Pollen Myriophyllum

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    The Pollen Morphology of theHaloragaceae with Reference toTaxonomyJ. Praglowski aa The Palynological Laboratory of the National Science ResearchCouncil ,Published online: 01 Sep 2009.

    To cite this article: J. Praglowski (1970) The Pollen Morphology of the Haloragaceae with Reference toTaxonomy, Grana, 10:3, 159-239

    To link to this article: http://dx.doi.org/10.1080/00173137009427397

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  • Grana 10: 159-239, 1970

    THE POLLEN MORPHOLOGY OF THE HALORAGACEAE WITH REFERENCE TO TAXONOMY

    J. Praglowski

    (Received December 12, 1970)

    ABSTRACT Praglowski, J. (The Palynological Laboratory of the Na- tional Science Rexarch Council). The po//efi niorphology oj the Haloragacrar with rrjereiice to taxorioriip. Grana 10: 159-239, 1970.-Pollen morphological in- vestigations of the family Haloragaceae were carried out using light microscopy supplemented by electron micro- scopical studies. An attempt was made to correlate the pollen morphology with the taxonomy of th: family. The pollen morphological data provide clear support in favour of the division of the family into two separate families, respectively, Ihloragaceae and Gunneraceae. noth subfamilies, i t . , Haloragoideae Schindl. (excluding a few species in the subsection Spongiocarpus Schindl.) and Gunnxoideae Schindl., are stenopalynous. Important arguments concerning pollen morphology, gross morpho- logy, ecology, and type of pollination, favour a regroup- ment of the Hnlorngis spxies belonging to the subsection Spongiocarpus, which is related to the genus Latidoniu Lindl. Eventual fusion of these IIalorugis species in th: subsection Spongiocarpus with the genus Lorrdoiria is worth consideration.

    The Haloragaceae is an interesting plant group from a palaeobotanical point of view. A review of the literature concerning fossil findings of the Haloragaceac and sporo- morphs of the Stemmn Normapolles Pf.,- some of which most probably could have bten related with the Halora- gaceae, point out th: interest of this family to palaeo- botany.

    The Haloragaceae, which have an antarctic origin, Seem to have been more widely distributed during the Tertiary than at the pres-nt time.

    INTRODU(XI0N

    The studies of the family Haloragaceae were undertaken in order to describe the pollen mor- phological features in a detailed and systematic way, as well as to correlate the pollen morpholo- gical results with the taxonomy of the family. Furthermore, it seemed desirable to determine the

    11 - i05800

    extent to which pollen morphological data could contribute to clarifying thc somewhat contro- versial ideas concerning thc taxonomy within ihe Haloragaceae. Initially the intention was to con- sider all species of the Haloragaceae, but diffi- culties in obtaining the polleniniferous material of certain species have prevented this. Thus, the examination has been limited to about 80% of the total species of the Haloragaceae.

    Sincc the beginning of the 19th century ihe Haloragaceae has been of interest to several botanists, see Schindler (1905) with references to older literature; Skottsberg (1928), hlattfeld (1932), Pokrovskaia (1958). Tardieu-Blot (1965), Raynal (1965 and 1967) and van der Meijden (1966 and 1969). The works of Raynal (1965 and 1967) and of van der hfeijden (1966 and 1969), which deal with different smaller groups of the family Haloragaceae, have been of much use to the author. The work of Schindler (1905), however, is comparatively modem and completc and therefore is used by the author as a basis for the taxonomical clrtssification and for the layout of th: present paper.

    The pollen morphology of recent Haloragaceae has also interested several scientists. Illustrations and pollen morphological descriptions of sporadic Halorngaceae species, based upon light-micro- scopic observations have appeared in the publica- tions of Erdtman (1952, for references to older literature), Bennet (1838), Petersen (1893), Schind- ler (1904), Erdtman ( 1 9 4 3 ~ . 1952, 1963, 1969), Cranwell (1953), Ikuse (1956), Fsgri & Iversen (1 950 and 1964), Pokrovskaia (1 955), Dyakowskn (1959), Erdtman et al. (1961 and 1963). Huang (1967). and Praglowski (1 969).

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  • 160 J . Praglowski

    As part of a detailed study of pollen of the subfamily Haloragoideae, it seemed worth-while to compare pollen of this subfamily with pollen of the Detulaccae, Corylaceae, hlyricaceae, Cas- uarinaceae, Oenotheraceae etc., since species be- longing to these families usually produce pollen, which to a certain extent, resemble pollcn en- countered in the Haloragoideae.

    Another aim of the present work is to em- phasize the role of the fossil Halaragoid sporo- morphs in the late Cretaceous and Tertiary floras and to attemt to judge their relation to the recent Haloragaceae.

    Sporadic fossil findings of Haloragaceae are known from the Tertiary and the Quaternary. Fossil sporomorphs resembling pollen of some Iinloragis species wcre also encountered in the Upper Cretaceous and in the Tertiary.

    T H E DISTRIBUTION

    The Haloragaceae contain seven genera with about 200 species. At the present time the distribution is in temperate and subtropical regions of all continents (a few species of Myriopliylhirn are found in the Arctic). The two largest genera Hnloragis Forst. (with about 90 species), and Grrnriera L. (with about 50 species) are mainly represented in the Southern hemisphere. These two genera constitute together a large regional palaeosubantarctic complex with a distribution concentrated around Australia, New Zealand and Tasmania; about 70% of the Haloragaceae occur in this area. The distribution of IIalorngis and Griririera points to an antarctic origin of the family (Schindler. 1905).

    Only 12 species of Haloragaceae, most of them belonging to the cosniopolitan genus Myriophyf- l i r r t i L., occur in the Northern hemisphere. The genus Ilalorrigis. with the exception of two species from Juan Fernandez Islands and one from New Calcdonia, occurs only in the Australia-New Zea- land-Tasmania area. The Ifaloragis specics are terrestrial, comparatively small (maximum height about one meter) herbs or subshrubs, which usu- ally grow in relatively humid areas.

    The species of the comparatively independent, terrestrial genus Gritiricra L. occur mainly in the Southern hemisphere. A few Girnriera species are found, however, north of the equator in South America and in the Philippines. Gritiriera plants Crana 10

    vary considerably in their gross morphology. The leaf size ranges from a few centimeters to about one meter in length. According to Bader (1961), the genus Girtittern can be divided, using gross morphology, into small-leaved subantarctic species, and the palaeo- or ncotropic species which are provided with large leaves. Subantarctic Girnrtem species arc concentratcd in New Zealand (9 spe- cies) but also occur in Chile, Argentina, and Juan Fernandez Islands. Girtinera is vertically distri- buted from sea level to an altitude of 4 500 meters in the South American Cordilliera.

    The most important centre of the cosniopolitan genus, hfyriopliyllrirri L., consisting of more or less submerged hydrophytic species, is to be found in Australia (about 15 species). About eight Myrio- pliylliirri species are found in temperate North Amcrica. The remaining species are spread al- most world-wide. A few species of hfyriopliylliirri are arctic and subarctic. Out of all the Halora- gaceae genera certain species of Myriophyllrirri have the most northerly distribution.

    The tcrrcstrial genus Loitdotiin Lindl. composed of three specics, occurs only in Australia.

    The genus Proserpiriacci L., to which only two small hydrophytic-helophytic spccies belong, is found in North America, hlexico and progres- sively southwards to Guatemala.

    The genus Lnirrcriibergia Berg., composed of about 20 species, occurs in Australia, New Zea- land, tropical Asia, Madagascar, KCunion, Africa, and in tropical America. Larirerribergin species are small, prostrate, stoloniferous herbs, which grow in humid, sunny areas. These species have an amphibic biology (Kaynal, 1965). i.c. each year they pass through a cycle of inundstion, followcd by an exundation, and so change from hydro- phytic to helophytic status.

    MATERIAL AND hlETHODS

    A. Lnboratory work Only polliniferous material from herbarium spe- cimens was used in this investigation. The material was obtained from the following herbaria: AD,

    BARIUhl, BISH, Bhl, BR, CHICO, CHR, GB, HO, JE. K, L, MASS, NSW, P, PERTH, S, U, UPS, us, w.

    An effort was made to ensure that the pollini-

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  • Pollrir iirorpho/ogy o/ Ildoragnccac 161

    were stained with I % aqucous uranyl acetaie and lead citrate (Rcynolds, 1963).

    ferous material used had, ;IS far as Possibk corn- and correct general information. material

    fro,,, unreliable Sources was omitted. In Tables I-VI, all species are provided with the data nec- essary to refer the species investigated to its ori_cinal herbarium sheet. h b s t Of thc species investigated were determined by botanists who have \vorked on the taxonomy Of the Haloraga- ccac, I was fortunate to be able to usc the pollini- fcrous material originating from herbaria spcci- mcns, which had been determined by Schindler, Skottsberg, St. John etc., as wcll as receiving an important part of the material determined by taxonomists presently active and interested in Ha- loragaceac (Dr hladame A. Raynal for African, Dr K. van der Meijden for Malaysian, D r B. Sparre for South American, and Dr A. Orchard for Australian species).

    Pollcn was acetolysed (Erdtman, 1960) and embedded in glycerine-jelly (melting point about 65C) composed of: phenol crystals 3.5 g, granular gelatine 25.0 g, H 2 0 dest. 87 g. and glycerine 75 g. Since thc Haloragaceae gencrally have minute flowers and usually very much reduced floral organs, they arc thus not easy objects to collect or pretreat as polliniferous material. Micro- acetolysis describcd by Punt (1963) was occasion- ally iised for samples which were extremely poor in pollen. Chlorination after acetolysis was not cmployed.

    The material used for electron-microscopic in- vestigations (sectioning, single-stage replicas, and scanning), with a few exceptions, was also acc- tolysed. The single-stage replicas were produced in the Botanical Institute, Univcrsity of Stock- holm. Carbon films according to the method for single-stage carbon replicas (Rowlcy 6: Flynn, 1966), wcrc produced in a Baltzers high vacuum evaporator. Polliniferous material dcstined for scanning electron micrographs was washed with water, treated with 50-959h ethanol, and dried. The dry pollen was transferred to aluminum ob- ject holders covcrcd by a thin film of Aqua Dag mixed with glue and acetone. The polliniferous niaterial for sectioning was dehydrated using an alcohol-acetone series, and embedded in Epon- Araldite (hlollcnhaucr, 1964). Epon was poly- mcriscd at 80C for about 24 hours. Ultrathin sections were cut with a diamond knife on a LKB Ultrotorne I. The sections were transfcrrcd on 200-mesh grids on a formvar membrane and

    B. Microscopy, photorriicrogr~plr)., atid polloi nrorphologicnl data Light microscopic studies were done using a Leitz-Laborlux microscope, equipped with an apo- chromatic oil immersion objective ( x 100 N.A. 1.32) and G.F. x 10 eyepieces. The slides were studied immediately after preparation. hleasure- ment errors caused by eventual swelling of pollen grains embedded in glycerinc-jelly were then avoid- ed. For each spccics a minimum of 10 measure- ments were taken for both the polar axis and the equatorial diameter. Small, morphologically aberrant, pollen grains occur in most species of Halorugis (Praglowski, 1969); thcse probably ster- ile and more or lcss abnormal pollcn grains, exhibiting a vcry wide range of morphological and size variation, have not been included in ihE size column in Tables I-VI.

    A Lcitz-Ortholux microscopc with an apo- chromatic oil immersion objective ( x 160, N.A. 1.40) and G.F. x 10 cyepiecc was used for the photomicrographic work. Photomicrographs were taken with Ilford orthochromatic G-30, or with llford panchromatic K-20 plates (9 x 12 cm). The original negatives were x 1 500.

    Transmission electron micrographs wcrc taken with a Zeiss EM 9 A electron microscope on Agfa-Gevaert 23 D 56 film. Scanning electron micrographs were taken with a Cambridge In- strument Co. LTD Stereoscan microscope.

    The results of the pollen morphological studies consist of:

    1. Generic diagnoses. 2. Tables I-VI, containing dctailed pollen mor-

    phological data for all species investigated with- in a genus (N.B. both points 1 and 2 are mainly based on light microscopical observa- tions).

    3. Pollen morphological commcnts for most of the genera, based upon the electron microscopic studies.

    4. Plates 1-14 illustrating pollcn grains of some species of each genus.

    Thc generic diagnoses predominantly describe all the pollen morphological features common to all species belonging to a given genus. These diagnoses are therefore comparatively short. The

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  • 162 1. Praglowski

    characters included in the generic diagnoses usual- ly have not been repeated in the Tables I-VI, dealing with the detailed pollen morphology of the individual species. (Decimal micron values employed for some measiircments in the generic diagnoses should not be considered as absolute values bot were used to give interpolated in- formation on the occasionally very minute dif- ferences existing between some pollen morpho- logical elements).

    Tables I-VI show the pollen morphological de- tails of cach species investigated. The tabular method was chosen in order to present the pollen morphological features in a concentrated form. Since the interspecific pollen morphological varia- tion in the Haloragaceac is vcry slight, the tabular method scemcd to be a more convenient and ra- tional form of dcscribing the specific pollen mor- phology than writing individual specific pollen diagnoses. The species in Tables I-VI have been grouped according to the taxonomical system of Schindler (1905). In addition to the taxa indicated by Schindlcr I.c., in sections and subsections in Tables I-VI, there are also species synonymous with those described by Schindler, and species dc- scribed morc recently than Schindlers publication. The latter species and synonyms are indicated by an asterisk in Tables I-VI.

    Tables I-VI, contain all general data necessary to locate the herbarium specimen from which the polliniferous material was sampled. These tables include also coded information for the NPC system (Erdtman & Stnka, 1961). several columns giving data on different pollen morphological features, and a column special morphological characteristics where typical features are briefly emphasized. Tables I-VI list the distribution of all species investigated, and are also to be con- sidered as the list of species investigated.

    The pollen morphological comments following cach of the six tables are based upon the elec- tron microscopic studies. They elucidate the pollen morphological data in a detailed way giving the size relation between different components of the exine, as well as an illustrative picture of its fine structure. The numerical figures obtained for the supratectal processes and tectal perfora- tions in the pollen morphological comments were calculated per 50 (standard area investigated) using the electron micrographic replicas and scanning electron micrographs. Grana 10

    As a result of comparatively difficult gross morphology of the Haloragaceae, many species have been published under two or more than two different names. A separate list of synony- mous species for each genus, based on the litera- ture, is given in alphabetical order following each of the six Tables.

    THE TAXONOhlICAL POSITION OF THE HALORAGACEAE

    Bentham 6: Hooker (1 862) referred the Halora- gaceae to the order Halorageac, one of the 11 orders of the Polypctalae. Haloragcae is placed among the orders Bruniaceae, Rhizophoreae, Combretaceae, and hlyrtaccae.

    According to Wettstein (1935) the Haloragaceac belong to the large order hlyrtales, as one of 20 families constituting this order. The Haloraga- ceae are preceded by the hlyrtaceae, Punicaceae, Melastomaceae, Lythracexe, and Oenothcraccae and are followed by Gunncraceae and Hippurida- ceae. A significant fact which is worth emphasiz- ing is the elevation of Gunneroideae to family status. Wcttstein did not believe that the Halora- gaceac could have a genetic origin among some of the families constituting the order hlyrtales. The Oenothcraceae and Lythraceae are most closely related to the Haloragaccae.

    Hutchinson (1959) reduced the relatively large order Myrtales by establishment of a small order Lythrales within which the Haloragaceae was placed together with the Lythraceae, Onagraceae, Trapaceac, and Callitrichaceae. Usually the Halo- ragaceae is considered to be a family of the Myrtales and in the case of Hutchinsons classifi- cation the author makes a rather peculiar excep- tion. The genera Halorcrgis, Giirtrtera, and Hip- pirris are included in the Haloragaceae.

    The superorder Myrtifloreae established by Takhtajan (1 959), included the orders hlyrtales and Halorngales. He placed the Haloragaceac, the Gunneraceae and the Hippuridaccae in the small order Haloragales, closely related to the Myrtales. The Haloragaceae are considered by Takhtajan as the most primitive family among the Haloragales, a view supported by the floral morphology and the embryology. Takhtajan strongly supports the segregation of the Gunnera- ceae and the Hippuridaccac from the Haloraga- ceae.

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  • Polleri rriorplrology of Haloragncccic 163

    Section b. Pleianthus Schindler Subsection u Trachyphylluni Schindler Subsection p Sclcrocalyx Schindlcr Subsection y Cercodia Schindler Subsection 3 Digyniuni Schindlcr Subsection E Trihalorrhagis Schindler Subsection C hlcioncctes (K. Br.)

    Subsection 71 Tetraptcryx Schindler Subsection 0 Tripteryx Schindler Subscction 1' Spongiocarpus Schindlcr

    Schindler

    ChadefaLid & Enibcrgcr (1960) placed the h- 1or;igaceac and the Gunncraceae within the order

    'hfyrtales as two c~osely related, but separate fa- milies. Among the 16 families included by these allthors in the hlyrtales, the families hiyrtaccae, Ocnotheraccac, Hippuridaceac, and Callitrichaceac are most closely related with the Haloragaceac and the Gunneraccae.

    hlelchior, in Engler's Syllabus (1964) subdivided the hlyrtales into three suborders, i x . , hlyrtineae, Hippuridineae, and Cynomoriineae. The Halora- eaccae is placed among 16 other families into thc hlyrtineae. This family is considered by hlcl- chior as reduced, but a t the same time, as being a niore developed fraction of the Oenotheraceae.

    Schindlcr (1 905) includcd the nionotypic genus hfezicllri Schindl. in the tribe Halorageae, con- sisting of fivc genera. This genus has not been placed within the Halorngaceae by any of the above mentioned authors.

    To summarize, it would seem desirable to placc the Haloragaccae within the order hlyrtales. The family undoubtedly should be considered as clos- ely related to the Ocnotheraceae, from which, however, it differs by lacking the intraxylary phloem and by having abundant cndosperni in the seeds. Wettstein's, Chadefaud's, and Emberg- er's as well as Takhtajan's classification, resulting in the separation of Grrririern from Ifnlorngis, by the establishment of a separate and more advanced family Gunncraccae, seems to be justified.

    SYSTEMA FAMILIAE accordirrg to Schiridler ( I 905)

    HALORRHAGACEAE

    Subfamily, I. HALORRHAGOIDEAE Schindler

    Tribe 1. HALORRHAGEAE Schindler

    Genus 1. LOUDONIA Lindl.

    Genus 2. HALORRHAGIS Forst. Subgenus I. Pseudohalorrhagis Schindlcr Subgenus 11. Euhalorrhazis Schindler

    Section a. hlonanthus Schindlcr Subsection u Rhagocalyx Schindler Subsection p Leptocalyx Schindler Subsection y Trachycalyx Schindler Subsection 3 hlacrogoniphus Schindler Subsection E Lamprocalyx Schindlcr

    Genus 3 . MEZIELLA Schindler

    Genus 4. LAUREMBERGIA Berg.

    Section a. Apopleura Schindler Section b. Ditetraplcura Schindlcr

    Subgenus I. Indolaurenibcrgia Schindler

    Subgenus 11. Afrolaurembergia Schindler

    Genus 5 . PROSERPINACA L.

    Tribe 2 . hlYRIOPHYLLEAC Schindler

    Genus 6. MYRIOPHYLLUhf L. Subgenus I. Eumyriophyllun~ Schindler

    Section a. Pentaptcris P. DC. emend. 0. Kizc.

    Subsection Q Spirophylluni Schindler Subsection p Pelonastes Hook.

    Subsection y Spondylophyllum Torr.

    Subsection 3 Leiocarpium Schindlcr

    Subsection u Trachycarpaeuni

    Subsection /I Spondylastrum Torr. e t

    Subsection y Ptilophylluni Nutt.

    f. emcnd. Schindler

    et G r a y

    Section b. Tcssaronia Schindlcr

    Schindler

    Gray

    Subgeniis 11. Brachytheca Schindler Subgenus 111. Dicarpium Schindler

    Subfamily 11. GUNNEROIDEAE Schindler

    Genus 7. G U N N E R A L. Subgenus I. hlilligania (Hook. f.) Schindler Subgenus 11. hlisandra (Comni.) Schindler Subgenus 111. Pseudo-Gunnera (Oerst)

    Schindler Subgenus IV. Perpensum (Burm.) Schindler Subgenus V. Panke (hlol.) Schindlcr

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  • colpatc pollen grains rare. NPC: 643 (543). - Polar axis 1622 /I, equatorial diameter 16-27 p . Pollen grains suboblate, oblate-spheroidal, or oc-

    Exine usually equal in thickness throughout, Gciteric dingnosis 1.5-2.5 p thick, or occasionally slightly thicker Pollen grains isopolar, radially symmetrical, per- at poles than at the equator. tectate, 6-colpate, usually tenuimarginate. Four Sexinc thicker than the nexine. Tectum ihin,

    THE POLLEN hlORPIiOLOGY OF THE GENERA

    LOUDONIA Lindl. PI. 1 casionally spheroidal, usually peritreme.

    Tables I-VI exhibit the pollen morphological data of the six genera investigated, bxed on the light micro- scopicsl observations. The distribution of species is presented in Tables: 11, 111, V, and VI. Following abbreviations are used in Tables I-VI:

    is0 isopolar irrtrrni pollen grains intermediary occ. occasionally para paraisopolar between goniotreme and corrrp. comparatively gonio goniotreme pollen grains peritrcme t. +- distinct prritr peritreme pollen grains up. aperture + comparativcly distinct

    prof. protruding - indistinct s. slightly

    Table I. LOUDONIA Lincll.

    Shape L Size Puncta Spccial morphologi- N P C PIE index i n p Colpi.(/i) in L.O. cal characteristics

    L. aiirrn Lindl.

    s.n. (US) W. Austr., A. hlorrison

    - S.W. Austr., Kock s.n.

    Det. Schindler (W)

    L. rod (Endl.) Schlechtd. W. Austr., G. E. Brockway sen. (PERTII)

    - S. Austr.. E. Pritzel s.n.

    Dct. Schindler (W)

    L. brhrii Schlccht. S. Austr., C. R. Alcock C 36 A (AD)

    - S. Austr., C. hl. Eardly s.n. (S)

    - Austr. Vict., No coll.

    Herb. hfus. Vindobon. 136560. Dct. Schindler (W)

    613 543

    643

    643 (543)

    543 (443)

    643 (543)

    543 (643)

    543

    so 82

    so 87

    so- OBL-SPH 88

    so- ORL-SPH 88

    so 84

    0 B L-S PI i 90

    SPH 100

    23 x 28

    22 x 25

    1 7 x 2 0

    17 x 20

    22 x 26

    1 8 x 2 0

    16x 16

    1 5 x 0 3 + Comp. thick exine (3 / i ) , grains s. gonio

    1 6 ~ 0 . 5 + +

    l o x 1 -

    8 x 0.5 -

    1 1 x 1 + + Comp. thick exinc (3 I0

    6 x 0 . 5 i-

    5 x 0 s -

    SYNONYhlOUS SPECIES

    Loirdonia arirea Lindl. L. f/aresccns Drummound L. cifrina F. v. hi.

    Glischrocaryon roei Endl. t. r o d (Endl.) Schlechtd.

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  • Iolleri rriorphology of Halorogrrcerte I65

    Plate 1. For explanation, see p. 187.

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  • 166 J . Praglowski

    provided with relatively distinct perforations and less distinct supratectal processes. Infratectal ba- c h a distinct in optical cross section.

    Nexine compact, its distal surface non-smooth. Colpi 5 X 1 p, slightly ingrooved.

    Polleti rriorphological coiriirietits bnseri iipoti clectrori microscopy

    for Loiirioriia aimxi I-indl. PI. 1, Fig. a-c.

    T h e exine-1 (ektexine seiisii Faegri 1956) com- prises about 95 % , whereas the exine-2 (endexine setisi~ Faegri I.c.) only about 5 % of the total of the exine thickness. Exine-2 is more electron dense than the cxine-I.

    l h e tectum (supratectal processes included) is about 0.6 p thick. Supratectal spinules (verrucae) are about 0.2 /i high, and about 0.2 fi wide at their bases. There are about 16 processes and 10 tectal pcrforations in Loitdot~i(i ruci per 50 p 2 . The perforations are cylindrical and their diameter is about 0.3 /i (half of the thickness of the tectuni).

    in dia- meter and about l long (twice as long as the thickness of the tectum). Infratectal horizontally elongated segments of a layer, fused niidway with the bacula, are encountered. These segments are probably not a continuous supplementary layer but should perhaps be considered as niore or less isolated elements of rudimentary character.

    The foot layer is about 0.4 thick, slightly thinner than the tectuni.

    The exine-2 seems to have no interruptions and is less compact in the apertural areas, where the exine is not thickened.

    The infratectal baciila are about 0.5

    HALORRHAGIS Forst. PI. 2-9

    Gerieric diagtiosis

    (with exception of H. rtioriosperitia F. V. hl., H . rncciriosa Labill., and I I . liicnsi hleiden in the subsection Spongiocarpus Schindl.)

    Pollen grains usually paraisopolar, or occasionally isopolar, radially symmetrical, pcrtcctatc, predom- inantly 4-5-colpate (about 8 0 % of species), or 4-5-porate (about 20 % of species), crassimarginate. frequently aspidote. Seven, or more than seven aperturate pollen grains occur rarely. Anomo- theme pollen grains are encountered in a number Gram 10

    of species. NPC: 443, 543, (643), (343)-(444), (544), (644), (344). - Polar axis 15-31 p , equa- torial diameter 18-40 p . Suboblatc pollen grains are most common (about SO%), oblate less com- mon (about 18 %), and oblate-spheroidal pollen grains are rare.

    With the exception of most species in the sub- section Spongiocarpus Schindl., all species usually have pollen grains provided with apertures which are never ingrooved. Goniotrenie pollen grains occur in about half the species, whereas peritreme pollen grains occur in about 1 0 % . The latter are usually found in the section Pleianthus. In about one third of the species of this genus pollen grains with an anib intermediate between that of gonio- trenie and peritreme pollen grains are encount- ered (cf. I I . ckpressa, PI. 4 , Fig. i).

    T h e paraisopolarity, which frequently occurs, depends upon three pollen morphological features:

    1. difference in convexity of the poles, which results in an almost plano-convex shape of pollen grain when observed in lateral view,

    2. difference in length of the polar axis on either side of the equatorial plane with the axis o n the more convex pole about twice the length of that on the less convex pole,

    3. a thicker exine at the less convex pole than at the more convex pole.

    T h e exine consists of the sexine, comprising the tectum, one or niore than one infratectal ba- cular layers and the nexine. The total thickness of thc exine at the centre of the niesocolpia in polar view, the supratectal processes and the in- nermost nexine details included, is 1.5-6 p . The tectum usually is as thick as the height of the in- fratectal bacula. It niay be smooth, slightly un- dulated, or occasionally provided with flattened, more or less irregularly shaped, supratectal in- sular protrusions (cf. H . heteropliyilri, PI. 5, Fig. k). hlinute supratectal processes (usually spintiles or verrucae) as well as tectal perforations were ob- served in several species by means of the L.O. analysis. Thcse processes, as well as the tectal pcrforations, have becn observed more or less distinctly in most species of Haloragis. Infratectal bacula are delicate, usually indistinct in optical cross section, and appear as a light bacillar zone.

    T h e nexinc varies in thickness. It is compact or, frequently, more or less loose, lamellated, or

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  • Pollett itiorphology of Haloragaccnc 167

    a sponge-like structure composed of short segnicnts with comparatively weak StIUCtUral cohesion. In several species the innermost nexine lamella (Iamellae) constitute n more or less com- pact inner sphere (cf. If. heterophylla, PI. 5 , Fig. g), which is usually attached in a few places to the surrounding nexine and seems to be ana- logous with the exine-2 (endexine seiisii Faegri 1965). In normally developed pollen grains, which usually lack a separated inner sphere, the inner- most nexine lamella (Ianiellae) is frequently slight- ly detached from the surrounding nexinc, but still adhcrcs to it in some places. In surface view, these separated lamellar fragments appear as ir- regular, polyangular scales, about 2-5 / i in dia- meter (cf. If. Iieferophyllu, PI. 5 , Fig. h-i). In most species the apertural margins are thickencd, due to the occurrence of a few short, compact, lamellar-like nexine zegments in this area.

    With the exception of a few species in the subsection Spongiocarpus, Ifaloragis has aspidote pollen grains with simple apertures. When ob- served in polar view and in optical cross section, the limits of the proximal part of the apertures, can be easily distinguished at the aperture bascs. The distal part of the apertures usually is smaller than the proximal part, the latter is also usually more circular.

    The most common type of aperture is a short colpus with a thickened margin. The thickenings of the aperture margins, when observed in polar view, are 2-6 p. The apertures, due to their thickened margins, frequently protrude and in these pollen grains thcre is ;i considerable aper- tural depth (i.e., the distance between the distal and proximal h i t of the aperture). The most common size of the colpi is 4-5 x 1.5 p. Loxo- colpate pollen grains i.e., pollen grains provided with colpi which in rclation to the equator are obliquely orientated, have been encountered in some species. They usually cut the equator a t about 45 and frequently alternate to thc right and left.

    The pori are usually slightly elliptical, 3 x 2 p. Circular, or comparatively elongated pores, bor- derline between colpiis and porus, frequently oc- cur. The shape of the pori may be more o r less irregular or occasionally elongated in the equa- torial direction. The pore margins resemble those of the colpi. They are occasionally provided with short fissures.

    Specific rlicigiiosis for Haloragis itiotiospcritia 1. V. hi. in the sub- section Spongiocarpus Schindler

    Pollen grains isopolar, radially symmctrical, per- tectate, 5-6-colpate, tenuimnrginate. NPC: 543, 643. - Polar axis 19 p, equatorial diameter 22 t i . Pollen grains suboblate, peritreme, in lateral view frequently slightly rectangular in shape (with flat- tened apocolpia).

    Exine of the samc thickness throughout, about 2 p thick.

    Sexine slightly thicker than the ncxine. T h e tectum thin, provided with comparatively distinct tectal perforations and less distinct supratectal processcs. Infratcctal bacula relatively distinct in optical cross section.

    Nexine compact, its distal surface non-smooth. Colpi about 5 x 1 ,I(, usually slightly ingrooved.

    Polleti itiorphological cottiitieiits bascrl iipoti electron rtiicroscopy (not including Haloragis species of the subsection Spongiocarpus)

    Two morphologically different pollen grain types occur simultaneously in flowers of most s p x i e s in the Ifaloragis. T h e degree of pollen wall cohe- sion, resulting in a compact exine in one iype, and in a more or less loose exine in the second type of pollen grains, is the main feature disting- uishing the pollen morphology of both types. (Cf. e.g., H . gossci, PI. 8). Pollen grains with a n exine exhibiting features intermediary between those mentioned above, are frequently encountered.

    1. Pollen grains with comparatively conipact ex- ine and simple infrabacular structure.

    Exine-1 comprises about 9 0 %, whereas the exine-2 about 10% of the total of the exine thickness.

    T h e tectum including supratectal dctails is about 1 .O ,I( thick. Supratectal processes, spiniiles or ver- rucae are about 0.2 ti high. They are about 0.2 p wide at their bases. The number of processes and tectal perforations vary in different species of Ifolorcrgis. Calculated per 50 p2, there are about 15 spinules and 10 tectal perforations in If. salso- loirles, 14 spinules and 16 perforations in H . iiiascitierratia, 16 spinules and 18 perforations in If. conferfifolia, and 19 spinules (perforations in- distinct) in If. heterophylla. As it appears from

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  • 168 J . Proglowski

    Table 11. HALORRHAGIS Forst. Haloragis polkn-types occuring in Table 11: ~ ~ ~ ~ - l = ~ ~ ~ d ~ ~ i ~ - t ~ ~ c (pollen grains similar to that Of Loudonia), Type-2 = T y p e 4 = pollen grains 4-apcrturate, TYpe-5 = pollcn grains 5-apcrturatc, Type-N - pollen grains not radially symmetrical, Of Type-1 and Type-2, are also found in Table v (hlyriophyllum).

    Pol a r- Shape & Size ity Amb N P C P/E index Type in It Colpi (11) Pori oc)

    Subgenus 1. Pseudohalotrhagis Schindler

    11. noditloso (Ncss) Walp. Para Interm W. Austr., C. A. Gardner 6481 (PERTH)

    Subgenus 11. Euhalorrhagis Schindler (non 0. Ktze.) Scctio a. hfonanthus Schindler

    Subsectio cz Rhagocalyx Schindler If. ronfcrfifolia F. v. hf.

    W. Austr., C. A. Gardner s.n. Det. C. A. Gardner (PEKTH)

    If. elnra A. Cum. S. Austr., Kraehen-

    buehl 1173 (AD)

    11. fcniris cap. schindl.

    If. pycriostnchyo P. v. M.

    Subsectio p Leptocalyx Schindler I / . scubrn (Koenig) Bcnth.

    China, Hook. 897. Dct. Schindler (WV)

    w r . clongafa Schindl. var. obbrcrigra Schindl.

    Austr. Victoria. Josephine 11. nrrziann Schindl.

    E. Tilden 919 (K) H. rcrbro Schindl.

    11. rilloso Schindl.

    If. fctrugyna (Labill.) Hook. f. Hongkong, K. L. Rechinger s.n. Det. Schindler (W)

    S. Austr.. 1. Opuel s.n. Det. Schindl. (W)

    var. grnirins Schindl. va r. dccrrmberrs Sc h i n d 1. var. hispidu Schindl. var. bnccobrn Schindl. var. serrnfa Schindl. var. birallosa Schindl. var. incona Kirk var. di//rcsa Hook. f.

    H. trucrioidcs A. Gray Tasmania, H. E. Combcr

    1721 (K)

    Grana I0

    Para

    Para

    is0 OCC

    Para or is0

    Para

    Gonio

    Gonio

    Gonio

    Intcrni

    443 (543)

    543 413

    443 (543)

    613 (543) (743)

    543

    so 81

    so 78

    0 71

    so-0 75

    so 77

    4 21 x26

    5 , 4 , 7 22x28

    15 x 21

    27 x 36

    22 x 29

    4 x 0.5

    4 x 0.5

    4 x 0.5

    6 x 1 5 x 0.5

    4 x 1

    Para Gonio 643 543 SO-0 6, 5.7 27x37 6 x 1 7s

    Para Gonio 643 543 SO 6 , 5 , 7 21x31 6x0.5 or 78 is0

    Para Interm 543 613 SO 5 , 6 31x40 6 x 2 i a 4 x 0.5

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  • Pollen rriorplrology of Hnlorngnccac I69

    hlejoncctes-typc (pollen grains similar to that of Haloragis brownii = hfeionectes brownii), Type-3 = pollen grains 3-aperturate. ~ ~ ~ ~ - p = p o l l e n grains only poratc, Type C=pollen grains only colpatc. All pollen grain types in Table 11, with the cxception

    -~ Exine thickness (10 Tectal details

    In polar view in LO.

    Supra- Aper- tcctal

    hleso- turcs At pro- Special morphological Austra- N. Zea- Tas- N. Gui- hlalay- colpia margins polcs Puncta cesses characteristics lia land manis nea sia Other

    + 2.5 3.5 2, 1.5 - -

    2.5 3 .54 2.5. 1.5 + - Teclum usually + undulated

    + 2 3.5 2. 1.5 - -

    2.5-3 5-6 2, 1.5 + -

    2-3 3

    ap. prot., colpi occ. arranged in a figure 8 around the equator

    2.5, 1.5 - - Ap. s. ingrooved, not + prot.

    2-3 4-6 2. 1 + - Ap. prot.. mcsocolpia + camp. short

    + - Ap. prot., mesocol- + pia comp. short

    +

    +

    +

    China

    China Japan

    China Japan

    2-3 4-5 2.5. 1.5 + - t +

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  • 170 J . Praglowski

    Table 11. (Cori/iriirmd.)

    Polar- Shape & Size ity Amb N P C PIEindex Type in/( Colpi (11) Pori (1')

    - S. Austr.. J. Jackson 388 (AD)

    H. reiccrioides DC. ex Schlechtd. S. Austr., A. E. Orchard 1810. Det. A. E. Orchard (AD)

    H. ufnea Schindl.

    H. aggrcgata Buchan. var. dffiisa (Ilook. 1.)

    var. incana (A. Cunn.) Schindl.

    Schindl. H. Iongi/alia Schindl.

    N. Guinea, hiaille s.n. Det. Schindl. (W)

    H. acanfhocarpa Brongn. Austr. Northern Terr..

    R. L. Specht 21 1 (PERTII)

    *H. elata Hook. 1. * H . gonocarpiis Sprcng *if. giitmii Hook. f . *if. incnna Walp. *H. isomcrn Parker * H . Ieptoflreca F. v. hi.

    H . holconensis hlerr.

    Lobl. 448 (K) Philippines,

    *- N. Guinea,

    R. Pullen 5035 (K) H . philippinensis hl err.

    Sumatra. J. A. LBmhp 9771 (L)

    Sumatra. C. G. G. J. van Steenis 9083 (L)

    *H. procunrbcns Cheesem. * H . chinensis (Lour.) hferr.

    Crutwell 1036 (K) *If. palaricnsis Tuyana. * H . secitnda Ridl.

    *-

    Papua. E. G.

    Papua, E. G. Crutwell I123 (K)

    *if. gjelleriipii \Vent. f. Grana I0

    P a n

    is0

    Para

    is0

    occ.

    OCC.

    Para

    is0

    Para

    is0

    occ.

    occ.

    Is0

    para

    Is0 o r para

    Para

    is0

    Para

    is0

    occ.

    occ.

    occ

    Para

    Gonio

    Gonio

    Gonio

    Gonio

    543 443

    543 643 (743)

    543 (443)

    543 (643)

    so 77

    so 78

    so 80

    so-0 75

    5.4 24x31 4.5~1

    5.6 28x36 5x1 4 x 0.5

    16x20 3x03

    24x32 4x1

    23x29 (5x 1.5) 4x 3 Pcritr- 414 (443) SO 4 Interm 79

    Peritr- 443 (444) SO 4 22x28 5x2 (3x2) Interm 79

    29x36 8x0.5 Interm 443 (543) SO 4 (643) (343) 81

    Peritr- 443 (543) SO 4 28x36 10x1 Interm 78

    Gonio 543 643 SO 5,6,7 24x32 4x1 76

    Para Interm 413 543 SO 4,5 23x29 6 x 1 occ. 79 5 x 0.5 is0

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  • Pollen tnorphology o/ iiulorogoceae 171

    Exi'ne thickness (p) Tectal details

    In polar view in LO.

    Supra- Aper- tectal

    hleso- tures At pro- Special morphological Austra- N. Zea- Tas- N. Gui- hlalay- colpia margins poles Puncta cesses characteristics lia land mania nea sia Other

    4-5 2.5, 1.5 - - 3

    - 4 5 3 ,2 +

    + +

    + +

    2 2-3 2, 1 - - grains comp. small. + exine comp. thick

    2-3.5 4-5 2, 1.5 + - 7-8 colpate and ano- + motreme grains occur, nexine much differen- tiated

    1.5-2 3-4 1.5, 1.5 + + - 1.5, 2

    2-2.5 3 4 2, 1.5 + - 2. 2

    2 3-4 2, 1.5 + -

    2-2.5 2.5-3.5 2, 1.5 + -

    2.5 4-4.5 2, 1.5 - -

    2 3

    Ap. not prot., 5-3 treme and anomotre- me grains occ. occur

    Ap. not prot., 5-3 trerne grains occ. occur

    tremc grains occ. occur

    Exine comp. thick, colpi long. ap. mar- gins thickened

    Tectum und., anomo-

    'rectum und., ap. prot., grains occ. loxocolpa te

    Exine comp. thin

    +

    + + Philip- pines

    + + Philip- pines

    + Philip- pines

    + Philip- pines

    + China

    4

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  • 172 J . Proglowski

    Table 11. (Contburd.)

    Polar- Shape 8; Size ity Amb N P C PIE indcx Type in11 Colpi (10 Pori @)

    ' H . neniorosa Went. f. N. Guinea, L. J. Brass

    I1672 (K)

    Para

    *If. fruricosa Went. 1. Pa 13 Papua, E.G. Cruttwell 1441 (K)

    *if. sangiiinea hlerr. & Perry

    'it. niicropli~llrrm Hoogl.

    *H. str//ri~ticosn Gibbs.

    Subsectio y Trachycalyx Schindlcr

    i f . reroirici/olia Sc h i nd I.

    Subsectio 6 hiacrogomphus Schindler II. cordigera (Endl.) Fenzl Para

    W. Austr., Dick & E. Pritzel 187 (PERTH)

    - Para W. Austr., E. Pritzel o x .

    s.n. Det. Schindl. (W) is0

    if. pirliyoidrs (Ness) Bcnth. Is0 Austr., Prciss 1224. or

    Det. Schindlcr (W) para

    H. scordioitlrs Uenth.

    H . pirsilla R. Br.

    H . solsoloides (Reichb.) Para Benth. N. Guinea, Sicber occ. 249. Det. Schindlcr (W) is0

    - Para Austr. N. S. W., hlajor ox .

    Vickary s.n. (K) is0

    if. inrricata Uenth. Para W. Austr., Drummond s.n. Det. Schindl. (W)

    H . tricliostacli~a Uenth.

    if. dffirsa Dick

    if. rotimdi;olia Benth. \V. Austr., A. hiorrison 9557 (K)

    W. Austr., R. D. Royce 100 (I'erth)

    if. r i d s Bcnth. W. Austr., Capt. A.

    D. Smith s.n. (K)

    Grana 10

    Conio

    Gonio- Interm

    Gonio

    Gonio

    Interm- Peritr

    Gonio

    Gonio

    Gonio

    443 (543)

    443 543

    543 (443) (643)

    443 (543)

    443 (343) (444)

    443 543 (544) (444)

    443 543

    543 443

    so 81

    so 77

    0 63

    so 78

    so 82

    so 78

    so 77

    0 73

    4 2 2 x 2 1 5 x 1

    4,5,7 2 3 x 3 0 6 x 1 4 x 1

    5 17x27 5x0.5

    4 2 4 x 3 1 9x0.5 G x 0.5

    4 2 4 x 2 9 5 x 1 (3x2) 4 x 0.5

    4,5,7 2 4 x 3 1 5 x 1 ( 2 x 1 3 ) 3 x 1

    4.5 2 7 x 3 5 8 x 2 6 x 1

    5,4 14x19 3x0.5

    2 2 x 2 8 3x0.5 4 x 3 Para Gonio 443 444 SO 4 occ. (343) (344) 78 is0 Para Conio 443 444 SO 4 2 5 x 3 2 4 x 1 4 x 3 occ. (343) (344) 78 is0 Para Interm 443 (444) SO 4 2 5 x 3 0 4 x 1 (3x2) occ. (543) (544) 83 is0

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  • PoUeri inorphology of Hnlorogaccne 173

    Exinc thickness 00 Tectal details

    In polar view

    Meso- turcs At pro- Special morphological Austra- N. Zen- Tas- N. Gui- hlalay- colpia margins poles Puncta cesses characteristics lia land mania nea sia Other

    in LO.

    Supra- Aper- tcctal

    1.5, 1 - - 3 4 4 Ap. prot., tectum und.

    k i n e comp. thick, + 2-3 4-5 2.5, 1.5 - - ap. prot.

    2-3 4-5 2.5, 2 - - Tectum occ. s. und., + ap. prot., grains distinctly goniotrcme

    + 2-3 4-5 2, 1.5 - -

    2.5-3.5 4-6 2.5. 2 + - Exine comp. thick, ap. + prot., anornotreme grains occur, pori rare

    2.5-3.5 5-7 2.5, 2 + - Exine comp. thick. ap. + prot., anomotreme grains occur, colpi long

    + 2 3 1.5, I - -

    2.5 4-5 2, 1.5 + - Exine comp. thick +

    3 4-6 2.5, 2 + - Exine comp. thick +

    +

    +

    2-2.5 3 4 2.5, 1.5 + +- Ap. not prot., ap. + margins comp. thin

    Crana 10

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  • 174 J . Proglowski

    Tablc 11. (Contintred.)

    Polar- Shape & Size ity Amb N P C PIE index Type in/[ Colpi (11) Pori 01)

    Subscctio t Lamprocalyx Schindler

    Japan lionshu, Iiiroe 12635 H . mirrontlra (Thunb.) R. Br.

    Det. Schindler (1)

    N. Zealand. Travers s.n. Det. Schindler (W)

    H. drpressi (A. Cunn.) Walp. Tasmania, Burbidge 3287

    (K)

    *

    Para or is0

    Para or is0

    Para

    Para N. Zealand, 11, E. Connor s.n. Det. H. E. Connor (CIiR)

    occ. is0

    var. serpyllijolin

    var. tnontatri (Hook. 1.)

    var. nggregatn (Buchanan) Kirk

    var. spicata (Petrie)

    (t1ook.l.) Benth.

    Benth.

    Schindl.

    Gonio

    Gonio

    Interm

    Peritr

    443 543

    543 643

    643 (543) (743) (843)

    543 443 (544) (444)

    so 4, 5 23x28 79

    so 5, 6 2 6 x 3 4 76

    so 6 30x39 77

    so 5.4 21x25 84

    5 x 1.5 3 x 0.5

    5 x I 4 x 0.5

    6 x 1 5 x 1

    5 x 1.5 (3x2) 4 x 1

    I I . tiniflorn Kirk

    var. genrrina Schindl. N . Zealand, King 549 (K)

    var. bibrnctrolata (Col.) Schindl. N. Zealand, Colenso s.n. (K)

    var. spicata (Petrie) Schindl.

    * H . ritriodora Walp.

    H. niininia Col.

    ' H . ttiontana Hook. f. Tasmania, Hook. 1. s.n.

    Det. Hook. f. lsotypc (UPS)

    *iI. rernicosa Walp. *H. scrpjlli/olia (Hook. f.)

    Wolp. Austr. NSW., Kaspicw 88

    * H . spicota Petrie * H . tenello Brongn. * H . pnrtcidcntata Hosakawa

    Gratin 10

    Pan Interm- 443 so 4 23x29 5 x 1 peritr 79 4 x 0.5

    Para Interm- 443 so 4 2 5 x 3 0 4 x 1 occ. peritr a2 is0

    Para Gonio 643 0 6, 7 21 x33 4x0.5 or 64 is0

    Para Gonio 643 (743) SO 6 26x34 5x0.5 occ. (543) 76 is0

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  • Pollen riiorphology of Haloragaceae 175

    Tectal details in LO.

    Exine thickness

    In polar view

    hieso- turcs At pro- Special morphological Austra- N. Zea- Tas- N. Gui- hlalay- colpia margins poles Puncta cases characteristics lia land mania nea sia Other

    Supra- Aper- tectal

    2-2.5 4 2, 1.5 -t + - Tecturn smooth c i- +

    2.5 4-5 2, 1.5 + + - 6-treme grains rare, -t- + + tcctum smooth

    2-3

    2

    3.5-6 2, 1 - -

    2.5 1.5, 1 + +

    - 3 3.5 2, 1 +

    2.5-3 4-6 2.5, 1.5 + -

    2-2.5 4 2. 1.5 + +

    i-

    Ap. margins not t thickened

    Ap. not prot.

    Ap. not prot. +

    Ap. distinctly prot., -f- momotrerne grains occur

    Anomotrcme grains + occur

    + +

    + +

    +

    c

    +

    + China India Japan Formosa

    + China India Japan Formosa

    I!? - 705890 Cratia 10

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  • 176 J . Proglowski

    Table 11. (Contirrited.)

    Polar- Shape & Size ity Amb N P C P/E index Type in/ ' Colpi (1') Pori (1')

    Sectio b. Pleianthus Schindler

    Subsectio a Trachyphyllum Schindler

    H. glaiira Lindl. Austr. NSW.. T. & J.

    White 1745 (K)

    H. hctcroplijllo Brongn. S. Austr.. C. K. Alcock C 17 B (AD)

    - S. Austr., A. Orchard 310. Det. A. Orchard (AD)

    S. Austr., A. Orchard 316. Det. A. Orchard (AD)

    S. Austr., A. Orchard 308. Det. A. Orchard (AD)

    tar. ccratoplijllo (Zahlhr.) Schindl.

    va r. roprcolicornis Sc h i n d 1. Queensland, Hubbard 4092. Det. Schindl. (W)

    Schindl. var. gluitci/olin

    var. ospcro (Lindl.) Schindl.

    w r . rioido Schindl.

    H. pirinoigida Gray

    H . r i d i s Schindl. H. jolioso Bcnth.

    s.n. (PERTH) W. Austr., N. T. Burbidge

    'If. crspcro Lindl.

    748 (AD) S. Austr., D. N. Kraehenbuehl

    ' H . ccrotoplijlla Zahlbr.

    H. jiliJortiiis Gray

    Subsectio /? Sclerocalyx Schindler

    H. prosfroto Forst. N. Caledonia, 11. S. hlc. Kee 5234 (K)

    H. plotjrarpa (hlurr.) Schindl.

    Grana 10

    Para or is0

    Para

    is0

    Is0 or para

    1 so or para

    Para or is0

    occ.

    Para

    is0 occ.

    Para

    Para

    Para

    is0 occ.

    Pcritr- intcrm

    Gonio

    Gonio

    Gonio

    Gonio

    Gonio

    Gonio

    Gonio

    Gonio

    443 (444)

    443 543 (444) (544)

    543 (643)

    443 543 444 544

    543 544 (443) (444) (643) (644)

    543 (643) (443)

    443 (543)

    443 543

    443 (444) (343) (344)

    so 4 86

    so 4, 5 76

    so 5 78

    so 4.5 78

    so 5 80

    0 5 73

    so 4 83

    so 4. 5 84

    0 4 74

    21 x 2 6

    25 x 33

    28 x 26

    25 x 32

    28 x 35

    5 x 1 4 x 1

    5 x 2 4 x 1

    5 x 0.5 4 x 0.5

    4 x 1.5

    5 x 1.5

    24x33 5xO.5 4 x 0.5

    25 x 35

    22 x 26

    4 x 1

    4 x 1

    (2 x 1.5)

    (4 x 3)

    4 x 3 3 x 3

    4.5 x 3.5

    23 x 32 4 x 1.5 (3.5 x 2.5)

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  • k i n e thickness Tectal details

    In polar view

    hieso- tures At pro- Special morphological Austra- N. Zea- Tas- N. Gui- hlalay- colpia margins poles Puncla ccsses characteristics lia land mania nea sia Other

    in LO.

    Supra- Aper- tectal

    1.5-2 2-2.5 2, 1.5 + + Ap. not prot. +

    2-2.5 3-4 2.5, 1.5 + - S. concave rerocolpia +

    2-2.5 4-5 2, 1.5 4- - Anoxotrcmc and -+ Ioxocolpatc grains occur

    2.5-3 3-4 1, 1 + + - THO pollen types, + (porate. exice comp. thin; colpatc, exine comp. thick)

    3 1.5, 1 + - Exine comp. thin, 7-8 + treme and anomo- treme grains occ. occur

    1.5

    2-2.5 3-5 2.5, 1.5 - -

    2

    I .5

    3 4 2, 1.5 - -

    3 4 1.5, I + -

    +

    Ap. not prot. +

    +

    i

    +

    1.5-2 3 -4 2, 1.5 + - N. Cale- donia

    Graria I0

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  • I78 J . Proglowski

    Table 11. (Confhrued.)

    Polar- Shape & Size ity Amb N P C PIE index Type in/ ' Colpi ( / I ) Pori @)

    Subsectio 7 Cercodia (hfurr.) Schindler

    11. erecta (hlurr.) Eichl. N. Zealand, L. D.

    H. exalata F. v. hi.

    hfoore s.n. (CIIR 87922)

    W. Austr.. Nepcau River. (K)

    H. Inccis Schindl. Austr.. NSW. noorman s.n.

    Det. Schindl. (W)

    H. pedicrllntn Schindl.

    H. enrtilaginea Cheesem. N. Zealand, Baglin s.n. (CHR 18171)

    '11. nlura Jacq.

    s.n. (S) N. Zealand. S. Berggren

    var. cartilqinra Cheesem.

    H. altrrn(folia \Valp. *H. rcrcorlin Ail.

    J. Fernandez Isl. Collector? ( S )

    *H. colcnsoi Skottsb. N. Zealand, L. I). hioore s.n. (K)

    '11. icsefolii Salisb.

    * H . itiass/itcrmis Skottsb. hias Afuera, Skottsb. 1216 Del. Skottsberg (S)

    * H . masatierrana Skottsb. hias 3 Tierra, Skottsb. 304. Dct. Skottsbcrg (UPS)

    * H . telrngonin I'Herit.

    Para

    is0

    Par3

    is0

    Is0 or para

    occ.

    occ.

    Para

    Para or is0

    Para o r is0 1 so

    Para

    Para or is0

    Subsectio 8 Digynium (Ktze.) Schindler

    H. digpi i Labill. Para

    Det. nentham (PERTH) is0

    H. scrra Brongn. Para

    SW. Austr., hiaxwell s.n. occ.

    Austr. NSW.. C. Stuart s.n. (K)

    Para

    is0

    H. scopsrii Fenzl Para W. Austr.. Dummong s.n. o r Det. Schindl. (W) is0

    - Austr. NS\V., J. hl. hiaiden sen. H L B 910 199-1402 (L)

    occ.

    Gratta 10

    Gonio

    Gonio

    Gonio

    Gonio- interm

    Interm

    Interm- peritr

    Peritr

    Gonio

    Interm

    Gonio

    Gonio- interm

    Interm

    Interm

    443 (543) (343)

    443 (543)

    444 443

    443 (543)

    543 541 (6-13) (644) (443) (444)

    443 (444) (543) (544)

    443 (543)

    413 543 (444) (544)

    444 (443)

    443 (543)

    444 (443)

    413 (444)

    343

    so 87

    so 70

    0 73

    so 81

    so 80

    so 8 1

    so 83

    so 82

    so 18

    0 13

    so 80

    so 83

    so 78

    4

    4

    4

    4

    5

    4

    4

    4, 5

    4

    4

    4

    4

    3

    2 7 x 3 1

    2 4 x 3 4

    24 x 33

    27 x 33

    28 x 35

    26 x 30

    25 x 30

    26 x 32

    27 x 35

    24x 33

    2 6 x 32

    24 x 29

    26 x 33

    4 x 1 3 x

    4 x

    3 x

    .5

    .5

    4 x 1.5

    3 x 1

    4 x 1

    4 x 0.5 3 x 0.5

    2.5 x I

    S X 1 4 x 1

    5 x 2

    5 x I 4 x 0.5

    3 x 2 2 x 2

    4 x 3 3 x 2

    (3 x 2)

    (3 x 2)

    4 x 3 4 x 2.5

    4 x 2.5 2 x 1.5

    (3.5 x 2)

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  • Pollen niorpliology of Hnloragnccac 119

    &c thickness (/o Tcctal details In p o h r view

    hieso- turcs At pro- Special morphological Austra- N. Zea- Tas- N. Gui- hialay- colpia margins poles Puncta cesses characteristics lia land mania nea sia Other

    in LO.

    Supra- Aper- tectal

    2 3-4

    2 4-5

    2 3-4

    2 3.5

    2 2.5-3

    2, 1.5

    2, 1.5

    2, 1.5 292

    2. 1.5

    1.5, 1

    Ap. prot., ap. shape occ. irregular

    Ap. not prot.

    Ap. not prot.. exine comp. thin

    Ap. not prot., ap. shape occ. irregular

    +

    +

    +

    +

    + +

    2 3 2, 1.5 + + Ap. not prot., exine + + comp. thin

    2 2.5-3 2, 2 + + - Dist. isopolar, ap. not + prot.

    2 3.5 2, 1 + - Ap. not prot.

    1.5-2 2.5, 3 1.5. 1 + + Ap. not prot. 1.5, 1.5

    2-2.5 4-5 2, 1.5 - -

    - 2-2.5 3-4 2, 1.5 -

    2 3 2, 1.5 - - hp. not prot.

    2 3-4.5 2, 2 + - 3-trcme only 2. 1.5

    Juan Fernandez Isl. Juan Fernandcz Isl.

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  • 180 J . Pragloivski

    Table 11. (Carilirtitcd.)

    Polar- Shape & Size ity Amb N P C P/E index Type in/( Colpi ( I t ) Pori ( I t )

    ii. acrileolara Bcnth. W. Austr., A. hlorrison

    11246 (K) ' H . nrrrcronafa Benth.

    Subsectio E Trihalorrhagis Schindler

    H. Izrandra F. v. M. W. Austr., hl. Koch

    var. scrrata Schindl.

    var. infcgrifolia Schindl.

    \V. Austr., Diels &,

    2259 (K)

    1. renrtifolia Uenth.

    Pritzel 189 (PERTH)

    - W. Austr., Kock s.n. Det. Schindl. ( \ )

    Para or is0

    Para

    is0 occ.

    Is0

    para

    Is0

    para

    occ.

    occ.

    Subscctio hfcionectes (R. nr.) Schindlcr I f . broirnii (Kook. f.) Schindl.

    Austr. Victoria, A. C. Bcau- glehole 5861. Dct. H. 1. Aston (L)

    H. brerilobg Schindl. n.sp.

    Subsectio 7) Tctrapterix Schindler

    H. coronopifolia Schindl. S. Austr., IIill 101.

    Det. Schindl. (W) H. acufangula E v. M.

    7543 (PERTH) W. Austr., C. A. Gardner

    ii. odontocarpa F. v. hf. Austr. NSW. Wanaaring,

    Glenfield Vcterin. Res. Station 39/1738 (NS\V)

    W. Austr. T. E. II. Aplin 2367 (PERTH)

    'ii. ciliata Black S. Austr., A. Orchard

    1871. Det. A. Orchard (AD)

    Subsectio 0 Tripterix Schindler

    S. Austr., T. R. N. Lothian If. gossci F. v. M.

    1706 (AD)

    Grarra 10

    Is0

    Para

    Para

    Para

    Para

    Para

    Para

    Peritr

    Interm

    Interm

    Interm

    Interm

    Gonio

    Gonio

    Gonio- intcrm

    Gonio

    Gonio- intcrm

    Interm- peritr

    443 (543)

    443 543

    443 (543) (444) (544)

    443 444 (543) (544)

    444

    443 543

    443 543

    443

    443 543

    443

    443 (543)

    0 69

    0 12

    so 88

    so 80

    so 76

    so 17

    0 12

    0 13

    so 18

    so 17

    so 19

    4

    4. 5

    4,2

    4.2

    4.2

    4, 5

    4, 5

    4

    4, 5

    4

    4

    22 x 32

    23 x 32

    29 x 33

    24 x 30

    25 x 33

    33 x 43

    18x25

    22x30

    23 x 29

    26 x 34

    27 x 34

    5 x 1 3 x 1

    5 x 1 4 x 0.5

    6 x 2 3 (5x 3) s x I (4x3)

    3x0.5 5 x 3

    3 x 2 2 x 2

    5 x 1

    2.5 x 0.5

    4 x 1

    6 x 1 3 x 1

    4 x 1 3 x 0.5

    6 x 2 4 x 1

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  • Pollen niorpliology of Halorngnceac 18 1

    Exjnc thickness 0') Tcctal details

    i n polar view

    hfcso- tures At pro- Special morphological Austra- N. Zea- 73s- N. Gui- Malay- ,-olpia margins poles Puncta ccsscs characteristics lia land mania nea sia Other

    in LO.

    Supra- Apcr- tectal

    2 2.5-3 2, 2 + - Ap. not prot. + 2. 1.5

    2-2.5 3-4 2, 1.5 + - Ap. not prot. +

    1.5-2 3 2, 2 + + Exine comp. compact, + 1.5, 1.5 convex mcsocolpiu,

    ap. as in Subsectio hfeionectes

    1.5-2 2.5-3 2, 2 + + Exine comp. compact, + 1.5, 1.5 convex rnesocolpia,

    ap. as in Subsectio hlcionectcs

    2 2 2 . 2 + 1- Exine comp. compact. + convex mesocolpia, ap. simplc, lacking cndoapertural part, dist. is0

    + 2.5-3 4-5 2.5-1.5 + -

    2 2-3 1.5. 1 - - Grains comp. small +

    1.5-2 2.5-3.5 2, 1.5 + - Ap. not prot. +

    2-2.5 3-4 2, 1.5 + - Ap. not prot.

    1.5-2 2-3 1.5, 1 + - Ap. not prot.

    +

    +

    +

    2 3.5 1.5, 1 + + + Exine comp. thin, ap. + not prot.

    Grnna 10

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  • 182 J . Piaslowski

    Tablc 11. (Confinrred.)

    Polar- Shape & Size ity Arnb N P C PIE index Type in11 Colpi (11) Pori (p)

    - W. Austr.. N. T. Bur- bidge 1052. Det. N. T. Burbidge (PERTH)

    If. tripnocarpa F. v. hi. W. Austr., A. S. George 693 (PERTII)

    W. Austr., C .A. Gardner s.n. (PERTH)

    Subscctio t' Spongiocarpus Schindler

    If. monosprrnia F. v. hi. Austr. NSW.. W. Baucrlcn s.n. Det. Schindler (W)

    - Austr. NSW., F. v. hlueller

    s.n. 903 364-188 (L)

    H. raceniosz Labill. W. Austr.. R. D. Royce 6225 (PERTH)

    SW. Austr.. T. E. H. Aplin

    var. angiirrifoliu Schindl.

    var. bartertenif (F. v. M.)

    2620 (PERTH)

    Schindl.

    H. stricla R. Br. Austr., R. Br. s.n.

    Dct. Schindl. (P)

    H. paniculata (R. Br.) Benth. W. Austr., A.

    hiorrison 141 18 (K) - W. Austr.. Morrison 133 (K)

    H. lanceolata (R. Br.) Bcnth. *H. baitcrlenii F. v. hi. *H. Iucasi hiaiden

    Austr., W. Blakcly s.n. (S)

    Para Intcrm 443 (543) SO 4 2 6 x 3 1 4 x 1 82 3 x 0.5

    Para Gonio 543 443 SO 5.4 2 4 x 3 0 4 x 1 82

    Para Gonio- 443 (543) SO 4 22x27 3x1.5 occ. interrn 81 i s 0

    Is0 Peritr 643 (543) 0 SPH 6, 1 2 0 x 2 2 Sx I 91 4 x 1

    Iso Peritr 543 643 SO 5.6.1 18x22 6x1.5 81 s x I

    Is0 eritr 543 643 SO 5, 6, 1 19x22 7 x 1.5 86

    Is0 Peritr 543 643 SO 5.6.1 17x 19 7 x 1 89 6 x I

    Is0 Gonio- 443 (543) SO 4 17x22 2x0.5 occ. interm 17 para Para Gonio 643 (543) 0 6.7 2 2 x 3 0 4 x 1

    (443) 73

    Para Gonio 543 643 SO 5 . 6 30x38 5xO.5 79 4 x 0.5

    Is0 Peritr 643 (543) SO-0 6, 1 2 0 x 2 6 4 x 1 (443) 7s 2.5 x 0.5

    Grana 10

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  • Pollen r,rorphology of Halotngtrceae 183

    Exine thickness ( p ) Tectal details

    In polar view in LO.

    Supra- Aper- tectal

    Meso- tures At pro- Special morphological Austra- N. Zea- Tas- N. Gui- hialay- colpia margins poles Puncta cesses characteristics lia land mania nca sia Other

    2-2.5 3-4 1.5, 1 + + Ap. not prot. +

    2 3 - 4 1.5, I + - Ap. not prot. +

    2 2.5 1.5. I + - Ap. not prot. +

    2 1

    2 I

    2 1

    2 1

    2. 2 + - Exinc compact, apo- + colpia flattened, grains tenuimarginate, of Loudonia-type

    colpia flattened, grains tcnuimarginate. of Loudonia-type

    tenuimarginate, of Loudonia-type

    tenuimarginate, of Loudonia-type

    2 . 2 + - Exine compact. apo- +

    2, 2 + + Exine compact, grains +

    2, 2 + + + Exine compart, grains +

    2 2-2.5 2, 2 - - Grains comp. small +

    3-4 5 3, 2 + + + Ap. prot., concave + mesocolpia, tectum undulated

    2-2.5 3.5 2.5, 1.5 + - Ap. prot., concave + mesocoIpia, tectum

    undulated. grains comp. large

    2.5-3 1.5 2.5, 2.5 - - Exine compact, grains tenuimarginate, of Loudonia-type, comp. large

    Craria I 0

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  • 184 J . Proglowski

    SYNONYhlOUS SPECIES

    Halorogis acantliocarpa Brongn.

    II: aggregota Buch.

    H. aggrcgata var. diffusa (Hook. 1.) Schindl.

    ti. aggregata var. incann (A. Cunn.) Schindl.

    II. alternif4lia \Valp.

    H. aspera Lindl.

    If. baucrlenii F. V. hi.

    H. broirnii (Hook. f .) Schindl.

    ti. cortilaginea Cheesem.

    H. cercodia Ait.

    H. chinensis (Lour.) hlerr.

    ti. citriodora Walp.

    H. depressa (A. C u m ) Walp.

    H. depressa var. scrpylli/olia

    ti. depressa var. iiiontana (Hook. f.) Benth.

    Ii. diffitsa Dick

    H. digyna Labill.

    (Hook. f.) Benth.

    II. erccta (hlurr.) Eichl.

    ti. filiformis Gray

    Ii. grrnii Hook. f .

    H. holconensis hlerr.

    if. heterophylla Brongn.

    H. Ieptotheca F. v. hl. ti. palartensis Tuyama

    H. tetragyna Hook. 1. I I . tetrafynn F. V. hl.

    Ii. tetragyna Hook. 1. var. diffiisa Hook. 1.

    H. tetragyna Hook. f . var. incana Kirk H . incana Walp.

    ti. alata Jacq.

    II . ceratophjlla Zahlbr. II. pinnatijida Hook. f . (non Gray) if. racciiiosa Labill.

    hfcionectes brownii Hook. 1. Afeioncctes brownii F. v. hl.

    ti. alata Jacq. var. cartila~inea Chccsem.

    If. alnta Jacq.

    H. scabra (Kocnig) Benth. H. scabro (Koenig) Benth. var. elongata Schindl. If. terragyna Benth. (non Hook. f.)

    II . micrantha (Thunb.) I

  • Polletr riiorphology of Ifalorogaccoc 185

    11. incana Walp.

    11. icaefolia Salisb.

    I{ . mczicma Schindl.

    H . ttiirrantha (Thunb.) R. Br.

    H . philippinensis hlerr. ---- _ - Ii. procumbcns Cheesem.

    H. srabra (Koenig) Benth. 11. teniri/olia Benth.

    Ii. teiragonia L'kkrit.

    Ii. tetragjns (Labill) Hook. f .

    H. tetrandra Schott

    11. teircrioides DC.

    H. unflora Kirk H . imflora Kirk var. genuina

    H. imffora var. bibracteolata (Col.) Schindl. 11. uni/lora var. spicata (Petrie) Schindl.

    H. cernirosa \Valp. H . rillasa Schindl.

    Schindl.

    H. ietragjna var. incana Kirk

    H . alata Jacq.

    H. teurrioides DC.

    If. minima Col. H . tenrlla Brongn. H. paircidentata Hosokasva H . srabra var. abbreriata Schindl. H . isoniera Parker ti. tcrragyna Clarke (non Hook. f.)

    H. te t rq jna var. dffitsa Hook. 1. H . Ji//rrsa (Hook. I.) Ckn. H. tetragjna F. V. hI. H . tenuifolia F. v. hl. H . alata Jacq. H. tetragyna F. v. hl. H. tetragyna (Labill.) Hook. f .

    var. dffusa Hook. 1.

    Laurembergia tetrandra (Schott) Kanitz ssp. brachypada (Wclw. ex Hiern.) Raynal var. brarlijpoda Raynal

    1

    {

    {

    H. teucrioides Gray H . elata Hook. f . H. tetragyna F. v. M. ti. dcpressa Hook. 1.

    H. depressa Hook. 1. (non Walp.) var.

    H . bibrarieolata Col. H. spirata Petrie

    ti. dcprcssa Hook. f. (non \Valp)

    Laurentbergia tetrandra (Schott) Kanitz

    scrpjlifolia Kirk (non Benth.)

    ssp. bmrhjpoda (Welw. ex Hiern.) Raynal var. brarhypoda Raynal

    the above data the supratectal processes in Ifalo- ragis approximately equal in number the tectal perforations. The latter usually are cylindrical and have a diameter of about 1.0 p. The processes and pcrforations also occur in the apertural areas, closc to the edges of the apertures. In addition to supratectal processes, delicate supratectal rnicro- processes about 1/10 of the size of the' latter, are encountered (cf. H. rtrasaticrrana P1. 7 , Fig. b). There are, however, species of ffulorngis provided with a smooth tectum. ff. rnicranrha (cf. Pi. 4 , Fig. l), may be quoted as an example illustrating this.

    The infratectal bacula are shorter than the thickness of the tectum. They are about 0.2 i t high and 0.2 p thick, although they are com- monly slightly thinner in the middle. A number of thicker bacula with a diameter of 0.5 it occur.

    Bacula extremely thickened at their bases and coalescing with more or less isolated elements of the underlying foot layer, were encountered (cf. 12. niicrantlia, 1'1. 4, Fig. I).

    The foot layer is about as thick as the tectum (processes included). It is relatively compact and its proximal and distal surfaces are smooth (cf. H. possei, PI. 8, Fig. f). The foot layer in some species seems to consist of spherical, ir- regular, comparatively large (diameter about 0.5 p) exine-1 elements. The latter coalesce more or less intimately into a rough structured foot Iaycr, which is perforated in numerous places by radial- ly orientated minutc channels. These channels can frequently be fused with thin interlamcllatcd areas in the proximal part of the foot layer (cf. H. rnicrorirhn, PI. 4, Fig. 1).

    The apzrture margins arc thickened and meas- Graiia 10

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  • 186 J . Praglowski

    ure about 4 11 in polar view. They consist of a few irregularly shapcd, thickened lamellae or lamellar segments.

    2. Pollen grains with a comparatively loose exine and complicated infratectal bacular structure.

    In this type of pollen grain the relation between the exine-1 and exine-2, is about the same as was the case for the previously described, simple type. Both types of pollen grains in Haloragis are also similar with regard to the structure of the tectum and the tectal details in general.

    In pollen grains of type-2, the infratectal bacula show a morphologically intimate contact with the underlying foot layer. This peculiar coalescence, which occurs in a very irregular manner, makes it difficult, if not quite impossible, to locate the limit between the bacula and the foot layer (cf. e.g., H. gossei, PI. 8, Fig. g-h, and H. ttimajiicratin, PI. 6, Fig. I-m). For interpretation of this prob- lem two alternatives seem to be worth discussing:

    a) either there are two, more or less irregularly shapcd, thin baculate layers (distinctly thinner than those in type l), which are divided in the middle by a zone of densely coalescent segments of the exine-I. These segments form there a com- paratively thick (about 1 p ) intrabacular exine-1 layer, provided with rough distal and proximal surfaces (cf. H. gossei, PI. 8 , Fig. g-h).

    b), or, according to the other alternative, the intrabacular-layer, suspended in the middle of the infratectal bacula could be considered as a distal zone of the foot layer. As a matter of fact the distal bacula layer supporting the tectum seems to be slightly, better developed than the second proximal bacula layer. If this concep- tion is true, the second alternative i.e., the exist- ence of a thick, split foot layer, involving also ihe exine-1 layer suspended in the middle of the bacula, seems to offer the most plausible inter- pretation of this complicated structure of the cxine-1. According to the first alternative, the thickness of the foot layer is about one third, and according to the second alternative two thirds of the total of the exine-I.

    The exine-2 has a loose structure. It consists of a few lamellae. The exine-2 coalesces with the proximal surface of the foot layer coniparatively weakly, by means of more or less densely spaced segments consisting of the exine-2 lamellae. In Grana I0

    wide areas of the pollen wall, the junction of the exine-2 with the innermost surface of the foot layer is formed by a sponge-net-like structure. This spongy zone, consisting of a complex of split Iamellae, surrounds the inner sphere. Both the spongy zone and the inner sphere are composed of the exinc-2 material which, however, in the latter seems to be more compact than in the spongy area.

    The margins of the apertures are thickened. They vary in thickness and measure about 4 1 4 in H . gossei, and about 5 p in H . tnicratitha. The lamellated segments of which the apertural thick- enings are composed, are found occasionally in the bacular zone as well as in the foot layer. The exine-2 does not occur in direct connection with the marginal thickenings.

    Iollcti tnorphologicnl coiiitnents based iipori electroti iiiicroscopy

    for Haloragis species of the subsection Spong- iocarpus

    The pollen wall in Haloragis species of the sub- section Spongiocarpus is comparatively compact.

    The exine-1 is much thicker than the cxine-2, the latter comprising only about 5-10% of the total of the exine thickness.

    The tectum including supratectal spinules or verrucae is about 0.5 p thick. The processes are about 0.2 p high. They are about the same (0.2 p) width at their bases. There are about 23 verrucae- like supratectal processes (tectal perforations in- distinct) per 50 p in H . Iiicasi. In I I . 1)iotiospcriiia the perforations are cylindrical, or slightly conical, with the proximal diamcter about 0.2 p, larger than the distal one (cf. P1. 9, Fig. e).

    The infratcctal bacula arc 0.5-1.0 r i long ix., longer than the thickness of the tectum. Jnfra- tcctal, horizontally elongated segments of a layer, fused midway with the bacula arc encountered (cf. H. iriotiospertiio PI. 9, Fig. c-k, and see also Loiidonia aiirca p. 166, PI. 1, Fig. a-c). This in- trabacular layer, which morphologically seems to be almost connected with the underlying foot layer, frequently does not occupy the entire pollen wall of a pollen grain, but is encountered in only part of it. In the remaining part of the exine simple bacula without any supplementary elements usually occur.

    The foot layer is about 0.5 p thick (intrabacular,

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  • Pollot r,torphology uf IIaloragaceae 187

    horizontally elongated segments not included). The distal surface ol the foot layer is not smooth. Small holes (about 0.1 p in diameter), or more or

    less distinct radial perforations have been observed in the foot layer (cf. H . liicasi 1'1. 9, Fig. k-m).

    The exine-2 is homogeneous, uninterrupted, and not thickened in the apertural areas.

    Explntintiotis to plntes 1-14 T h e magnification of all pollcn grains taken ni th a Leitz-Ortholux light microscope in Plates 1-14 is 1300.

    Plate 1. LOVDONIA. - (a-c) I-. airrea; (d-e) I.. behrii; (f-I) t. roei. - Note that pollen grains are isopolar. tenuimarginate, and have slightly ingrooved colpi. (c) thin section, nonacetolysed pollen grain, x 30 000; thick arrow shows elongated segmrnts of an intrabacular "layer". (k) thin section through the colpal area, acetolyscd pollen grain, x 10000. (1) single stage replica x 12000. p = tectal pxforations, s = supratcctal processrs, and ex.-2 = exine-2 (endexine sensu Faegri).

    Plate 2. IIALORAGIS. - (a-c) 11. tiodirlosn; (d-i) II. cotifertifolia; (k-m) H. tiie:iurra. - (a-m) pollcn grains paraisopolar, (0 tectum undulated. (m) exine with "spongy zone" (thick arrow). (g, h, i) scanning electron micro- graphs showing verrucae-like supratcctal processes (tectal pxforalions indistinct). (g) x 1700. (h) cdpus area, x 8 OOO. (i) the surface of the apocolpium x 17 000.

    PIote 3. IIALORAGIS. - (a-d) 11. tetragytro; (e-g) II. Ieircrioides; (h-i) H. halconeiisis; (k-m) 11. philippiiierisis. - (a) apcrtures protruding, concave mesocolpia, "spongy zone" (arrow). (e) exine lamellation distinct at the aperture margin. (g) "inner sphere" (thick arrow). (h) apxtures not protruding. (i) pore oval, irregular in shape. (k) tectum undulated. (m) comparatively long colpus. (d) thin stction, acetolysed pollen grain, x 10 000, tectal per- forations distinct, cxine-2 p3rtiaIly interrupted.

    Plate 4. IIALORAGIS. (a-d) I I . salsoloides; (c-g) 11. rirdis; (h-k) 11. depressa; (I) 11. rtiicrorttho. - In (a, e, f ) tectal perforations (dark dots) distinct. (i) pollen grain with an amb intermediate between that of goniotrem: and peritreme. (d) single stage replica x 14 000. S = supratectal processes, p = tectal perforations. (I) thin section, non- acetolysed pollen grain, x 16 000. Note smooth tcctum, complicated structure of the foot layer with lamellation in its proximal part, and radial channels. ch. = channcls in the foot layer, ex.-2 = exint-2, int. = intine.

    Plate 5. IIALORAGIS. - (a-c) If. iriottratia; (d) If. serpyllifolia; (e-k) H . Iieteropliylla; (I) 11. nspera. - (b-c) pollen grains in lateral view, one pole more flattened than the oppxi tc and more convex pole. (g) pollen grain with distinct "inner sphere" (arrow in.). (h-i) pollen grains in polar view, showing irregularly shaped nexine scales (ar- rows s), slightly detached from the nexin:. (k) scanning electron micrograph x 6 OOO, showing supratectal spinules. (I) single stage replica x 10 OOO. p- tectal pcrforations, spinules distinct.

    Plate 6. IIALORAGIS. - (a-d) 11. prostrata; (e-f ) 11. alata; (g) 11. cercodia; (h-m) H. iiiasajireratia. - In (c, f ) " innx sphere" (thick arrow). (d) abnormal pollen grain. (k) paraisopolar pollen grain. (1 oblique s:ction

    The magnification of thin sections, scanning elcctron micrographs. and single stage replicas in Plates 1-14, is given in the figure descriptions.

    through an acetolysed pollen grain x 8 000. showing: tectal perforations (white dots in tht tectum), the intra- tectal "layer" (arrow), and partially interrupted exine-2. (m) thin section through the aprrturc, acetolysed pollen grain, x 7 000, the lamellation consists of 4-5 Iamrllac. Plate 7 . HALORAGIS. - (a-b) 11. rrtasatiermrio; (c) 11. scoparia; (d-f) If. tetiiiifolia; (g-k) 11. browtii. - (a, b) scanning electron micrographs, x 1 800, and x 18 000 respectively. In (b) P - tectal perforations, S = supratectal processes, and hi - supratcctal micro-processes. (c) the innermost ncxine lamella (arrow) separated from the nexine in some places. (d and g) ncxinous "bottom of the ap:rtures" is lacking, and is prescnt in (f and i). (h) isopolar p ~ l l c n grain. (k) thin section, acetolysed pollen grain, x 10 OOO. tectal pxforations distinct, foot layer with loose structure. Plate 8. HALORAGIS. - (a-h) il. Rsssei; (b) pollen grain with "inner sphere" and a thick "spongy zone". Note distinct exine lamellation. (e-h) thin sections, acctolyscd pollen grains, (e) x 12000, the margin of the apcrture, note thickened area (thick arrow) consisting of the foot layer material. (r) x 7 OOO. exine comparatively compact, simple structure of the bacula. (g) x 8 500, exine compara- tively loose, complicated structure of the bacula, in- trabacular "layer" (thick arrow), foot layer (thin arrow). (h) x 5 000 "inner sphere" (thick arrow), note the pre- s:nce of "spongy zone" in thc apertural areas. Plate 9. IIALORAGIS. - (a-f) H . ttiotiospcrriia; (g-i) 11. raceriiosa; (k-m) H. Iircasi. - Black dots in (a, c. and g), are tectal perforations at high focus in LO analysis. (b and g) pollen grains peritreme, tenuimarginate. (c, d, and h) isopdar pollen grains in lateral view. (e, f, and k) thin sxtions, acetolysed pollen grains. (c, f) x 12000. (e) tcctum with distinct spinules and perfora- tions. ( f ) the margin of the aperture. observe that the exine lamellation does not appcar in this area. (k) x 7 000. thz segments of the intrabacular "lapr" can be s x n , the foot layer is perforated by delicate channels (thin ar- row). (i, I, and m) scanning electron micrographs, (i) x 6000, the margin of the colpus. (I) x 1500. (m) x 7 000. Plate 10. (a-b) LAUREAlnEKGlA cocciriea; (c-c) I.. r f - petis; (f) L. tetratidra; (g, h) Serpicitla itidica; (i-1) S. atigoletisis; (m-p) P roserpiriaca paliistrir. - In (3-1) pol- len grains exhibit a similarity to the pollen of Ilabragis, (b and d), however, they are distinctly isopolar. (e) thin section, nonacetolysed pollen grain, x 11 000, supratcctal processrs and tectal perforations distinct, the intrabacular "layer" (thick arrow) can bt seen, delicate channels are present in the foot layer (thin arrow), exint-2 inter- rupted.

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  • 188 1. Praglowski

    Platc 2. For exp!anation, see p. 187.

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  • Pollen morphology of Hnlorognccrre 189

    Plate 3. For explanation, see p. 187. Gmna 10

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  • 190 1. Praglowski

    Plate 4. For explanation, see p. 187.

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  • Plate 5. For explanation, see p. 187.

    13 - i O Z S 9 0

    Pollen morphohgy of Hnlorngacene 191

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  • Plate 6. For explnnaiion, see p. 187.

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  • Pollcti morphology of Ilalorcigciccne 193

    Plate 7. For explmation, see p. 187.

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  • Phtc 8. For explanation, scc p. 187.

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  • Pollen morphology of iialoriignccne 195

    Plate 9. For exp!anation, sec p. 187. C r a m 10

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  • 196 J . Prnglowski

    Table 11I. LAUREMBERGIA Berg.

    Polar- Shape & Size ity Amb N P C P/E index i n 11 Colpi 01) Pori ( p )

    Subgenus 1. lndolaurembergia Schindler

    Scctio a. Apopleura Schindlci

    L. inrlic:, (Thwait.) Schindl.

    L. wurrgcrinii Schindl.

    L. hirsrrtn (W. & A.) Schindl.

    var. rjpica Schindl. var. rorrutdifolio Schindl. var. anpstifolin Schindl.

    L. grundifolin Schindl.

    Sectio b. Ditelrupkrira Schindler

    L. hrcrites (W. & A.) Schindl.

    L. glnbcrrins Schindl.

    L. zcj/nnirn (A.) Schindl. L. coccincu (nlume) Kan.

    Java, van Stccnis 4245 (L)

    Java, Olurne 41 I. Det. Olume. Isolectotype

    L. jncnnicn (Miq.) Schindl.

    -

    H L B 903 364-361 (L)

    Is0

    Is0

    Subgenus I I . Alrolaurembtrgia Schindler

    L. rcpens Uerg. Is0 or S. Africa Capc Prov.. para

    L. /nodQgoscnrlerrJis Schindl. Is0 or hfadagascar, Herb. Hort. para

    I.P.H. Acocks 17892 (UPS)

    Bot. de Tananarive 4016. Det. J. Uosser (A. Raynal) (P)

    L. rcrticillnta Schindl.

    L. opposiri/oliu Schindl.

    L. rrronicifolis (Bory) Schindl. Reunion. hi.

    Frappicr 348. Det. A. Raynal (P)

    L. nngolrnsis Schindl. Rhodesia, R. E. Fries 610.

    Det. R. E. Fries (UPS)

    Gonio 543 643 so 2 6 x 3 3 3 x 1 3 x 2 544 644 78 2 x 0.5 2.5 x 2

    Gonio 543 (643) so 25x33 4 x 1 (3 x 2) (544)(644) 76 3 x I (2 x 2)

    Gonio 444 544 so 22 x 28 (413) (543) 79

    Gonio 544(644) SO 2 1 x 2 6 80

    (3x1) 3 x 2 2 x 2

    2 x 2 2 x 1.5

    Is0 or Interm 413 414 so 2 0 x 2 6 2 x 0.5 2.5 x 2 slightly 76 2 x 1.5 para

    Is0 or lnterrn 443 543 so 2 2 x 2 7 3x0.5 ( 2 . 5 ~ 2 ) para (444) (544) 81

    L. engleri Schindl. Is0 or Uganda. Y. E. Symes 500 (K) pan.

    Gonio- 443 543 SO 21 x 27 interm 78

    3 x 1

    L. rillosa Schindl.

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  • Pollen morphology of Haloragncene 197

    Exine thickness

    In polar view

    bleso- Apertures At Special morphological S. \v. s. Central gas- colpia margins poles characteristics America Africa Africa Africa car India Other

    East RC hlad3-

    I .5 3 4 I , 1.5

    I .5 3-3.5 I , 1.5

    1.5-2 3-3.5 2, 1.5 Ap. occ. irregularly shaped

    +

    1 4 1.5, I hlesocolpial exinc comp. compact, aspidia > than mesaspidia, ap. prot.

    +

    1.5 2 1.5, 1 Ap. not prot. Reunion hladagascar

    1.5-2 3 1.5, I Two pollen types occur: Iargcr with not prot. ap., and smaller with prot. ap. and more differentiated nexinc

    1.5, 1 Innermost nexine lamella occ. separated into indi- vidual fragments

    + +

    1.5 3 + +

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  • 198 J . Proglowski

    Table 111. (C'm/i/iired.)

    .

    Polar- Shapc 6; S i x ity A m b N P C I'/E index in 11 Colpi 01) I'ori (/))

    ' Larirciirbcrgicr brcripcs (W. 6; A.) Schindl. L. glabrrriiira Schindl. L . grmidifolia Schindl. L. lrirsirto (W. & A.) Schindl. L. inilicn (Thw.) Schindl. I-. jnrcrnicn (hliq.) Schindl. L. icnrigcrinii Schindl. I.. zcjlunica (A. ex Clarke) Schindl. L . agnstjoiiialcrjniin I Ienry Epilithcs cocci/lcn 111. ffnlorogis oli,yuiirha Arm (non \V. & A.) Scrpicrrla brcriprs \V. 8: A. S. hirsutn W. & A. S. c.pilirlirs BI. S. jmaniccr hliq. S. indica Thw.

    t. rrrraiidra (Schott) Kanitz Colombia. N. C. Fassctt 25898. Det. L. B. Sm. (US)

    Is0

    ~

    Pcri 543 643 SO 86

    ~ ~~

    25 x 29 3 x 0.5

    I so Uruguay, Herter 99860.

    Det. Hcrtcr (S)

    Is0 Brazil. G. Iiatschbach

    5133 (US)

    ssp. braclijpoila (Welw. ex Hicrn.) A. Raynal var. brnclijpodn Guinte Francaisc, J. G. Adam 14648. Del. A. Kaynal (P) -

    GuinCe Francaise, A. Chevalier 12189. Det. A. Raynal (P)

    var. niiiriidicn (Duricu ex. Uatt. B; Trab.) A. Raynal Tangnnyika. H. hl. Richards 17043. Dct. A. Raynal (P)

    (Schindl.) A. Kaynal va r. iiiildbrncdii

    Is0 or slightly para

    Is0 or slightly para

    Is0 o r para

    Interm

    Interm

    Gonio- interm

    Interm- peri

    Gonio- interm

    541 444 so (344) (543) 81 (443)

    543 (443) so (544) (444) 76

    543 so 8G

    443 543 so (434) (544) 85

    543 (443) so 79

    26 x 32 3 x 3 3 x 2

    2 3 x 3 0 2.5x 1 ( 2 x 2 )

    18 x 21 2.5 x 0.5

    2 1 x 2 4 3 x I (2 x 1.5) 2 x 0.5

    2 2 x 2 8 6 x 1 3 x 0.5

    Lniirciiihergio brcripcs (W. R: A.) Schindl.

    L. cocrinca (UI.) Kanitz

    L. glahcrrinra Schindl.

    L. Iiirsrrtn (W. 8: A.) Schindl.

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  • Pollett morphology of Hnlorognccnc 199

    Exine thickness ( / I ) - In polar view

    hlcso- Apertures At Special morphological S. W. S. Central gas- colpia margins poles characteristics America Africa Africa Africa car India Other

    East Rc hlada-

    2.5-3 4 2

    2 4 2

    1-13 3 4 I .5

    2 2 2 Ap. not prot., amb Ulmus + like, innermost nexine lamella separated into individual fragments

    Ap. not prot. +

    Ap. not prot., ap. shape + occ. irregular

    1.5 2

    2 3

    1.5

    I .5

    Ap. usually prot.

    Ap. not prot.

    Ap. not prot.. tcctum occ. undulating, colpi comp. long

    +

    +

    +

    N. Africa

    N. Africa

    N. Africa

    N. Africa hladagascar Reunion

    L. indica (Thw.) Schindl.

    L. jaranira (hliq.) Schindl.

    L. repcns Berg.

    L. tctranilra (Schott) Kanitz ssp. tetrandra

    ssp. brnchypoda (Wclw. ex Hiern) A. Raynal var. braclt.vpoda

    var. ntildbraedii (Schindl.) A. Raynal

    var. ntintidica (Durieu ex Batt. 6r. Trabb.) A. Raynal

    L. reronici/olia (Bory) Schindl.

    L. reylmica (A.) Schindl.

    ~~ ~~~ ~

    Scrpicrtla indicn Thw.

    Serpicrila jarmica hfiq.

    Serpicrtla rcprns L. S. scrpylli/olia Uurm. S. capensis Racusch.

    Scrpicrila brasilierrsis Carnbcss. Ifaloragis tctra