Polen Pared

THE
VOL. 4 9
BOTANICAL
OCTOBER-DECEMBER, 1983
REVIEW
NO. 4
Comparative Morphology of Monocot Pollen and Evolutionary Trends of Apertures and Wall Structures
M I C H A E L S. Z A V A D A l
Biological Sciences Group, U-43 The University of Connecticut Storrs, Connecticut 06268
Ab stract .................................................................................................................................................................................... S o m m a i r e ................................................................................................................................................................................ A6cTpaIcT ................................................................................................................................................................................. I n t r o d u c t i o n .......................................................................................................................................................................... Materials a n d M e t h o d s ................................................................................................................................................. T e r m i n o l o g y U s e d to Describe Pollen M o r p h o l o g y a nd Wa l l Structure ..................... Classification and Phylogeny of M o n o c o t s .................................................................................................... I. A l i s m a t i d a e....................................................................................................................................................... A l i s m a t a l e s ....................................................................................................................................................... B u t o m a c e a e ........................................................................................................................................... L i m n o c h a r i t a c e a e .............................................................................................................................. A l i s m a t a c e a e ......................................................................................................................................... H y d r o c h a r i t a l e s ............................................................................................................................................. H y d r o c h a r i t a c e a e ............................................................................................................................... N a j a d a l e s ............................................................................................................................................................ A p o n o g e t o n a c e a e ............................................................................................................................... Scheuchzeriaceae ............................................................................................................................... Ju ncaginaceae ....................................................................................................................................... P o t a m o g e t o n a c e a e ............................................................................................................................ R u p p i a c e a e ............................................................................................................................................. N a j a d a c e a e ............................................................................................................................................. Z a n n i c h e l l i a c e a e ................................................................................................................................. P o s i d o n i a c e a e ....................................................................................................................................... C y m o d o c e a c e a e .................................................................................................................................. Zosteraceae ............................................................................................................................................. 333 333 334 334 335 335 336 336 336 336 337 338 338 338 339 339 339 339 340 340 340 340 340 340 341
'Presen t address: D e p a r t m e n t o f Biology, J o r d a n Hall 138, I n d i a n a U n i v e r s i t y , B l oomington, I n d i a n a 47405. Copies o f this issue [49(4)] m a y be o b t a i n e d from: P u b l i c a t i o n s Office, The N e w Y o r k Botanical Gard en, Bronx, N Y 10458 USA. P R I C E (Includes postage a n d h a n d l i n g fee. Valid until 31 D e c e m b e r 1983.): U.S. Orders: $10.25. Non-U.S. Orders: $11.00. ( P a y m e n t in U.S. currency d r a w n on a U.S. b a n k m u s t a c c o m p a n y order. T h a n k you.)
The Botanical Review 4 9 : 3 3 1 - 3 7 9 . Oct.-Dec., 1983 Cc) 1983 The New York Botanical Garden 331
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THE BOTANICAL REVIEW
T r i u r i d a l e s ....................................................................................................................................................... P e t r o s a v i a c e a e ............................................................................................................. T r i u r i d a c e a e ........................................................................................................................................ II. A r e c i d a e .............................................................................................................................................................. A r e c a l e s ............................................................................................................................................................... A r e c a c e a e ............................................................................................................................................... C y c l a n t h a l e s ..................................................................................................................................................... C y c l a n t h a c e a e . ...................................................................................................................................... P a n d a n a l e s ...................................................................................................................................................... P a n d a n a c e a e .......................................................................................................................................... A r a l e s .................................................................................................................................................................... A r a c e a e ...................................................................................................................................................... L e m n a c e a e ............................................................................................................................................ I I I . C o m m e l i n i d a e . . .............................................................................................................................................. C o m m e l i n a l e s ............................................................................................................................................... R a p a t e a c e a e ......................................................................................................................................... X y r i d a c e a e .............................................................................................................................................. M a y a c a c e a e ............................................................................................................................................ Commelinaceae .......................................................................................................................... E r i o c a u l a l e s .................................................................................................................................... E r i o c a u l a c e a e .................................................................................................................................... R e s t i o n a l e s ........................................................................................................................................................ F l a g e l l a r i a c e a e ...................................................................................................................................... J o i n v i l l e a c e a e ................................................................................................................................. R e s t i o n a c e a e ................................................................................................................................... C e n t r o l e p i d a c e a e ................................................................................................................................ J u n c a l e s ......................................................................................................................................................... J u n c a c e a e . ............................................................................................................................................ T h u r n i a c e a e ........................................................................................................................................... C y p e r a l e s ............................................................................................................................................................ C y p e r a c e a e .............................................................................................................................................. P o a l e s .................................................................................................................................................................... P o a c e a e ..................................................................................................................................................... H y d a t e l l a l e s ...................................................................................................................................................... H y d a t e l l a c e a e ....................................................................................................................................... T y p h a l e s .............................................................................................................................................................. S p a r g a n i a c e a e ....................................................................................................................................... T y p h a c e a e ............................................................................................................................................... I V . Z i n g i b e r i d a e ..................................................................................................................................................... B r o m e l i a l e s ..................................................................................................................................................... B r o m e l i a c e a e . . ................................................................................................................................... Z i n g i b e r a l e s ...................................................................................................................................................... S t r e l i t z i a c e a e ....................................................................................................................................... H e l i c o n i a c e a e ................................................................................................................................... M u s a c e a e ................................................................................................................................................. Z i n g i b e r a c e a e ........................................................................................................................................ C o s t a c e a e ................................................................................................................................................. C a n n a c e a e ......................................................................................................................................... Maranthaceae .....................................................................................................................................
341 341 341 341 341 341 343 343 343 343 344 344 346 346 346 346 346 348 348 350 350 350 350 350 351 351 352 352 352 352 352 352 352 353 353 353 353 353 353 353 353 354 354 354 354 354 354 355 356
COMPARATIVE M O R P H O L O G Y OF MONOCOT POLLEN
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V. Liliidae ................................................................................................................................................................. Liliales .................................................................................................................................................................. Philydraceae .......................................................................................................................................... Pontederiaceae ..................................................................................................................................... Haemodoraceae .................................................................................................................................. Cyanastraceae ....................................................................................................................................... Liliaceae ................................................................................................................................................... Iridaceae ................................................................................................................................................... Velloziaceae ........................................................................................................................................... Aloeaceae ................................................................................................................................................. Agavaceae ............................................................................................................................................... Xanthorrhoeaceae ............................................................................................................................. Hanguanaceae ...................................................................................................................................... Taccaceae ................................................................................................................................................ Stemonaceae .......................................................................................................................................... Dioscoreaceae ...................................................................................................................................... Orchidales .......................................................................................................................................................... Geosiridaceae ....................................................................................................................................... Burmanniaceae .................................................................................................................................... Corsicaceae ............................................................................................................................................. Orchidaceae ........................................................................................................................................... Evolutionary Trends of Pollen Aperture Types in the Monocots ................................................ Evolutionary Trends of Pollen Wall Structure in Monocots ........................................................... Literature Cited .................................................................................................................................................................. Acknowledgments ............................................................................................................................................................. Appendix I .............................................................................................................................................................................
356 356 356 356 356 357 357 359 359 361 361 361 361 361 363 363 363 363 363 365 365 367 371 373 373 378
Abstract
D a t a o n p o l l e n a p e r t u r e a n d wall u l t r a s t r u c t u r e are r e v i e w e d f o r t h e m o n o c o t s . N e w u l t r a s t r u c t u r a l d a t a o n 30 t a x a r e p r e s e n t i n g 18 m o n o c o t f a m i l i e s are also p r e s e n t e d . T h e e v o l u t i o n a r y t r e n d s o f a p e r t u r e s a n d p o l l e n wall s t r u c t u r e are d i s c u s s e d a n d it is p r o p o s e d t h a t t h e e v o l u t i o n a r y t r e n d s o f p o l l e n a p e r t u r e s a n d wall s t r u c t u r e in t h e m o n o c o t s p a r a l l e l t h o s e p r o p o s e d b y W a l k e r f o r t h e dicots. H o w e v e r , t h e i m p o r t a n c e o f t h e s e t r e n d s differ f o r m o n o c o t s , suggesting t h a t t h e s e l e c t i v e p r e s s u r e s affecting p o l l e n wall a n d a p e r t u r e e v o l u t i o n in d i c o t s a n d m o n o c o t s h a v e been similar, but with different emphasis.
Sommaire
D o n n 6 e s e x p 6 r i m e n t a l e s sur l ' o u v e r t u r e d e p o l l e n et la u l t r a s t r u c t u r e d u m u r des M o n o c o t y l 6 d o n e s s o n t r e v u s . Les n o u v e l l e s d o n n 6 e s ex p 6 r i m e n t a l e s sur 30 taxa, r e p r 6 s a n t a n t e s 18 f a m i l l e s d e M o n o c o t y l 6 d o n e s s o n t aussi present6s. Les c o u r s 6 v o l u t i o n n a i r e s d es o u v e r t u r e s et s t r u c t u r e d u m u r d e p o l l e n s o n t discut6s et c ' e s t p r o p o s 6 q u e les c o u r s 6 v o l u t i o n n a i r e s d es o u v e r t u r e s d e p o l l e n et s t r u c t u r e d u m u r en M o n o c o t y l 6 d o n e s
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THE BOTANICALREVIEW
se mettent en parall61e de ceux proposes par Walker pour les Dicotyl6dones. Toutefois l'importance de ces cours sont diff6rents pour les Monocotyl6dones, donnent que la selection naturelle touchant l e m u r de pollen et les cours 6volutionnaires des ouvertures en Dicotyl6dones et Monocotyledones ont 6t6 semblables mais avec des accentuations diff6rantes. A6cTpaKT KOHCneKT. OJIHO]lO.rlbHbIe oro3peBaJOTC~ Ha ]laaable y31bTpacTpyKTypbl nhI21blleBOfi anepTypbl n CTeHbl. HOBbIe yJ1bTpacTpyKTypHbie jlaHHbIe 30 TaKCOHOB, npe~IcTanslalomnx 18 polos o~no~o~bnhlX, Tagx~e npe~cTaa~eHbl. O6cy~IaroTcn 3aoa~otmoI-inh~e nanpaB31eHna anepTyp n cTpyKTypbI nbI3161IeBOI~ CTeHbl, 14 npe~aaraeTcn, qTO 3BO.IIIOU14OHHble HanpaaJIeHnu nblJIbtleBbIX anepTyp 14 cTpyKTypbI CTeHbl ~lJln OJIHOj102IbHhIX napa.rIJIeJlbHbIMTeM, KOTOpb~e Ba31Kep npe~Jmran nasa ~ByJ1oJmnbIX. O~HaKO, Ba)KHOCTb 3TI4X HanpaBnen14fi pa33mqaeTca ~lan ORHO~IOSIbHblX
B TOM, qTO OTrOpHble )laB.rleH1451, BJll, l~IBIn14e Ha 3BOJIIOUHIO nblJlblleBOfi CTeHbl 14 anepTypbl R2Ia ]IBy]lOJIbHblX 140]IHO]lO.rlbHblX, 6bUlB FIOXO)K14
~pyr Ha ~Ipyra, riO OT3114qaJl14CbB Ba)KHOCT14. Introduction Interpretation of the evolutionary significance of fossil pollen has relied heavily on comparative morphological studies of extant pollen. Comparative morphological studies of extant pollen have, to some extent, elucidated evolutionary trends in a variety of dicot pollen characteristics, particularly in the ranalean dicots (e.g., Walker, 1974a, 1974b). Comparative morphological data on monocot pollen that can help determine the phylogenetic relationships among extant monocots and the primitive dicots, however, are lacking. The paucity of data on monocot pollen represents a considerable handicap to paleobotanists and evolutionary botanists interested in studying the evolution ofmonocots and, in general, the early evolution of the angiosperms. This paper summarizes what is currently known about pollen morphology and wall ultrastructure of monocot pollen, and includes morphological and ultrastructural data for 30 previously uninvestigated taxa representing 18 monocot families. Descriptions of pollen are treated primarily at the familial and subfamilial levels and in some cases at the tribal and generic levels. My goals are to provide an overview of the evolutionary trends of pollen aperture and wall structure in the monocots that will aid in interpretation of the fossil pollen record, and to more clearly illustrate the similarities and/or differences between monocot and dicot pollen.
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Materials and Methods All pollen was removed from dry herbarium material or vouchered live material grown in The University of Connecticut greenhouses. Live material was prepared for transmission electron microscopy (TEM) by fixation in cacodylate HCl-buffered gluteraldehyde-formaldehyde, followed by fixation in osmium tetroxide, dehydration in an ethanol series and embedding in Dow Epoxy Resin-334 (DER-334). Pollen removed from dry herbarium specimens was acetolyzed and prepared for TEM by dehydration in an ethanol series and embedding in DER-334. Sectioning was done on an LKB-1 ultramicrotome; the sections were post-stained in uranyl acetate and lead citrate for 15 minutes respectively, and viewed on a Philips EM-300. Pollen was prepared for scanning electron microscopy (SEM) by mounting the pollen on stubs with the high vacuum wax Apiezon W-100 and coated with gold-palladium. A Coates and Welter field emission scanning electron microscope was used for viewing. A list of the specimens used in this study and the herbarium data are presented in Appendix I. TERMINOLOGY USED TO DESCRIBE POLLEN MORPHOLOGY AND WALL STRUCTURE The terminology used to describe aperture type, tectal and supratectal ornamentation, and pollen wall structure follows Faegri and Iversen (1964). Faegri and Iversen consider the pollen wall to consist of three primary layers: the ektexine, endexine and intine. The ektexine and endexine are the outer sporoponeninous, acetolysis-resistant layers of the wall preserved in dispersed fossil pollen and are distinguished by their staining properties with basic fuchsin, the outer ektexine staining dark red and the inner endexine pink. The distinction between the ektexine and endexine is further substantiated by their differential stainability with the EM stains uranyl acetate and lead citrate. The intine, a non-acetolysis resistant layer, is not preserved in fossils and is not considered here. The ektexine is further divided by Faegri and Iversen (1964) into tectum, columellate layer and footlayer. The tectum is the outermost layer of the ektexine and may be perforate or imperforate and may exhibit supratectal ornamentation. The middle layer of the ektexine, the infratectal layer, may be columellate, consisting of radially directed columns, anastomosing rods, or spherical granules. The innermost layer of the ektexine, the footlayer, is the unsculptured layer of the ektexine. The unsculptured endexine underlies the ektexine, when present. Recent studies have shown endexine will corrode on treatment with acetolysis solution, giving anomalous results as to the extent ofendexine in certain species (Walker, 1976;
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Zavada, work in progress). Thus, its presence, absence, or extent must be interpreted with some reservations in both extant acetolyzed and fossil material. In addition to the differential stainability of the ektexine and endexine, Faegri and Iversen's terminological distinction between these wall layers is also substantiated by developmental data. These layers can be considered separate wall units based on their timing of development. Pollen wall development generally has three phases, the premeiotic phase, tetrad phase and free spore phase (Heslop-Harrison, 1971). The ektexine develops wholly within the tetrad phase which is characterized by the isolation of the meiotic tetrad in a callose wall. The free spore phase begins after the callose wall breaks down and the microspores are free in the anther locule. At this time, endexine development takes place. Thus, the Faegri and Iversen terminology used here is appropriate because it is based, not only on the differential stainability of the ektexine and endexine, but also on differences in the timing of the development of these wall layers. Although the following descriptions of monocot pollen walls are based on morphology and staining properties with the EM stains uranyl acetate and lead citrate, developmental data are included whenever possible.
Classification and Phylogeny of Monocots

There are numerous schemes to describe the classification and phylogeny of monocots. All are in a state of flux. This is, in part, due to the fact that monocot development, morphology, anatomy, chemistry and palynology have not received the same attention as certain important groups ofdicots (e.g., the ranales). Although all current taxonomic and phylogenic systems dealing with monocots have weaknesses, for the sake of convenience, only one scheme is followed in the present study, the widely respected scheme of Cronquist (1981). It is used here because it takes into account a broad range of morphological, biogeographical and anatomical data. The Cronquistian system deals primarily on and above the familial level. Where it is necessary to describe the palynological characteristics of taxonomic units below the familial level, monographic works dealing with the individual families are consulted.
I. ALISMATIDAE ALISMATALES
Butomaceae The family is monotypic (Butomus umbellatus L.) and native to temperate Eurasia. Pollen is monosulcate with reticulate sculpturing (Fig. 1). Pollen wall structure is tectate-columellate, with a well developed footlayer and no endexine (Fig. 2) (Argue, 1971).
COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN
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Figs. 1-5. Fig. 1. Butomus umbellatus L.: SEM showing reticulate exine sculpturing. x 1298. Fig. 2. Butomus umbellatus: TEM showing tectate-columellateexine structure, 7950. Fig. 3. Aponogeton echinatus Roxb.: SEM showing foveolate and spinulose exine sculpturing, x 2528. Fig. 4. Aponogeton echinatus: TEM showing tectate-columellate exine structure, 5400. Fig. 5. Aponogeton echinatus: TEM showing tectate-columellate exine structure and the thick columellae which are not firmly fused to the thin continuous footlayer, x 11,813.
Limnocharitaceae T h e family consists o f three genera a n d 7 - 1 2 species d i s t r i b u t e d t h r o u g h the tropical a n d subtropical regions o f the O l d a n d N e w W o r l d . T w o genera a n d t w o species h a v e b e e n investigated using light a n d t r a n s m i s s i o n electron m i c r o s c o p y (Argue, 1973). Pollen o f this family is p e r i p o r a t e a n d pore n u m b e r varies f r o m 3 - 7 in L i m n o c h a r i s f l a v a Buch. to 10-11 in
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T H E B O T A N I C A L REVIEW
Hydrocleis nymphoides (Willd.) Buch. Sculpturing in L. flava is fine to coarsely granular and pollen wall structure is tectate-columellate with irregularly shaped and branched columellae. The tectum is thin and imperforate. The footlayer varies in thickness and is fibrillar and/or lamellated. No endexine is evident. In contrast, pollen wall structure of Hydrocleis nymphoides is granular with a thick rectum which is infrequently perforated by minute channels. Hydrocleis nymphoides lacks a well developed footlayer and no endexine is evident. Exine sculpturing of this taxon consists of broad-based spines. Pollen of Tenagoclaris latifolia (Don) Buch. was studied (Argue, 1973) with light microscopy and appears similar to H. nymphoides pollen.
Alismataceae This family has a cosmopolitan distribution and consists of 12 genera and 75 species. Pollen of this family has been studied in detail with light, scanning and transmission electron microscopy (Argue, 1972, 1974, 1976; Hesse, 1980; Punt and Reumer, 1981). Argue recognizes two pollen groups in this family based on pore number and pollen amb. Type 1 pollen, represented only by the genus Caldesia, has three or fewer irregularly distributed pores, pollen amb is lenticular to ovate and exine sculpturing is reticulate. Pollen wall structure is tectatecolumellate with thick, irregularly shaped columellae which rest on a footlayer of varying thickness. No endexine is evident. The tectum is traversed by conspicuous perforations. Type 2 pollen is spheroidal to polyhedral and periporate. The number of pores varies from 9-29 and sculpturing is granular to spinulose. Pollen wall structure is tectate and the infratectal layer has irregular, branching columellae fused to a thin footlayer. No endexine is evident. The genera with type 2 pollen are: Alisma, Baldellia, Damasonium, Machaerocarpus, Echinodorus, Limnophyton, Lophotocarpus, Sagittaria and Wisneria.
HYDROCHARITALES
Hydrocharitaceae The family has a cosmopolitan distribution and consists of 15 genera and about 100 species. The pollen of this family has been little studied with scanning and transmission electron microscopy. Pollen is monosulcate or inaperturate and often filiform with psilate, echinate or clavate sculpturing. Meyer (1966) reported the ultrastructure of a variety of Helobiaen taxa (the Helobiae encompasses the Alismatales, Hydrocharitales and Najadales) including the monosulcate pollen of the Hydrocharitalean genus Stratiotes. It appears Stratiotes aloides L. is tectate with supratectal
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clavae. There is a thick tectum and a granular infratectal layer which rests on a thin but continuous basal layer. Meyer (1966) calls this basal layer endexine, but it exhibits none of the characteristics associated with endexine (e.g., differences in staining properties from the granular and tectal layers). In addition, endexine is unknown from all of the alismatidean taxa investigated thus far, making Meyer's interpretation questionable. The pollen wall structure of Halophila decipiens and Thalassia hemprich ii Aschers. was studied by Pettitt and Jermy ( 1975) and Pettitt ( 1981 ), who found the pollen to be filiform, with a thin unstructured and unsculptured exine, underlain by a relatively thick intine.
NAJADALES
Aponogetonaceae This monogeneric family has about 40 species distributed in the tropics of the Old World and South America. The pollen ofAponogeton echinatus Roxb. is the only one studied using scanning and transmission electron microscopy. Pollen of this family is monosulcate, foveolate, the foveola are funnel-shaped interruptions of the ektexine and sculpturing consists of spinules (Fig. 3). Pollen wall structure is tectate-columellate; the tectum is relatively thick and the columellae are broadly attached to the tectum and taper toward the footlayer (Figs. 4, 5). The columellae rest, but are not firmly fused to the thin footlayer (Fig. 5). Scheuchzeriaceae This monotypic family (Scheuchzeria palustris L.) is distributed in the temperate Northern Hemisphere. Pollen is inaperturate and occurs in dyads. Pollen wall sculpturing consists of a reticulum (Miyoshi, 1982). Pollen wall structure is tectate-columellate with a thin footlayer (=Meyer's endexine, 1966). Meyer (1966), once again, describes this basal layer as endexine, but, this layer has all the characteristics of footlayer, i.e., fusion to the baculate sculpturing elements and the same staining properties as the baculae. Juncaginaceae The family consists of five genera and 20 species distributed throughout the temperate regions of the Northern and Southern hemispheres. Pollen is inaperturate and reticulate. Pollen wall structure is only known in Triglochin palustre L. (Meyer, 1966) and is tectate-columellate, with a thin footlayer. Meyer (1966) indicates again the presence of an endexine, but no unequivocal endexine appears on his electron micrograph.
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Potamogetonaceae This monogeneric family has about 100 species and a cosmopolitan distribution. Wall structure TEM studies have been published for the following species: Potomogeton perfoliatus L. (Schwanitz, 1967b), P. pectinatus L. (Pettitt and Jermy, 1975; Schwanitz, 1967b) and P. natans L. (Meyer, 1966; Pettitt and Jermy, 1975). Pollen wall structure in all of the taxa investigated is tectate-columellate with a thin footlayer and no endexine. Exine sculpturing is reticulate. Ruppiaceae This monotypic family (Ruppia maritima L.) has a cosmopolitan distribution. Pollen is heteropolar, bilateral, multiaperturate, with reticulate exine sculpturing. Pollen wall structure is tectate-columellate with a thin footlayer and no endexine (Pettitt and Jermy, 1975; Schwanitz, 1967a, 1967b). Najadaceae This cosmopolitan, monogeneric family has about 35 species. Pollen is inaperturate, ellipsoidal and lacks exine sculpturing. The pollen wall, as revealed by TEM, consists of a thin homogeneous sporopolleninous layer, underlain by a relatively thick intine (Pettitt and Jermy, 1975; Shpakov and Kolesnickov, 1970). Zannichelliaceae This family consists of four genera and 7-8 species. The genus Althenia occurs in the Mediterranean region, Lepilaena in Australia, Vleisia in South Africa and Zannichellia is cosmopolitan. Pollen is predominantly inaperturate and wall sculpturing can be reticulate to verrucate (Zannichellia palustris L.; Schwanitz, 1967b). Pollen wall structure of Zannichellia palustris is the only one known. It consists of a thin continuous basal layer on which are irregular verrucae, which often coalesce forming an imperfect reticulum (Pettitt and Jermy, 1975). Posidoniaceae This monogeneric family has three species and is distributed in Australia and the Mediterranean. Pollen is inaperturate and filiform in shape. Little is known of its wall structure, but it is generally thought to be exineless (Erdtman, 1952). Cymodoceaceae This family includes five genera and 18 species distributed along subtropical and tropical seacoasts. Pollen of two genera (Halodule wrightii
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and Thalassodendron ciliatum Aschers.) have been investigated with TEM. The exine of both genera consists of a thin unsculptured and unstructured sporopolleninous layer, underlain by a relatively thick intine (Pettitt, 1981; Pettitt and Jermy, 1975). Pollen is inaperturate. Zosteraceae This family includes three genera and about 18 species distributed along seacoasts in temperate and tropical regions of the world. Pollen is inaperturate and filiform. Transmission electron microscope studies of the pollen of Zostera have revealed a very thin, unsculptured, unstructured sporopolleninous layer underlain by an intine (Pettitt and Jermy, 1975; Schwanitz, 1967b).
TRIURIDALES
Petrosaviaceae This is a monogeneric family with two species. Plants occur in Southern China, Japan, Malaysia and Borneo. Little is known of the pollen wall structure of the monosulcate pollen of this group. Triuridaceae This family includes seven genera and about 70 species distributed throughout the tropical and subtropical regions of the world. Pollen is predominantly inaperturate, except for Sciaphila which is monosulcate. Pollen wall ultrastructure has not been investigated in any of the taxa from this family. II. ARECIDAE
ARECALES
Arecaceae The Arecaceae consists of 206 genera and about 2700 species distributed throughout the tropics. Moore (1973) recognizes 15 informal groups within the Arecaceae and his treatment of the family is foUowed. Pollen has been studied in detail using light microscopy (Kedves, 1980; Mahabale, 1967; Sowunmi, 1968, 1972; Thanikaimoni, 1970a, 1970b) and in a few taxa with SEM (Parthasarathy, 1970). However, considering the size and importance of the family, too few studies have been initiated using TEM. 1. Coryphoid palms, 32 genera, 322 species Pollen is monosulcate and exine sculpturing is reticulate. Pollen wall ultrastructure has been studied in Chelyocarpus, Rhapidophyllum and Thrinax, and is tectate-columellate with a well developed footlayer and no endexine (C. P. Daghlian, pers. comm.).
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2. Phoenicoid palms, one genus (Phoenix), 17 species Pollen is monosulcate with reticulate exine sculpturing. Transmission electron microscope studies of the pollen wall have revealed a tectatecolumellate structure with a well developed footlayer. No endexine is evident (C. P. Daghlian, pers. comm.). 3. Borassoid palms, seven genera, 56 species Pollen is monosulcate with reticulate exine sculpturing. Little is known of the pollen wall structure in this group. 4. Lepidocaryoid palms, 22 genera, 664 species Pollen morphology and wall structure are not known for all the genera of this group. The aperture types that are prevalent in the taxa investigated are: monosulcate (Oncocalamus, Pigafetta, and Raphia), ulcerate (some species of Mauritia and Raphia), zonasulcate (Korthalsia and Plectocomia), dicolpate (Calamus, Ceratolobus, Cornera and Daemonorhops) and diporate (Eleiodoxa). Sculpturing is also variable and can be reticulate, verrucate, baculate, echinate or clavate. 5. Nypoid palms, monotypic (Nypa fruticans Wurmb.) Pollen is zonasulcate and wall sculpturing is finely reticulate with supratectal echinae. Pollen wall structure as revealed by TEM is tectatecolumellate with a thick footlayer and no endexine (Thanikaimoni, 1970a). 6. Caryotoid palms, three genera, 35 species Pollen is monosulcate. Pollen wall structure is known for only one genus and species (Caryota urens L.; Thanikaimoni, 1970a) and is tectate-columellate with a thick footlayer. No endexine is evident. 7. Pseudophoenicoid palms, one genus (Pseudophoenix), four species Pollen wall structure is not known in this genus, however, pollen is monosulcate to trichotomosulcate with reticulate exine sculpturing (Moore, 1973). 8. Ceroxyloid palms, four genera, 30 species Pollen is monosulcate to monoporate and exine sculpturing is reticulate and/or echinate. Pollen wall structure is not known for any of the genera. 9. Chamaedoreoid palms, six genera, 146 species Pollen is monosulcate and exine sculpturing is reticulate to finely reticulate. Pollen wall structure has not been investigated. 10. Iriarteoid palms, eight genera, 52 species Pollen is monosulcate and exine sculpturing can be reticulate, clavate or echinate. Pollen wall structure is unknown for all genera in this group. 11. Podococcoid palms, one genus (Podococcus), 1-2 species
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Pollen is monosulcate with finely reticulate exine sculpturing. Pollen wall structure has not been investigated. 12. Arecoid palms, 88 genera, 760 species Little is known of the pollen wall structure from this rather large group. However, pollen is monosulcate, trichotomosulcate to ulcerate. Exine sculpturing is also variable and runs the gamut of supratectal ornamentation. 13. Coccoid palms, 28 genera, 583 species Pollen is monosulcate, ulcerate or trichotomosulcate. Exine sculpturing is scabrate to finely reticulate and the tectum can be perforate (Nair and Sharma, 1963; Punt and Wessels Boer, 1966b). Pollen wall structure is only known from the genus Coccos nucifera L. and is tectate-columellate with a thick footlayer. No endexine is evident (Thanikaimoni, 1970a). 14. Geonomoid palms, six genera, 92 species Pollen is monosulcate and exine sculpturing is finely reticulate (Punt and Wessels Boer, 1966a). Pollen wall ultrastructure is not known. 15. Phytelephantoid palms, 3-4 genera, 15 species Pollen is monosulcate, ulcerate or trichotomosulcate. Exine sculpturing is reticulate. Pollen wall structure is unknown.
CYCLANTHALES
Cyclanthaceae This family consists of 11 genera and about 180 species distributed throughout tropical America. Pollen is monosulcate in Asplundia, Dicranopygium, Evodianthus, Ludovia, Pseudoludovia, Sphaeradenia, and Stelestylis; ulcerate in Choanopsis and Cyclanthus; monoporate in Carludovica and diporate in Thoracocarpus. Exine sculpturing varies from psilate to finely reticulate (Harling, 1958). The pollen wall structure of Sphaeradenia irazvensis (Cuf.) Had. and Stelestylis stylaris (Gleas.) Harl. was examined in the present study. Pollen of both genera is finely reticulate without supratectal ornamentation (Figs. 6, 8). Pollen wall structure is tectate-columellate with a thick footlayer and no endexine (Fig. 7).
PANDANALES
Pandanaceae This family consists of three genera and 500-600 species distributed throughout the tropical and subtropical regions of the Old World. Pollen is inaperturate (porate-ulcerate?) and exine sculpturing consists of small
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Figs. 6-10. Fig. 6. Sphaeradenia irazvensis (CUE) Having: SEM, X2378. Fig. 7. Sphaeradenia irazvensis: TEM showing tectate-columellate wall structure, X6225. Fig. 8. Stelestylis stylaris (Gleas.) Had.: SEM, X2820. Fig. 9. Arisaema triphyllum (L.) Schott: SEM showing echinate sculpturing and inaperturate condition, X2753. Fig. 10. A risaema triphyllum: TEM of a collapsed pollen grain showing "spongy" wall structure and faint echinae, which exhibit the same "spongy" structure (arrows), X7500.
spines, or verrucae (Huynh, 1980; Huynh and Stone, 1981). Nothing is known of pollen wall structure in this family.
ARALES
Araceae
This family consists of 110 genera and about 1800 species predominantly distributed in tropical and subtropical regions of the world. Pollen of the Araceae, like that of the palms, has been little studied using SEM
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and TEM. Thanikaimoni (1969) has investigated a variety of taxa with light microscopy. Eight subfamilies are commonly recognized and the pollen morphology of each of these subfamilies is summarized below. 1. Pothoideae Aperture types in this subfamily vary considerably. Monosulcate pollen occurs in Pothos and Pothoidium of the tribe Pothoeae and the Acoreae. Dicolpate in the tribe Heteropsideae, zonasulcate in the Zamioculcaseae, periporate in the Anthurieae and inaperturate in the Culcasieae. Exine sculpturing also varies, but is predominantly reticulate and often echinate. Wall structure is unknown for all the genera in this subfamily. 2. Monsteroideae Pollen is zonasulcate in the Monstereae and inaperturate in the Spathiphylleae. Exine sculpturing is reticulate in the Monstereae and for the most part striate in the spathiphylleae. Trevisan (1980) investigated the wall structure of Spathiphyllum wallisii Hort. and found it to be tectatecolumellate (possibly granular?) with a very thin footlayer which is underlain by a "sponge-like" endexine. 3. Calloideae Pollen is dicolpate and exine sculpturing can be foveolate or reticulate. Hesse (1980) investigated the pollen wall structure of Lysichitum americanum Hulten & St. John and found it to be tectate-granular with a thin continuous footlayer and no endexine. 4. Lasioideae Pollen is monosulcate (Lasieae) and inaperturate (Amorphophalleae, Nephthydieae, Montrichardieae). Exine sculpturing can be reticulate (Lasieae), psilate (Amorphophalleae), striate (Nephthydieae) or scabrate (Montrichardieae). Pollen wall structure of all the genera are unknown. 5. Philodendroideae Pollen is inaperturate and exine sculpturing is psilate, scabrate or echinate. Pollen wall structure is unknown. 6. Colocasioideae Pollen is inaperturate and exine sculpturing is predominantly echinate. Pollen wall structure is unknown. 7. Aroideae Pollen is inaperturate and exine sculpturing is psilate or echinate. Hesse (1980) investigated the pollen wall structure ofArum maculatum L. (Areae) and found it to be homogeneous with no tectum or infratectal structure (intectate). No endexine is evident. During the present study I investigated the wall structure of Arisaema triphyllum (L.) Schott. Pollen is inaperturate and exine sculpturing consists
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of short echinae (Figs. 9, 10). Pollen wall structure is tectate, imperforate with a "spongy" infratectal layer (Fig. 10). No footlayer or endexine is evident (Fig. 10). 8. Pistioideae Pollen is monosulcate and exine sculpturing is striate. Wall structure of all of the genera in this subfamily is unknown. Lemnaceae This family consists of six genera and about 29 species with a cosmopolitan distribution. Pollen is monoporate and exine sculpturing is echinate (Lemna minor L.; de Sloover, 1961). Pollen wall structure is not known. III. COMMELINIDAE
COMMELINALES
Rapateaceae This family consists of 16 genera and about 100 species distributed throughout tropical South America and West Africa. Carlquist (1961) studied the pollen morphology of this family using light microscopy. Pollen is monosulcate in the tribes Saxofridericieae, Schoenocephalieae and Monotremeae. In the tribe Rapateae pollen is zonasulcate (Cephalostemon, Duckea and Rapatea) and disulcate in Spathenthus. Exine sculpturing can be reticulate, psilate or scabrate. The pollen wall ultrastructure was studied in Saxofridericia regalis R. Schomb. and Stegolepis angustata Gleas. (Saxofridericieae) and Schoenocephalium cucullatum Maguire (Schoenocephalieae). In all three taxa pollen wall structure is tectate-columellate with a thick footlayer (Figs. 12, 14, 16). The tectum is occasionally traversed by minute channels (Figs. 12, 14, 16) and exine sculpturing consists of minute scabrae (Figs. 11, 13, 15), which are underlain by the columellae (Figs. 12, 14, 16). Endexine is present in Saxofridericia regalis as a thin granular layer (Fig. 12), and is similar to endexine reported in Zea mays (Skvarla and Larson, 1966). Xyridaceae Consists of four genera and more than 20 species distributed throughout subtropical and tropical regions of the world. Pollen is monosulcate (Xyris) or inaperturate (Abolboda). Carlquist (1961) reports Xyris pollen is similar in many respects to pollen of the Rapateaceae, particularly in aperture type and characteristics of exine sculpturing. Pollen wall structure is not known.
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Figs. 11-16. Fig. 11. Saxofridericia regalis Schumb.: SEM of a collapsed pollen grain, x 1050. Fig. 12. Saxofridericia regalis: TEM showing the tectate-columellate wall structure and the thin granular endexinal layer (arrow). Note that the columellae underly the small scabrae, X 12,000. Fig. 13. Schoenocephalium cucullatum Maguire: SEM showing scabrate wall sculpturing and orbicules, x945. Fig. 14. Schoenocephalium cucullatum: TEM showing tectate-columellate wall structure, note columellae underly the small scabrae, Fig. 15. Stegolepis angustata Gleason: SEM, X945. Fig. 16. Stegolepis angustata: TEM showing tectate-columellate wall structure, X6750.
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Mayacaceae Consists of a single genus (Mayaca) and about four species distributed throughout tropical West Africa and warm temperate America. Pollen is monosulcate and exine sculpturing is finely reticulate (Fig. 17). Pollen wall structure of Mayaca aubletti Michx. is tectate to intectate with branching, anastomosing columellae, which often become fused at their distal ends to form a tectum or remain free giving the appearance of being intectate (Fig. 18). The columellae are fused at their bases to form an irregular footlayer (Fig. 18). Beneath the fused bases of the columellae is a thin granular endexinal layer similar to endexine reported in Zea mays (Skvarla and Larson, 1966) and Saxofridericia regalis (see above). Pollen wall structure of M. aubletti exhibits many similarities to wall structure found in some Commelinaceae (e.g., Commelinanta). Commelinaceae This family consists of 50 genera and about 700 species distributed throughout subtropical and tropical regions of the world. Pollen is predominantly monosulcate, however, triaperturate pollen is known in the genera Commelinanta and Tinantia. Exine sculpturing varies considerably. Pollen wall structure has been reported for 11 taxa: Callisia, Rhoeo, Setcreasea, Spironema, Zebrina (Rowley, 1959), Coleotrype, Commelina, Tinantia (Poole and Hunt, 1980), Tradescantia (Horvat, 1966; Mepham and Lane, 1969, 1970; Rowley, 1959, 1975), Commelinanta (Rowley, 1959; Rowley and Dahl, 1962) and Tripogandra (Mattsson, 1976). The pollen wall structure of Dichorisandra thyrsiflora Mikan. is described in the present study. The pollen wall structure of Callisia, Coleotrype, Commelina, Rhoeo, Setcreasea, Spironema, Tinantia, Tradescantia and Zebrina is similar: tectate-columellate with irregular to branching columellae. The tectum is periodically traversed by small perforations, the footlayer is thick to very thin and may be interrupted in some taxa by endo-perforations. No endexine is evident in any of these taxa. Pollen TEM studies of Dichorisandra reveals a wall structure similar to that of the above genera (Figs. 19, 20). It has a thick tectum, with stout, irregular, occasionally branching columellae and a thick footlayer, and no endexine (Fig. 20). One notable difference is the lack of tectal perforations (Figs. 19, 20).
Figs. 17-22. Fig. 17. Mayaca aubletti Mich.: SEM showing reticulate exine sculpturing. 1905. Fig. 18. Mayaca aubletti: TEM showing unusual exine structure and thin granular endexine (arrow), X9750. Fig. 19. Dichorisandra thyrsiflora Mikan.: SEM showing scabrate
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exine sculpturing, 1253. Fig. 20. Dichorisandra thyrsiflora: TEM of nonacetolyzed pollen, showing thick tectum, columellae, footlayer and thin intine. Note abundant rough endoplasmic reticulum, also characteristic of Tradescantia, 13,350. Fig. 21. Centrolepis exserta Rts.: SEM showing scabrate-rugulate exine sculpturing, ulcerate aperture and tectal perforations, 1270. Fig. 22. Centrolepis exserta: TEM showing granular infratectal wall structure and one tectal perforation which does not communicate with the infratectal area,
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Pollen of Commelinanta has three ill-defined apertures. Pollen wall structure is tectate-columellate, the columellae are irregularly shaped, branched at times and fused at their bases to form a thick footlayer which has dentate endosculpturing. The wall structure is similar to that found in Mayaca pollen, however, no endexine is evident. Pollen of the genus Tripogandra is dimorphic (Lee, 1961) and correlates well with filament length. Pollen of the short filament anthers is spheroidal, monosulcate and exhibits a wall structure similar to Tradescantia. Pollen of this type is viable at maturity. Pollen of the long filament anthers is ellipsoidal and non-viable at maturity. Exine sculpturing is striate with the striae running parallel to the long axis of the pollen grain. Pollen wall structure is also similar to that of Tradescantia. The endexine is absent from both types of pollen (Mattsson, 1976).
ERIOCAULALES
Eriocaulaceae Consists of 13 genera and about 1200 species distributed throughout subtropical and tropical regions of the world, with a few species occurring in temperate regions. Pollen morphology was, studied using light microscopy by Thanikaimoni (1965), who considers the pollen of this family to be spiraperturate. Exine sculpturing is echinate. Little is known of the pollen wall structure.
RESTIONALES
Flagellariaceae The family consists of one genus (Flagellaria) and about three species distributed throughout the tropical regions of the Old World. Pollen is spheroidal, monoporate with a weakly annulate graminoid pore. Exine sculpturing is verrucate-subgranulate and minutely punctate (Chanda, 1966; Ladd, 1977). Pollen wall structure is unknown. Joinvilleaceae Consists of one genus (Joinvillea) and about two species native to the Pacific Islands. Pollen is spheroidal and monoporate with a graminoid pore. Exine sculpturing is verrucate and minutely punctate (Chanda, 1966). Pollen wall structure is tectate-granular with a thick footlayer and no endexine (Chanda and Rowley, 1967). Pollen of this family is similar to pollen of the Flagellariaceae.
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Restionaceae Consists of 30 genera and about 400 species distributed throughout the Southern Hemisphere. The family is generally divided into two tribes, the Anarthrieae and Restioneae. 1. Anarthrieae This exclusively Australian tribe has four genera: Anarthria, Ecdeiocolea, Hopkinsia and Lyginia. Pollen has been studied using light microscopy (Chanda, 1966), SEM (Ladd, 1977) and in one genus TEM (Chanda and Rowley, 1967). Pollen of this tribe is monoporate with graminoid, annulate pores. Exine sculpturing can be verrucate, psilate and the rectum is usually perforated with minute punctae (called scrobiculi). Pollen wall structure of Ecdeiocolea monostachys F. Muell. is tectatecolumellate with thin columellae. The footlayer is thick and no endexine is evident. 2. Restioneae This tribe has about 26 generz Pollen was studied in detail using light microscopy (Chanda, 1966), SEM (Ladd, 1977) and TEM (Chanda and Ferguson, 1978; Chanda and Rowley, 1967). Pollen is considered to be monoporate with an irregularly outlined ulceroid pore (Centrolepidoidtype pore) which lacks an annulus. Exine sculpturing can be verrucaterugulate and the tectum is usually perforated by minute channels (scrobiculi). Pollen wall structure in Restio and Meeboldina is tectate-granular with a thick footlayer and no endexine. Pollen wall structure in the genera Calorophus, Empodisma and Staberoha differs from the above genera in being tectate-columellate with a thick footlayer and no endexine. Staberoha pollen is interesting in that the spinulose exine ornaments are underlain by the columetlae, a situation thus far only observed in the Rapateaceae and Poaceae. Centrolepidaceae The family consists of four genera and about 35 species distributed throughout Australasia. Pollen of this family has been studied using light microscopy (Chanda, 1966) and SEM (Ladd, 1977). Pollen is monoporate with a centrolepidoid pore. Exine sculpturing is verrucate and the tectum is perforated by small punctae (scrobiculi; Fig. 21). The pollen wall ultrastructure of two species of Centrolepis (C. aristata Roem. and C. excerta Rts.) was investigated. Pollen wall structure is tectate-granular with a thick footlayer (Fig. 22), and no endexine. The tectal perforations in these two species do not open into the infratectal layer, but are interruptions of the exine (Fig. 22).
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JUNCALES
Juncaceae Consists of eight genera and about 300 species distributed throughout the temperate regions of the world. Pollen is monoaperturate and often united in tetrads. Exine sculpturing can be psilate to scabrate. Hesse (1980) investigated the pollen wall ultrastructure of two species of Luzula and found the sporopolleninous wall to be thin, atectate, composed of globular ektexinal units appressed to one another laterally. No endexine is evident. Thurniaceae This monogeneric family (Thurnia) has only three species, all native to the Amazon basin. Pollen is monoaperturate (ulcerate) and often united in tetrads. Pollen wall structure is little known.
CYPERALES
Cyperaceae Consists of 70 genera and about 4000 species, cosmopolitan in distribution. Pollen is usually monoaperturate, ulceroid, but Calyptrocarya has been reported to have four ill-defined pores. Exine sculpturing can be psilate, baculate or minutely scabrate. Hesse (1980) studied the wall structure of three species of Carex, and Dunbar (1973) the pollen wall development and structure of Eleocharis spp. They found the wall structure to be tectate-columellate with a perforate tectum and a relatively thick footlayer. No endexine is evident. Nilsson et al. (1977) studied Scirpus maritimus L. using SEM and Carex caespitosa Good. with SEM and TEM. Carex caespitosa is tectate-columellate with a discontinuous footlayer, otherwise it is similar to exine sculpturing in Scirpus maritimus and the other taxa studied in this family.
POALES
Poaceae This family consists of 500 genera and about 8000 species cosmopolitan in distribution. Pollen aperture type is monoporate, annulate. Exine sculpturing can be psilate, spinulose or minutely scabrate. Pollen wall development is remarkably constant and is tectate-columellate with a thick footlayer (Rowley, 1962a, 1962b, 1963, 1964). The tectum is traversed by minute channels. In pollen with spinulose supratectal ornamentation the spinules are subtended by columellae, a situation thus far observed in the tribe Restioneae of the Restionaceae and the Rapateaceae (Grant,
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1972; Hesse, 1980; Larson et al., 1962; Liem and van Andel, 1968; Nilsson et al., 1977; Rowley, 1960). Skvarla and Larson (1966) report the presence of a thin, granular endexine in Zea mays, however, the other taxa investigated with TEM lack this wall layer.
HYDATELLALES
Hydatellaceae Consists of two genera (Trithuria and Hydatella) and seven species native to Australia and New Zealand. Bortenschlager et al. (1966) investigated the pollen morphology of these two genera with light microscopy and Ladd (1977) studied Trithuria using SEM. Pollen is monoporate with an ill-defined ulceroid aperture. Exine sculpturing is microverrucate to scabrate with minute punctae. Pollen wall structure is little known.
TYPHALES
Sparganiaceae Consists of one genus (Sparganium) and 13 species distributed throughout the north temperate regions. Pollen is monoporate with an ulceroid pore, the exine sculpturing finely reticulate (Nilsson et al., 1977). Pollen wall ultrastructure is little known. Typhaceae This family consists of one genus (Typha) and 10 species cosmopolitan in distribution. Pollen is monoporate, ulceroid and is usually united in tetrads. Exine sculpturing is finely reticulate. Nilsson et al. (1977) investigated the pollen wall structure of Typha latifolia L. and found it to be tectate-columellate with a thick footlayer and no endexine.
I V . ZINGIBERIDAE BROMELIALES
Bromeliaceae Consists of 45 genera and about 2000 species distributed mostly through North America. Pollen of 39 genera and 217 species was studied with light and scanning electron microscopy (Ehler and Schill, 1973). Pollen of the subfamilies Pitcairnioideae and Tillandsioideae is primarily monosulcate and pollen of the Bromelioideae can be monosulcate or multiaperturate. Exine sculpturing is reticulate (Fig. 23). Pollen wall structure is little known, however, one species of Billbergia (Bromelioideae) was
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investigated in the present study. Brighigna et al. (1981) studied the wall structure of Tillandsia caput-medusa! Morr. (Tillandsioideae). Both taxa are tectate-columellate with a thin footlayer and no endexine (Fig. 24).
ZINGIBERALES
Strelitziaceae This family consists of three genera, Phenakospermum, Ravenala and Strelitzia distributed through South America, Madagascar and South Africa, respectively. Pollen is inaperturate and sporoderm sculpturing can be psilate or scabrate to spinulose. Pollen wall structure is little known. Heliconiaceae This family consists of one genus (Heliconia) distributed throughout tropical and subtropical South and Central America. Pollen is inaperturate, however, the pollen tube consistently emerges from a polar papilla. Sculpturing is psilate to spinulose. Pollen wall structure and ontogeny has been studied by Kress et al. (1978) and Stone et al. (1980). Pollen is exineless except for a few sporopolleninous spinules. The intine comprises the majority of the outer wall being greatly elaborated into an alveolarlike structure on the papillate pole, and homogeneous to fibrillar on the opposing pole. Musaceae This family consists of two genera, Ensets and Musa, distributed through the tropical and subtropical regions of the Old World. Pollen is inaperturate and exine sculpturing is psilate. The sporopolleninous exine is very thin, however, little else is known about the pollen wall structure. Zingiberaceae This family consists of 47 genera and about 1200 species distributed throughout southern and southeastern Asia. Pollen is generally inaperturate (monosulcate in Zingiber) and exine sculpturing is spinulose or psilate. Pollen appears to be exineless, however, pollen wall structure has not been studied in detail. Costaceae Consists of four genera and about 150 species of pantropical distribution. Pollen aperture types are monosulcate (Dimerocostos), spiraperturate (Tapeinochilos), or polyporate or polyulcerate (Costus and Monocostus) (Punt, 1968). Exine sculpturing is psilate. The pollen wall structure
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Figs. 23-26. Fig. 23. Billbergia sp.: SEM showing reticulate exine sculpturing, X1058. Fig. 24. Billbergia sp.: TEM of a nonacetolyzed pollen grain in the region of the aperture, showing tectate-columellate wall structure and elaborated intine (arrow), Fig. 25. Cyanastrum cordifolium Oliver: SEM ofnonacetolyzed pollen; tapetal substances deposited on the exine obscure the finely reticulate nature of the exine sculpturing, X1095. Fig. 26. Cyanastrum cordifolium: TEM showing tectate-columellate wall structure and the relatively thick footlayer, >(9938.
of Tapienochilos has been investigated (Stone et al., 1981) and is considered a two-layered structure. The outermost region of the sporopolleninous exine is composed of tangentially layered "rodlets" and the innermost region is composed of radially oriented "pegs," giving the appearance of a granular wall structure (Stone et al., 1981). The relationship of these two wall layers to Faegri and Iversen's ektexine and endexine is unknown. Cannaceae This monogeneric family (Canna) consists of about 50 species distributed throughout the tropical and subtropical regions of the New World. Pollen is inaperturate and exine sculpturing is spinulose (Nair, 1960). Pollen for the most part is exineless, except for the sporopolleninous spinules. The intine is a thick, elaborated, alveolar-like structure similar
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to that found in Heliconia pollen (Rowley and Skvarla, 1975; Skvarla and Rowley, 1970). Maranthaceae This family consists of 30 genera and about 400 species, pantropical in distribution. The predominantly inaperturate, psilate pollen of this family has been little studied.
V . LILIIDAE LILIALES
Philydraceae This family consists of four genera and five species all occurring in Australia. Pollen is monosulcate or triaperturate. Exine sculpturing is primarily reticulate. Pollen wall strutcture is unknown. Pontederiaceae Consists of nine genera and about 30 species distributed throughout the tropical and subtropical regions of the world. Pollen is disulcate and exine sculpturing is minutely scabrate to finely reticulate. Little is known of the pollen wall structure. Haemodoraceae The family is composed of about 16 genera and 100 species distributed mostly throughout the Southern Hemisphere and the Eastern United States. The family is usually divided into two tribes, the Haemodoreae and the Conostyleae. Pollen was studied with light microscopy by Radulescu (1973a). 1. Haemodoreae This tribe consists often genera and pollen of all the taxa is monosulcate. Exine sculpturing can be scabrate, gemmate or psilate. In the present study the pollen wall structure of three genera and three species was investigated. Pollen of Lachnanthes caroliana (Lam.) Dandy is monosulcate and exine sculpturing is scabrate (Fig. 27). Pollen wall structure is atectate and no endexine is evident (Fig. 28). Pollen of Wachendorfia paniculata Burm. is monosulcate and exine sculpturing on the proximal and distal faces of the pollen grain is gemmate to pustulate; a conspicuous psilate ridge separates the proximal and distal faces of the pollen grain (Fig. 31). Pollen wall structure is intectate-granular
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in the proximal and distal regions, and tectate-granular in the region of the psilate ridge (Fig. 29). No footlayer or endexine is evident. Pollen of Xiphidium caeruleum Aubl. is also monosulcate and exine sculpturing is psilate with minute punctae (Fig. 30). Pollen wall structure is tectate-granular with no footlayer or endexine (Fig. 32). 2. Conostyleae This tribe has six genera. Pollen is 2-8-porate, except Lophiola aurea Ker-Gawler pollen which is monosulcate. Exine sculpturing is predominantly scabrate (reticulate in L. aurea). The pollen wall structure of three genera and three species was investigated in the present study. Pollen ofAnigozanthosflavus D.C. ex Red. is diporate and exine sculpturing is scabrate (Fig. 33). Pollen wall structure is atectate and no endexine is evident (Fig. 34). Pollen of Conostylis setosa Lindl. is triporate and exine sculpturing is scabrate (Fig. 35). Pollen wall structure is atectate. No endexine is evident (Fig. 36). The pollen wall structure of this taxon is identical in all respects to A. flavus. Pollen ofLophiola aurea is monosulcate, which is unique for this tribe. Exine sculpturing is reticulate (Fig. 38) and pollen wall structure is tectatecolumellate with a thin footlayer"and no endexine (Fig. 37). There has been some question as to the inclusion of this taxon in the Haemodoraceae. It exhibits many morphological and palynological similarities to taxa allied with the Tecophilaeaceae (Zavada et al., 1983). Little is known of the pollen wall structure of the remaining taxa in the Haemodoraceae. Cyanastraceae Consists of one genus (Cyanastrum) and about seven species native to tropical Africa. Pollen is monosulcate to trichotomosulcate and exine sculpturing is finely reticulate. Pollen wall structure of Cyanastrum cordifolium D. Oliver was investigated and TEM has revealed a tectatecolumellate structure with a relatively thin footlayer. No endexine is evident (Figs. 25, 26). Liliaceae This family consists of 280 genera and about 4000 species, cosmopolitan in distribution. There have been a number of studies with light microscopy (Nair and Sharma, 1965; Radulescu, 1972, 1973b, 1973c, 1973d), however, for the size and importance of this family it is little known palynologically. Pollen can be inaperturate, monosulcate, trichotomosulcate, disulcate
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Figs. 27-32. Fig. 27. Lachnanthes caroliana (Lam.) Dandy: SEM, X 1680. Fig. 28. Lachnanthes caroliana: TEM of a nonacetolyzed pollen grain showing atectate wall structure with a thin intine, X9750. Fig. 29. Wachendorfiapaniculata Burm.: TEM ofnonacetolyzed
pollen showing granular infrastructure in the region of the ridge which separates the proximal and distal faces of the pollen grain; no footlayer is evident, the granules rest on a thin intine, x7950. Fig. 30. Xiphidium caeruleum Aubl.: SEM of distal face showing psilate exine sculpturing and small perforations, x2100. Fig. 31. Wachendorfiapaniculata: SEM showing
359
or 4-porate. Exine sculpturing is extremely variable and runs the gamut of sculpturing types. Pollen wall structure in a few genera has been investigated with SEM and TEM. Pollen wall structure is tectate-columellate with a relatively thin footlayer and no endexine in Clivia miniata Regel (Afzelius, 1956; Gullvag, 1964), Endymion non-scriptus (L.)Garcke (Angold, 1967), Disporum spp. (Takahashi and Sharma, 1980), Hemerocallis sp. (Takahashi, 1980), Hymenocallis sp. (Larson, 1965), Litium longiflorum Thunb. (Heslop-Harrison, 1968a, 1968b), and L. humboldtii Rozel & Leichtl. (Southworth and Branton, 1971). The pollen of Crinum americanum L. was studied and was found to be monosulcate, the exine sculpturing consisting of spinules and spines (Fig. 39). Pollen wall structure is unique, consisting of two distinct layers: an outer layer of closely packed granules in which the spines and spinules are embedded and beneath this layer a thick homogeneous layer traversed by minute channels (Fig. 40). These two layers appear to comprise the ektexine. No endexine is evident. Takahashi (1982) investigated 17 species of Trillium ultrastructurally. Pollen of Trillium is inaperturate and exine sculpturing can be echinate, granulate, verrucate or spinulate. The pollen wall is comprised of a very thin outer sporopolleninous wall and an inner thick and often elaborately channeled intine. Iridaceae Consists of 80 genera and about 1500 species, cosmopolitan in distribution. Pollen is usually monosulcate or inaperturate and exine sculpturing is predominantly reticulate (Radulescu, 1970a, 1970b; Rodionenko, 1956). Pollen wall structure was investigated in Sisyrinchium californicum L. and Iris pseudacorus Ait. and was found to be tectatecolumellate with a thin footlayer (Figs. 42, 44). No endexine is evident. Both taxa are monosulcate and exine sculpturing is reticulate (Figs. 41, 43). Velloziaceae This family consists of six genera and about 250 species, distributed throughout Africa, South America and Madagascar. Pollen is predominantly monosulcate and exine sculpturing is reticulate (Barbacenia) or verrucate (Vellozia). Vellozia pollen usually occurs in tetrads (Maguire,
geminate exine sculpturing on the proximal (P) and distal (D) faces of the pollen grain, note the psilate ridge, X953. Fig. 32. Xiphidium caeruleum: TEM showing small perforation (arrow) and granular infratectal wall structure, X9750.
360
Figs. 33-38. Fig. 33. Anigozanthosflavus D.C. ex Red.: SEM showing diporate condition, x 1095. Fig. 34. A nigozanthos flavus: TEM of nonacetolyzed pollen showing atectate wall structure and thick intine, X6225. Fig. 35. Conostylis setosa Lindl.: SEM showing triporate condition, 1868. Fig. 36. Conostylis setosa: TEM of nonacetolyzed pollen showing atectate wall structure and the intine, X 11,625. Fig. 37. Lophiola aurea Ker-Gawler: TEM of nonacetolyzed pollen showing tectate-columellate wall structure which is unlike all other wall structure types known from the Haemodoraceae, XI 1,625. Fig. 38. Lophiola aurea: SEM showing the monosulcus and finely reticulate exine sculpturing, X2003.
361
1969). Pollen wall structure of Barbacenia is tectate-columellate with a thin footlayer and no endexine. Vellozia pollen can be tectate-columellate (e.g., V. flavicans) with a thin lamellated footlayer and no endexine, or granular with the granules (or somewhat distorted columellae) resting on a thin footlayer (e.g., V. granulifera). No endexine is evident in either taxon (Ayensu and Skvarla, 1974). Aloeaceae Consists of five genera and about 700 species, distributed in Africa, Madagascar and the Arabian peninsula. Pollen is predominantly monosulcate and exine sculpturing is reticulate (Majumbar, 1972). Yuhl and Maj umbar (1981) investigated 14 species of Haworthia pollen using SEM and TEM. Pollen wall structure of Haworthia is tectate-columellate with a thin footlayer. No endexine is evident. Agavaceae Consists of 18 genera and about 600 species distributed throughout the tropical and subtropical regions of the Old and New World. Pollen is monosulcate, exine sculpturing is reticulate (Panchukshurappa, 1965). Meyer and Yaroschevskaya (1973) investigated the wall structure of Dra~cena, which is tectate-columellate with a thin footlayer. No endexine is evident. Xanthorrhoeaceae Consists of nine genera and 55 species, distributed throughout Australia, New Guinea and New Caledonia. Pollen has been studied in detail using light and scanning electron microscopy by Chanda and Ghosh (1976). Pollen is monosulcate in Acanthocarpa, Chamaexeros, Kingia, Xanthorrhoea, and some Lomandra; spiraperturate and multiaperturate pollen is found in Lomandra. Exine sculpturing can be reticulate, echinate, psilate, verrucate or scabrate. Pollen wall structure is little known. Hanguanaceae One genus (Hanguana) and one to two species distributed throughout Malesia and Ceylon. Pollen is inaperturate and exine sculpturing is spinulose. Pollen wall structure is not known. Taccaceae Consists of one genus (Tacca) and about 10 species, pantropical in distribution. Pollen is monosulcate and exine sculpturing is verrucate to
362
Figs. 39-44. Fig. 39. Crinum americanum L.: SEM showing spinulose exine sculpturing and the monosulcus, X1230. Fig. 40. Crinum americanum: TEM showing the spinules embedded in a granular layer which rest on a thick basal layer traversed by minute channels, X 18,750. Fig. 41. Iris pseudacorus L.: SEM showing reticulate exine sculpturing and monosulcus, X1635. Fig. 42. lris pseudacorus: TEM showing tectate-columellate wall structure, X7500. Fig. 43. Sisyrinchium cafifornicum Air.: SEM showing finely reticulate exine sculpturing and monosulcus, X2708. Fig. 44. Sisyrinchium californicum: TEM of nonacetolyzed pollen showing tectate-columellate wall structure and thin intine,
363
somewhat striate (Fig. 45). Pollen wall structure of the two species investigated [T. macrantha Redonte and T. integrifolia (non-Ker-Gawler) Schrank] is tectate-columellate with a thick footlayer (Fig. 46). No endexine is evident and the tectum is traversed by minute perforations (Fig. 46). Stemonaceae Consists of three genera (Croomia, Sternona, Stichoneuron) and about 30 species, distributed in Australasia, except for one species of Croomia that occurs in the southeastern United States. Pollen is monosulcate and exine sculpturing often appears finely reticulate (Fig. 47). The pollen wall structure of Sternona japonica (B1.) Miq. consists of three layers, a thin discontinuous tectum (Fig. 48), an alveolar infratectal layer, reminiscent of the alveolar walls of some fossil pteridosperms and extant cycads (Fig. 48), and a thin footlayer (Fig. 48). Dioscoreaceae This family consists of six genera and about 630 species distributed throughout the tropical and subtropical regions of the world. Pollen is monosulcate or multiaperturate. Exine sculpturing can be reticulate or spinulose (Clarke and Jones, 1981). Transmission electron microscope studies of the pollen wall of Dioscorea polygonoides Uline have revealed a tectate-columellate exine structure with a footlayer and a thin granular endexine (Fig. 50), much like the endexine observed in Zea mays (Skvarla and Larson, 1966), Mayaca aubletti and Saxofridericia regalis. Pollen of D. polygonoides is monosulcate with reticulate exine sculpturing (Fig. 49).
ORCHIDALES
Geosiridaceae This monotypic family (Geosiris aphylla Baillon) occurs in Madagascar. Little is known of its sulcoidate-ulcerate pollen. Burmanniaceae Consists of 20 genera and about 200 species, pantropical in distribution. Pollen is monosulcate or 1-2-porate (Fig. 51). Exine sculpturing is psilate (Chakrapani and Raj, 1971). The wall structure of Burmannia bicolor Hort. was investigated and TEM studies have revealed a thick tectum traversed by minute channels (Fig. 52), and an infratectal structure consisting of numerous, thin, irregular columellae which are fused to a thick footlayer (Fig. 52). The innermost portion of the footlayer is lamellated (Fig. 52). No endexine is evident.
364
Figs. 45-50. Fig. 45. Tacca integrifolia (non Ker-Gawler) Schrank: SEM showing rugulate exine sculpturing and monosulcus, 1778. Fig. 46. Tacca macrantha Redonte: showing tectate-columellate wall structure, which is identical in all respects to T. integrifolia, 9750. Fig. 47. Stemonajaponica (B1.) Miq.: showing monosulcus, 1538. Fig. 48. Stemona japonica: TEM showing thin tectum which is often absent (arrow), granular infrastructure, and thin footlayer, Xl 1,625. Fig. 49. Dioscorea polygonoides Uline.: SEM of a collapsed
365
Corsiaceae This family consists of two genera (Arachnit& and Corsia) and about nine species, distributed throughout Chile and New Guinea. Pollen is monosulcate and exine sculpturing is finely reticulate. Little is known of the pollen wall structure. Orchidaceae This family consists of about 1000 genera and possibly 15,000-20,000 species of cosmopolitan distribution. Five subfamilies are commonly recognized; pollen data for each subfamily are summarized below. 1. Apostasieae Pollen is monosulcate, in monads and exine sculpturing is usually reticulate. Pollen wall structure is tectate-columellate with a thin fooflayer and no endexine (Newton and Williams, 1978; Schill, 1978). 2. Cypripedieae Pollen of this subfamily can be sulcate, ulcerate or porate, and occur in monads or tetrads. Exine sculpturing can be reticulate, verrucate or scabrate. Pollen wall structure appears tectate-columellate as revealed by SEM (Newton and Williams, 1978; Schill and Pfeiffer, 1977). 3. Neottieae Pollen can be ulcerate, porate or inaperturate in monads, but usually occurs in tetrads or pollinia. Pollen wall structure, as based on SEM, appears tectate-columellate, and exine sculpturing is predominantly reticulate (Ackerman and Williams, 1980, 1981; Balogh, 1979). 4. Orchideae and 5. Epidendreae Pollen of these subfamilies is similar and can be porate, or ulcerateinaperturate, and usually occurs in pollinia. Exine sculpturing may be psilate, reticulate, verrucate-scabrate or the pollen may be exineless (Caspers and Caspers, 1976; Dulieu, 1973; Williams and Broome, 1976). Pollen wall structure can be tectate-columellate (Chardard, 1958, 1969; Heslop-Harrison, 1968c) or exineless in the interior of the pollinium with the sporopolleninous walls restricted to the most distal pollen in the pollinium (Figs. 53, 54). In the two genera investigated in this study [Cleisostoma racemifera (Lindl.) Garay and Laelia automnalis Londl.]
4---
pollen grain showing monosulcus and finely reticulate exine sculpturing, Fig. 50. Dioscorea polygonoides: TEM showing tectate-columellate wall structure, thin footlayer and granular, thin endexine (arrow), X9750.
366
367
the sporopolleninous wall is restricted to the outermost pollen grains in the pollinium (Figs. 53, 54) and it is tectate-granular, lacking a footlayer and endexine (Fig. 54). The innermost pollen grains in the pollinium are exineless and have large cytoplasmic connections between the individual pollen grains (Fig. 55).
Evolutionary Trends of Pollen Aperture Types in the Monocots

Palynologists agree that the monosulcate aperture is the most primitive aperture type among angiosperms. This is supported by its prevalence in the primitive ranalean complex (Walker, 1974b), by the fact that it characterizes the first unequivocal fossil-dispersed angiosperm pollen (Doyle, 1969), and by its widespread occurrence in the most primitive taxa of each monocot subclass (Kuprianova, 1948, 1967; Walker and Doyle, 1975; present study). The most notable evolutionary trend among aperture types in the monocots is the loss of the sulcus (Fig. 56). This is usually accompanied by reduction in the thickness and complexity of the exine. Two subclasses exhibit these trends; Alismatidae and Zingiberidae. In both subclasses the presumably primitive families (Butomaceae and Bromeliaceae, respectively) have monosulcate pollen. The more advanced families of each subclass have inaperturate pollen, including the Najadaceae, Zannichelliaceae, Posidoniaceae, Cymodoceaceae and Zosteraceae of the Alismatidae, and almost all families in the Zingiberales (Strelitziaceae, Heliconiaceae, Musaceae, Zingiberaceae, Costaceae and Cannaceae). In many of these families the sporopolleninous exine is either absent or is a thin unstructured and unsculptured layer (Pettitt and Jermy, 1975; Schwanitz, 1967b). The Orchidaceae exhibits a similar trend toward the loss of the sulcus, but the transition from sulcate to inaperturate, exineless pollen occurs via the ulcerate type (Fig. 56). This is concomitant, not only with the loss of
Figs. 51-55. Fig. 51. Burmannia bicolor Hort.: SEM showing psilate exine and monoporate condition, Fig. 52. Burmannia bicolor. TEM showing thick tectum, columellate layer and the thick lamellated footlayer, x 11,625. Fig. 53. Cleisostoma racemiferum (Lindl.) Garay: TEM showing the sporopolleninous wall which surrounds the entire pollinium (SW) and can be found surrounding some of the pollen grains peripheral in the pollinium (P), X 13,125. Fig. 54. Laelia automnalis Lindl.: TEM ofnonacetolyzed pollen showing thick sporopolleninous wall which surrounds the pollinium, note granular structure underneath the thick rectum, x9000. Fig. 55. Cleisostoma racemiferum: TEM of nonacetolyzed pollen showing cytoplasmic connections (CC) between pollen grains in the interior of the pollinium, x6375.
368
Inoperturote Monods-Polyods
Multiaperturote eg. Triporates, Polyforotes
Monoporate ~V
,,,,
Ulcerote
/
Trichotomosulcote
Monosulcote
Fig. 56. Major evolutionary trends o f apertures in monocots. I. Alismatidean trend: monosulcate ~ inaperturate. II. Zingiberidean trend: monosulcate ~ inaperturate. III. Orchidacean trend: monosulcate ~ ulcerate ~ inaperturate. IV. Commelinidean trend: m o n o sulcate ~ ulcerate ~ monoporate. V. Arecidean trend: monosulcate ~ multiaperturate forms. VI. Liliacean trend: monosulcate ~ disulcates, trichotomosulcates, zonasulcates ~ multiaperturate. VII. Alismatalean trend: monosulcate ~ multiaperturates (polyforates).
the exine, but also with the aggregation of pollen grains into tetrads, massulae and pollinia. The most primitive orchid tribes, the Apostasieae and Cypripedieae, have monosulcate pollen usually shed in monads. In the intermediate tribes, the Orchideae and Neottieae, both monosulcate and ulcerate pollen occur and the mealy (sectile) grains are usually shed in tetrads or loosely packed pollinia (except in Cephalantherawhere they are shed in monads; Dressier and Dodson, 1960). In the Epidendreae, the most advanced orchid tribe, pollen occurs in tightly packed, waxy pollinia and is inaperturate. Pollen is exineless toward the interior of the pollinium, but
369
MONOCOTS
EXINELESS
eg. Cannoceae Orchidoceoe
/
,~.. ~...~.~,.;~..~:.~..;~
Stemono- type
eg.-H~modoroc~e
M_ayoca-type
:~..,:;.'l..V.-' %.:r162
.'$(.:r. ;~ f .S...l~, -I,.
.".'r
Arisoema-type
eg. Centrolepidaceae
Burmonnio - type
J
4. ~,.: :..::.,.; ",:.." ,'.,.:.~:: ~,.': :.'. :'
~,~ectote-imperforate
tectate-perforate/}
e.g. Butomoceae, Arecaceae, Apostasieae

Fig. 57. Major evolutionary trends of wall structure types in the monocots. The primitive tectate-columellate (perforate or imperforate) wall structure type, possibly derived from a Nymphaeacean-like ancestor with atectate or granular walls, gives rise to monocotyledonous atectate or granular walls and finally extreme reduction of the exine, in which it may be completely absent.
the pollen grains located peripherally in the pollinium may have a tectategranular sporopolleninous wall (Figs. 56 and 57). The reduction of the sulcus to a well defined pore via the ulcerate aperture type is another major trend and can be observed in the subclass
370
Commelinidae. The monosulcate aperture prevails in the Commelinales with the ulcerate aperture common in derived orders (e.g., Eriocaulales, Restionales, Typhales and Hydatellales; Cronquist, 1981). Monoporate pollen is observed in the most advanced orders of the subclass; Juncales, Poales and Cyperales. The trend from the monosulcate aperture via the ulcerate type, and subsequently to the monoporate condition, correlates with a shift from insect (e.g., some Commelinaceae) to wind pollination (e.g., Poaceae; Walker and Doyle, 1975) in this subclass. The last major evolutionary trend in the monocots is the increase in the number and types of apertures. This trend is exhibited in the subclass Arecidae, the order Liliales and in the order Alismatales. As in the primitive orders and families mentioned above, the monosulcate aperture characterizes the most primitive taxa, and the other monosulcate-derived aperture types (i.e., zonasulcate, trichotomosulcate, disulcate, and multiaperturate) are common in the advanced taxa; especially in the Liliales and Arecidae. Triaperturate pollen, a widespread aperture type in the dicots, is thus far known from only few monocot taxa: Sclerosperma mannii (Palmae) and Conostylis setosa (Haemodoraceae). The Alismatales is unique in the Alismatidae because multiaperturate forms prevail in a subclass where reduction of the sulcus culminating in the loss of the aperture is most common. Another notable exception is the genus Trillium (Liliaceae). This taxon is inaperturate in a family where multiaperturate forms commonly occur. The evolutionary trends in monocot aperture types parallel many of the trends observed in dicots, but different trends are predominant in the monocots. A conspicuous feature of dicot aperture evolution is the tendency towards an increase in the number and types of apertures, particularly the prevalance oftriaperturate pollen. This trend is observed in all of the dicot subclasses (Walker, 1974a). The Liliales, Arecidae, and A1ismatales exhibit parallel trends; however there is a notable dearth of triaperturate pollen types (known from only few monocot taxa, see above). Reduction of the sulcus is the most common trend among many monocot subclasses and families (e.g., Alismatidae, Zingiberidae, Commelinidae, and Orchidaceae). In many cases, the reduction or loss of the aperture, with the concomitant reduction in the thickness and complexity of the exine, occurs in hydrophilous taxa (e.g., Alismatidae) or in groups with elaborate or highly specialized pollinating mechanisms (e.g., Zingiberidae, Orchidaceae). The structure and chemical nature of the exine is ideally adapted for preventing desiccation of the gamete and for accommodating harmomegathic changes due to water loss (Muller, 1979). Adverse environmental factors influencing water relations of pollen are greatly diminished in hydrophytes and in pollen (or pollinia) that are efficiently transferred to a receptive stigma by a faithful pollinator. Thus a reduction
371
of the thickness and complexity of the exine and aperture might be expected to occur in these taxa where desiccation is no longer a problem. Pollen of the Commelinidae also exhibits a tendency toward reduction of the aperture, but in this case it is correlated with the shift from insect to wind pollination (Walker and Doyle, 1975), a situation in which the pollen is exposed to the desiccating effects of the environment. Pollen of the anemophilous commelinidean taxa have thick, imperforate, psilate exines (e.g., Poaceae), and the aperture is a small, well defined, operculate pore. This combination of apertural and exinal features is well adapted to prevent water loss during air transport.
Evolutionary Trends of Pollen Wall Structure in Monocots The paucity of data on monocot wall structure has prevented the observation of evolutionary trends. New data on pollen wall structure described above (18 families, 30 taxa) plus reports from the literature provide enough information to allow certain trends in pollen wall structures to be suggested. There are three unique aspects of pollen wall structure and evolution in monocots. First, in contrast to dicots (Walker, 1976), the tectate-columeUate wall structure is observed in the most primitive families and orders of monocots. Second, endexine is very rare in monocots and when present is different from the endexine observed in dicots. Third, in derived taxa in which the sporopolleninous exine is absent or reduced to a thin, unstructured layer, the intine is often thick and highly elaborated -- a situation not observed in dicots. Based on morphological data (Cronquist, 1981), monocots are thought to be derived from dicotyledonous nymphaeacean-like (ranalean) ancestors that diverged from dicots early in the evolutionary history of angiosperms (Daghlian, 1981; Muller, 1981). The occurrence of the monosulcate aperture cannot be construed as strong palynological evidence in favor of a common origin of monocots and dicots because there are a number of pre-Cretaceous and extant gymnosperms which also have monosulcate pollen and are not necessarily phylogenetically related to the angiosperms. Among the primitive families and orders of the five subclasses of monocots, the tectate-columellate wall structure predominates (e.g., Butomaceae, Arecaceae, Bromeliaceae, some Liliaceae, and the Apostasieae of the Orchidaceae). This situation might be interpreted as supporting an independent origin of the monocots. It might be expected that the palynological characteristics of the most primitive dicots would also be shared by the most primitive monocots i.e., the granular infrastructure, much
372
like the granular wall structure found in the Nymphaeaceae (Roland, 1965, 1968; Rowley, 1967; Ueno, 1962; Ueno and Kitaguchi, 1961), and similar to the wall structure observed in other primitive ranalean taxa (e.g., Magnoliaceae, Annonaceae). However, a transition from a nymphaeaceanlike ancestor to the primitive monocot can be palynologically envisioned as a shift from the granular or atectate wall structure to the tectate-columellate wall type. This occurs concomitantly with the shift from the nymphaeacean-like morphology to a primitive monocotyledonous morphology. The tectate-columellate wall structure, although considered derived by Walker (1976), occurs even among ranalean taxa and appears to be an early palynological development among dicotyledonous angiosperms (Doyle, 1969, 1973). Thus the hypothetical derivation of the tectatecolumellate wall structure in primitive monocots from the granular walled nymphaeacean-like ancestor parallels evolutionary developments that occurred relatively early in dicots. This places the primitive tectate-columellate monocots on the same evolutionary level as the derived tectatecolumellate ranalean dicots, lending support to the dicotyledonous origin of monocots. Another interesting aspect ofmonocot exine evolution is that a number of more advanced taxa have granular or atectate walls (e.g., Centrolepidaceae, Haemodoraceae, respectively), as well as a variety of other wall types (Fig. 57). These wall types are derived from the primitive tectatecolumellate walled monocots (Fig. 57). This trend coupled with the proposed origin of the primitive monocot tectate-columellate wall type from a nymphaeacean-like ancestor with granular or atectate walls, is identical to the evolutionary trends proposed by Walker (1976) for dicots. He also considers the granular or atectate walls in some morphologically advanced dicots to be secondarily derived from the tectate-columellate type. In the monocots, however, reduction and simplification of the exine is even more extreme and culminates in certain orders with exineless pollen (e.g., the Najadales, Zingiberales and the Orchidaceae; Fig. 57). It is interesting to note, however, that in many of the taxa with exineless pollen or pollen with a thin, unstructured sporopolleninous wall, the intine is often very thick and greatly elaborated, possibly assuming many of the functions of the exine. Another major difference between dicot and monocot pollen wall structure is the rarity of endexine in monocots, and in those instances where presence of the endexine is reported it is different from dicotyledonous endexine. Skvarla and Larson (1966) note the presence of a distinct granular endexine in Zea mays pollen and similar endexinal layers have been observed in Saxofridericia (Rapateaceae), Mayaca (Mayacaceae), Dios-
373
corea (Dioscoreaceae) and possibly Spathiphyllum (Araceae). In all cases this wall layer is thin and inconspicuous, and unlike endexine in dicots it does not become thickened in the apertural region. Instead, it is the footlayer that often exhibits apertural thickening in monocots (e.g., Skvarla and Larson, 1966; Figs. 22 and 23). Until more is learned about the functional significance of this proposed monocotyledonous endexine, and about the selective pressures operating on pollen wall structure in monocots, the differences in morphology between dicot and monocot endexine will be open to speculation. It is interesting that endexine is found in the advanced taxa of both monocots and dicots (absent from most primitive taxa) and a homologous wall layer has yet to be demonstrated in gymnosperms (Zavada, ms. in preparation). In summary, the evolutionary trends observed in wall structure and aperture types in the monocots parallel those proposed for dicots. This suggests that the selective pressures affecting pollen wall and aperture evolution in these two groups have been similar. However, the frequencies of the different wall and aperture types in monocots differ from those in the dicots, further suggesting that there are differences in the organismenvironment relationships.
Acknowledgments
I wish to express my thanks to William L. Crepet, Ralph Collins, Henry N. Andrews (University of Connecticut), Thomas N. Taylor (Ohio State University), and Charles Daghlian (University of Oklahoma) for their many helpful suggestions and discussion.
Literature Cited
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APPENDIX
List o f s p e c i m e n s s t u d i e d Collector/# Aponogetonaceae

Aponogeton echinatus Roxb.
Herbaria
Locality
Saldanha, 16082 Eames, May 25, 1939 Zavada, 512 Wolfe, 209 Parker, 66.65 Pritzel, s.n. Specht, 737 Zavada, 511 Zavada, 510 Tillett & Tillett, 43904 Standley, 42130
US 2616535 CONN CONN CONN CONN US 406021 US 2094678 CONN CONN US 2621046 US 52664
India Connecticut Connecticut Surinam New York Australia Australia Connecticut Connecticut British Guiana Costa Rica
Araceae Arisaema triphyllum (L.) Schott Bromeliaceae Billbergia sp. Burmanniaceae Burmannia bicolor Hort. nutomaceae Butornus umbellatus L. Centrolepidaceae Centrolepis aristata Roem. Centrolepis exserta Rts. Commelinaceae
Dichorisandra thyrsiflora Mikan.
Cyanastraceae Cyanastrum cordifolium Oliver Cyclanthaceae Stelestylis stflaris (Gleas.) Had. Sphaeradenia irazvensis (Cuf.) Having
379
APPENDIX I
Continued Collector/# Dioscoreaceae Herbaria Locality
Dioscorea polygonoides Uline.

Haemodoraceae
Pfeifer, 2746
CONN
Puerto Rico
Anigozanthos flavus
D.C. ex Red.
Edwards, s.n. Wolfe, s.n. Weatheley, s.n. Edwards, s.n. Christenson, 435 Wolfe, 234
CONN CONN CONN CONN CONN CONN
Connecticut Australia Connecticut New Jersey Connecticut Surinam
Conostylis setosa Lindl. Lachnanthes caroliana

(Lam.) Dandy
Lophiola aurea Ker-Gawler Wachendorfia paniculata Burro. Xiphidium coeruleum Aubl.

Iridaceae
Iris pseudacorus L. Sisyrinchium californicum Air.

Liliaceae
Collins, June 12, 1 9 2 7 Zavada, 503
CONN CONN
Rhode Island Connecticut
Crinum americanum L.
Mayacaceae
Curtiss, 5052
CONN
Connecticut
Mayaca aubertii Mich.

Orchidaceae
Bartlett, 1160
CONN
(Not Given)
Cleisostoma racemiferum
(Lindl.) Garay
Christenson, 365 Christenson, 365
CONN CONN
India Mexico
Laelia automnalis Lindl.

Rapateaceae
Schoenocephihum cucullatum
Maguire
Saxofrederica regalis Schumb. Stegolepis angustata Gleason

Stemonaceae
Maguire, 41901 Williams, 14389 Steyermark, 92599
US 2279340 US 859466 US 2584557
Venezuela Venezuela Venezuela
Stemonajaponica (BI.) Miq.

Taccaceae
Coll. ig. n.
US 1310132
Japan
Tacca macrantha Redonte Tacca integrifolia

(non Kew-Gawler) Schrank
Zavada, 506 Zavada, 507
CONN
CONN
Connecticut Connecticut

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THE BOTANICAL REVIEW

COMPARATIVE M O R P H O L O G Y OF MONOCOT POLLEN

COMPARATIVE M O R P H O L O G Y OF MONOCOT POLLEN

THE BOTANICAL REVIEW

Classification and Phylogeny of Monocots

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

THE BOTANICAL REVIEW

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

THE BOTANICAL REVIEW

COMPARATIVE M O R P H O L O G Y OF MONOCOT POLLEN

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

THE BOTANICAL REVIEW

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

THE BOTANICAL REVIEW

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

THE BOTANICAL REVIEW

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

THE BOTANICAL REVIEW

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

THE BOTANICAL REVIEW

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

THE BOTANICAL REVIEW

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

THE BOTANICAL REVIEW

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

THE BOTANICAL REVIEW

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

Evolutionary Trends of Pollen Aperture Types in the Monocots

THE BOTANICAL REVIEW

Multiaperturote eg. Triporates, Polyforotes

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

eg. Cannoceae Orchidoceoe

.'$(.:r. ;~ f .S...l~, -I,.

e.g. Butomoceae, Arecaceae, Apostasieae

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

THE BOTANICAL REVIEW

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

THE BOTANICAL REVIEW

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

THE BOTANICAL REVIEW

List o f s p e c i m e n s s t u d i e d Collector/# Aponogetonaceae

COMPARATIVE MORPHOLOGY OF MONOCOT POLLEN

Dioscorea polygonoides Uline.

CONN CONN CONN CONN CONN CONN

Connecticut Australia Connecticut New Jersey Connecticut Surinam

Conostylis setosa Lindl. Lachnanthes caroliana

Lophiola aurea Ker-Gawler Wachendorfia paniculata Burro. Xiphidium coeruleum Aubl.

Iris pseudacorus L. Sisyrinchium californicum Air.

Collins, June 12, 1 9 2 7 Zavada, 503

Rhode Island Connecticut

Mayaca aubertii Mich.

Christenson, 365 Christenson, 365

Laelia automnalis Lindl.

Saxofrederica regalis Schumb. Stegolepis angustata Gleason

Maguire, 41901 Williams, 14389 Steyermark, 92599

US 2279340 US 859466 US 2584557

Venezuela Venezuela Venezuela

Stemonajaponica (BI.) Miq.