Table Of ContentNumber
Volume 79, 4 & Howard
Goldblatt 903
1992 Alophia
resemblance goes no and
but the further, C. kin- in the genus. reportedly has pendent
It lilac -violet
probably does not have a
tonii particularly close flowers with a yellow center, connate filaments,
Alophia
relationship to or Ainea, pandurate anthers, and deeply forked branch-
style
mm
Little attention has been paid to the broad con- es, the arms of which are 14-15 long. In the
nective condition in Iridaceae, and although seems article in which he described A, intermedia, Ra-
it
intuitively to be apomorphic, the character venna (1964) included A. Alophia^
is dis- trijiora in a
two now
tributed in of three genera of Maricaeae and species regarded as conspecific with Tigridia
several species of CjKpeZ/a, possibly the basal genus meleagris (Molseed, 1970). Examination of the
of Tigridieae. Thus, a broad connective probably type of A. intermedia^ not available to us, nec-
is is
synapomorphous
for the clade that includes Mari- essary before status and generic position can be
its
ceae and
Tigridieae (see Goldblatt, 1990) but pie- assessed.
siomophic for Ainea and Alophia, although the Alophia veracruzana seems most closely re-
pandurate shape in Alophia seems unique. There lated to A. drummondii, which resembles ui the
it
no longer seems any reason to place Alophia in size and general form of the flower (Figs. lA, 2A),
Tigridiinae. The shared polyploid base number Alophia drummondii normally a taller plant,
is is
almost
certainly convergent, and /i/op/ija does not and the flowers are violet to red-purple with the
i
have marked cream
either the united filaments or bisulcate pollen well-diflFerentiated tepal claws to
grains that are apomorphic Beyond yellow and densely speckled throughout with dark
for Tigridiinae.
a possible close relationship with Ainea, the red-purple. The major diff"erence be ween
affin-
ities of Alophia seem more uncertain than ever. species in the shape of the inner tepals. In A,
lies
drammondiij the inner tepals are narrower than
the outer tepals, and the limbs are sharply bent at
Distribution and
Review
Species
base 2 The edges of the lower part of
the (Fig. A).
Alophia occurs across a wide portion of tem- the inner tepal limbs are closely pinched together
perate and tropical America. The most known below where they are whitish to yeUow and arched
well
and
widespread species, A. drummondii (Graham) over the median lower part of the limb. This fold
R. Foster (synonyms Eustylis purpurea (Herbert) covers a narrow median zone of stalked glandular,
&
Engelm. The
part of
Gray, Ravenna), presumably nectariferous tissue. distal
E, />artc/a/a (Herbert)
extends from limb dark red-purple and as-
the southern United States (Arkansas, the outer tepal is
Louisiana, Oklahoma, and Texas) southern Mex- cending. Alophia veracruzana (Fig. lA) lacks a
to
«co limb
(including Vera
Cruz, Chiapas, and Tabasco)
&
(Goldblatt, 1974; Henrich The mner
1992).
Goldblatt,
species, or one closely related to also occurs in undulate margins, have no nectariferous band.
it,
Guyana
Maas
aL 1774, seems
(cf. et U). This to
have
SystematICS
slightly smaller flowers, filaments united in
-
the
lower
and
half, style branches that are shorter -
veracruzana
^nd do
not A
arch over the stamens. second species,
Mexico.
TYPE:
Ae
smaller-flowered
A, (Loesener) Gold-
silvestris
Missouri
ted
Watt
(synonyms
&
Nemastylis
hequaertii Standley,
IJouard
Garden, June 990, Goldhlatt
ical 1
rotacea
Ravenna), Amer-
^. Central
restricted to MEXU;
is K, MO),
9070
i.^otypcs,
(holotype,
ica, where
extends from southern Mexico Costa
to
it L
Figure
A
'^ica.
medusa
third species, A. (Baker) Goldblatt
(basionym
Chlamydostylis medusa tiini
Baker), ap-
o mm
c
_' , . ,- . 5-0 i
4-5, laminis phcatis, latis,
FPa^rreenmiilyv m g-. •' cis atrobrunneis, foliis
pr.or^o^r-.li,y known, .
i occurs Goias, centrali '__
^f/;;.°"^4"-"5^ spatha iin„t,e^rri\oLr^e 3-x2o 3-x8iK mm,i,u. llnonncg.ap,
florum,
rhipidiis
orazil.
It is particularly distinctive in large flow-
its interioribus marulis albis vel luteis
tepalis
floribus iilacinis
ers and
cm
spathes 2.5-3
(tepals long, inner spathes obscure unguiculatis. exlerioribua
obovatis
tepalis
notatis,
mm
16-19
mm
*^^)j a conspicuous band of glandular tissue in 17-20 longis, ca, 10 latis, interioribus
mm
mm
staminibus atropnrpurffb,
10
^he median of the inner tepal limbs, and terete longis, ca. latis,
mm
angubtc
contiguis anlheris
2 longis infra
eaves. To filamentis ca.
now
these the new A. veracruzana can mm
^ 1.1
loiigts*
added.
This has a restricted distribution in north- mm
2 supra.
ca.
eastern
Mexico, known from Vera
currently only
25-30 cm
Bulbs nar
geophyte high.
Perennial
Leaves 4-5,
One dark brown.
Ravenna rowly ovoid, the scales
species, Alophia intermedia
±
shrath-
entirely
(1964) except the uf>permosl
(known laminate
to us only from the description)
from abc as
may and elamtnate, blad<^s pllr.ae, iiK^-ily t
Sinaloa western Mexico, be misplaced ing
in
^
904 Annals of the
Garden
Missouri Botanical
8
long to slightly longer than the stems, the lower begin to separate at about A.M., and by 9
A.M.
2-3
inserted below the ground, the upper two they have spread sufficiently to allow access
to the
inserted well above ground, the lower of these near style and stamens; by this time the anthers have
dline and short- dehisced, revealing the bright yellow pollen. At
mm
5-8
est, narrowly lanceolate (to almost linear), about A.M. the inner tepals reach their open
1 full
Stem The
at the widest, apices attenuate. terete, gen- position. tepals begin to collapse at about 2:30
erally with one branch produced in the upper third, P.M., and by 3:30 P.M. are closed together. Like
drummondii^
A. A. veracruzanaisstM-
relative
its
Infl>
4-5
rhipidia, single and terminal, usually -flowered; compatible. Self-fertilization often occurs In one or
spathes green, attenuate, the apices generally dry two flowers of a rhipidium with resultant capsule
mm
32-38
mner and seed production. As the stigmas are not
in
secund, contact with the anthers while the flowers are open,
inner spathe, pollination presumably occurs after the flowers col-
curved outward at the apex; perianth actinomor- lapse, when the parts of the flowers are pushed
phic, pale to dark lilac-purple (rarely whitish), paler together by the inward-falling tepals. Outcrossing
toward the center and lightly flecked toward the presumably the normal mode of reproduction for
is
mner with the species in view of the colorful and fragrant
lis
undulate margins and each The presumably serves
with a pair of large perianth. occasional selfing
white to yellow marks near the base of the limbs, to ensure seed production in the absence of insect-
±
can
sweet ob- mediated Plants of ^. t;eracr«za/ia
pollination.
,
ovate, claws weakly and growing
differentiated spreading be raised from seed to flowering in a single
mm
17-20
outward, outer tepals long, the limbs season. Seeds sown in the greenhouse
in late fall
mm
spreading, ca. 10 wide, inner tepals 16-19 in St. Louis grew to flowering size in about six
limb months.
right angles to the claws and with the lower
lateral
areas and median
raised the MORPHOLOGY
portion recessed. Sta- Karyology AND POLLEN
mm
mens
dark purple, filaments 2
ca. long, broader
veracruzana
Chromosome number Alophia
in
and
contiguous below, narrow
in the upper
half;
pair
one
chromosomes,
Four
28. long
anthers coherent
at their bases, with a broad nar- ^^
conlras
submetacentric and the other acrocentric,
rowly pandurate connective narrowest
in the mid-
much« mailer
^^^
mm ^^^'^^^ remaining s
mm the 12
die %, weakly ascending, 4 LI
ca. long,
the e -
mm The karyotype accords with that of
pairs.
wide at the base, nearly 2
at the apex,
spo-
IV
-.
members (Goldblatt,
raploid of Tigridiinae
rangia marginal and
nearly
latrorse; pollen yellow,
&
Benton Heywood,
1984).
Ovary
broadly sulcate. narrowly 4
obovoid,
ca.
mm
^^^
long, exserted from P°"^" ^^""^ ^ '""^'^ 7
the spathes; style 5.5 S'"^"'"
ca. j^r
IHblatl
mm expanded^oo
when
^^"^ as the grains fully
^
long, dividing opposite the middle
of the an-
* those^^
^^ Thomas, and resemble
in prep.)
thers into three thickened branches,
the branches
^^
mm drummondii '^^^^!'^'"'
(Goldblatt, 1 974).
erect, ca. 1 .5 long, each dividing into two long
°„,ity
of the
mm correspond with those
J
ascending-recurved arms, '^^^"^ ^^"^^
these 3
ca. lone
^^
^' (7I^"d'«'
- of Tigridieae excluding Tigridiinae
tubular, stigmatic
apically, arching
over
the an-
Cobana—Uo\see
^
and
teria, Sessilanthera,
Capsules
thers. obovoid
-oblong, trigonous and
grams-
have bisulcate
1970; Ravenna, which
1974),
truncate
obovoid-pyrlform,
narrowest
mi
at the
cropylar end, the chalazal end concave, Literature Cited
Ughtly
brown
CaLDERON
DE RZEDOWSKI, G. 1987-
'^'S'f^^^^ie
&
28 Howard
(Goldblatt
"^'^
9070).
una nueva de iridaceas
nezii, especie „
^
Mexico
^^[^
Hidalgo (Mexico). BoL Soc. Bot.
'^^
maa
bulbous
Goldblatt. 1974. Revision of the
P.
Flower Phenology
and
Compatibility 373-385.
of North America. Brittonia 27:
^^
Chromosome cytology "^^^
1982.
As
.
in Tigridieae, ^^^^y^.
all the fugacious
flowers
last Iridace
neotropical
suprageneric systematics of
befo
Bot 186-198.
7:
t^^^.
r
.
oi
fuU -. 1990. Phylogeny and classificatmn
during
Ann. Missouri Bot
Herbert
unfold, but until 7 A.M, the inner
tepals remain
n^-
on
(Iridaceae-Tigridieae): notes t\lh^ixisi
closed
c/vli^
Salp^^b
nus and synonymy of
reduction
to "
Number 4 & Howard 905
Volume Goldblatt
79,
Alophia
1
Cardiostigma, Ilysa, and Catila. Ann. Missouri Bot, Ravenna, P. 1964. Notas sobre Iridaceae. Revista Inst.
504-511. Mun. 51-60.
Card. 78: Bot. 2:
&
Henrich, E. P. Goldblatt. 1992. Iridaceae. In: 974. Cobana, a new genus of Central Amer-
1
J. .
Flora Mesoamericana 1. (In press.) ican Iridaceae. Bot. Not. 127: 104 108.
&
Kenton, A. C. A. Heywood. 1984. Cytological 1979. Ainea, a new genus of Iridaceae from
.
467-469.
studies in South American Iridaceae. PL Syst. Evol. Mexico. Bot. Not. 132:
&
146: 87-104. RuDALL, P. A. Wheeler. 1988. Pollen morphology
Kew
693-701.
MoLSEED, E. 1968. Fosteria, a new genus of Mexican in Tigridieae. Bull. 43:
Iridaceae. Brittonia 20: 232-234. Shinners, L. 1964. Tigridia purpurea (Herbert) Shin-
1970, The genus Tigridia (Iridaceae) in Mex- ners, comb. nov. (Iridaceae). Sida 1: 295.
.
t
and
ico Central America. Univ. California Publ. Bot.
54.
!
I
-I
I
^
CHROMOSOME NUMBER
IN Snow^ and
Neil Peter Goldblatt^
TICODENDRON
(FAG ALES,
TICODENDRACEAE)
Abstract
number of Ticodendron incoe:nitum reoorted for the time 26, and the phylogenetic
is first
number
ignificance of this within
among Fagus, Castanea,
After exhaustive collaboration several Castanopsis^ Cyclobalanopsis,
&
Qu
(Hammel
authors Burger, 1991) the monotypic i
&
family Ticodendraceae was described (Gomez- Hans, 1969; Raven, 1975; Morawetz Samuel,
&
Gomez x 21 22
Laurito 1991) accommodate Trigonobalanus has or
P., to the 1989), while
Mesoamerican tree Ticodendron incognitum Go- (Hou, 1971; Soepadmo, 1972). Colombobalanus
&
mez-Laurito Gomez P. (1989). Evidence from and Formanodendron, segregated from Trigono-
&
wood anatomy remain un-
(Carlquist, 1991), leaf anatomy balanus by Nixon Crepet (1989),
&
may
which
(Hickey Taylor, 1991), anatomy and mor- counted Nothofagus,
floral the family.
in
=
x 13
phology (Tobe, 1991), pollen (Feuer, 1991), and merit own family (Nixon, 1982), has
its
&
Within
sieve element characters (Behnke, 1991) strongly (Armstrc ig Wylie, 1965; Ono, 1977).
—
We n.^ J
/" DOcs/trr-iy/onpm/s.us form
suggests its inclusion in Fagales. report for the Betulaceae Carpin• us, Ostrya, and
= o
x
first time the chromosome number of Ticodendron a evidently paleodiploid group with
distinct,
=
&
incognitum 2n 26 and Betula,Alnus
as discuss the possible (Raven, 1975; Love Love, 1982);
1984,
number (Kovanda,
phylogenetic significance of this within Fa- (Raven, 1975), and Dusckechia
==
may have
gales. Pashuk, 1987) have x Corylus
1 4;
= =
14 equire
(records of 2
2fi
ac
=
genus as
Materials and Methods 11 the
but there are records of in
re
& monogeneric
The
L5ve 1982).
well Liive,
(e.g.,
Root were
tips collected at approximately 9:30
(Morawetz
20
x
B^l^^oP^^^ae appear to have
from specimen
A.M. a single growing in the green-
21
in
n ,
count of
Samuel, 989) (the sn^gle
1 J^
houses at the Missouri Botanical Garden (Voucher: f
may erroneous^
be
W„™™./^ R.w„ 1979)
Hammel & 17R^A MH^ Af.J fi.au5^raZma(Goldblatt,
Rivera 17834, MO).
After
pretreat-
Our
Fagales.
Qearly, x 3 infrequent in
1 is
ment
with aqueous -bromonaphthalene
4
1 for hours, x
=
suggests
count ot 2n 26 from a single plant
the were
tips fixed in 3:1 ethanol (95%)-gIacial
=
order
the
3 Ticodendraceae, so within
for /^
1
10%
acetic acid. After hydrolysis HCl
in for 7 no
with
codendron number only
shares
this
minutes at 70*C, root tips were squashed FLP
in
fagus, a genus from which distinct.
is
it
orcein (Jackson, 1973).
basic
x
Although Raven proposed
(1975)
^^
scenario for
for Fagales, a plausible
Discussion
^^^^^-''''^'J^,
x
on
based
evolution can be suggested
-
The
=
din 26 and small, suming x R- nremimed base numbe' '"^
& ^^n-
uniform chromosomes
relatively
of Ticodendron weU (Goldblatt
related Hamamelidales as
corrobo &
^ n.cii dress, 1977; Morawetz Samuel, l^^^^^*"^*
.v.
J
o^^
indicate the distinctness of taxon from reduction
this other members dysploid
early of Fagales,
families of Fagaceae. = and sut*^*!"^^
4
7 followed by a doubling to 1
,
The
Fagales (sensu Cronqust, 1981) have base seems us^^^
reduction and II »« ^^
to 13, 12,
=
numbers x e
and 4 withm
8, 1 1 , 1 2, 1 3, 1 (Raven, 1 975). reasonable explanation for change
=
13,
Fagaceae x
In 12
predominates, numbe
occurring The
in ibers of the order. base
De{u.rt,nent of Biology. Washington
• University,
St. Louis. Missouri 63130. U.S.A.
63166
B. A. Krukoff Curator of African Botany. Missour.
Missouri
Botanical Garden, P.O. Box 299, Louis.
St.
Missouri Bot
907.
1992.
_^^j~ -L-^i.n^#^i~\^ > J-
Number Snow
4 &
Volume 79, Goldblatt 907
Chromosome Number
1992 Ticodendron
in
&
common W.
to Ticodendron and Nothofagus, was most Hickey, L. D. Taylor. 1991. The leaf ar-
J.
=
chitecture of Ticodendron and the application of
achieved independently from x
likely 14, a hy-
foliar characters in discerning its relationships. Ann.
supported by wood
pothesis significant differences in
105-130.
Missouri Bot. Card. 78:
anatomy
(Carlquist, 1991), reproductive mor-
Hou, D. 1971. Chromosome numbers of 7a^(>«o6a/a-
phology (Tobe, 1991), leaf architecture (Hickey nas Forman (Fagaceae). Acta Bot. Neerl.
t;er/fci7/a/a
We
& 543-558.
Taylor, 1991) and other features. note, 20;
Chromosomal Haplo-
J^^^^^^' ^- evolution in
comment
however, the by Tobe (1991: 141) that 1^^.^:
^ pappus Evo-
gracilis: a centric transposition race.
Nothofagus the best candidate comparing
in
is
243-256.
lution 27:
Fagaceae with Ticodendron, which at least su- Kovanda, M. 1984. Chromosome numbers selected
is in
289-301.
perficially similar in a number of important char- angiosperms Preslia 56:
(2).
&
lOPB chromosome
Love, A. A. Love. 1982. In:
acters.
number LXXIV. Taxon 31: 120-126.
reports
&
lUPB chromosome
Mehra, Hans. 1969.
P. A. S. /«;
LrrERATURE Cited XXL
number Taxon 310 315,
reports 18:
&
MoRAWETZ. W. M. Samuel. 1989. Karyo-
& R. A.
Armstrong, M. Wyue. A new
A. P. 1965. basic
J,
129 154
Pp.
patterns the Ilamainelltlae.
chromosome number logical in
the family Fagaceae (Noth-
in
&
Crane Blackmore Evolution,
in P. R. S. (editors),
ofagus), Nature 205: 1340-1341.
Ilamamclidae,
and History of the
Systematics, Fossil
Behnke,
H.-D. 1991.
Sieve-element characters of
Ti-
and 'Lower Hamamelidae'. System-
Introduction
codendron. Ann. 131-134. 1:
Missouri Bot. Card. 78:
Volume 40A, Clarendon
Association Special
Carlquist, Wood atics
S. 1991. and bark anatomy of
Ti-
Oxford.
Press,
codendron: comments
on Ann. Mis-
relationships.
Nixon, K. 1982. In support of recognition of the family
souri Bot. Card. 78: 96-104.
Kuprianova. Misc. Ser. Bot, Sci,
Nothofagaceae
Cronquist, An
a. 1981. Integrated System of Clas-
Amer. 162: 102.
New
sification of Flowering 2nd York
Plants, edition.
&
W. Crepet. 1989. Trigonobalanus (Fa-
L.
New
Bot, Card, Bronx, York.
taxonomic status and phylogenetic relation-
gaceae):
rKUER,
S. 1991. Pollen morphology and the systematic
828-841.
Amer. Bot. 76:
ships. J.
relationships of Ticodendron incognitum. Ann. Mis-
Chromosome some South
numl>er
Ono, M. 1977. in
souri Bot. Card. 78: 143-151.
agaceae
>TI
t.
^^LDblatt,
p. 1979. Miscellaneous chromosome counts
m Mag. Tokyo 90: 313 316.
Angiosperms including new family and generic
II,
Chromosome numt)ors species
Pashuk, K. T. 1987. in
records. Ann. 856-861.
Missouri Bot. Card. 66:
Chernogora (Ukrainian Carpa-
& of subalpine belt of
P. K. Endress. 1977. Cytology and evo-
1069-1074.
Zurn. 72:
Bot.
thians).
67-
lution in Hamameiidaceae. Arnold Arb. 58:
J.
The angiosporm phylog-
1975. bases of
Raven, H.
P.
71.
724-
Card. 62:
Ann. MLssouri Bot.
& eny: cytology.
ComezLaurito,
Gomez
1989. Ticoden-
L. D. P.
J.
dron: new
a tree from Central America. Ann. Mis-
265- nora
103
1972. Fagaceae. Pp. in
SOEPADMO,
E.
souri 1148-1151.
Bot. Card.
76:
& Malesiana ser. 1, 7.
199L new
Ticodendraceae: a
. anatomy,
morphology,
Reproductive
Tobe, H. 1991.
family
of flowering plants. Ann. Missouri Bot. Card.
Ticodendron, Ann. Missouri Bot.
and relationships of
78: 87-88.
135-142.
Hammel, & Card. 78:
W.
B. G. Burger. 1991. Neither oak nor
alder, but nearly: the history of Ticodendraceae. Ann.
Missouri 89-95.
Bot. Card. 78:
^-J^r
- -