Pairs of
Rachi
leaflets Pubescence Peduncle
IS Position of
Variety length (cm) per pinna (mm)
of rachis' length^ nectary on
rachis'
caven 5-7 4-6
A
5 15
or absent
spkaerocarpa 3-4 8-14
5-15 A
or absent
2-4
dehiscens
6 12 5-8 A
macrocarpa 6-10
3-4(-5) 3-5 A
microcarpa 3-7 4-6
10-15 A B
or
4-8
stenocarpa 4-8
10-25 A
B
or
—
±
usually glabrous;
variable.
'
Not shown
Figure
^ in
1
'A- —
at or near junction of first pair of pinnae; B midway from A to base of rachis.
estimated as pod length x xr^ Seeds were Chaco
re- the that periodically inundated for several
is
moved,
coimted, and weighed
separately from pods. months each
year. Variety dehiscens limited to
is
Average
seed weight
varied within popula-
little the rather dry, isolated range of mid-altitudinal
tions, as seen in a number of preliminary scale ranges between San and Cordoba,
stretching Luis
To
checks. determine average
seed weight each
for west-central Argentina; variety spkaerocarpa De-
population, seeds from each sample were
all pooled, curs only in high-rainfall areas of eastern Argentina
and random
three subsamples 100
of seeds were and caven
adjacent Uruguay. Variety
portions of
weighed. The average
weight of a single seed was the most widespread, demarking the outer pe-
is
calculated from mean
the resulting values. Data riphery of the geographic range of the whole spe-
were analyzed by Duncan
Range
Multiple
s test cies, except in Bolivia and Paraguay.
ANOVA
and two-way
A
to evaluate the caven
relative con- gap occurs in the distribution of A. in
tribution of population and
variety to overaU due
vari- the middle of the Chaco, undoubtedly to aridity.
ance.
Moreover, the other than variety ca-
five varieties
ven seem an pattern
to spread out in irregular
Results
remniscent
around
this natural gap. This situation
is
Africa
of that found Acacia karroo southern
Table shows in in
the comparison
of selected mor-
1
chemotypic
(Brain, 1989). In that species, three
among
phological
traits the six putative
varieties
around the tree-
allopatricaUy
Although some ''^''^^ °''*'"^ ^^"'""^
diagnostic
differences occur, veg-
Drakensberg The geographic distn-
'^^^ massif.
etative organs do
clearly not display the most
..--
dis-
.. —
.*.-
•*• witn
-^ +k^c-^ ^^^^r^ ar^pwp^^e^ar^r^sc correlated
V^»"ii^t^;*^-i"K^» -^~f^cf *^^^^ ^^^^^
tinctive infraspecific variation. Table 2 shows
in-
precip-
changes and of
in the quantity distribution
pod
polymor
m
^^^
17 "^*'''"- '" specimens of ^. caven,
populations
representing
the
six putative
been
phism has
a peroxidase banding pattern
in
varieties pod
was found be-
determined and
pod a clear distinction
(Brain
tween Chilean and non-Chilean populations
tinguished: caven, spkaerocarpa,
and
(i)
dekis-
&
needed
are
Aronson, Further studies
cens, macrocarpa, and in prep.).
microcarpa
(ii) and
(iii)
stenocarpa, which in thiIS area.
further
is distinguished by
nar-
row pod shape and
small seed
size.
Taxonomic
Treatment
Figure shows
2
collection
localities for the
six
Acacia caven Stor. r<la
varieties. Partial geographic (Molina) Molina, in Sag.
isolation occurs some
in
varieties, notably variety dehiscens ^nd 163; 299, 1810.
^^^^*' ed.,
in west-central
Argentina, and
variety microcarpa + much
m
variety
Shrub usuaUy
or tree to 10 high, but
p"""'° more
TrZ7T T' "' ""'^'^^^^ «--"-^ 5-20 or
smgle-stemmed or with
'.h"
"i
" ^^76"^y-
" Geographic
T,,T sep- on history
densely crowded depending
basal stems,
Cr?.rn
nrat on com^lated
.s to ecological
differentiation mottled
in
of cutting and you^g branchlels
burning;
most Thus,
cases.
variety
nu^crocarpa
not only
is JJs, pron,ine»|.
g.ay, sparingly puber not
lenticels
ZL^Z:Xo^rtr' ^''' "-''^^
^-^^^ ^^ Stipllarspin'ela4-2.5cmlong,slende;.not.wol-
above
2,000 m.
s.tes Both
variety murocarpa
and often
brown,
]«„ at the base white or gray rarely
variety stenocarpa
are
restricted
to tho
portion of borr Leaves bi-
mottled, ;airrafev;ry nlle.
in
Number
Volume 79, 4 Aronson 963
1992 Evolutionary Biology Acacia caven
of
4
Table 2. Means and ranges of four morphometric fruit and seed traits in six putative varieties of Acacia caven.
N =
For each variety, 240, from three populations. Means in a column followed by the same letter are not significantly
=
{P
different 0.05).
Fruits Seeds
I
Dry weight Seeds Dry seed
Variety Length (cm) Width (cm) Volume (cm^) per pod weight
(g)'
(g)
caven
5.1 a 1.5a 9.0a 5.4a 21.3a 1.23a
Range
(4.6-8.4) (1.4-2.3) (7.0-14) (3.7-9.7) (9-39) (1.2-1.3)
sphaerocarpa L15a
4.0b 2.9ab 6.3b 3.5a 22.9a
Range
(2.1-6.5) (1.6 4.8) (5.4-68) (1.5-6.0) (11-31) (1,0-1.3)
dehiiscens 5.5a 1.6a 11.9ab 6.3ab 18.4a 0.91b
Range
(3.3-7.5) (0.9-2.2) (7.1 18) (5.2-8.3) (4-28) (0.90-0.93)
macrocarpa
8.4c 2.2b 31.9c 9.8b 30.2b ].26a
Range
(6,2-11) (1.8-2.5) (26-38) (8.4-12.7) (16-41) (1.25-1.27)
mirrocarpa 0.47c
2.3d 1.2c 2.6d 0.9d 11.2c
Range
(1.7-3.1) (2.2-3.0) (0.7-1.4) (7-15) (0.4-0.5)
(0.5 1.5)
t
stenocarpa 3.7a 0.8c 2.4d 1.4d 12.4c 0.51c
Range
(3.0-6.5) (0.6-1.1) (1.9-3.6) (0.8-2.4) (8-17) (0.4-0.56)
With
seeds
pmnate 2 cm often with 2. Pod elongate-fusiform, more than twice as broad
long,
beak 4
as long, excluding
prmiary
Pod 2.0-3.0 X 1.0-1.5 cm; mesocarp
usually
3(2).
cm
2-5(-8)
rachis
long, sparsely puberulous; pinna mm
somewhat peduncle more than 15
reddish,
A
cm
pairs 4-8(- 0.7-1
typically rachiUae long; micracarpa
1 4); long var.
...
4 mesocarp
leaflets white,
lin 3.
mm
mm, peduncle than 5 long
^ 0.5 glabrous; midrib and nerves not less 1
lateral
sphaerocarpa
var.
„
readily
distinct. Peduncles usually pubescent but
Pod 3.0-4.0(-6.5) x 0.6- cm, narrowing
1.1
4(2).
mm
sometimes
3-15(-25) and
glabrous, long, rel-
both ends, sometimes subtorulose, often light
at
atively
constant in length within populations. Flow- brown, occasionally reddish purple, peduncle
mm
mm
er head 5-10 one usually more than 15 and rarely up to
spherical or long;
ellipsoid,
mm
stenocarpa
30 var.
or
usually „
several per highly prior
leaf axil, visible
Pod 5-11 X 1.5-2.5 cm, elongate, acumi-
4.
to the emergence mm
of Flowers yellow,
leaves. sessile,
peduncle than 15
nate, not torulose, less
mm
highly
fragrant. Calyx 0.5-1.0 glabrous, 5
long,
long
mm
1-3 30-
t-orolla slightly
approximately
long, glabrous, 5(4)
5-8 cm
with one butural ridge, this
55 44-58 pie, lung,
flowers per head; stamens per flower,
„
caven
mesocarp evanesrnit var.
often indistinct;
with
filaments free from the base and anthers eglan-
more
than
Pod usually reddish purpi«', usually
5.
atilate; pods brownish black or purplish brown, cm
8 long, with thrtf ilistinct sutural ridgeft,
oblong-elongate mesocarp resinous
or subglobose, straight or slightly these often sharply distinct;
mncrornrpn
var.
lalcate, frequently basaUy and/or acu-
stipilate
^>ttale, extremely volume and
variable in size,
weight;
indehiscent, except in variety dehiscens; Acacia caven var. caven, Mfmosacai^ea Molina,
±
glabrous, usually not striate. Mesocarp spongy Sag. Stor. Nat. Chili .t rd., 174. 1782. TYPE:
1
5-
or
Ran-
Seed ©Dip Neotype here designated:
tic, known.
not
mm.
Seeds 1828, Bertero (SCO).
cagua, Chile, Oct. s.n.
35
per pod red
Figure
la.
fully
ripe pods.
&
206.
Am.,
adennpa Hook. Mi^cell. 3:
Acacia Elot.
183.3.
Key
to
Fruiting Specimens of Acacia caven a.) WilW. var. brachicnrpn 0. Kuntw.
Acacia farnesiaiia
farnrsianail.)
1:156. 1891 Iraciti
Rev. Gen.
1. Mature dehiscens PI.
pod dehiscent var.
&
Am.) O.Kuntze, Rev.
cavenia (H(K)k
1. Mature 2 Willd. var,
pod
indehiscent
Vachelha farnesiann (L)
Gen. n. 3:47. 1898.
*^l). Pod
broad as
subglobose, than twice as
Ies6
& Acad.
Wight cairma (Molina) !^g., B<>I.
3 Arn.
f.
long, excluding beak and peduncle
J
•
964 Annals
of the
Garden
Missouri Botanical
\
\
Nac. Cs, C6rdoba 26(2): 298. 1923. Acacia far- entes: ca. 27^27'S, 58°46'W, 60 m, ^'alrede-
nesiana cavenia Min.
(L.) Willd. f. E. C. Clos, Bol. dores de la ciudad de Corrientes, antiguo cami-
Nac. 455. 1930. m
Agric. 28(4):
500
no Matadero, de 17
a la ruta, Feb.
&
m
Shrub 10 much 1989, 5. G. Tressens A. Radovancich 3539
or tree to high, but usually
\
smaller; single-stemmed or with 5-20 or more (holotype, K; isotype, CTES). Figure Id.
crowded young
densely basal stems; branchlets
A
omnibus microcarpa
varietatibus var. sola ex-
aliis
mottled gray, sparingly puberulous. Stipular spines
^epta legumine subgloboso a var. m/crocar/^a legu
differt,
cm
0.4-2.5
long, slender, white or gray, borne in mine maiore pedunculum crassum baud excedente differt.
pairs at every node. Leaves bipinnate, deciduous;
when
Fruits are nearly spheroid small (ca. 15
mm
5-15
peduncle long, usually with one nectary
X mm) when 25-
15 or slightly ovoid larger (ca.
5-7 cm
at midpoint of petiole; primary rachis long,
30 X 20 mm), often with a sharp acuminate
tip.
sparsely pubenJous; pinna pairs 4-6; rachiUae 0.7-
(814
numerous and leaves are
Leaflets are pairs)
cm 4-6
1.0 long; leaflets pairs per pinna, linear, ori^fcv^inic: I
m
.
, . . 30-45 x lU-lD
'arger than variety caven:
n ^ AC. J^^.:uLju*..„i
mm,
X ^i.u ....
11.0 0.5 glabrous.;. midrib and nerves
lateral mm
mm;
stipular spines mostly less than 5 long.
not readily distinct. Peduncles usually pubescent,
mm
3-15
and more humid part
long, relatively constant in length Distribution (Fig. 2). In the
within populations. Flower head spherical or of the Chaco and adjacent regions of Corrientes
ellip-
cm
soid, 0,5-1.0 long; one or usually several per and Entre Rios, Argentina, rarely in Santa Fe and
leaf axil, highly visible prior to the emergence of Cordoba provinces, Argentina, and in western Par-
leaves. Flowers sessile, yellow, highly fragrant. Ca- aguay and Uruguay. In damp fields where livestock
mm
mm
0,5-1.0 1-3
lyx long, glabrous. Corolla roam,
30-55
long, glabrous, approximately flowers per
ARGENTINA. COR-
Representative specimens.
44-58
head; stamens per
flower; filaments free
RIENTES: Empedrado, Estado La Yeia, ca. 27°53 S,
from
the base; anthers eglandulate; pods brownish
km NE Krapovickas
black or purplish brown, oblong-elongate or subglo- 10 of Colonia C. Pelligrini, Route 14,
29427 (CTES, entre Depto Islas del
rigs:
bose, straight or slightly falcate, frequently «' ^l. SI),
stipitate
^
km N Buenos
with
± Ibicuy, 14.5 of the interprovince line
and/or acuminate 5-7 x 10-13
basally cm,
8-
indehiscent, glabrous, usually not
1 g; striate. Tronson'7648 (M0).7anta RIo Nah Tuli Piague. 9
fe:
± ±
Mesocarp Laguna
spongy. Seeds brown, Uruguay.
olive Ragonese 2444
elliptic, 1937,
Jan. (SI).
5-7 X 4-6 mm, areole 4-5 x 3-4 mm; 9-39 Guayaca, Apr. 1906, Berro 4002 (MVFA).
±
per pod in 2 rows in young pods but scattered
sheet
herbarium
one
In 1947, Burkart labeled
in fully ripe pods.
„ (Ragonese 2444, "forma sphaerocarpa n.f."
SI)
Habit and, d,istrihuUon r^
(Fig. In disturbed
2). morphological
^ut never described formaUy. In
all
m it
sites northern Argentina, southeastern
Bolivia,
^ . shows affinity
^le^.ly
^^^jj^ ^j^^p^^ ^j^i,
and south-central Paraguay, occasionaUy an
inte- from carpologi-
^-^^^ ^^^ety caven. However, apart
m
grated element xerophytic Chaco woodlands and
distribution
geographical
^^j differences, limited
along former river courses in adjacent areas. Also under
growth rate
and comparatively rapid
^y^ 2)
vaUeys
in the central of Chile (36-27°S), parts of
(Aronson et
eontrofled environmental conditions
Uruguay and southern
EspeciaUy
Brazil. frequent va-
^^g^^ -^^ as a distinct
^,^ recognition
m
much-burned, overgrazed cow
pastures and -^^ u^^^ "
in -y.- ^,J-^^, ,^^_, .^ unusua ' "
abandoned
fields.
leaves, regardless of water availability.
Representative specimens. CORDOBA
km
E
of C6rdoba, ca. 3in9'S, 64°57'W, 280 m,
11
exv fial-
Acacia caven dehiscens rb>urki art*
1988. Aronson 7642 (CONC. K MO^ var.
Oct. RnirviA
Bolivia
Ap
km TYPE:
1984.
CHUQUISACA: 1 9 S of Cainargo, on road to Tarija ca deUa. Darwiniana 25: 76.
20°41'S, 65''15'W, 2,350 m, 26 Feb. 1989, Aronson 22 Sep. 1
Cordoba: Ascochinga,
gentina.
km
7799 (CONC, MO); 53 S
of Palos
Biancos, ca. 21°45'S
962
Nicora Figure f
E. G. (SI). 1
63''38'W, 850 m, 28
Feb. 989, Aronson 7827
(MO)'
1
km
E
Lo Cor-
Chile. 4 of Ovalle, ca. 33»02'S, 71°22'W, 220 of
Distribution. The mid-altitudinal hills
m, 18 Nov. 1988, Aronson 7692
MO);
(K, Cuesta de j Ko j c AArrggentina.
km N ^""^^ ^""^ ^^" ^t "^•' provmces, ww«e<s.tt«e.rrnn ^
Chacabuco, 45
of Santiago, ca. 70»42'W
32<>58'S.
o
Andean slopes
750 m, 23 Nov. 1988. Aronson 7696 ^^""^ ''^''^ "* ^^^ '<'^®'" pre-
(K, MO). Uruguay!
km i5al
E (1984),
2 of Gauleguaychu, tamarca and, according Cialdella
ca. 33°10'S, 58''22'W, 45 m. to •
frequen
SCO)
disturbed
Acacia caven sphaerocarpa human
var.
Burkart permanent
ex
proximity
to large,
Aronson,
var. nov. type.
Argentina.
Corri-
ulations
(Fig.
2).
Number
Volume 4 Aronson
79, 965
1992 Evolutionary Biology of Acacia caven
Representative specimens, ARGENTINA. CATAMARCA: purple pericarp, and the spongy, resinous mesocarp
m
CORDOBA
of the pod are unusual Acacia caven and are
San Roque, Apr. 1945, Hunziker 5984 Ascochinga,
(SI);
some
reminiscent of forms of A, farnesiana. Nev-
km
405 E
Giardelli (SI); 2 Villa de Totoral Rte. 9
(jet.
W
some
ertheless, Chilean, Bolivian, and one Uru-
and Rte. ca
17),
1988, Aronson 7644 (MO, SCO). SAN Bajo Grande, guayan specimen of variety caven show morpho-
LUIS:
Larca, A, Maldonado 102 (SI); Larca, Hunziker 2097 logically intermediate pods that suggest possible
km
Embalse La 2 E
(SI); Florida, of Trapiche, ca. 33°06'S,
gene macrocarpa.
flow with variety
66*^2' W, 930 m, 27 Mar. 1990, Aronson 79(97 (MERL,
SCO).
Acacia caven microearpa
Burkart
var. (Speg.)
Fruiting material generally required key
is to ex Darwiniana 25(1-4): 77. 1984.
Cialdella,
out this variety. Variety dehiscens displays well- Formosa.
TYPE: Argentina. Depto. Patino,
opened pods on
(see Fig. If) the tree, especially Fortin Soledad, A. Krapovickas 1283
(SI;
Embalse La
at Florida. Special caution required
Figure
is isotype, LIL). Ic.
with specimens from Lago San Roque, near Cor-
&
Wight microearpa
Vachellia farnesiana Arn.
(L.)
doba and from San f.
Luis, since natural hybridization
Speg., Bol. Acad. Nac. Ci. Cordoba 26: 301. 1923.
appears to occur between variety and A. atra-
this
20f,
f fig. 3.
tnentaria. Moreover, pods on some herbarium
Compared
to other varieties except variety
sheets appear dehiscent due crushing of all
to speci-
stenocarpa^ the pod here much smaUer (2-3 x
mens. is
much
1-2
cm) and the peduncle longer (usually
>
15 mm). The variety differs from variety steno-
macrocarpa much somewhat
carpa by shorter, spherical or
its
TYPE
subglobose than twice as long as broad.
fruit, less
19 Oct. 1948, Burkart 17577 Figure Mesocarp white and evanescent.
ril, {SI). is
le.
On
heavy
Distribution. soils in the seasonally
A
Paraguay and
varietatibus leguminibus multo maioribus inundated portions of central north-
aliis pilis
niinutis
glandulosis rubropurpureis omnimoobsitis et su- Argentina (Formosa and Chaco provinces).
eastern
turis prominentibus
differt.
proximity
been
Mesocarp
spongy and
whitish.
Chaco. Found under
Brazilian portion of the rel-
Distribution. This varietv has onlv been found
pristine
at high altitudes (2,000-3,200 m) on the eastern
^^.^^^ -^ ^^^-^^
^^^^
flank
^nd Representative specimens, ARGENTINA. CHACO: Pto.
Jujuy and
provinces, northwestern Argentina
MVM).
Dec, 1939, Poiraberi 77 (LP,
Barrangueras, 5
»n Chuquisaca, Cochabamba, and La Paz depart-
201698
10 1957, Burkart
FORMOSA: R. Juarez, Jan.
nients
of southern Bolivia* This variety generally
(CTESy
is
only found
in dry streambeds and alluvial plains salta: between Colonia Castelli y Rivadavia, Mar. 1967,
CROSSO DO
^vhere
groundwater
near the surface
relatively
is
ao Fazenda
Mun. Porto Murtinho, estrada Jerere,
SUL:
(Fig.
2).
Dec
GROSSO
Representative specimens. Argentina. CATAMARCa:
&
Furtado 7287 (NY). PARAGUAY. PRESI-
Andalgala, Jorgensen 960 (MVM); Depto. Capital, "La 1 980, Pires 1
near Puerto Ma.stinho, Dec.
Vuebrada," DENTE HAYES: Chicas,
Krapovickas 1748 between Paliiias
SALTA:
(SI).
Nueva,
Guachipas 65°37'W, 1957, 1937, Rojas 7694 (SI); Colonia, M«-nno. Mi-sion
y Alemania, 25°38'S, Jan.
ca.
Armas 200 BoyOERON: Ma-
59°40'W,
Job 1511 (LP); ToIoinb6n, 20 Oct. 1948, Real 12084 ca. 23''05'S, (SI).
&
Casas Molera
23 1980,
(MERL); Oct. F.
Estigarribia,
Quebrada de Conchas, 14 Aug. 1936, Ca- riscal
las
brera 3758 4431 (MO).
Mark, Sprg-
Santa Los Arcos,
(SI); Valle
gazini 5983
(SI). jujUY: Villa El Perchal, ca. 23''20'S,
65°30'W. 7677 (MO,
2,490 m. 7 Nov. 1988, Aronson
slenocarpa Burkart
Acacia caven (Speg.)
SCO). var.
BouviA. LA PAZ: Prov. Murillo, Mallasa. 16°32'S,
1984. Va-
Darwiniana 25: 78,
68»08'W. 3,200 m, 7 Aug. 1986. Solomon 15483 (LBP, ex CialdeUa,
&
MBM).
Wight
COCHabaMBA: Prov. Capinota, Santibanez to Capi- chellia farnesiana (L.) Arn. f. sleno-
iota
road, 66°17'W, 2,500 m, Feb. 1985,
ca. 17''45'S, 1 carpa Acad. Nac. Ciencias Cor-
Speg., Bol.
km
Pedrotti et 87 (LPB). CHUQUISACA: 27 S of Las
al.
doba 262: 301. 1923. 20. TYPE: Argentina,
fig.
Careras, ca. 21°16'S, 65n7'W, 3.080 m, 26 Feb. 1989,
Santa Ana, Burk-
Candelaria,
Aronson 7808 Misiones. Dept.
MO).
(K,
14734(51). Figure lb.
art
Although prepared
Burkart had drawing for
a
primarily
This variety ^•'rn in
Distribution. is
ti«s variety 1949 he did not write a
in (Fig. 2e),
and
Argentina
Formosa and provinces, in
Mi.sionef*
description. The the reddish
three sutural ridges,
966 Annals
of the
Garden
Missouri Botanical
adjacent Paraguay, especially within the area pe- nan (1957) for A. nilotica (L.) Willd., the type
riodically flooded by the Rio Pilcomayo. However, species of Acacia, In A. nilotica, nine subspecies
the few specimens from the Rio Uruguay collected were delim
1930
before suggest that its former distribution, in the shape, size, and pubescence of the pods.
and possibly that of variety microcarpa^ formerly Only fruiting specimens
Uruguay and
included parts of possibly the Rio subspecies, but geographic separation tends to con-
Parana region as well. Both of these areas are now firm the validity of the infraspecific system pro-
transformed by agriculture and urban sprawl. There posed. Brenan's (1957) elaborate system for A,
also exists one specimen from Tucuman, Argentina, nilotica has been maintained by Ross (1979, 1981).
km &
800 On
nearly to the west of current area of a higher taxonomic level, Britton Rose's
its
distribution. (1928) subdivision of Bentham's (1875) series
Gummiferae into 12 genera, partly on the basis of
Representative specimens, ARGENTINA. CHACO:
characters, has been rejected by most sub-
^^^^^
1987
Colonel Benitez, 7 July 1937, Schinini COR-
(SI).
RIENTES: 75 km N of Merced, Laguna Trinidad, Culan- sequent workers. Moreover, for generic and tribal-
trillar, Schinini et ah 11761 (MBM). FORMOSA: between level revisions throughout the Leguminosae, has
it
Formosa and Mosou de Fieri 7 Jan. 1945, Ragonese much
o, been argued that far too emphasis in the
&
2658
Cosso ISl); Clorinda, banks of the Rio Pilcomayo,
T loono P^st. was Di, aced, on ob, vi. ous Irrui-t^ treat^ures, ar.n^aA itVniaait
M^,
114A INov, 11 o9/4i/4i, 7. n/?oj*as 12298 /(TLiTLT)\, E^ N.T^^RE ^RI.O^S^: 5c k1 m r r j
i
i^ the future these should be replaced, or at least
S of Colon, ca. 32**20'S, 58°06'W, 20 m, Aronson 7882
(CONG. K, MO). Paraguay, Trinidad, Bahia Caballero, supplemented, by consideration of flowers, inflo-
&
Depto. Central, 11 Nov. 1950, Sparre Vervoorst 57 and attempts to pro-
rescences, seed, seedlings in
PREsmENTE
(K). HAYES: Mariscal Estigarribia, Dec,
1
1 j nnaattuurraall ciassmcation ssyvsstteemmss^r(Puoilihill et al.,
classification
&
1987, Schinini Palacios 25934 (CORD, CTES, ^^^,
G);
However, and particu-
^^^^'' the specific
25). at
Presidente Hayes, Dec. 1877, Royas 6979
Capitan
1 (SI);
m
Acacia caven, fniit
Lopez de Filippis, Oct. 1938, Rojas 8419 Bajo larly the subspecific level
(SI);
km
Chaco, 30 from Rio Aguaray Guazu, 4 Jan. 1980, and seed parameters seem more conservative with-
P
Arenas 1571
(CTES). caaguazu: Hassler 9085
(LIL).
-^ populations than leaf pubescence, size of leaflets,
Uruguay.
PAYSANDU; Santa Banado
Lucia, del Rio Uru-
^api- tuilum d,i- ameter, andi st-ipuli ar sp•me il^ern^rgrtflhi, all
/
guay, ca. 34-28'S, 56^22'W, Nov. 1919, Schroder
com-
8865 (MVM)- of which have been used to split up species
within
categories
plexes or to erect infraspecific
This variety distinguished by narrow
is its short, Mimosoide
mm)
pod 10 and
long peduncle mm).
15 most
many
( ( YoT A. caven, and related species,
Fresh pods are usually brown and
light often bear
parameters
pj^^^^^ j^^g^^^ ^^j spinescent stipule
m
three strongly delineated suture
ridges as variety even
and
^j^^^ considerable infraspecific,
^^^j^
m
macrocarpa much
(and
material of A. fame- taxc
gain
they
within-population, variation that
x
siana). Leaves are big (80 50 mm),
with pinnae
greenh
mm
up 30 As
to long. in variety stenocarpa,
trees when they are
^^ common-garden
experiments, or
tend he weak and spmdly compared
to
to other Aronson,
^^own
to be geneticaUy fixed traits (cf.
Some
varieties. introgression apparently
takes place
1992)
between variety stenocarpa and
variety micro- a nd
legumes
By mature
contrast, the shape of
m
&o r^
carpa , . Sc^parre Much
Vervoorst
(e.g., 57, K). developed
^^e number and average weight of fully
more
coUect.on needed PUcomayo
is in the Rio withm
^^^^^ conservative
be
,^
^^
m
basm and
western Paraguay.
contrasted
though
^^^^
populations of A. caver.,
The
few, rather specimens
old of variety
steno- sys-
^ and can play a role in
^^^eties, thus
car;.a coUected from Uruguay &
bear witness to the EUas
Lewis
7 ^^^^.-^ ^^j
^ ^^^^^ „f studies.
1^
wider distribution of variety
this in earlier times. ex-
provide
(iggi) ^^^^^^^ characters
^^^^ ^^^-^
They
also raise the possibUity that A. caven
var. Mimoseae.
^^jj^^^ ^^^^^^^ ^^hin the
-^
-i
W
Stenocarpa
populations
that existed
in eastern Ar- -
gentina and western Uruguay were
seems
eradicated, only
aspectrofTIriatL"withinl7ada'cat;en, it
be more
replaced
to recently by A. caven more
var.
to
subject
^^at and seed characters are
fruit
caven.
vege
than other
intense pressure
selective
d'^pers^
seed
ones. At least four different types of
Discussion
o
may varieties
strategies have evolved in the
An
infraspecific revision of a widespread and
described
polymorphic Acacia stenofor
based
in large part on carpa, microcarpa, and variety
differ- variety
disper
pod and pa may seed
seed weight have undergone selection for
1
ibe
inund
gularly
Number 4
Aronson
Volume 967
79,
1992 Evolutionary Biology of Acacia caven
(
i
occupy. By contrast, high in the Andes, the un- Arechavaleta, 90 Flora Uruguayana, 436-
J. 1 1 III:
.
Museo
437. Anales Nac. Montevideo, Montevideo.
macrocarpa
large pods of variety could
usually
Aronson, 1992, Description and distribution oi Aca-
J.
have been selected for long distance dispersal by
&
macracantha Humb.
cia Bonpl. (Leguminosae,
mega vertebrates. Finally, variety dehiscens has
Mimosoideae) in northern Chile. Gayana, Bot. 48:
possibly undergone selection for seed dispersal by 81-87.
&
C. OvALLE AvENDANO. 1991. Early
rodents, ants, or avian granivores (Aronson, in J.
growth and 44
rate nitrogen fixation potential in
prep.).
legume grown an and
species in acid a neutral soil
on number
Studies variation in polyad grain
from Managem. 225*
central Chile. Forest Ecol. 47:
among Acacia
varieties of caven, those of
e.g., 244.
&
Kenrick Knox (1982) for A. retinodes, reveal Arroyo, M. T. K. 1981. Breeding systems and polH-
723-769
between number and nation biology in Legumlnosae. Pp. in R.
a correlation pollen grain
&
M. H. Raven Advances
Polhill P. (editors), in
maximum number no
seed per pod. Yet, there
is
Legume Volume
Systematics, Royal Botanic Car-
1
.
obvious correlation between seed number and av-
Kew.
dens,
erage seed size in A. caven, such as predicted by Atchison, E. 1948. Studies in the Leguminosae.
IF.
Harper et (1970) and others, A related question Cytogeography of Acacia (Tourn.) L. Amer. J. Bot,
al.
651-656.
35:
number
the possible correlation between ovule
IS
&
Barneby, Zanoni. 1989. Las Acacias
R, C. T. A.
in Acacia caven flowers and the number of seeds
{Acacia, Mimosaceae) de Espanola: dos nuevas,
la
&
borne per pod (Lewis Elias, 1981; Ph. Guinet, una mejor descrita, y una clave para todas, las in*
4-27.
Moscosoa
pers. comm.). digenas asf conio las cultivadas. 5:
& Bentham. G. 1842. Notes on Mimosaceae, with a syn-
In Hispafiola, Barneby Zanoni (1989) used
Kew
Hooker's Bot. Card. Misc,
opsis of species.
J.
pod dehiscence one
as of the diagnostic features
318-392.
1:
macracan-
for distinguishing A. tortuosa from A.
Mimoseae,
1875. Revision of the suborder
.
At London 335-667.
tha. a higher taxonomic the occasional Trans. Linn. Soc. 30:
level,
Bernardi, 1984. Contribucion a Dendrologia Pa-
occurrence of pod indehiscence considered a L. la
is
L 1-341.
raeuaya. Boissiera 35:
r
A\^r.^^ *• ^ r 4 - '
diagnostic feature of su1bgenus Acacia, since no-
\gQ^ ^cacm
rmg
^^^^^^^ Genetic races a species,
in
where
in subgenus Aculeiferum or Phyllodinae 181-185.
karroo, S. African Sci. 85;
J.
can
be found 1981), Within subgenus Brenan, M. 1957. Notes on Mimosoideae:
it (Elias, P. III.
J.
Kew
75
96.
Acacia, legume Bull. 12:
as well as various other taxa,
in
&
BrittoN, N. L. N. Rose. 1928. Mimosaceae.
J.
It is observed that once pod indehiscence has arisen,
23 1-194.
North Arner. (Part
Fl. 2):
remains comm.).
It firmly fixed (B. Verdcourt, pers.
Bucher, E. 1987. Herbivory in arid and semi-arid
Ine
question arises whether A. caven variety de- regions of Argentina. Rev. Chilena Hist. Nat. 60:
265-273.
hiscens
most
the primitive of the six varieties
is
igummosas Argentinas
BURKART, A. 1952.
or a case of evolutionary "reversion." This latter
Acme
2nd Agency,
Cultivadas, edition.
vestres
y
hypothesis seems more view of the fact
likely in
Buenos
Aires.
that seed arrangement in variety dehiscens is poly- 1967. Flora de Provincia de Buenos Aires,
la
INTA. Buenos
seriate, Volume 3A, Leguminosae. Aires.
just as in the other varieties. l\\
all
&
Willink. 1973. Biogeograffa de
The Cabrera, A. L. A.
significance of polyseriate seeds in a given
Americas,
Organization of the
America Latina.
pod
should be noted context. In contrast to
in this
Washington, D.C.
uniseriate seeded pods, pod length of ^. caven (and
M. 970. Cranos de polen de Leguriuno-
CaCCAVARI, A.
1
of no Tribu
its close relative A. farnesiana) bears cor-
144-174.
relation with seed number. Thus pod volume and cicue. Darwiniana 16;
11...
doce
Tastronovo 1945. Estudio carif>logico
m -j^u a. dt;
fdtrrxy, ,weighi_t. . , . pod1 width
are required, addition to
^^^f„^^^^^^^^^ de Legurnlnosas. DDaarrwwiinniiaannaa 77:: 3388 5577..
especies
and
length, order adequately describe vari-
in to genero Arnria (Legumi-
M. 1984. EI
CiALDELLA, A.
ation in reproductive output and therefore in ev- Darwiniana 25: 59 111.
en Argentina.
nosae)
la
Segunda contribucion conuci-
olutionarily As Vassal 1930. al
significant phyletic groups. Clos, E. C.
en
arbustos cultivados
miento de arboles y la
los
(1972) put 'The genus Acacia endowed with
it: is
(Leguininobae-MimosoiHeae). Bol. Minist.
Argentina
great has not
evolutionary dynamism; apparently
442468.
it
Nacion 28:
de
Agric. la
yet exhausted potentialities.'' Systematic 1981. Mimosoideae. Pp 143 152 in R.
all its Elias, T. S.
& Advances
ti'eatments within the genus need to take this dy- M. Polhill P. H. Raven (editors), in
Vdume
Royal Botanic Gar-
Legume Systematics,
'^amism I.
into account.
Kew.
dens,
&
PopuLtjon
AbHTON. 1978.
H.
D.
Farrell, T. p.
Literature Variation
Cited on Acacia melanoxylon R. Br.
studies 1.
Austral. Bot.
XXXVIL seed and vf (jetative characteristics. J.
ARata, .4ca- in
P. N. 1891. Ciencias Naturales
365 380.
26:
cia cavenia Hook Am. Anales Dept. Nac. (\r
y
&
AvnK> Securu. 1989. Land.
A.
FUENTF5. E. R-, R.
43^44.
Hygiene
(Argentina)
1(2):
"^i
CO
9 Annals of the
• I*
Garden
Missouri Botanical
scape change under indirect effects of human use: Pedley, L, 1986. Derivation and dispersal of Acacia
the savanna of Central Chile. Landscape Ecol. 2: (Leguminosae) with particular reference to Australia,
73-80. and the recognition of Senegalia and Racosperma,
&
219-254.
GuiNET, Ph. Vassal. 1978. Hypotheses on the Bot. Linn. Soc. 92:
J. J.
&
differentiation of the major groups the genus Aca- Peralta, L, G. RODRIGUEZ M. T. Kalin Arroyo.
in
J.
Kew
cia (Leguminosae), Bull. 32: 509-527. Breeding systems and aspects of pollination of Acacia
&
Gutierrez, Armesto. 98 El rol del ganado caven (Mol.) Mol. (Leguminosae: Mimosoideae) in
J. J. J. 1 1
.
en la dispersion de las semillas de Acacia caven the mediterranean-type climate zone of central Chile.
&
3-8.
(Leguminosae). Ci. Invest. Agric, (Chile) 10: Bot. Jahrb. Syst. (in press)
& &
Harper, L., P. H, Lovell K. G. Moore. 1970. PoLHiLL. R. M., P. H. Raven C. H. Stirton 1981.
J.
The shapes and sizes of seeds. Annual Rev. Ecol. Evolution and Systematics of the Leguminosae. Pp.
&
327-355. 1-26 M. H. Raven
Syst. 1: in R. Polhill P. (editors),
Hassler, E. 1909. Contribuciones a la Flora del Chaco Advances in Legumes Systematics, Volume I. Royal
Kew,
Argentino-Paraguayo. Parte. Florula pilcomayensis. Botanic Gardens,
I
&
Trab. Mus. Farmacol. No. 21. Buenos Aires, Ar- Ramella, L. R. SPICHIGER. 1989. Interpretacion
gentina. preliminar del medio fisico y de la vegetacion del
Heusser, C. 1971. Pollen and spores of Chile. Univ. Chaco Boreal. Contribucion al estudio de la flora y
J.
639-
of Arizona Press, Tucson. de vegetacion del Chaco Candollea 44:
la I.
&
W. W.
Hooker, Arnott. 1830-1841. The
G. A. 680.
J.
&
La
Botany of Captain Beechey's Voyage. Henry G. Bohn, Ragonese, a. E. C. Castiglioni. 1970. ve-
J.
London. getacion del Parque Chaqueiio. Bol. Soc. Argent. Bot.
&
HuECK, K. P. Seibert. 1972. Vegetationskarte von 133-160.
11:
Sudamerika. Gustave Fischer, Stuttgart. Rico Arce, L. 1984. The genus Acacia in Mexico.
&
50-59.
Kenrick, B. Knox. 1982. Function of the polyad Group Study Mimosoideae 12:
J. Bull. Int.
.
&
1984.
in reproduction of Acacia, Ann. Bot. (London) 50: Rodriguez, M. Quezada O. Matthei.
R.,
721-727. Concepcion,
Arborea Universidad de
Flora de Chile.
KUNTZE, 0. 1898. Revisio Generum Plantarum 3: 47. Concepcion.
&
Lewis, G. P. T. S. Elias. 1981. Mimoseae, Pp. Ross, H. 1979. A conspectus of the African Acacia
J.
&
155-168
M. Raven South Africa
in R. Polhill P. H. (editors). species. Memoirs of the Bot. Survey of
Advances Legume Volume
in Systematics, Royal 44: 1-155.
I.
Acacia
Botanic Gardens, Kew. An African
1981. analysis of the
LOMBARDO, A. 1982. Flora Montevidensis, Volume origins and rela-
species: their distribution, possible
I:
126-127.
Intendencia Municipal de Montevideo, 89-413.
tionships. Bothalia 13:
Acad.
Uruguay.
Spegazzini, C. 1923. Acacieas Argentinas. Bol.
Molina, G. 1782. Saggio Sulla Storia Naturale del Nac. Cordoba 26: 161-334.
I. Ci.
John
Chili, 1st edition Bologna. Tatsuoka, M. M. 1971. Multivariate Analysis.
&
1810. Saggio SuUa Storia Naturale Wiley New York.
del Chili, Sons,
.
Argen-
2nd Bosques
ed. Bologna. Tortorelli, L. a. 1956. Maderas y
OVALLE, 1986. Etude du Systeme
C. Ecologique Sylvo- Acme, Buenos Aires.
tinos. Editorial
ontogemques
Pastoral a Acacia caven (Mol.) Hook, et Arn.: ap- Vassal, 1972. Apport des recherches
J.
taxono-
plications a la gestion des ressources renouvables et seminologiques a Petude morphologique,
boc.
dans Taire climatique mediterraneenne du Chili. Ph.D. mique phylogenetique du genre Acacia, Bull.
et
11
Universite de Sciences Techniques du Languedoc,
et Hist. Nat. Toulouse 108: 15.
mR.M.l^ol-
Montpellier, France. 169-171
1981. Acacieae. Pp.
.
& Legume
Aronson. Del Pozo &
a. AvendaNo. Raven Advances in
, J. J. hiU P. H. (editors).
Key-
The
1990. Gardens,
Espinal: agroforestry systems of the Volume Royal Botanic
Systematics,
I.
Prodromus
&
mediterranean-type climate region W. 1834.
of Chile: state of Wight, R. G. A. Arnott.
Lon-
and Volume
the art prospects for improvement. Agroforest. 1.
Florae Peninsulae Indiae Orientalis,
Systems 213-239.
10:
don.
Parsons, D. 1976.
Vegetation
structure med-
J. in the
iterranean scrub communities
of California and
Chile
EcoL 435-447.
64:
J.
DOCUMENTING
SCIENTIFIC
J
Peter
Goldblatt,^
I
THE NEED and
FOR Peter Hoch,^
C.
DATA:
M, McCook^
Lucile
VOUCHER
SPECIMENS
many
we
Like botanists have been following the sources; few gardens have the personnel or the
developments
in molecular systematics with inter- library and herbarium resources to verify the names
and One many
est excitement. aspect of most current of all the plants in their collection. Yet mo-
work
in nucleic acid sequencing and restriction lecular studies cite botanical garden accessions as
enzyme
analysis, C-value estimation, and natural they were sources of accurate
identifications,
if
products chemistry is, however, a growing source Some authors assume that any samples sent to
of concern. This the question of voucher them are adequately vouchered and documented,
infor-
is
mation for taxa studied. Our experience demands caution in this area. Mis-
A
survey
of papers on nucleic acid sequence identifications and mixing of labels occur with suf-
data recently published has revealed a lack of con- frequency that no systematist should be
ficient
sistency in the citation of voucher data (Cantino, with procuring a sample from a botanical
satisfied
1992; Gianassi et 1992; Hufford, 1992; 01m- garden, or any other source, without being certain
al.,
&
stead et al,, 1992; Downie Palmer, 1992; Don- that a proper voucher exists that corresponds to
&
oghue
et 1992; Olmstead Palmer, 1992), the plant being sampled and that the plant prop-
al.,
is
Some
papers neither cite nor allude to any voucher erly identified. Original locality data should also be
data. Others
refer to previous papers or individuals established possible.
if
as documentation or sources of experimental ma- Absence of a voucher makes impossible to
it
Yet To
tenal. voucher data are not necessarily in- reassess identity. avoid the potential for future
eluded we
in the referenced papers. problems suggest that leading scientific journals
Vouchers are central to any serious questioning insist on the citation of an existing herbarium
or reexamination of data and conclusions. An un- voucher specimen and an indication of its location.
expected result may be due to convergence or to Absence of a voucher for critical taxa should be
somehow
past misinterpretation of morphology and other explained so that readers are assured of
A
characters, but may be caused by misiden- the identity of study materials, statement that
also
it
may
tification of the plant examined. The most prudent an author be consuhed personally for voucher
Are we
step to take before accepting an unexpected ex- data not satisfactory. This rarely works.
is
perimental result to question the possibility of simply going to have to discard dozens or possibly
is
huge
misidentification of experimental material or mis- hundreds of sequences, obtained at cost in
m
labeling The because of inadequate voucher
the laboratory. next step to du- research dollars,
is
seems
plicate the result. Because of the absence of vouch- documentation? This likely,
We
ermaterial, chromosome strongly urge that molecular systematists
thousands of early
literally
counts, published over the past 50 years, have had take the time and make the effort, sometimes not
proper documentation of
discounted, and we suspect that fully inconsiderable, to assure
We
urge
counts experimental subjects. also that jour-
their
on the inclusion of direct citation
questio nal editors insist
cannot be of voucher material in all papers presenting original
Annals Missouri
the
Most As an example, the of
systematists have had the experience of data.
Garden the following
seeing know Botanical instituting
species they well misidentified in bo- is
Papers presenting original data
voucher will
tanical garden Not seeds policy.
infrequently
collections.
not be accepted for consideration unless they in-
botani
mm-
At
be species other than the one named on the pack- elude citation of voucher information. the
herbarmm
Imum means of an extant
age. Just as commercial nurseries, botanical this citation
in
and number) and an
gardens receive seed and plants from commercial specimen (minimally collector
Box Louis
Garden, P.O. 299, St.
B. A. KrukofF Curator of African Botany, Missouri Botanical
'
U.S.A.
63166, U.S.A.
Box Missouri
= Missouri Botanical Garden, P.O. 299, St. Louis,
63K
Garden, P.O. Box 299, St. Louis, Mi880ur»
'Horticultural Taxonomist, Missouri Botanical
96< 970. 1992.
Gahik
Ann. Missouri Box. 79: