A monograph of Eucharis and Caliphruria (Amaryllidaceae)

MISSING IMAGE

Material Information

Title:
A monograph of Eucharis and Caliphruria (Amaryllidaceae)
Physical Description:
viii, 497 leaves : ill. ; 28 cm.
Language:
English
Creator:
Meerow, Alan W
Publication Date:

Subjects

Subjects / Keywords:
Eucharis   ( lcsh )
Caliphruria   ( lcsh )
Amaryllidaceae   ( lcsh )
Horticultural Science thesis Ph. D
Dissertations, Academic -- Horticultural Science -- UF
Genre:
bibliography   ( marcgt )
non-fiction   ( marcgt )

Notes

Thesis:
Thesis (Ph. D.)--University of Florida, 1986.
Bibliography:
Bibliography: leaves 450-469.
Statement of Responsibility:
by Alan W. Meerow.
General Note:
Typescript.
General Note:
Vita.

Record Information

Source Institution:
University of Florida
Rights Management:
All applicable rights reserved by the source institution and holding location.
Resource Identifier:
aleph - 000898244
oclc - 15625180
notis - AEK6943
sobekcm - AA00004867_00001
System ID:
AA00004867:00001

Full Text










A MONOGRAPH OF Eucharis AND Caliphruria (AMARYLLIDACEAE)





By

ALAN W. MEEROW


A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN
PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF DOCTOR OF PHILOSOPHY



UNIVERSITY OF FLORIDA


1986















































Copyright 1986

by

Alan W. Meerow















ACKNOWLEDGMENTS


Many individuals have facilitated the execution of this

dissertation. Grateful appreciation is extended to my graduate

committee chairman, Bijan Dehgan, for his unwavering support throughout

my graduate program. Thanks are also extended to the remaining members

of my graduate committee, Charles L. Guy, Walter S. Judd, Thomas J.

Sheehan, and Norris H. Williams. In particular, I thank Walter Judd for

his advice and assistance throughout the course of my work, and Charles

Guy for the generous use of his laboratory and materials for my

electrophoretic investigations. Grateful appreciation is extended to

the curators of the herbaria cited in Chapter XII for the loan of

specimens, and the individuals and institutions, also cited in Chapter

XII, who provided living material of various genera of Amaryllidaceae.

Bart Schutzman provided fellowship, comraderie, and much assistance with

computer problems throughout the past five years. Kent Perkins,

collections manager at FLAS, dealt effectively and patiently with a

complex herbarium loan history. Much of the work detailed herein was

supported by National Science Foundation Dissertation Improvement Grant

BSR 8401208, and a Garden Club of America/World Wildlife Fund Fellowship

in Tropical Biology. I thank both granting organizations for this

material support. Gratitude is also extended to the Florida Federation

of Garden Clubs, and the Garden Writers Association of America, for

their respective scholarship awards. Above all, I thank my wife, Linda








Fisher-Meerow, for the love, support, and patience that has sustained me

during the completion of this work; for her excellent illustrations of a

number of Eucharis species; and her welcome companionship in the field.




















TABLE OF CONTENTS


ACKNOWLEDGMENTS .

ABSTRACT . .

CHAPTERS

I INTRODUCTION ..

II TAXONOMIC HISTORY .

III VEGETATIVE MORPHOLOGY


Materials and Methods
Results and Discussion


IV FLORAL MORPHOLOGY .

Materials and Methods
Results and Discussion

V POLLEN MORPHOLOGY .

Materials and Methods
Results .
Discussion .
Conclusions .

VI PHENETIC ANALYSES .

Materials and Methods
Results ..
Discussion ..
Conclusions ..

VII CHROMOSOME CYTOLOGY

Materials and Methods
Results ..


Discussion and Conclusions


VIII ELECTROPHORETIC ANALYSES OF ISOZYME VARIATION

Materials and Methods . .


Page

iii


vii


1

5


8


8
9


46


46
46

73

73
74
78
82

95

96
99
105
107

142

142
144
149

186

188


ooooo

ooooo


ooooo


oo
ooooo


ooooo


ooooo
ooooo

ooooo

oooeo
ooooo
oooo
ooooo

ooooo

ooooo
ooooo
ooo
oooeo

ooooo


oooo
ooooo
. .

. .


. .


. .
. .


. .


. .
. .

. .

. .
. .
. .
. .

. .

. .
. .
. .
. .

. .

. .
. .


. 0 .











Results . .
Discussion . .
Conclusions . .


IX ECOLOGY, PHENOLOGY,


AND P


H'


Ecology .
Phenology .
Dispersal .
Phytogeography .

X REPRODUCTIVE BIOLOGY .

Pollination Biology .
Breeding System .

XI PHYLOGENETIC RELATIONSHIPS

A Review of Urceolina .
Phylogenetic Analysis .

XII TAXONOMIC TREATMENT .

Materials and Methods .
Taxonomic Treatment .

LITERATURE CITED .

APPENDIX . .

BIOGRAPHICAL SKETCH .


YTOGEOGRAPHY .

. 4







. *



. 4

. 4

. I
. 4

AND EVOLUTIONARY


. .
. .

. .

. .
. .

. .

. .
.N .


* .

. .





HISTORY
* .









HISTORY

. .
. .

. .

. .
. .

. .

. .

. .


195
202
211

239

239
241
244
245

255

255
258

262

263
266

303

303
307

450

470

497














Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy


A MONOGRAPH OF Eucharis AND Caliphruria (AMARYLLIDACEAE)

By

Alan W. Meerow

December, 1986

Chairman: Bijan Dehgan
Major Department: Horticultural Science

Eucharis and Caliphruria are neotropical genera of petiolate-

leaved, white-flowered Amaryllidaceae found in the understory of primary

tropical rainforest. Together with the Peruvian endemic Urceolina,

Eucharis and Caliphruria form a monophyletic group on the basis of leaf

and seed morphology and ecological specialization. Sixteen species and

two natural hybrids within two subgenera are recognized in Eucharis.

Subgenus Eucharis, marked by its crateriform flowers, curved perianth

tube, well-developed staminal cup, and unicellular stigmatic papillae,

is distributed from Guatemala to Bolivia, chiefly in the western Amazon

basin and adjacent lower slopes of the eastern Andes. Subgenus

Heterocharis represents three relict species with many ancestral

characters of the genus. Caliphruria (4 species, 3 of which are endemic

to Colombia) has funnelform flowers, straight perianth tube, reduced

staminal connation, and multicellular stigmatic papillae. Abaxial leaf

surfaces of both genera have dense cuticular striations. Undulate

anticlinal cell walls are characteristic. A distinct palisade layer is









absent from the mesophyll. Eucharis subg. Eucharis has the least

derived pollen morphology, with characteristics in common with other

putatively ancestral genera of pancratioid Amaryllidaceae. Caliphruria

exhibits reduction trends in pollen grain size and exine sculpturing.

With the exception of two tetraploid species, all species are

characterized by 2n = 46. Karyomorphological change may be an important

factor in species divergence. Phenetic analyses achieve only fair

results in resolving phenetic relationships among Eucharis species, many

of which are highly variable morphologically. Analysis of isozyme

variation within two species complexes of Eucharis indicates high levels

of heterozygosity. Founder effects and hybridization are respectively

considered two important factors in the speciation of these groups.

Modern-day distribution of Eucharis and Caliphruria is related to

Pleistocene refugia theories. Phylogenetic analysis supports certain

species relationships hypothesized on the basis of phenetic data, but

indicates possible paraphyly for Eucharis if Caliphruria and Urceolina

are segregated as a distinct genera. Acceptance of paraphylly in

Eucharis is argued on the basis of degree of divergence of Caliphruria

and Urceolina. The relationship between these genera is paralleled

within other lineages of "infrafamily" Pancratioidinae. Keys and

descriptions are provided for all species of Eucharis and Caliphruria.


viii















CHAPTER I
INTRODUCTION



The closely-related genera Eucharis Planchon and Caliphruria

Herbert (Amaryllidaceae), the Amazon lilies, comprise, respectively, 16

and 4 species of bulbous, rainforest geophytes, adapted to the low-light
conditions of the forest understory. Together with the Peruvian endemic

Urecolina Reichb., Eucharis and Caliphruria form a monophyletic group

delimited by petiolate leaves with distinctive cuticular striation; a

turgid seed with a lustrous, usually black, testa; and complete fidelity

to the rainforest understory niche. The species are distributed from

Guatemala to Bolivia. The major center of diversity for Eucharis is

located in the western Amazon basin (inclusive of major tributary

systems, e.g., the Napo, Pastaza and Huallaga) and the adjoining lower

slopes of the eastern Andean cordillera. With the exception of single

Peruvian species, Caliphruria is restricted to the Cordilleras

Occidental and Central of Colombia. The species of both genera are no

where abundant, and are found growing only in primary, rarely secondary,

forest from ca. 50-1800 m elevation on soils of high fertility. The

latter factor is probably important in limiting their distribution in

the wild, and may also account for the highly localized population

demographics of many of the species. Large scale deforestation has

proven catastrophic to these plants. The plants are unable to adapt to

the higher light intensity of the clearings and soon perish. At least

several species are likely near extinction.










The Amazon lilies are marked by their evergreen, petiolate leaves;

white, often pendent, sometimes fragrant flowers with a frequently

conspicuous stamina cup or false corona formed by the basal connation

of the staminal filaments; obtusely tri-lobed stigma; and large, turgid,

ellipsoid seeds with a black, brown or metallic blue testa. A single

species of Eucharis, E. amazonica Linden ex Planchon, is widely known in

horticulture [erroneously as E. grandiflora Planchon and Linden (Meerow

and Dehgan, 1984a)], but neither genus has never been critically treated

in the taxonomic literature. Baker (1888) provided a key and

descriptions for all species known at the time in his Handbook of the

Amaryllideae, and Macbride (1936) treated the known Peruvian species of

Eucharis for the Flora of Peru. Though species of Eucharis have

continued to be described well into the present decade, the delimitation

of these species from previously described taxa has consistently

remained vague. No assessments of variation at either the population or

species level have been attempted.

My study of these genera began in 1980. Both were combined with

the closely related Urceolina by Traub (1971) without any supporting

data, the investigation of which formed the basis of my unpublished

master's thesis (Meerow, 1983). I refuted Traub's combination, and

Eucharis was re-established as a distinct genus with three subgenera:

Eucharis, Caliphruria (Herbert) Meerow ined. and Heterocharis Meerow

ined. On the basis of my continuous work since that time, I now believe

that Caliphruria is best retained as a distinct genus as well. This is

discussed in Chapter XI. Species delimitations and associated

systematic studies form the basis of this present work.









Neither Eucharis nor Caliphruria is not well represented in

herbarium collections, and critical morphological characters are often

obscured by the drying process. Consequently, a living collection of

over 100 accessions representing one dozen species was accumulated from

botanical gardens, individuals, and field collections by myself and

various colleagues. Study of living material not only clarified aspects

of floral and vegetative morphology, but allowed detailed study of

vegetative and seed anatomy, chromosome cytology, and electrophoretic

analysis of allozyme variation, as well as permitting the start of a

hybridization program. Populations were observed and sampled in

Colombia, Ecuador and Peru.

As two of only three genera of neotropical Amaryllidaceae (the

other being the related Urceolina) exhibiting complete fidelity to a

primary rainforest niche, systematic understanding of Eucharis and

Caliphruria offers important information relating to the evolution of

the pancratioid Amaryllidaceae ["infrafamily" Pancratioidinae sensu

Traub (1963)], centered in the central Andean region of South America.

As coadapted plants of the world's most complex and threatened

ecosystem, their scarcity in the wild will only increase to the point of

extinction if tropical deforestation continues at the present rate.

No taxonomic scheme can encompass all of the information about the

group of organisms under study (Ornduff, 1969; Raven, 1976; Holsinger,

1984, 1985). In genera such as Eucharis and Caliphruria, rare in the

wild and often accessible only with difficulty, and which often exhibits

cryptic patterns of variation, this observation becomes that much more

acute. By accumulating data from as many diverse sources as possible, I

have attempted to construct a classification of the Amazon lilies that







4

reflects their phylogeny and inter-relationships as accurately as these

data allow.















CHAPTER II
TAXONOMIC HISTORY



Herbert (1844) described the new genus Caliphruria from

collections made in New Granada (Colombia) near Guaduas by Hartweg,

placing Caliphruria in the "section" Pancratiformes of "suborder"
Amaryllideae. Genera of this section were united by the single

character of stamina connation. The small, white, funnelform flowers

were marked by the presence of a bristle or slender tooth at either side

of the filament. Herbert (1844) made no mention of basal connation of

the filaments. The orthography of the name, a single '1' in the Greek
stem "calli-," was emended by subsequent workers (e.g. Baker, 1877), but

was returned to Herbert's original by Meerow and Dehgan (1984b).

Planchon (1852) introduced another new pancratioid genus,

Eucharis. The first species described, E. candida Planchon and Linden,

collected in New Granada by M. Schlim, was characterized by its

crateriform flowers with a conspicuous staminal cup and a widely

spreading perianth limb. If Planchon was aware of Caliphruria, he did

not note any relationship between it and Eucharis.

Noting the relationship between the two genera, Bentham and Hooker

(1883) placed Caliphruria and Eucharis in the tribe Cyathiferae.

Caliphruria subedentata Bak. was combined with Eucharis in their

treatment, while C. hartwegiana Herb. was retained. Baker (1888)

accepted this treatment, and described another new species as C. tenera.

He described Caliphruria as "nearly allied to Eucharis."










Baillon (1894) described the species C. castelnaeana and declared

it precisely intermediate between Caliphruria and Eucharis. He further

suggested that Eucharis should be treated as a section of Caliphruria.

Macbride (1931) later transferred C. castelnaeana to Eucharis.

Nicholson (1884) transferred C. hartwegiana (the type species of

Caliphruria) to Eucharis, ignoring the fact of the former's

nomenclatural priority. Traub (1967) made the formal transfer of the

remaining species of Caliphruria, C. tenera Baker, to Eucharis and also

combined the monotypic genus Plagiolirion Baker (1883) with Eucharis.

He had previously listed Caliphruria, Plagiolirion, and Mathieua

Klotzsch [another monotypic genus (Meerow, MS in subm.)] as synonyms for

Eucharis in his Genera of the Amaryllidaceae, citing Baillon (1894) as a

"special reference" (Traub 1963, p. 74). The nomenclatural priority of

Caliphruria Herbert was overlooked. No proposal for the conservation of

Eucharis Planchon over Caliphruria Herbert has ever been proposed

previous to that of Meerow and Dehgan (1984b).

Traub (1971) later combined Eucharis with Urceolina Reich. (nom.

cons.), a small Andean genus with petiolate leaves and markedly

urceolate, brightly colored flowers. He designated five subgenera:

Urceolina, Eucharis, Caliphruria, Mathieua, and Plagiolirion. Traub

(1971) offered no explanation for the combination of Eucharis and

Urceolina, but presumably his decision was prompted in part by reports

in the literature of two intergeneric hybrids between Eucharis and

Urceolina: X Urceocharis clibranii Masters, an artificial hybrid, and X

U. edentata C. H. Wright, putatively discovered in the wild state in

Peru.






7


In my preliminary work (Meerow, 1983; Meerow and Dehgan, 1984a,

b), I refuted Traub's unsupported combination, re-establishing Eucharis

(including Caliphruria), Plagiolirion, and Mathiuea as distinct genera.

Plagiolirion and Mathieua are monotypic genera allied to Hymenocallis

Salisb. and Stenomesson Herbert respectively (Meerow, MS in subm.). On

the basis of the work detailed in the following chapters, I prefer to

treat Caliphruria as a distinct genus as well.














CHAPTER III
VEGETATIVE MORPHOLOGY



Materials and Methods



Leaf Surface Morphology [Scanning Electron Microscopy (SEM)]

Fresh material was fixed in FAA for a minimum of 24 hrs. Leaf

sections were always taken from midpoint of the lamina. Samples were

EtOH-dehydrated, critically point dried with a Denton DCP-1 apparatus,

mounted, and coated with 600 X of gold-palladium mixture with a Technics

Hummer V sputter coater. Specimens were observed and photographed on an

Hitachi S-450 scanning electron microscope at 20 kv.



Anatomical Studies

Petioles were freehand sectioned with teflon coated razor blades,

and lightly stained with toludidine blue. Leaf clearings were prepared

by immersing fresh, medial lamina sections in 85% lactic acid at 450 C

for 5-7 days. Abaxial and adaxial epidermal layers were then separated

using diluted Jeffrey's (1917) solution. Epidermal tissue was stained

in 1% safranin, and mounted in 1:1 glycerol 95% EtOH mixture. Leaf

tissue (medial lamina) fixed in FAA was processed for paraffin block

sectioning with an American Optical T/P 8000 tissue processor.

Dehydration began with 50% EtOH, and continued through 50-100% tert-

butyl alcohol. Dehydrated material was then transferred to a 1% solution

of safranin, followed by a 1:1 mixture of absolute tert-butyl alcohol










and mineral oil, a 3:1 mixture of Paraplast and mineral oil, and finally

100% Paraplast. Embedded material was sectioned with an American

Optical No. 820 rotary microtome set at 10 um. Sections were further

stained with toluidine blue. Sections were mounted using Pro-Texx

mounting medium. All material was examined and photographed on a Nikon

Labophot photomicroscope with AFX-II photographic attachment.

Terminology is referable to Stace (1965) and Dilcher (1973).



Statistical Procedures

Statistical tests were performed with SAS Release 5.08 (SAS

Institute, INC.) on the Northeast Regional Data Center (NERDC) of the

University of Florida.



Results and Discussion



Bulbs

The globose or subglobose bulbs of Eucharis and Caliphruria,

composed of concentric and modified leaf bases (scales), are

characteristic of most members of the Amaryllidaceae. The outer scales

of Eucharis and Caliphruria bulbs are modified into a papery tunic,

either brown or tan in color. On the whole, little systematic

information can be derived from bulb morphology. Size of mature bulbs

is variable and proportional to the whole plant size range of each

species.

The bulb scales of most species of Eucharis and Caliphruria are

apically articulated into a neck or pseudostem of variable length.

Distally, the pseudostem grades into the petiole of individual leaves.









Unlike species of Hymenocallis subg. Ismene, the neck of Eucharis and

Caliphruria bulbs always remains below the soil surface. Length of the

neck may therefore be more a factor of bulb depth in the substrate

rather than of any phylogenetic significance.

Van Bragt et al. (1985) found flower initiation will not occur in

bulbs of E. amazonica less than 35 mm diameter. This species

characteristically produces among the largest bulbs of any in the genus.

Irmisch (1850) recognized the existence of both monopodial and

sympodial shoot structure in bulbs of Amaryllidaceae. Pax (1888) and

Troll (1937) followed these same concepts, but other workers, (e.g.

Church, 1919; Peter, 1971; Brunard and Tulier, 1971; U. and D. MUller-

Doblies (1978), Dzidziguri, 1978; Akensova and Sedovi, 1981; and Arroyo,

1984) have not always been in agreement as to which shoot structure was

applicable to various taxa. Most recently, Arroyo (1984) characterized

only four South African taxa as possessing true monopodial organization.

She is now in accord (Arroyo, pers. comm.) with MUller-Doblies and

Mlller-Doblies (1972) who suggest that only sympodial structure occurs

in bulbs of Amaryllidaceae.

Bulbs of Eucharis and Caliphruria are, without controversy, true

sympodia (Arroyo, 1984; van Bragt et al., 1985). Each bulb has a single

terminal vegetative meristem. After its transformation to a floral

apex, a new vegetative growing point is formed laterally.

Bulbs of most species of Eucharis and Caliphruria offset regularly

and vigorously, forming sizable clumps in time if undisturbed. Offset

bulbs are at first usually tightly joined at the basal plate to the

parent bulb. Only very rarely does an elongated rhizome develop from

which a new shoot system can arise at some distance from the parent









plant, a mode of vegetative reproduction characteristic of some species

of Hymenocallis (pers. obs.; T. Howard, pers. com.).



Leaf Gross Morphology

Leaves of Eucharis and Caliphruria are uniformly long-petiolate,

with a well-developed elliptic, ovate or lanceolate lamina. Petiolate

leaves are characteristic of a number of genera of Amaryllidaceae,

either completely or in part, and have undoubtedly evolved independently

several times from the linear or lorate leaf morphology typical of the
family. In most of these cases, the petiolate leaf appears to be

primarily an adaptation to reduced levels of light concurrent with

colonization of forest understory (e.g. Eucharis, Caliphruria,

Urceolina, Eurycles Salisb., Scadoxus (Raj.) F. Nordal, Hymenocallis

tubiflora Salisb. and allied species.). Foilage of petiolate-leaved

genera occurring in more open situations (e.g. Phaedranassa Herbert,

Rauhia Traub), is generally marked by increased succulence and/or

pruinosity [leaf surface wax is capable of reflecting light and heat

(Cutler et al., 1980)].

The petiole of the leaf of Eucharis and Caliphruria is usually as

long or longer than the lamina. It is subterete in cross-section (Fig.

1-2), rounded abaxially, and flattened adaxially, becoming slightly

channelled proximal to the sinus. The petiole is winged proximal to the

sinus by the attenuation of the lamina. The midrib is pronounced

abaxially along the entire length of the lamina, and slightly channelled

adaxially, continuous with the petiole.

Leaf shape only rarely provides taxonomically useful information.

Length : width ratios are subject to considerable variation even among









the leaves of a single bulb. Herbarium specimens will frequently

include only a single leaf, with no indication of its developmental age.

Taxonomic consistency of leaf shape is exceptional, but useful in the

few cases where it occurs. For example, leaves of E. ulei are

consistently narrowly elliptic. Eucharis amazonica (leaf length/width

ratio greater than 2) may be delimited from from E. anomala (1 : w less

than 2).

The leaves of Eucharis and Caliphruria are completely glabrous and

non-glaucous with a single exception. Eucharis bonplandii (Kunth)

Traub, a rare tetraploid species from central Colombia, develops a

glaucous bloom in strong light that gives the leaf a blue cast.

The leaves of Caliphruria are slightly thicker than those of subg.

Eucharis and Heterocharis. This may reflect adaptation to slight water

stress, as the species of Caliphruria sometimes inhabit slightly drier

forest associations than characteristic of the Eucharis (see Chapter

IX). Eucharis bouchei Woodson and Allen and E. bonplandii (both subg.

Eucharis), however, have thickened leaves, possibly a consequence of

their tetraploid constitution (see Stebbins, 1950).

The leaf apex of all species is shortly acuminate, the base

attenuate. Coarse undulation of the margin will sometimes make the

lamina appear cordate at the base. Leaf margins of Caliphruria are

uniformly non-undulate.

Venation of the leaf of Eucharis and Caliphruria is

parallelodromus (Hickey, 1973), with a great number of transverse,

commisseral veins inter-connecting the primary vasculature. Whether the

leaf is plicate along the primary veins can be a taxonomically useful

character. Unfortunately, this, and many other leaf characters (e.g.,









marginal undulation, the color and luster of the epidermis), readily

observed in live material, are completely obscured in herbarium

specimens.

All species of Caliphruria have smooth, non-plicate leaves.

Eucharis is variable for this character, but the majority of species of

have plicate leaves.

The adaxial epidermis of most species of both genera is a

lustrous, dark green; the abaxial surface appears lighter, or silvery-

green. Only E. astrophiala (Ravenna) Ravenna has diverged markedly from
the typical morphology, and has a uniquely non-lustrous, bullate-

pustulate leaf texture.



Leaf Surface Features

Cuticle. Cuticular striation is prominent on the abaxial leaf

surfaces of most Eucharis and Caliphruria species (Fig. 3-7, 9-13, 15-

18). Striae are thickest in C. subedentata (Fig. 6). Arroyo and Cutler

(1984) recognized eight cuticular sculpturing classes in a survey of 25

genera of Amaryllidaceae. The most common cuticular morphology of

Eucharis and Caliphruria fits their class VII: "thick striae, parallel

or not, interlocking, + transverse" (Arroyo and Cutler, 1984, p. 471), a

type they reported only for the few species of Phaedranassa, Scadoxus,

and Griffinia Ker-Gawl. that they examined, all three genera with

petiolate leaves, but not closely related. Phaedranassa is, however,

rather variable in its cuticular morphology (Meerow, unpubl.). Eucrosia

Ker-Gawl., a close ally of Phaedranassa, also has cuticular striation

similar to that of Arroyo and Cutler's type VII (Meerow and Dehgan,

1985), but differs in the orientation, thickness and pattern of the









striae. Urceolina, a small genus very closely related to Eucharis and

Caliphruria, has cuticular morphology much like that of the latter

genera.

In a few species of Eucharis (E. amazonica, E. anomala (Fig. 3; E.

bouchei, Fig. 18) the striation is much less pronounced. Caliphruria

korsakoffii (the sole representative of Caliphruria outside Colombia)

has the most aberrant cuticle morphology (Fig. 7), corresponding more or

less to type V of Arroyo and Cutler (1984): "central, thick axial

striation with less pronounced striae running from it, directly to

anticlinal walls" (Arroyo and Cutler, 1984, p. 471). The adaxial

cuticle of Eucharis and Caliphruria is either smooth or rarely much more

finely striate than the abaxial surface, the striations entirely axial.

The adaxial cuticle of C. korsakoffii (Fig. 8) has several, thick,

transverse striations across each cell, and the epidermis is unusually

flat in topography.

Stomata. Leaves of Eucharis and Caliphruria are predominantly

hypostomatic. Stomata occur adaxially only along the midrib and

vicinity (Fig. 19A), and also occasionally in the proximity of primary

veins. Stomata are usually absent from the abaxial midrib (Fig. 19B).

Intercalary stomata were regularly observed only in E. cyaneosperma

Meerow (Fig. 14 & 21C). A survey of leaf surfaces in "infrafamily"

Pancratioidinae (Meerow, unpubl.) suggests that loss or reduction of

adaxial stomata frequently accompanies the evolution of petiolate leaves

in Amaryllidaceae. Most linear or lorate-leaved genera are

amphistomatic. The stomata of Eucharis and Caliphruria are anomocytic,

as is typical for Amaryllidaceae (Arroyo and Cutler, 1984; Dahlgren and

Clifford, 1982), though E. astrophiala exhibits at least slight









differentiation of cells neighboring the stomata (Fig. 9-10) from other

epidermal cells. These cells are more densely and regularly striate

than other epidermal cells, as well as slightly more upraised. The

guard cells of Eucharis and Caliphruria are oriented with their longest

axis parallel to that of the leaf. Wide variation in stomatal index

{[no. of stomata / (no. of stomata + no. of epidermal cells)] X 100

(Salisbury, 1927)} is evident (Table 2). Infraspecific variation in SI

can be as wide as that between species, however, and seems to have

little taxonomic significance. Correlations between SI and leaf width,
length and length:width ratios were tested for all plants examined.

Pearson correlation coeffcients for the three comparisons were 0.249

(width), 0.421 (length) and 0.232 (length:width). Greatest correlation

of SI was with leaf length, but none of the three tested factors are

very significant. Salisbury (1927) reported that humidity affects

stomatal index, and other workers (Yapp, 1912; Gupta, 1961) have

suggested that SI may not be as invariant as has been claimed. The

great morphological variation of Eucharis species (subg. Eucharis in

particular) in characters of floral morphology (Chaper IV) is also

present in vegetative characters.

Epidermal cells. The epidermal cells of all species of E. subg.

Eucharis have strongly undulate anticlinal walls (Fig. 20-23A), as as

noted by Asatrian (1984) for the few species he surveyed. Abaxial

cells are more strongly undulate than those of the adaxial surface.

Abaxial epidermal cells of Caliphruria (Fig. B-C) are more weakly

undulate, and the adaxial cells of C. korsakoffii (Fig. 24C) are

completely straight. Eucharis subg. Heterocharis is polymorphic for

anticlinal wall morphology. Eucharis amazonica (Fig. 23A) and E.








sanderi (not illustrated) have strongly undulate walls, while E. anomala

(Fig. 23B) has essentially straight walls. I have surveyed the leaf

surface morphology of all genera in "infrafamily" Pancratioidinae with

the exception of Pucara Ravenna (Meerow, unpubl.). Strongly undulate

anticlinal walls are very rare among these genera. Arroyo and Cutler

(1984) report similar findings for the genera of pancratioid

Amaryllidaceae that they surveyed. Arroyo and Cutler (1984) and

Artushenko (1980) consider undulate anticlinal walls to be primitive for

the family. No detailed reasons are given by these authors for this
assessment, though reference is made to Scadoxus, a putatively primitive

bulbless genus of African Amaryllidaceae with undulate anticlinal walls

and Type VIII striation. This genus is often considered close to the

ancestral complex that gave rise to the Amaryllidaceae (Arroyo, 1982;

Arroyo and Cutler, 1984; Nordal and Duncan, 1984). Yet Scadoxus has a

baccate fruit, "brush" type inflorescence morphology, and petiolate

leaves (Nordal and Duncan, 1984), all derived characters in relation to

the rest of the family (Meerow, 1985a). Consequently, there seems

little evidence to suggest that the undulate anticlinal walls of

Scadoxus represent the primitive condition for the Amaryllidaceae.

Eucharis anomala, putatively the most primitive species of Eucharis (see

Chapter XI) has straight anticlinal walls (Fig. 23B). Taking this fact

into consideration, along with the relative rarity of undulate

anticlinal walls throughout the Pancratioidinae, I believe the undulate

condition is more likely the derived state. End walls of the both the

abaxial and adaxial cells of all species range from oblique to rounded.

Abaxial epidermal cells range from rectangular to irregular in

shape. Adaxial epidermal cells are, in almost all cases, rectangular.









Eucharis astrophiala (Fig. 20A) has the most irregularly shaped cells of

both the abaxial and adaxial surfaces. In the vicinity of the midrib on

both surfaces of the leaf (Fig. 19), epidermal cells become

conspicuously elongated, and anticlinal walls are straight. Epidermal

cells of the midrib are extremely long and narrow.



Leaf Anatomy

In petiolar transverse section, a single arc of vascular bundles

is usually observed (Fig. 1-2). Median bundles are the largest. In
petioles of E. anomala Meerow (Fig.2B) and the closely related E.

amazonica (Fig. 2C), both in subg. Heterocharis, small secondary bundles

were observed near the adaxial surface. These bundles are most

conspicuous in E. anomala; they are markedly smaller in E. amazonica.

These secondary vascular traces disappear above the middle of the

petiole. Asatrian (1984), who reported on petiole anatomy of three

Eucharis and Caliphruria species, did not observe these bundles in E.

amazonica (cited as E. grandiflora).

The internal morphology of leaves of Eucharis and Caliphruria

(Fig. 25-31) is largely invariant across both genus. No well defined

palisade layer is evident, a characteristic of most genera of

"infrafamily" Pancratioidinae (Arroyo and Cutler, 1984; Meerow, unpubl.

data). Mucilage cells, common throughout the family (Arroyo and Cutler,

1984), are often present near the leaf surface, and raphides are

occasionally observed in epidermal cells. The mesophyll consists of

several layers of chlorenchyma both ad- and abaxially, and a thicker

region of spongy, slightly aerenchymous tissue. Small air cavities

occur regularly only directly below stomata. Vascular bundles are






18


surrounded by a sheath of 1-2 layers of parenchymous cells. The only

xylem elements present are tracheids with annular thickenings (Fig. 28).


















CL
3
4-)

Qv








LL a
i- 4-
0 0








x0a
U
0 *-

au


LL.




0
a(fl r
*-o

I- <




0









f V
0 S









4-)





4- (V
00
>

















-C Z
*i- 0
<0 -o
ad)
E U-
0
+J +J









aQ)
V) a)
O 0.
C 0







a,)
0 0



30





4.)





0a)



















a)
XU -
.0
r0













I-
2c,
**E




r-a
^= u







*-


mt CM m .-.1 00 lc" LO

* O ( *-
0M -4 r-4 r-4 -4 r-4









o r.. 0 In 0 0 Co )
o lO LO m 1a, LO cn
O n IC CM m n c)
cv) C) ('.2 ('2 v) cv) C


CM. 0 ':a*
C- ~i> F-.

o ct li.










CO I0 C')
C) 0 C
CM) C) 0M









0 LO 1LO

COCO CO






0 In 0
C ) *

-4 i-4 CM


x




-J
z

























-Jr




I.- I--- --
CD3
















-J
I
L_(- -
-t QE
UJZU <





-1X ^



Ll -
<
LUZ
-IJO
_1











Q:
UJ

=>


o C) O-4 m
* *







0 aLO
* *
c d 2


3 LO LO PO

C ID LO 1-0 ii m m -I

) M 0 1, Ca) a) 0 t
o0 0 0 0 C = 0 0 0 C
L L L S. S. LC
c ( 0 0c
S- S 3 S- 0











U) .C 3 v-) 0> C U) 0
0w I a 0
00 4

0 U 0j h > 0 E
0) rS.. 0I U 01 (0 C S

Lag al LalO ialO(0 al








































O 3 CM

C0 CM C4

















-4 4 -4
LO C>J CD












SCO 0
* *-





U0 0 0


LiLI

6-4














-.J




U-

<
LI- u





-J L
=



















..LL J
W-JU





















I-I


m ,o ,--4
-4 10 -4













0'i m C0






C4 C4 y










o In o


mO CM C0
00 CM C


n 4


S L Lm L


0 0 0 0
o 1' s






014 -i


C u- (O 0 C
If) 0 0 .C : (0


<0 r-C 3


*-I T- O '-


(U (U
C C 0 0

0 0u (U (




0. 0.


00 40-(
=0



U (J U >
-II-l010 01w











*r- I (U
0.0Q. 0.

LjL | L;|




































.-4 r-4 ,--









u) r A0

L- CM CP0 0
10 No 0 -,









CO O 0 L
00 co CO nO
* *l 9


x
UJ
0



'--

I-









-r
0







3




















-
<=

U-3--









L"J "
_j0 2







<
U-I---.-

-1










LU


=
0















0
x


oCD 0 M

e-4 e1 H M

2 3 3 3
0 0 0 0
0- S- 1- 0-

v) a> 0) a)






*r- 0U
*r-
0 +- 4-
*r- 0 C

UI I (>
.= S- .0
0. 0 3
*a -a 01

0 0| )


r-





I-
O


0





5-
x
0










e-
4r-












0 -

4r-J






0 o
C
*Qe


















O 0
U0






4- *- C-
r4-
0 0 0















L 0
o

e x x









0 0


L) r-

lA LA
i-i i-

















-I



CD











L 0)





CL
0 U









0 S




o -
u l












((
0.
0 tf.























(- 0






U 01U

ro x
0










0<
O-
< o





























0







V LLJ

o .
.- (U


C II















00


*O
0C,,<







-r-



















-Q







23











































3
'3 \






























o I 1
3 3



3 '3
'3
6:: '7
o Lr c>~ t :'

'3 '
a 3



CY '3~
















'3'3'
C,: C,
aC
a 9 ,
a C,





























e-
r *






0
0*











0 *o
S0













ja


"rI
< --.









a) LL
ke-







0
4*-
id
r- E





U ,
0) WI



In



Q. E











Q..
II



L *r

C *i



-J

I* -




310





*-I

zi-
*1)







25







































4 4
as

























Ia



r
: p J .^ t y




















Figures 3.3-3.8. SEM photomicrographs of Eucharis and Caliphruria leaf
surfaces. 3-7. Abaxial surfaces. 3. E. anomala (Meerow 1141,
FLAS). 4. E. X grandiflora (Madison et-al. s. n., SELL~. T
Calicharis 'utcheri (Meerow 1TIJT FAST.-6.C.- subedentata
(Meerow 1109, FLAS). 77. korsakoffii (Meeriw T10, FAST. 8.
Adaxial surface of C. korakottri (Meerow Tt, T-AS). All scales
= 25 num.












~ar~x"i~~PaE~"3~c~s~ts~
Lli~~v~ *r*I.....YIEIJI~i,



















Figures 3.9-3.14. SEM photomicrographs of Eucharis leaf surfaces. 9-
13. Abaxial leaf surfaces. 9-10. E. astrophiala (Meerow 1111,
FLAS). 11-12. E. plicata subsp. pTicata (Meerow 10 5FTLAST.
13. E. cyaneosperma (Meerow 1032, FIAS). 1T. axiaT leaf
surface, E. cyaneosperma (Meerow 1032, FLAS). All scales = 25 pm.









-LIE-I




















Figures 3.15-3.18. SEM photomicrographs of Eucharis abaxial leaf
surfaces. 15. E. bakeriana (Meerow 1108, FLAS). 16. E. formosa
(Meerow 1103, FTAS). 17. bonpland1iT(Bauml 686, HUITT). 18.
E. bouchei ar. dressleri TMeerow 11077 FLAS). T scales = 25
/Mm.




31










I
7-7-



W-- % esi




















Figure 3.19. Leaf epidermal cell configurations of representative
Eucharis species in the vicinity of the midrib. A. E. formosa
(Schunke 14174, FLAS), adaxial surface. B. E. plicata subsp.
plicata (Meerow 1025, FLAS).


































0.2MM
II




















Figure 3.20. Leaf epidermal configurations of Eucharis species in the
inter-costal area of the leaf. A. E. astrophiala (Madison 3792,
SEL). B. E. bonplandii (Bauml 686,-HUNIJ. C. E. bouchne var.
bouchei (MFeerow I125, FLAST-











































0.2MM
B ------



















Figure 3.21. Leaf epidermal configurations of Eucharis species in the
inter-costal area of the leaf. A. E. candida (Meerow 1144, FLAS).
B. E. castelnaeana (Schunke 14156, FLAS). C. E. cyaneosperma
(MeTrow 1032, FIS).




37




ab _-ad













A














0.2MM B












C
.......C...... ...... ...... ......




















Figure 3.22. Leaf epidermal configurations of Eucharis species in the
inter-costal area of the leaf. A. E. formosa (Schunke 14174,
FLAS). B. E. plicata subsp. brevidentata (Meerow 1143J,FLAS). C.
E. ulei (ScFunke 14153, FLAS).





39





ab ad
'I












A
















B .2MM















C




















Figure 3.23. Leaf epidermal configurations of Eucharis species in the
inter-costal area of the leaf. A. E. amazonica (Schunke 14179,
FLAS). B. E. anomala (Meerow 1141)7 C. E. X grandiflora Merow
1127, FLAS).




41







ad













A














B -. ,0.2MM




















Figure 3.24. Leaf epidermal configurations of Eucharis and Caliphruria
species and hybrid in the inter-costal area of the leaf. A.
Calicharis butcheri (Meerow 1110, FLAS). B. C. subedentata
(Meerow 1123, FLAS). C. C. korsFkoffii (Meeroi 1096, FLAS.





43



ab ad












A














B 0.2 M













C



















Figures 3.25-3.31. Transverse sections of Eucharis and Caliphruria
leaves. 25. E. bonplandii (Bauml 686,- HUNT) 26. E. astrophiala
(Madison 3792, SEL). 27-8E.-To.rmosa (Meerow 1107, FLAS). 28.
Tracheid wit" annular thickenTngs. 29. EE. ouchef1var. dressleri
(Meerow 1107, FLAS). 30. C. subedentata-(Meerow 1123, FLAS). 31.
C. korsaIToTii (Meerow 1097, FLAS). AlT scales ="--~7 jm except 25
pm- in Fig. 28.











































L -(


trtr2

^- A. *^^ r- .-

-h- -I




27


, ", -J --
_* ~- L ~ *- --, _-- S



;-f- --I














CHAPTER IV
FLORAL MORPHOLOGY


Materials and Methods


Scanning Electron Microscopy (SEM)

Stigmas and seeds preserved in FAA were prepared and examined as

described in Chapter III.



Anatomical Studies

Seeds preserved in FAA were prepared for parafin block sectioning

as described for leaves in Chapter III. Scape sections were prepared

freehand as described for petioles in Chapter III.



Results and Discussion



Inflorescence

The inflorescence of Eucharis and Caliphruria is a naked scape

typical of Amaryllidaceae. The scape is sub-terete in cross-sectional

outline (Fig. 1), and has a solid pith. Vascular bundles are

distributed in several concentric rings within the pith (Fig. 1). A

layer of collenchyma cells occurs just below the epidermis of the scape.

The scape is terminated by two valvate-imbricate, ovate-lanceolate

bracts that enclose several secondary bracts and the flower buds before

anthesis. These bracts vary from green (E. subg. Heterocharis) to

greenish-white (most species of subg. Eucharis) and are soon marcescent










after opening and spreading laterally. Each flower is subtended by a

linear-lanceolate bracteole.

The inflorescence of the Amaryllidaceae is traditionally described

as "umbellate". Developmental work by Mann (1959) on Allium, and Stout

(1944) on Hippeastrum suggests that the superficially simple umbel of

Amaryllidaceae actually represents a complex series of reduced, helicoid

cymes. Anthesis occurs in a strict sequence within each cyme from the

developmentally oldest flower to the youngest. The peripheral cymes

flower first; the central cymes flower last.

Flower number varies in Eucharis and Caliphruria from 2-10, rarely

as many as 12 (C. korsakoffii). Number of flowers is often a

taxonomically useful character, though any species characterized by 8-10

flowers is capable of producing a depauperate inflorescence with fewer

florets. An increase in flower number generally does not occur. In

some species of subg. Eucharis (E. astrophiala, E. bouchei, E. ulei), a

flower number of 5 has become virtually fixed. Reduction in flower

number is usually considered the derived state in Amaryllidaceae (Traub

1962, 1963).



Flower Size and Fragrance

Flower size. The largest flowers in Eucharis are found in subg.

Heterocharis, flowers of which average 7-8 cm in length. Flowers of

Caliphruria are the smallest, never exceeding 4 cm in length. Subgenus

Eucharis, the largest of the two subgenera of Eucharis, is variable,

with flowers ranging from 3-7 cm in length. Within a fairly broad

range, flower size can be used to distinguish phenetic species complexes

within subg. Eucharis (see Chapters VI and XII), however, most species









of this subgenus are quite variable in size. Flower size may also be a

factor of plant vigor and soil fertility. I have repeatedly noted

differences from year to year in the size of flowers of greenhouse

collections, depending on the relative health of the plant.

Floral fragrance. Subgenus Heterocharis is the only subgenus of

Eucharis that is uniformly fragrant. The fragrance of all species of

subg. Heterocharis is intense and sweet. Flowers of Caliphruria do not

emit any detectable fragrance. Most species of E. subg. Eucharis are

also without noticeable fragrance. In the few species of this subgenus
that are fragrant (E. bakeriana, E. castelnaeana, E. formosa, and E.

plicata subsp. brevidentata), the odor is not intense. In one case (E.

formosa), the fragrance is slightly fetid. The significance of floral

fragrance in Eucharis is discussed further in Chapter X.



Perianth

The perianth of Eucharis and Caliphruria consists of six tepals in

two whorls, basally connate into a tube of varying length and

morphology. The tube of E. subg. Eucharis (Fig. 2E) is cylindrical for

almost its entire length, abruptly dilating near the perianth throat.

The tube of subg. Eucharis is also strongly curved, either abruptly just

above the ovary (E. bakeriana, E. cyaneopserma), or gradually throughout

the proximal half of its length (all other species). The curving of the

tube results in the pendent habit of most species of subg. Eucharis.

The tube is white for its entire length.

The tube of subg. Heterocharis (Fig. 2C, D) is tinted green

proximally (for at least half its length). The tube is curved, though

not as markedly as that of subg. Eucharis, and the habit of the flowers









is either declinate (E. anomala, E. sanderi) or sub-pendulous (E.

amazonica). The tube is cylindrical for 1/2 to 2/3 of its length; it

abruptly dilates in the distal half to 1/3. The tube morphology of X

Calicharis butcheri (Fig. 28), putatively an inter-subgeneric hybrid

between E. sanderi and C. subedentata, is intermediate between

Caliphruria (Fig. 2A) and E. subg. Heterocharis (Fig. 2C, D).

The tube of Caliphruria (Fig. 2A) is straight, and dilates

gradually from base to throat. It is either sub-cylindrical (C.

korsakoffii) or funnelform in shape (all other species). The tube is

tinted green proximally (in C. subedentata, for 1/2 to 2/3 of its

length).

The tepals of Eucharis and Caliphruria flowers are white. Those

of the outer series are almost always longer and narrower than the inner

tepals. The outer tepals are apiculate. The apiculum frequently has a

small, papillate horn on the adaxial surface in E. subg. Eucharis. The

inner tepals vary from acute to obtuse, sometimes minutely apiculate, at

the apex.

The tepals of most species of subg. Eucharis spread at an angle of

900 or more from the throat. Perianth morphology of subg. Eucharis is

thus predominantly crateriform. At times the tepals may be reflexed

strongly above the midpoint of their length, or rarely for their entire

length. Tepal habit varies even among flowers of the same inflorescence

and shows no taxonomic consistency. If exposed to strong light, the

abaxial midrib of the tepals of some species of subg. Eucharis may be

lightly pigmented yellow.

The perianth of Caliphruria is infundibular. The tepals remain

imbricate for half their length and spread distally at an angle of only









45-600. The tepals of subg. Heterocharis are also, for the most part,

imbricate proximally, and spread at 45-600 from the throat. The

perianth is more or less campanulate in morphology. One species, E.

amazonica, has the crateriform perianth characteristic of subg. Eucharis

with a wide-spreading (ca. 900) limb.


Androecium

Staminal connation is one of the major characteristics of

"infrafamily" Pancratioidinae. Some taxonomic workers have mistakenly
considered the staminal cup of pancratioid genera homologous to the

corona of Narcissus (e.g., Pax, 1888). The corona of Narcissus is of

perianthal origin (Eichler, 1875; Arber, 1937), while the stamina cup

of pancratioid taxa is composed entirely of androecial tissue (Arber,

1937; Singh, 1972).

The stamens of Eucharis and Caliphruria are variously connate

proximally. In most species of subg. Eucharis and several species of

subg. Heterocharis (E. anomala and E. amazonica), a conspicuous staminal

cup or false corona is present (Fig. 3-4). In Caliphruria, the cup is

reduced to a short, membranous, connate portion of the filaments near

the perianth throat (Fig. 5). Eucharis sanderi (subg. Heterocharis) has

a reduced staminal cup similar to that of Caliphruria.

Stamens of Eucharis and Caliphruria may be dentate, edentate or

irregularly toothed. Both types of stamina morphology may occur in the

same species, and variation may occur even among flowers of a single

clone. The presence or absence of stamina dentation has frequently

been overweighted in the alpha-taxonomic literature relating to these

genera (e.g., Ravenna, 1982), but only occasionally has profound









taxonomic significance [e.g., E. astrophiala (Fig. 3), the only species

of subg. Eucharis that always has an edentate staminal cup].

A variable pattern of green or yellow pigmentation is present in

the androecium of all species of subg. Eucharis and Heterocharis.

Stamens of Caliphruria are completely white. In subg. Heterocharis, the

green (rarely yellowish) pigmentation is largely restricted to the

interior of the cup, and extends into the dilated portion of the tube as

well (Fig. 4). The coloration is concentrated along the filamental

traces, but the tissue between the traces is suffused with green as

well. In subg. Eucharis, pigmentation is present on both the exterior

and interior surfaces of the cup, does not extend into the dilated

portion of the tube, and takes the form of either broad spots below each

free filament, or a uniform band of color at the basal 1/2 to 1/3 of the

cup. In subg. Eucharis, the pattern is of limited taxonomic

significance. Whether this pigmentation functions as nectar guides for

pollinating animals is unknown.

The stamens of most species of subg. Eucharis constrict distally

into a broadly subulate portion (> 1 mm wide for most of its length) of

varying length. Only in two species, E. astrophiala (Fig. 3) and E.

bouchei (in part), do the stamens constrict gradually from the rim of

the staminal cup to the apex of the filament. The free filaments of

Caliphruria are narrowly subulate (< 1 mm wide for most of their length,

Fig. 5). The free filaments of E. sanderi (subg. Heterocharis) are

narrowly subulate and slightly incurved. Those of E. anomala and E.

amazonica are broadly subulate.

Anthers of Eucharis are introrse, dehiscing longitudinally and

either dorsifixed or sub-basifixed in attachment. They are most









frequently oblong in shape, but are linear in subg. Heterocharis. At

anthesis, the anthers of Caliphruria and E. subg. Eucharis are erect,

but become versatile as they age. In subg. Heterocharis the anthers are

versatile at anthesis.



Gynoecium

Stigma and style. The flowers of almost all Eucharis and

Caliphruria species are protandrous. Stigma receptivity does not occur

until the second or third day following anthesis. In some cases, the
stigma does not fully expand until the perianth has begun to senesce.

The styles of Eucharis and Caliphruria are usually exserted beyond

the anthers, most frequently from 0.5-1 cm. In subg. Heterocharis, the

styles are somewhat assurgent away from the stamens, and are exserted

well over 1 cm past the anthers. In two species of subg. Eucharis, E.

castelnaeana and E. plicata, the style is included within the cup. In

the former species, autogamy seems to occur with regularity, and stigma

receptivity coincides with anthesis.

The stigma of Eucharis and Caliphruria (Fig. 6, 8-9, 11-12) is

obtusely triblobed. Trilobed stigmas are relatively rare in the

Pancratioidinae, and Urceolina, sister group to Eucharis and

Caliphruria, has a capitate, entire stigma. Traub and Moldenke (1949)

and Traub (1963) considered a trilobed or trifid stigma the ancestral

state in the Amaryllidaceae.

The stigmas of Eucharis and Caliphruria are papillate. The

papillae of Eucharis are unicellular (Fig. 7, 13-16), while those of

subg. Caliphruria (Fig. 10) are multicellular, consisting of both a

stalk cell and globose head cell. X Calicharis butcheri, putatively a









natural hybrid of E. sanderi and C. subedentata has the multicellular

stigmatic papillae (Fig. 17-18) characteristic of Caliphruria.

Heslop-Harrison and Shivanna (1977) characterized the stigmas of

Eucharis and Caliphruria as dry-type, and suggested a correlation

between this type of stigma morphology and sporophytic self-

incompatibility. According to a number of workers (Heslop-Harrison,

1976; Kress, 1983; Larsen, 1977), however, gametophytic incompatibility

is characteristic of monocots. At present, the incompatibility system

of Eucharis and Caliphruria, though apparently present, is unknown (see

Chapter X).

Ovary and ovules. The ovary of Eucharis and Caliphruria is

inferior and contains septal nectaries. It is green, with the exception

of two species, E. astrophiala and E. castelnaeana (subg. Eucharis) in

which the ovary is white at anthesis. Ovaries of Eucharis and

Caliphruria range from oblong-ellipsoid (subg. Heterocharis) to globose

or sub-globose (subg. Eucharis and Caliphruria). The ovary of subg.

Heterocharis is both trigonous and rostellate after senescence of the

perianth. Ovaries of Caliphruria and subg. Eucharis are non-rostellate

and smooth, with three exceptions: Eucharis bouchei var. bouchei, var.

darienensis, and E. cyaneosperma have a trigonous ovary at anthesis.

The ovules of Eucharis and Caliphruria are globose, anatropous,

and axile in placentation. Ovule number is quite variable throughout

both genera. Within limits, however, ovule number is characteristic of

species or species complexes. Subgenus Heterocharis has the largest

ovule number in Eucharis, generally 16-20 per locule, but occasionally

as low as 7 in E. sanderi (which otherwise has 16-20 throughout most of

its range) and 9-12 in E. amazonica. In both subg. Eucharis and









Caliphruria, ovules do not number more than 10 per locule. Eucharis

astrophiala, E. bouchei, E. bonplandii, E. cyaneopserma and E. ulei

characteristically have 2 ovules per locule, but rarely as many as 5.

In these species, there is a positive correlation between reduction in

flower number and ovule number.

Traub and Moldenke (1949) and Traub (1962, 1963) considered

numerous ovules an ancestral character in the Amaryllidaceae. In the

Pancratioidinae, an ovule number of ca. 20 per locule characterizes the

putatively ancestral complex of genera with typical, crateriform,
pancratioid floral morphology, heavy floral fragrance and well-developed

stamina cups (Meerow, 1985). Reduced ovule number is therefore likely

a derived character state.



Fruit and Seed

Fruit. The mature fruit of Eucharis and Caliphruria is a tri-

loculicidal capsule typical of the non-baccate fruited Amaryllidaceae.

In fruit, the pedicel elongates to 2 or more times its length at

anthesis. In Caliphruria and E. subg. Heterocharis (E. anomala), the

capsule is thin-walled and green, sometimes turning yellow or brown at

dehiscence. In subg. Eucharis, however, the capsule is leathery and

bright orange (Fig. 19), contrasting vividly with the shiny black or

blue seeds at dehiscence. It is probable, though unsubstantiated, that

the combination of fruit and seed color functions mimetically to attract

avian dispersal agents (sensu van der Pijl, 1982). This type of fruit

morphology is unique among neotropical Amaryllidaceae. There is a

single known exception to this characteristic fruit morphology in subg.

Eucharis. Eucharis castelnaeana (Fig. 20) produces a capsule much like









that of Caliphruria. The fruit of this species is often tardily

dehiscent, and sometimes abscises before opening, though the seeds

within are ripe. The infructescence of E. castelnaeana bends to the

ground (in all other species it remains erect), a habit noted in many

Crinum species (Hannibal, 1972). In this manner, an indehiscent fruit

might rot in contact with the substrate, thereby releasing the seeds.

Seed. Regardless of the number of ovules per locule in any

species of Eucharis and Caliphruria, all but a few abort as the fruit

matures. Generally 1-2 seeds are present per locule in mature capsules,

but as many as four have been observed.

The seed of both Eucharis and Caliphruria is usually globose or

ellipsoid and turgid, the consequence of copious endosperm and a high

moisture content. Left at room temperature, the seeds will shrink away

from the testa somewhat as moisture is lost, but are still capable of

germination in this condition. Long-term viability has not been tested.

The seed of subg. Eucharis (Fig. 21) is characteristically

ellipsoid, and has a shiny, smooth black (blue in E. cyaneosperma)

testa. The single exception so far known is again E. castelnaeana (Fig.

22). The seed of this species is wedge-shaped by compression in the

capsule, is less turgid than seeds of con-subgeneric species, and has a

dull, rugose testa. The seed of E. anomala (subg. Heterocharis) is

globose to very slightly compressed, and has a brown, slightly rugose

testa.

In Caliphruria, the seeds of only C. korsakoffii and C.

subedentata are known. Seeds of C. korsakoffii are globose, turgid,

and have a smooth, lustrous brown testa. Seed of C. subedentata is

slightly compressed, with a lustrous black, but rugose, testa.









Seed surface morphology (Fig. 23-28) does not reveal much

taxonomically useful information. The testa is alveolate in all species

examined. In E. bouchei var. dressleri (Fig. 24), abundant wax

extrusions are found across the surface.

The testa of Eucharis and Caliphruria seeds is composed of

phytomelan (Huber, 1969), a simple, largely inert, carbonaceous compound

characteristically present in the seed coat of non-baccate fruited

Amaryllidaceae (Huber, 1969; Darlgren and Clifford, 1982). Werker and

Fahn (1975) reported the occurrence of phenolic quinones in the
phytomelan layer of Pancratium seeds. In most species of Eucharis and

Caliphruria, the phytomelan layer is all that remains of the integuments

(Fig. 30, 36). In E. bouchei, however, there is an additional layer of

integument tissue, ca. five cells thick, interposed between the

phytomelan and the endosperm (Fig. 34). Whether this may be a

consequence of the tetraploid condition of this species is unknown.

Most of the seed body of Eucharis and Caliphruria is taken up by a

copious quantity of endosperm characterized by abundant transfer cells

(Fig. 35). At maturity, no remnants of the nucellus were observed.

Most workers (e.g., Baker, 1888; Traub, 1963; Hutchinson, 1959;

Dahlgren et al. 1985) have allied Eucharis and Caliphruria with

Hymenocallis, Eurycles and Calostemma (i.e., tribe Euchareae) on the

basis of "fleshy seeds." The latter three genera do indeed have fleshy,

bulbiform seeds that are sometimes viviparous, but they are not

homologous structures.

The large, green seed of Hymenocallis is unique in a number of

respects. The bulk of the seed body consists of two large, fleshy

integuments with a well-developed vascular system and abundant









chlorenchymous tissue (Rendle, 1901; Whitehead and Brown, 1940). The

embryo contains a large amount of stored starch (Whitehead and Brown,

1940). Whitehead and Brown (1940) characterized the seed, which does

not undergo any period of dormancy, as intermediate between true

vivipary and dormancy. Additionally, polyembryony has been observed

frequently in seeds of Hymenocallis (Bauml, 1979; Rendle, 1901; Traub,

1966).

The seeds of Calostemma and Eurycles superficially resemble seeds

of Hymenocallis, though they never achieve the size of the latter.

According to a much-overlooked review of bulbiform seeds in

Amaryllidaceae by Rendle (1901), the propagule of these two closely

related Australasian genera is not actually a true seed, but represents

an adventitious vegetative growth. After fertilization, at the chalazal

end of a normal ovule, adventitious shoot and root growth occur and a

true bulbil is formed. The integuments and the remnants of the nucellus

form the bulbil's outer coat.

The turgid seed of Eucharis and Caliphruria, despite a high

moisture content when first ripe, cannot be accurately described as

fleshy. This becomes evident if the seed is allowed to dehydrate

slightly at room temperature, and is most apparent in the hard seeds of

E. castelnaeana, which, at capsule dehiscence, are less turgid than

seeds of other species of subg. Eucharis. Seeds of Eucharis and

Caliphruria have a reduced integument, represented in most cases only by

the compressed phytomelan layer, and have never been observed to

germinate viviparously. Phytomelan is absent from the testa of the

pseudoseeds of Eurycles and Calostemma. It is present in only a single

species of Hymenocallis, H. quitoensis Herbert (and possibly H.









heliantha Ravenna), which has been segregated into a separate genus,

Lepidochiton Sealy (1937), on this basis.

Seeds of Pancratium are structurally most similar to those of

Eucharis and Caliphruria. Though variable in morphology (Werker and

Fahn, 1975), several species of Pancratium have a hard, turgid,

compressed seed body with copious endosperm (Meerow, unpubl. data;

Werker and Fahn, 1975). All species of Pancratium that I have examined

have a phytomelanous testa with an alveolate testa. Seeds of Eucharis

and Caliphruria do, however, have a higher moisture content than those
of Pancratium, all species of which occur in xeric to seasonally dry

habitats.




















Figure 4.1. Serial tranverse sections through the scape of Eucharis
castelnaeana (Schunke 14156, FLAS). p = proximal, m = medial, d =
distal.





60






























m
Ow
d











IN M








9 0 P
P^1^
















"a c






a 0
r*-




LL. 41
<. *'
* 10
-0
0u. I

*r .
CLa


0- <-0





=r,,
* .J


S- 0






*U






4- M N
0
LW
0 00


O to










0.

- aN) (
*a* *=








'I)
-C LL
L.





L.
CO







*r-
&.(0 0
Q)**> -
Q.( 0_
v) u-
CULO
1)L




0>














E
U


VL


















Figures 4.3-4.5. Androecial morphology of Eucharis and Caliphruria
species. 3. E. astrophiala (Madison 3792, SEL). 4. E. amazonica
(Schunke 14177, FLAS. 5. C. subedentata (Meerow 110-, FLAS).


























V..r
4. ; #.>


1CM
I----<



















Figures 4.6-4.18. SEM photomicrographs of Eucharis and Caliphruria
stigmas. 6-7. E. astrophiala (Meerow 1111, FLAS). 8. E. plicata
(Plowman 13941, FLAS). 9-10 C. Csub-edentat (Meerow 1152). 11.
C. korsakof-iT (Meerow 1Q96, FTAS). 12-1 E. X granifTTora
TMeerow f1127,FL 3T-. 1TT. anomala (Meerow' 1141, FLAS).- 5. E.
sanderiT (Cuatrecasas 16350, FT. 16. E. amazonicia (Schunke 14177,
FLAS). 17-18. X Calicha's butcheri,,Meerow 111_0, FLAS). 17I
scales = 50 um.



































".. 1.- 14.
g.^,


GbCI~jiaL^2^ ^I' -


S:7iog, A



















Figures 4.19-4.22. Fruits and seeds of Eucharis subg. Eucharis. 19-20.
Mature capsules. 19. E. formosa (Schunke 14174, FLAS). 20. E.
castelnaeana (Schunke T417, FAS). 21-22. 7 needs. 21. E.
boucnei var. bouchei (Meerow 1125, FLAS). 22. E. castelnTe ana
(Schunke 14157"T1S).








































22


















Figures 4.23-4.28. SEM photomicrographs of Eucharis and Caliphruria
seed surfaces. 23. E. astrophiala (Meerow 1111, FLAS). 24. E.
bouchei var. dressleri (Meerow 1107, FLA)T. 2. E. formosa -
(Meerow 1103) 267. ~ castelnaeana (Schunke 14155, TFAS). 27.
C. korsaloFii (Meerow 1096, FLAS)8.28. C. subeentata (Meerow
T152, FLAS).




70





















25 126
4 JS '




















Figures 4.29-4.37. Photomicrographs of Eucharis and Caliphruria seed
anatomy. 29-33. C. korsakoffii (Meerow 1096, FLAS). 29.
Longitudinal sectTon through whole seed. 30". Transverse section
through testa and part of endosperm. 31. Longitudinal section
through radicle of embryo. 32. Longitudinal section through apex
of embryo. 33. Longitudinal section through vascular initial of
embryo. Scale = 40 Aim. 34-35. E. bouchei var. bouchei (Meerow
1125, FLAS). 34. Transverse section through testa. Note several
layers of additional integument cells below outer phytomelan
layer. 35. Endosperm. Note transfer tissue with pitted walls and
plasmodesmata. 36-37. E. castelnaeana (Schunke 14156, FLAS). 36.
Transverse section through test and part of endosperm. 37.
Tranverse section through embryo. All scales = 100 jm unless
otherwise indicated, em = embryo, en = endosperm, t = test.





72
















eem

-TI- W-4.

je
















32. 33 34 3
...



;Y. .
.... .,;% -em -, .
.. o ,. ., .,














CHAPTER V
POLLEN MORPHOLOGY



Materials and Methods



Scanning Electron Microscopy (SEM)

Fresh, dehisced anthers were removed from living collections,

fixed in FAA, and pollen extracted. Pollen from herbarium specimens was

treated according to the process of Lynch and Webster (1975). Samples

were treated for and examined with SEM as described for leaf surfaces in

Chapter III. Measurements of muri and lumina were derived from SEM

photomicrographs.



Transmission Electron Microscopy (TEM)

Pollen grains were fixed for 12 hr in 3% glutaraldehyde in 0.1 M

Na-cacodylate at pH 7.4, washed three times for 10 min with 7.5%

solution of sucrose in 0.1 M Na-cacodylate at pH 7.4, post-fixed for 1

hr in 2% Os04 in 0.1 M Na-cacodylate, washed as above three times for 10

min, and brought through an EtOH dehydration series. Dehydrated pollen

was placed through two pure propylene oxide baths, then placed in 1:1

propylene oxide:epon for 1 hr, 1:2 propylene oxide:epon for 12 hr, and

pure epon for 2-3 hr. Pollen was polymerized for 48 hr at 600 C,

sectioned, and viewed on a Zeiss 10A electron microscope at 80 kv.










Light Microscopy

Pollen size measurements were averaged for twenty grains examined

with a Nikon Lapophot photomicroscope.


Pollen Viability

Pollen was stained with Alexander's (1969) stain for 24 hrs at 500

C. Percentages given are based on the number of grains staining from a

200 grain sample.


Statistical analysis

Correlations of pollen size with style length were performed with

SAS release 5.08 on the Northeast Regional Data Center (NERDC) of the

University of Florida.


Terminology

Terminology follows Erdtman (1969) and Walker and Doyle (1975).



Results



Pollen grains of all species of Eucharis and Caliphruria (Fig. 1-

19) are boat-shaped elliptic, monosulcate, heteropolar, and bilateral in

symmetry. The germination furrow (sulcus) runs the length of the

presumed distal face of the grain (Fig. 12, 15). Exine sculpturing is

semi-tectate-columellate and reticulate in all species examined (Fig. 1-

19), composed of a network of muri reticulumm walls) and lumina

(intervening gaps).









Pollen Grain Size (Table 1)

Pollen grain size is quite variable in Eucharis and Caliphruria ,

and a notable size class (sensu Walker and Doyle, 1975) differential

occurs between Eucharis and Caliphruria. Pollen of Eucharis falls into

the large size class of Walker and Doyle (longest equatorial diameter

50-100 pm). Pollen of Eucharis has average longest equatorial diameters

greater than 60 ,um, with two exceptions: E. castelnaeana and E. plicata

subsp. brevidentata. Pollen of Caliphruria falls into the medium size

class of Walker and Doyle (1975) with average longest equatorial

diameters of near 50 pm.

The greatest number of species of Eucharis have pollen grains with

longest equatorial diameters between 65 and 75/im. Eucharis astrophiala

(subg. Eucharis) has the largest pollen grains in the genus, with

longest equatorial diameters of 83-86/um.

Polar diameter of pollen of Eucharis ranges from (39-) 45-60.6,/m.

Polar diameter less than 401um is rare in these subgenera. Polar

diameter of pollen of Caliphruria is always less than 40 um.

Considerable infraspecific variation pollen size is evident in

some species of Eucharis (Table 1). Eucharis formosa is a wide-ranging

and morphologially variable species (see Chapter XII). Longest average

equatorial diameter among the populations sampled of this species shows

a 12.7% difference between the smallest and largest value. The two

subspecies of E. plicata show a 15% differential in pollen size. Other

species are much more uniform in pollen grain size. Eucharis

astrophiala is a narrow endemic restricted to western Ecuador with

distinctive leaf and androecial morphology that is consistent among all

populations. Three populations of this species sampled show only a 3.8%









difference. Eucharis bouchei, a tetraploid species also of limited

distribution, but highly polymorphic, shows only a 2.4% difference

between the largest and smallest values.

The smallest pollen grains in Eucharis and Caliphruria are found

in species with the smallest flowers (Table 1), i.e., all species of

Caliphruria and, in E. subg. Eucharis, E. castelnaeana. Nonetheless,

one of the largest flowered species, E. sanderi (subg. Heterocharis) has

small pollen grains relative to other large-flowered species. The

largest pollen grains in the genus are found in E. astrophiala, a
species at the smaller end of flower size range in the genus. Since

style length is directly correlated with perianth size in Eucharis and

Caliphruria, style length was tested for correlation with longest

equatorial diameter of pollen of species in Table 1. Pearson

correlation coefficient for style length with pollen size of 29 Eucharis

and Caliphruria collections representing 16 species was only 0.379, and

therefore not significant (significance is indicated by proximity of

value to 1.000).



Exine Sculpturing (Fig. 1-19, Table 1)

The semi-tectate, reticulate exine sculpturing pattern of Eucharis

and Caliphruria may be subdivided into three classes on the basis of

lumia width. The first, designated Type 1 in Table 1, is characteristic

of most species of Eucharis (Fig. 1-11, 13, 15-16).. The reticulum of

Type 1 exine is coarse, with largest lumina widths equal to or greater

than 5/um. Type 1 exine can be further subdivided on the basis of muri

width. In Type 1-A (Fig. 3-11, 13, 15-16), the muri are equal to or

greater than 1,um wide. This is the most common exine morphology of









Eucharis. In Type 1-B exine, the muri are less than 1/im wide. This is

characteristic of a single species of subg. Eucharis: E. astrophiala

(muri ca.0.6 jum wide, Fig. 1-2).

In Type 2 exine, lumina are 2-3 jum wide, and a marked reduction in

reticulum coarseness occurs at the meridional faces of the grain. Only

two species of Eucharis have Type 2 morphology, E. oxyandra (subg.

Eucharis, Fig. 12), E. sanderi (subg. Heterocharis, Fig. 14), and one

species of Caliphruria, C. korsakoffi (Fig. 19). Width of the muri,

however, is variable among the species with Type 2 sculpturing, ranging

from less than 0.4jum in E. oxyandra, to ca.0.75jum wide in E. sanderi,

and ca. 1 um wide in C. korsakoffi.

Type 3 exine sculpturing is only characteristic of the Colombian

species of Caliphruria (Fig. 17-18). Type 3 sculpturing is finely

reticulate with lumina only 1-2 pm wide, and the muri 0.5-0.6 pm wide.

As in Type 1 sculpturing, the reticulum is predominantly consistent in

coarseness throughout the grain surface.


Pollen Wall Ultrastructure (Fig. 20-31)

Eucharis and Caliphruria pollen grains are remarkably uniform in

their exine stratification patterns. They are completely ektexinous in

composition. The columellae arise from a thin foot-layer (usually ca.2

Aim thick), and the intine is as thick or thicker than the exine. The

tectum is quite fragile, and usually ca.5 um thick. No channelling is

apparent in either the exine or intine.








Discussion



Large, boat-shaped-elliptic, monosulcate pollen grains with

reticulate exine morphology are the most common type of pollen found in

the Amaryllidaceae (Erdtman, 1952; Meerow and Dehgan, 1985; Walker and

Doyle, 1975). Similar morphology has been reported for many Liliaceae

sensu lato (Erdtman, 1952; Walker and Doyle, 1975; Zavada, 1983), and

conforms to the fossil form genus Liliacidites Couper, one of the major

angiosperm pollen types described from early Cretaceous deposits (Doyle,
1973; Walker and Walker, 1984). This type of pollen morphology appears

to be basic to the monocotyledonous orders in general (Doyle, 1973).

Among the Amaryllidaceae, only one group of genera show a radical

departure from this basic pollen morphology. Crinum and its allies

[tribes Crineae (Pax) Traub and Strumarieae Salisb. sensu Traub (1963)],

all have bisulculate pollen and spinulose exine sculpturing (Dahlgren

and Clifford, 1982; Erdtman, 1952; Nordal et al., 1977; Meerow, unpubl.

data). With the exception of Crinum, these genera are restricted to

Africa, many of them endemic to South Africa. In a remarkable example

of convergence, Donoghue (1985) reported a similar divergence in

Caprifoliaceae.

The Type 1 exine morphology that is characteristic of most

Eucharis pollen seems to have phylogenetic significance within

"infrafamily" Pancratioidinae (Meerow, 1985; Meerow and Dehgan, 1985).

All or some of the species of each of the genera with putatively

primitive pancratioid floral morphology (i.e. Eucharis, Hymenocallis,

Pamianthe Stapf, Pancratium, and Paramongaia Velarde) have large to very

large, coarsely reticulate pollen. The pollen of related genera with









divergent floral morphology shows reduction trends in both size and

reticulum coarseness (Meerow, 1985; Meerow and Dehgan, 1985).

Reduction in size and reticulum coarseness have been considered

evolutionary trends for angiosperm pollen in general (Walker and Doyle,

1975). Colombian species of Caliphruria (Fig. 17-18) show the greatest

degree of divergence for these pollen characters in comparison with

Eucharis.

The differentiation of the reticulum into coarse and fine areas,

characteristic of species with Type 2 exine, is restricted to monocot

pollen (Doyle, 1973; Walker and Walker, 1984), and has been observed in

some Liliacidites pollen from the early Cretaceous (Walker and Walker,

1984). The evolutionary polarity of this character is unclear, however.

Meerow and Dehgan (1985) described a transformation series from

auriculate pollen through dimorphic reticulum to homogeneous reticulum

among the subgenera of Hymenocallis (sensu Traub, 1962, 1980), which

would suggest that the homogeneous reticulum is an advanced character

state. The three species with dimorphic exine sculpturing (E.

oxyandra, Fig. 12; E. sanderi, Fig. 14; and C. korsakoffi, Fig. 19)

each represent isolated taxa of their respective genus or subgenera (see

Chapter XI). The dimorphic reticulum in these three species may thus be

symplesiomorphous. On the other hand, each of three species differ in

muri width, thus the Type 2 exine morphology may have had an

independent, and thus derived, origin in each of the three.

In width of both muri and lumina, the pollen of E. oxyandra (Fig.

12) resembles that of Urceolina, sister group to Eucharis, though pollen

of the latter genus fits the medium size class of Walker and Doyle, and

does not exhibit a substantial differentiation of the reticulum into









coarse and fine areas (Fig. 1 in Chapter XI). Eucharis oxyandra is a

problematic species morphologically as well, with certain characters of

intermediacy between Eucharis and Urceolina, particularly in androecial

morphology (see Chapter XII). I have suggested that E. oxyandra may

represent a relict taxon related to the ancestor of Urceolina, or a

possible intergeneric hybrid (see Chapter XII), but this species is at

present too poorly known to confirm any of several hypotheses concerning

its origins.

Zavada (1984) associates reticulate exine sculpturing with
sporphytic self-incompatability (SSI). Though the SI system of Eucharis

and Caliphruria, if present, is unknown, two morphological characters of

the genus-- pollen sculpturing, and stigma type (Heslop-Harrison and

Shivanna, 1979)-- have been correlated with sporophytic SI, despite the

fact that only gametophytic SI has been reported for monocots (Heslop-

Harrison, 1976; Kress, 1981; Larsen, 1977).

Kress and Stone (1982) reviewed pollen wall ultrastructure of

monocots. The lack of endexine in the pollen grain wall appears to be a

virtually universal characteristic of monocot pollen. The thin foot-

layer and columellate structure of the exine found in Eucharis and

Caliphruria is common to all other genera of the Pancratioidinae that I

have examined (Meerow, unpubl. data; Meerow and Dehgan, 1985), and may

be basic to the Liliflorae in general (Doyle, 1973; Walker and Walker,

1984). The pattern of exine stratification in the pancratioid

Amaryllidaceae thus appears highly conserved.

Pollen grain size and style length has been correlated in some

investigations (Baker and Baker, 1979; Lee, 1978; Plitmann and Levin,

1983; Schnack and Covas, 1945; Taylor and Levin, 1975) but not in others









(Cruden and Miller-Ward, 1981; Darwin, 1896; Germeraad et al., 1968;

Ganders, 1979; Hammer, 1978). Cruden and Lyon (1985) observed that all

studies which showed a strong correlation involved related species,

while non-correlating studies involved unrelated taxa. They tested

correlations between both style length and stigma depth (an

approximation of the distance a pollen tube must grow to reach exogenous

resources in the transmission tissue of the style) and pollen grain

volume among species of several genera in several families. Cruden and

Lyon concluded that style length has little correlation with pollen

size, while stigma depth was highly correlated with style length. Where

style length and pollen grain volume do correlate, i.e., among related

species, they suggest that phylogeny, rather than function, is

represented. They further conclude that pollen grains need not contain

sufficient endogenous resources to reach the ovules, but only enough for

pollen tubes to grow through the stigma and reach exogenous substances

in the stylar transmission tissue.

In Eucharis and Caliphruria as a whole, little correlation between

style length and pollen grain size (as represented by longest equatorial

diameter, rather than volume) is evident (Table 1). Stigma depth, in so

far as I understand Cruden and Lyon's determination of this measure,

does not seem to vary appreciably among species of Eucharis. The stigma

of E. astrophiala, the species with the largest pollen grains in

Eucharis, is no larger or "deeper" than that of E. plicata, the species

of subg. Eucharis with the smallest pollen grain.









Conclusions



In characteristics of pollen grain size (medium size class), and

exine sculpturing (Type 3), Caliphruria shows the greatest degree of

divergence from the putatively ancestral, large, coarsely reticulate

pollen grain characteristic of most species of Eucharis. The Type 1

exine sculpturing of Eucharis shows relationship to the pollen

morphology of other genera of infrafamily Pancratioidinae with similar

floral morphology, i.e., Hymenocallis, Pamianthe, Pancratium and
Paramongaia (Meerow, 1985; Meerow and Dehgan, 1985).

Pollen grain size in Eucharis does not demonstrate any obvious

correlation with flower size (= style length). The large amount of

variation in pollen grain size in a few species of Eucharis may suggest

that this character, under certain conditions, is subject to as much

infraspecific variation as characters of vegetative and floral

morphology.
















UC
0
E





C.




C-






S-



u.
4)
u






Q..

0.

I-









e=,,
0












L
0








- 0

0
r- 4-3





- s





O
0 *-
3



o -o
v-











i-a
I-3













u-
0





i- 0 1-
Q. 1


T--f








t0


Z



LJ -
-I E

CD




>-
o,








LUJ

.W-








PI-

Ljj





CD
U 0U





























0

















0

i-


I I
-4 -4


C% 0 C) 4 C mn c .- o0
ror~a 0r> (3) LO f mm un ra- m C ) o m-
0 CO CO C') *
'. -1 C'j CO o
*D *
co +I c +I o1 +1 +-I +' 0+ +| + + +| + o +1




m ) o C) 04 0 0 m -4
u-co Co- m OC oo o0 or 0 L-
o0 to U) rf. .* .0
*: *. *M *.t ** *.-4
0 CO 0 0 m C) O m
t0 +| U +| ko + O m +| + + d +1 +I




-> CM

-i o-







000
O rOO CO
,--II c c-4


i V- o 0 0


S- .^ U I- fi E S- S- S-
a(1) 0 3a a 0




a) r a)- to 0.















UJ iiI UJ I I JI J LI
go "0i

LI cc0 0
cFI ) -r 0 -0[,O

ui c P L

























A- 0
E


S* t- '- o nf o o0 *
*cNJ *-I *C I *CN *M *4 *C
SO CM Im tu L u om m
10+| + ++|I + I( % + I L +I "o +| ko +| 10 +1 r-- +1






*-I *4 *C *CM .. *C% C %J *-4







oco



o Lo o ~ o o o
(0 0 0 0 0
d-





-- -(2.) (2.)C)

3 3 1 3 3 3
0 0 0 E 0 0 0 0
SL L ( 3 S..
i aw -0 41 4 a ) a )
a) w U U U w wU w
X: X: I= r V S M X


(z



,-0J









LUt-






0 0


















=
I- -i







- 0-




cc"
< u j


(0
41 0


>1 0
U E
SI O
U| Lij







85















I-
w O L CO O

SM J M
Ln n C t cLA c' ( -1 3-
>-
,-








,, 0- I I I
UJ -g



-1

i-) a: a- O C Co o n a


0- 0 L 0



XC z c *cO ccO n *
d. +0 r +I O++I +I +1 1o+1 M+I -It +1







a-.< 1^+| +1 i +l +1 +| +1 +1




U, m E U 0 0 -


u- m 3 0
( 4-) (*) o ) 4) w



c-u u. r- o W 0
4 -i 4 I o l S


0 to u u u > i
0 CL U 3 i <




03
F'- I- -.

+U 4l l4 .




I-. 0 < (0 00 0.






















I-
CD



-

>-
--


0
a.



*4-
0





U-)




cr
4.3
cr



e-





I 41



cd C- m ,,









LO CO Ln C) O" C
0



























4 & .)C' .O 0 M

LX)+ +I + C +) +I I-
C,
0
U
4I-






















lI C
3.-I CMJ mM W .*-

9-

















N -
r. o.9
T c- r .0



L L9 L 4 r U)0+ a)+ 0 U)


























0 0 0
o +| I-
r. C l .I r,-) .. (


0I LOG LllO 0 LI 0 0o






O c c
f0

I0 0

al 0- 4I M-


0 0
*S- S 0 L. 0 0 C *
V). o t 4) St S- Cx

0 0 0 0 00 0
IV 0 U)

0U 0 0
. I- 4.)4


* 0

0 0a '- 0 0 o .c
3 U 0 4c- 4 4- 4.)
U) *r 0 r- *t 0 C> .
C i L I- I u 0I L 4-I 3


0 2 C 0O 0. 0 0 u i- 0u 0o
Ld WI 1I WIJ | U-'I U|I 0 .


(a
ZLUJ
x>-
Ui-



,--I

<,) I-- A
uJ 0 UJ E

-jcr 9=








-UJ E

P-4

















s-




















Figures 5.1-5.6. SEM photomicrographs of Eucharis pollen grains. 1-2.
E. astrophiala (Madison 3792, SEL). 1. Whole grain, proximal
polar view. 2. Exine sculpturing. 3-6. Whole grains, proximal
polar views. 3. E. bonplandii (Bauml 686, HUNT). 4.. E. bouchei
var. dressleri (MeTerow 1107, FLAJT-. 5 E. candida (Asplu-nd
19120, S). 6E. EcasteTniaeana (Schunke 1TF156W-T ~ ). All scales
= ca. 5 um.



























^LiL4-


L"1


qmmww 4PP

fcOA
ia Aww


4^k-7




















Figures 5.7-5.12. SEM photomicrographs of Eucharis pollen grains. 7-8.
Whole grains, proximal polar view. 7. E. corynandra (Ravenna
2090, K). 8. E. cyaneosperma (Seibert 7145, US). 9-1I0.T .
Tormosa (MeeroW 113, FLAS). 9. Whole grain, proximal polar view.
10. Exine sculptu'Tring. 11-12. Whole grains, proximal polar view.
11. E. plicata (Meerow 1025, FLAS). 12. E. oxyandra (Hutchison et
al. S98J, UL), ob 11que Tr~tal polar view.- ATIT sles = ca 5 )Um.





90





















L 4 p






















Weir. f. s-
h 1 1, I.













12



















Figures 5.13-5.19. SEM photomicrographs of Eucharis and Caliphruria
pollen grains. 13-17. Whole grains. 13. E. amazonica (Asplund
13214, S), proximal polar view. 14. E. sanderi (Killip 35401,
TS), oblique lateral longitudinal vie. 1. X Calicharis butcheri
(Meerow 1110, FLAS), oblique distal polar view. 16. E. X
grandiflora (Meerow 1127, FLAS), lateral longitudinal view. 17-
18. C. subedentata (ex short s. n., K). 17. Distal polar view.
18. Exine sculpturing. 19. T. Torsakoffi (Meerow 1096, FLAS).
All scales = ca 5 um.












^M^J"=
tt


let
.



Pd.-
&&<