Morphological and Anatomical Considerations of the Grass

59 Pages·2002·8.11 MB·English

S M I T H S O N I A N C O N T R I B U T I O N S T O B O T A N Y N U M B E R 1 1
Morphological and Anatomical
Considerations of the
Grass Subfamily Bambusoideae
Based on the
New Genus Maclurolyra
Cleofe/ E. Calderon
and Thomas R. Soderstrom
SMITHSONIAN INSTITUTION PRESS
City of Washington
1973
A B S T R A C T
Caldeoh, Cleoee E., and Thomas R. Soderstrom. Morphological and Anatomi-
cal Considerations of the Grass Subfamily Bambusoideae Based on the New
Genus Maclurolyra. Smithsonian Contributions to Botany, number 11, 55 pages,
24 figures, 1973.-hPaclurolyra tecta, a new genus of grasses from Panama, is
described. Features of its leaf anatomy and epidermis, seedlings, inflorescence
morphology, floral structure, and cytology, indicate that it is a member of the
tribe Olyreae of the subfamily Bambusoideae. A description is given of the
“bambusoid” type of leaf anatomy, as well as comments on the vascular bundle
sheaths in grasses, and chloroplast structure and photosynthetic pathways as new
criteria in grass taxonomy. The phylogenetic position of Maclurolyra is discussed
and a list of genera comprising the Bambusoideae is presented.
OFFICIAPLU BLICATION DATE is handstamped in a limited number of initial copies and is recorded
in the Institution’s annual report, .“tr~ithsotiinn Year. SI PRFSSX uliBER 4782. SERIES COVER
DESIGN: Leaf clearing from the katsura tree Cercidiphjllum jafionicutn Siebold and Zuccarini.
Library of Congress Cataloging in Publication Data
Calderbn, Cleofe E.
Morphological and anatomical consideration5 of the grass subfamily Bambusoideae based on
the new genus Maclurolyra.
(Smithsonian contributions to botan), no. 11)
Bibliography: p.
1. Maclurolyra. 2. Bamboo. 3. Botanj-Morphology. 4. Botany-Anatomy. 5. Bamboo-Panama.
I. Soderstrom, Thomas R., joint author. 11. Title. 111. Series: Smithsonian Institution.
Smithsonian contributions to botany, no. 11
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Contents
Page
Introduction .................. .................................. 1
Acknowledgments . . . . . . . ...................... 5
Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
iMacluiolyia tecta, new genus and species ............................... 6
Seedling . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Torsion of the Leaf and Inflorescence . . . . . . .
Morphology of the Inflorescence ................ ................... 15
Floral Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Lodicules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2.0
S taminodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21
Gynoecium , , , . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2 2
Cytology . . . . . . ............................................ 27
* , , ,
Starch Grains ..... ...............................
Leaf Anatomy . ......................... 28
, , , ,
The Epidermises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
The Transverse Section of the Lamina . . . . . . . . . . . .
Olyroid Type of Siliceous Cell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3 6
The Bambusoid Type of Leaf Anatomy ......................... . . 36
,
The Vascular Bundle Sheath in Grasses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Chloroplast Structure and Photosynthetic Pathways ...................... 40
Phylogenetic Position of Macluiolyra . . . . . . . . . . . . . . . . . ........ 44
Literature Cited ............ ................................ 46
Appendix 1 : Genera of the Subfamily Bambusoideae Aescherson and
Graebner . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Appendix 2: List of Material Studied ............................ 52
Index to Grass Genera and Species . . ......................... 54
iii

Morphological and Anatomical
Considerations of the
Grass Subfamily Bambusoideae
Based on the
New Genus Maclurolyra
Cleofe' E. Calderon
J
and Thoma s R. Soderstrom
Introduction opportunity to reach areas of virgin forest previ-
ously inaccessible in Panama and Maclurolyra is
Accounts of the great diversity of species that
only one of many new plants found in recent years
are found in tropical rain forests are common,
in such forests.
but mostly these are based on inventories of the
Santa Rita is an area of primary forest situated
trees that are the conspicuous element of this
on an undulating terrain, which is traversed by
biome. Less attention has been paid to the
numerous streams and creeks, and ranges in eleva-
herbaceous vegetation of the understory where the
tion from about 200 to 450 meters. From April
decrease in illumination, change in light quality,
through December heal J rains occur almost daily.
and relati1 ely high humidity create conditions
Recent collections made in late October and early
which are hostile to most plants. The majority of
Noleniber, at the peak of the rainy season, showed
herbaceous plants that thrive under such condi-
all plants to be in flower, with some in fruit, and
tions in the rain forests of tropical America belong
with the presence of many seedlings around the
to relatively few families-Araceae, Bromeliaceae,
parent clumps. The type-collection was made in
Commelinaceae, Alarantaceae, Musaceae, Orchida-
earl) hlaich, at the end of the dry season. At
ceae, Zingiberaceae. Although they do not consti-
this time the plants were in flower and many
tute a significant part of this understory vegetation,
old inflorescences were also found. Apparently
either in number of species or in individuals, rep-
Mnclurolyra is in flower throughout the year with
resentatives of the grass family also occur here.
the most profuse flowering occurring during the
The new genus that we are describing here
rainy season.
occurs in Panama where it has so far been located
Plants of Moclzirolyra grow in small clumps
in forests of two areas-Santa Rita, on the Atlantic
scattered throughout the forest, usually around or
slope, and Cerro Jefe, on the Pacific slope (Figure
near trees or under plants of larger size. In some
1). Lumbering trails have afforded botanists the
places they grow in association with grasses of the
related genus, Cryptochloa (Figure 2a,c). All of
Cleofe E. Caldeidn and Thomas R. Soderstrom, Department
of Botany, Sattonal Museum of Natural History, Smithsonian these are found in reddish soil and apparently
Institution, Washington, D.C.2 0560. thrive only where it is shaded and humid. Plants
1
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY
FIGURE1 .-Map of Panama showing localities where Maclurolyra tecta has been collected:
[SR=Santa Rita, CJZCerro Jefe, cross-hatched area=Canal Zone].
of Maclurolyra left exposed in nearby cut.over morphology and anatomy of members of this tribe
areas of forest were found to be stunted and with see Calderon and Soderstrom, 1967.) Recently we
few, small, poorly developed inflorescences-pre- have discussed the pollination biology of some
sumably in response to the intense illumination grasses of the Olyreae, with comments on the
and decrease in humidity. Maclurolyra was found relationships of some of these herbaceous grasses to
also, but in less abundance, in some areas of Cerro the woody bamboos, all of which we refer to in a
Jefe, a forest region on the Pacific slope of Panama. general way as “bambusoid grasses” (Soderstrom
The higher elevation (ca. 800 meters), where and Calderh, 1971).
cooler and less humid conditions are encountered Our studies on the new genus have been made
than at Santa Rita, possibly accounts for the re- from the standpoint of its morphology and
duction in numbers of plants. anatomy, with a view not only to elucidating its
Studies of the morphology and anatomy of this systematic position within the grass family, but
peculiar new genus have revealed that it is related with the objective of clarifying and defining more
to Olyra and grasses of the tribe Olyreae (“olyroid precisely the “bambusoid type” of leaf anatomy.
grasses”), which tribe we include in the subfamily This allows us the opportunity to present some
Bambusoideae, as had Roshevitz (1946) and general considerations on the morphology, anatomy,
Parodi (1961). (For a brief account of the and taxonomy of the subfamily Bambusoideae,
NUMBER 11 3
FIGUREP .--Maclurolyra tecta in the field (Santa Rita forest, Panama): A, Mature plant (indi-
cated by arrow) growing in association with plants of Cryptochloa; B, close-up of adult plant;
c, close-up of adult plant with a plant of Cryptochloa in the background.
including remarks on the tribe Olyreae. We have colleague, Floyd A. McClure (1897-1970). We are
felt it useful to present at the end of this report a indebted to him for the countless hours of con-
list of all genera which we consider to be members sultation over the years regarding problems of
of the subfamily Bambusoideae (Appendix 1). bamboo morphology. He became familiar with the
The genus is named in honor of our late new genus as we studied it and shared with us, in
FIGURE3 .-InHorescence of dlnclurolyin tecta in the field (Santa Rita forest, Panama) : A, Newly
emerging inflorescence as seen from abo1e; n, inflorescence in front of the blade; C, inflorescence
beginning to twist; D, inHoi.escence at a late1 stage in back of the blade; E, old inflorescence
bent downward in back of a blade (see arrow), and one still in front of the blade; F, inflores-
cence axis bent downward (see arrow).
SUMBER 11
its interpretation, the wisdom he had acquired institutions. We would like to extend our thank3
during a lifetime dexoted to the study of bamboo. to Dr. C. R. Metcalfe (Jodrell Laboratory, Royal
The specific name derives from the Latin word Botanic Gardens, Kew, England) for his review
for “cover,” in allusion to the uppermost blade on and criticisms of the anatomical studies of the new
the flowering culm rvhich covers the inflorescence genus; and at the herbarium of the Royal Botanic
in umbrella-like fashion, shielding the flowers from Gardens, Kew, to the renowned agrostologist, Dr.
the rain (Figure 3). C. E. Hubbard, for his taxonomic suggestions. The
\\'bile lumbering trails such as those at Santa moi phological discussions concerning grasses, and
Rita allow us access to such genera as iWaclurolyra, particularly the inflorescence of the new genus,
these trails also signal the imminent destruction were made Tvith specialists at the Universitat Mainz,
of these same forests. The actual site of the type- LYest Germany, and particular acknowledgment is
collection of Maclzirolpra-primar) forest in 1968- made to Prof. Dr. H. Weber (Director, IInnstsittiuttu tf ufurr
was revisited in 1971 and found already cleared-a Spezielle Botanik) and his colleagues at the same
part of the ecosystem that took so long to evolve Universit), Profs. Drs. D. Hart1 and S. Vogel. We
destroyed forever. It is incumbent upon us, as ivould like to express our very deep gratitude to
biologists, to encourage that parts of these forests the eminent morphologist, Prof. Dr. Wilhem Troll,
be protected so that in the future plants of genera also of hlainz, for the numerous hours he devoted
such as i+fuclziro/yru can still be found in their to Calder6n in his laboratory, discussing the
natural habitat rather than in herbaria as mere inflorescence morphology of Maclurolyra. His
dried records of the past. advice, suggestions, and encouragement to continue
.~CKNO~\’LEDC1MENTS.-wef eel our deepest grati- in the difficult study of grass inflorescence morphol-
tude to our major professors who were responsible ogy, are all deeply appreciated. TYe are grateful
for our basic training in agrostology, Professor John as well for the advice given by Dr. H. J. Conert
R. Reeder (Laramie, T\”yoming) and the late Profes- (Natur-Museum und Forschungs-Institut Sencken-
or Ingenieio Lorenzo R. Parodi (Buenos Aires). berg, Frankfurt), Dr. H. Jacques-Felix (Museum
The present study was possible only because of National d’Histoire Naturelle, Laboratoire de
the support and facilities mofafenrye di nbsyt itu- Phanerogamie, Paris), Dr. F. Bugnon (Faculte des
tions and offices, and the personal assistance, Sciences de Dijon, France) , Dr. G. Bocquet (Institut
cooperation, and advice rendered to us by colleagues fur Spezielle Botanik, Eidg. Technische Hochschulle,
in the United States, Latin America, Europe, India, Zurich), and Dr. E. Mora-Osejo (Instituto de
and Ceylon. Primary credit is to be given to the Ciencias Katurales, Bogotzi). \Ye would like to
Smithsonian Institution, LVashington, D.C., for thank Dr. V. Puri (Meerut University, Meerut,
grants from the Smithsonian Research Foundation India) for his advice on embryological problems in
to the junior author which have provided for the the Gramineae and his generosity in allowing the
laboratory studies to be carried out. Support for use of his laboratory and facilities in India. Dr.
Calder6n to traLel to Central and South America Pierre Morisset (Universitie Laval, Quebec) was
during 1967-1968 Tvas provided by the Smithsonian’s kind enough to study the chromosomes of the new
Office of Systematics and Office of Ecology, and for genus and pro\ ide the photograph which appears
this trip a traiel grant was awarded by the Office in Figure 14d.
of Scientific Affairs, Organization of American IVe are grateful to llrs. Nina Smith (Hunt Bo-
States. The field work in Panama, during which tanical Library, Pittsburgh), who assisted in the
period the new genus was collected, was possible transliterations of the Russian titles and in trans-
onl) because of the assistance given by Dr. Robert lating passages from various Russian and German
L. Dressler (Smithsonian Tropical Research In- papers.
stitute, Balboa, Canal Zone) to whom we are \Ire appreciate the suggestions and assistance
especially indebted. A grant from the Smithsonian’s provided by several colleagues at our own institu-
Office of International Activities allowed Calder6n tion-Dr. E. S. Ayensu, Dr. Jose Cuatrecasas, Dr.
to spend some time in Europe on the way to India, Mason E. Hale, Jr., the late Mr, Conrad V. Morton,
and provided the opportunity to discuss matters Dr. Lyman B. Smith and Dr. William L. Stern.
relating to the new genus with specialists in various TVe were fortunate to have the habit drawings
6 ShIITHSOA‘IAN CONTRIBUTIONS TO BOTANY
pi epared by our illustrator, Mrs. Gesina Berendina the material needs no further transferring after
Threlkeld (GBT) of Delta, Alaska, and a few of fixing in the field. Flowers treated in this way are
the sketches by Mr. Christopher Reinecke (CR) of the most suitable for studies under phase contrast
SVashington, D.C. and polarized light. Flowers so treated were used
MATERIALASN D MEmoDs.-Plants were collected in the studies of the venation of glumes, lemmas,
in the field and herbarium specimens prepared in and paleas, and the vascular traces of the gynoecium.
the usual fashion by placing them between news- These parts were mounted in the same clearing
papers in a plant press and drying with supple- agent. The ovule structure was also studied from
mentary lieat. Living plants were collected in the material in this preservative.
area of Santa Rita in October 1971 and taken to Herbarium material, when used in dissection,
Sl’ashington, D.C., where they are under cultivation was treated with “Aerosol OT Solution” (Fischer
in a greenhouse. Laboratory no. SO-A-292). Spikelets were softened
Field photographs were taken with a Nikon F by treatment with a few drops of this solution for
camera (equipped with a Nikon Photomic-TN a few minutes and kept moist by drops of water
Finder), using the Micro-Nikkor 55mm lens and during dissection.
Kodak Tri-X film (ASA 400), without an addi- For studies of starch the single mature caryopsis
tional light source. A tripod was used in all available was soaked in a mixture of equal parts
instances. Aerosol OT Solution and water for one and a half
Material for morphological, anatomical, and hours. Cross-sections were made by hand, cutting
cytological studies was fixed in the field at time of with a razor blade the material positioned between
collection. For cytological studies young inflores- two pieces of pith. The sections were stained with
cences were fixed in a mixture of three parts 95 a drop of IKI for about 20 seconds, washed in
percent EtOH to one part glacial acetic acid, and water, and mounted in glycerine.
transferred within 24 hours to 70 percent EtOH Anatomical observations were made with the
and stored under refrigeration. Inflorescences, Leitz Ortholux microscope, equipped with plano
leaves, and seedlings were fixed in FAA (5 cc objectives, and photomicrographs were taken using
formalin: 5 cc glacial acetic acid: 90 cc of 50% this microscope and Kodak Panatomic-X film
.
EtOH) The blade of the first or second completely (ASA 32). Bright field, phase contrast, polarized
developed leaf from the uppermost part of the light, and dark field were used. Dissections were
culm was selected for preservation. Young in- studied under the Wild M 5 Stereo-microscope and
florescences were also fixed, in a mixture consisting drawings were made with the aid of the Wild draw-
of equal parts of glycerine and lactic acid (Bersier ing tube, Anatomical drawings were made using
and Bocquet, 1960), the SVild M 20 microscope, also with the aid of a
Studies of the leaf anatomy were made on SVild drawing tube, Illustrations of the habit of
material preserved in FAA. After washing, the the plant were made from herbarium specimens
sections were cut by hand with a razor blade from and field photographs.
the middle portion of the blade, and mounted, Voucher specimens of the plants reported in this
without staining, in glycerine or glycerine-lactic paper are filed in the United States National
acid. Herbarium, Smithsonian Institution. The collec-
Preparations of epidermises of the leaves were tion of material in liquid preservative is also
made by the standard technique of scraping and maintained at the same location.
mounted, without staining, in glycerine. Some
preparations were stained with a weak solution of
Maclurolyra tecta, new genus and species
safranin, without dehydration, to facilitate the
observation of the siliceous cells and microhairs. FIGURE4S- 7
The gynoecia were dissected and mounted, with-
DEscRIpTroN.-Gramen perenne sylvarum um-
out staining, in glycerine-lactic acid and studied brosarum, usque ad 48 cm altum. Culmi erecti,
with phase contrast and dark field illumination.
sine ramis, plerumque 20-48, cm alti, nodis 4-6,
The glycerine-lactic acid mixture acts not only as foliis 1-3. Foliorum vaginae cum setis ca. 1.5 mm
a good clearing agent but preservative as well and longis e marginibus superioribus emanentibus;