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(Paracryphiaceae) Monotypic, Arborescent, Dicotyledonous Plant

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(Paracryphiaceae) Monotypic, Arborescent, Dicotyledonous Plant BLUMEA 23 (1977) 417—438 The morphologyand relationships of Paracryphia (Paracryphiaceae) William+C. Dickison & Pieter Baas Department of Botany, University of North Carolina, Chapel Hill, U.S.A. & Rijksherbarium, Leiden, Netherlands Contents Summary 417 Introduction 417 Material and methods 418 Descriptive part 4 20 Vegetative Morphology and Anatomy 420 Leaf 420 Node 421 Axis 421 Bark 421 Wood 421 Reproductive anatomy 422 Inflorescence 422 Flowers 422 Pollen 424 Fruit 425 Seed 425 Discussion 425 General 425 Placement of Paracryphia in the General System 426 Acknowledgements 432 References 437 Summary A of floral and of the New Caledonian comprehensive study vegetative anatomy monotypic genus Baker in the Paracryphia was initiated an attempt to help clarify the evolutionary relationships of genus. of Detailed descriptions leaf, axis, nodal, wood, floral, pollen, and fruit morphology and anatomy are presented. In general, most vegetative characters are distinctly primitive whereas those of the reproductive The organs are regarded as advanced or specialized. present study confirms the opinion that Paracryphia merits familial All ofthe in favor of that status. previously suggested relationships genus are rejected a view and from envisions Paracryphia as an independent early divergence the Thealean, Ericalean, and/or Celas- with tralean lines of evolution. This view is based on similarities of Paracryphia Sphenostemonaceae, Actinidi- aceae and Theaceae in a number of characters. Introduction Paracryphia is a monotypic, arborescent, dicotyledonous genus endemic to New Cale- donia. The in habit from shrub 18 in and plant ranges a to a tree meters height, available data indicate that humid forestsbetween and meters altitude. it grows in 750 1500 Although it and the systematic relationships of the genus are obscure, has received little attention 418 BLUMEA VOL. 23, No. 2, 1977 than from classification. The as a result is more often not excluded general systems of genus P. suaveolens Baker was initially described by Baker (1921) with the name f.; however, alticola the plant had been previously described by Schlechter (1907) as Ascarina Schltr, a member of the Chloranthaceae. combined the two alticola In 1950 Van Steenis as P. (Schltr) Steen. Baker ( loc. cit.) originally included Paracryphia in the Eucryphiaceae, noting, however, that it differed from Eucryphia in a number of characteristics. Both Gilg (1925) and Bausch the vesselless (1938) were of the opinion that its affinities were probably near primitively families Trochodendraceae. This of was also Guillau- Winteraceae or point view expressed by the and floral characters of min (1948). Swamy (1953), following a survey of vegetative excluded from the Chloranthaceae and denied that it related the genus, it emphatically was familial to Winteraceae, Trochodendraceae, or Illiciaceae. Paracryphia was elevated to status that related Trochodendraceae and by Airy Shaw (1965) who believed it was closely to the be included that Shaw also the perhaps should even in family. Airy pointed out superficial between the similarity gynoecium of Paracryphia with that of another isolated genus, and have the of Medusagyne. Agababyan Zavaryan (1971) supported recognition an independent family, Paracryphiaceae, on the basis of pollen morphology. Takhtajan the and while ( x 973), in 311 uncertainmanner, recognized family Paracryphiaceae noting that its affinities it were very obscure, placed near Eucryphiaceae. and anatomical This study was undertaken to provide a more thorough morphological of with of affinities. description the genus the objective clarifying its natural MATERIALS AND METHODS Both liquid preserved (FAA) material and dried herbarium specimens were examined. boiled and thickness Wood samples were in water cut on a sledge microtome at a of sections stained with Heidenhain's and safranin. 20—30 /xm. Resulting were haematoxylin wood obtained least Data relating to cell length were by making at 25 measurements from macerations prepared using Franklin's method. Cell diameters were measured from sections. flowers embedded in transverse Preserved leaves, stems, and were paraffin and microtome. Sections obtained the of cut on a rotary were throughout length petioles. Flowers and nodes were cut serially. Sections were subsequently stained with a safranin leafand fast-green combination. Details of floral venation were observed using cleared tissues prepared in 5% hot NaOH followed by safranin staining. Epidermal leaf features were examined using free hand sections. Dried specimens were initially re-expanded in 5% NaOH, placed in FAA, and then treated like preserved materials. Fruits were em- celloidin bedded in and sectioned on a sledge microtome. Slides are deposited in the Rijksherbarium, and the DepartmentofBotany, University of NorthCarolina at Chapel Hill. Pollen was acetolysized and mounted in glycerine jelly. Acetolysized pollen as well as wood and seed samples were gold coated and observed with the Cambridge All Scanning Electron Microscope at the Geological Institute, University of Leiden. For transmission electron from herbarium microscopy, nonacetolysized pollen grains a specimen were soaked for which one week in 0.1% glutaraldehyde in 0.1 M cacodylate buffer to was added in The 2% glucose, and subsequently fixed 1% uranylacetate. material was em- and bedded in epon sectioned at 800 A with a diamond knife. The sections were counter- stained to with lead. Observations made with according Reynolds were a Philips EM 300 at the Zoological Laboratory of Leiden University. W. G. Dickison & P. Baas: Paracryphia 419 Line Camera lucida ofnodal and — Fig. 1—6. drawings petiole anatomy. equals 1 mm. 1 —5. Transverse sections from the base to the distal of the in the vascular (1) region (5) petiole showing changes system (Schlechter 15326).— 6. Transverse section ofthe node showing trilacunar, three-trace condition; M=medi- leaf L=lateral an trace, leaf trace ( McKee 31070). 420 BLUMEA VOL. 23, No. 2, 1977 Wood terminology corresponds to the Multilingual Glossary of Terms used in Wood Anatomy (Committee on Nomenclature, International Association ofWood Anatomists, 1964). Terminology relating to leaf venation and bark anatomy follow suggestions of Hickey (1973) and Zahur (1959) respectively. The following specimens obtained from the Museum National d'Histoire Naturelle, Paris (P) and the Rijksherbarium, Leiden (L) were studied: Paracryphia alticola (Schltr) Steen., New Caledonia: H. S. McKee 26820 (P), pickled flowers, driedleaves; H. S. McKee and 31070 (P), pickled leaves, stems, fruits, mature wood, bark; H. S. McKee 27400 (P), dried leaves; H. S. McKee 31490 (P), dried leaves and young fruits; H. S. McKee 4993 (P), dried dried Schmid leaves, immature wood; H. S. McKee 5380 (P), leaves; M. 1573 ( p) dried dried leaves and leaves; L. Bernardi 10362 (P), dried leaves; R. Schlechter 15316(L), twig. DESCRIPTIVE PART Vegetative morphology and anatomy Leaf Plate & (Figs 1—5, 7—9; I, I 2) in Leaves simple, petiolate, exstipulate, verticillate to subverticillate arrangement. with Blades coriaceous an acute to acuminate apex, an attenuate base, and finely serrate margins. Leafprimordia protected by well developed bud scales. Young leaves densely pubescent with unicellular, unbranched hairs. Indumentum progressively lost in more mature foliage. Venation pinnate with a straight primary vein. Secondary veins brochido- dromous orders with a series of 2 or 3 of ascending marginal loops (fig. 7). Secondary veins fairly thick, diverging at an occasionally variable moderate angle from the midvein, and abruptly curved. Composite intersecondary veins frequently originating from the veins in midvein. Tertiary a predominantly orthogonally reticulate pattern. High order which veins forming a reticulum in which the veins arise in an orthogonal manner and in well and areolation is developed, orientated quadrangular (fig. 8). Veinlets present or absent, mostly linear or curved, occasionally branched. Marginal ultimate venation Teeth vascularized medial vein the tooth looped. by a single extending to apex (fig. 9). Vein Teeth often cells. and order veins endings expanded. capped by opaque Low high sheathed by lignified parenchyma of irregularly shaped cells. Specialized terminal cells absent. unbranched. Lamina in surface view: Trichomes very sparse, unicellular, Adaxial sided with curved anticlinal epidermal cells 4 —7 straight to slightly walls, strongly elongate over veins. Abaxial epidermal cells 5 —6 sided with straight to curved anticlinal confined the abaxial cell walls, elongate over veins. Stomata to surface, anomocytic; guard long, 21 wide (Plate I, 1). pairs 23 —33 —39 /im —25—30 fim bifacial. Adaxial cuticle In transverse section: Lamina 9—12 jum thick; abaxial cuticle to thick midrib. Adaxial cells to 3 —4 jum, up 13 jum over epidermal upright cells. procumbent (mainly square), larger than abaxial Epidermal cells over midrib with dome-shaped outer periclinal walls. Stomata in level with the unspecialized epidermal cells, with well developed outer and small inner cuticular ledges. Mesophyll composed of of 2(—3) layers of elongate palisade cells with dark-staining contents and spongy region which has raised central portion thick-walled, lignified cells (Plate I, 2). Midrib with a abaxial and adaxial with vascular surface, supplied a simple to complex (fig. 5) system, of midrib sheathed by sclerenchyma fibres. Ground tissue collenchymatous at periphery, W. C. Dickison & P. Baas: Paracryphia 421 vascular with
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