BLUMEA 23 (1977) 417—438

The morphologyand relationships of

Paracryphia ()

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 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 . 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 . 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., : 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 thin-walled parenchymatous near system, interspersed very brachysclereids. with adaxial Major veins surrounded by thick-walled, lignified parenchyma extensions to

surface, usually not reaching abaxial surface; minor veins also with lignified parenchyma- but tous bundle sheaths embedded in mesophyll. Styloid crystals, up to 250 ium long,

scattered thick throughout mesophyll. Petiole provided with a cuticle (10—15 jum), the base with three vascular bundles which dissected supplied at distinct give rise to a strand that the ends crescent-shaped subsequently invaginates at distal to produce a closed,

in or nearly closed vascular system the form of a ventral arc and a flattened dorsal region with Whole a large centrally positioned collateral medullary bundle (figs 1 —4). system

sheathed by sclerenchyma in distal end. Ground tissue compact at periphery, lacunate around with and cells vascular system, interspersed thick-walled brachysclereids contain- ing styloid crystals.

Node (Fig. 6) from Trilaminar, three-trace in both young and older stems. Collateral traces diverging

the other. and cauline system at same level and close to each Median trace C-shaped larger than laterals.

Axis Plate (young twigs, 3—5 mm in diameter; I, 4) Unicellular hairs older material. unbranched abundant on young stems, absent on

Cuticle to 18 thick. cells with dome-shaped outer walls in up /.fin Epidermal erect, young twig, flattened rectangular in older material. Periderm arising in subepidermal layer. Cortex of cell with intercellular c. 14 layers parenchymatous, compact at periphery, large Vascular spaces in inner portion, rarely interspersed with brachysclereids. tissue in a with fibres. complete cylinder, a well developed ring of perivascular sclerenchyma Secondary

phloem composed of sieve tubes, companion cells, and parenchyma, without mechanical tissue. traversed and with similar in Secondary xylem by narrow rays angular vessels;

qualitative characters to mature wood. Pith c. 2 mm in diameter, composed of paren-

chyma, which becomes lignified from periphery inwards in older material, occasionally

interspersed with thick-walled brachysclereids. Styloid crystals in cortex, pith, and phloem

with smaller diameter than the parenchyma (Plate I, 4); those of secondary phloem a others.

Bark (of mature stem; Plate I, 3 & 5) Bark well thin-walled developed, 4 mm thick. Periderm of cells, rectangular in trans-

cells. verse section, up to 20 layers thick, occasionally interspersed with sclerified Secondary phloem composed of sieve-tube elements, companion cells, phloem parenchyma, and

dilated abundant thick-walled fibres, traversed by heterogeneous rays which are slightly and thick partly sclerified in the older phloem. Fibres in interrupted, I—5 cells tangential bands or clusters, differentiated close to the cambial zone. Parenchyma sclerified in older phloem. Sieve tube elements scattered with compound sieve plates in highly or in groups, end walls Plate abundant the oblique (Zahur's type I—II; I, 5). Styloid crystals throughout

bark; prismatic crystals observed in periderm of McKee 4993 only.

Wood (Plate II)

When McKee and die two mean figures are given, the first relates to a mature wood sample ( 31070)

second to an immature sample (McKee 4993). 422 BLUMEA VOL. 23, No. 2, 1977

2 Growth extremely faint to absent. Vessels diffuse, (27—)39(—5i)/mm solitary rings , radial rounded (57%) or in multiples of 2 —6, angular to in transverse section (Plate II, 1),

& & tangential diameter (35 —)47 64(—84) ;«m, vessel element length (1050—) 1760

20oo(—■:2620)/j.m. Perforations scalariform in almost vertical end walls, with (58-) 111 bars and (—203) (Plate II, 3 & 4). Bars very narrow closely spaced, occasionally reticulate.

Intervessel transitional pits scalariform or (Plate II, 3). Vessel-parenchyma and vessel-ray halfbordered. helical pits opposite to scalariform, Tyloses, vessel contents, or thickenings absent. Ground tissue composed of fiber-tracheids in tangential diameter, , 24—32 /an

& walls thick with bordered (890—)2o6o 230o(—3490) /im long; c. 7 //m numerous pits radially and tangentially. Pits c. in diameter with extended slit-like 4 /urn apertures.

Helical thickenings absent. Parenchyma diffuse and diffuse-in-aggregates, apotracheal as well as scanty paratracheal, strands 4—6 cells long. Rays heterogeneous I (Kribs, 1968), of

two distinct sizes; uniseriates of erect cells 10/mm; bi- and triseriates with long uniseriate and cells marginal portions of erect cells central portion ofprocumbent to square 1—2/mm absent. (Plate II, 2). Styloid crystals sparsely present in axial parenchyma. Silica bodies

Reproductive anatomy

Inflorescence

The multiflowered, branched, terminal inflorescence is paniculate although the flowers sessile are (i.e., a compotmd spike). Inflorescence axes are covered with unicellular, un- branched trichomes.

Flowers Plate (Figs 10—13; III, I) General organography and anatomy: The flowers of Paracryphia have are either bisexual, or occasionally exclusively staminate. These staminate flowers no abortive and levels gynoecium are borne at lower on the inflorescence axes. The un-

differentiated consists four perianth of caducous, decussate, concave, free segments. ofthe Aestivation perianth parts is cochleate, i.e., having one large helmet-shaped segment

that encloses or the others Perianth covered with covers (fig. 10). parts are externally unicellular (rarely bicellular), unbranchedtrichomes of the same type as occur on vegeta- tive the parts of plant. Lignified, highly pitted macrosclereids are scattered throughout the

perianth tissue. The apostemonous androecium is usually composed of 8 borne

a Each Each in single cycle. consists of a filament and a basifixed anther (fig. 13). is of anther composed a connective and four locules, i.e., shows the tetrasporangiate

condition. Before longitudinal dehiscence the partition between the two locules of the

same theca breaks thus has a down, forming a single chamber. Each microsporangium well developed endothecium with lignified thickenings in the form of bands. The endo-

thecium is interrupted in the region of the connective and locular partition. Filaments do not have a central lacunaassociated with the vasculation. Stamens ofmale flowers develop

Fig. 7—13. Camera lucida drawings ofleaf and floral morphology ofParacryphia. Line equals 1 mm. — 7.

Leaf showing pattern oflow order venation ( McKee 26820). — 8. Details of high order leaf venation ( McKee

26820). — 9. Details ofmarginal leaf venation including teeth vasculation (Bernardi 10362).— 10. Transverse

section of flower showing pattern of vasculation; DB=dorsal carpellary bundle, VB=ventral carpellary

bundle ( McKee 26820). — 11a, b, 12. Cleared stamens from male flowers showing expanded filament and branched venation 11b filament — from bisexual system, in transection (all McKee 26820). 13. Stamen flower ( McKee 26820). W. C. Dickison & P. Baas: Paracryphia 423 No. 424 BLUMEA VOL. 23, 2, 1977

swollen filaments while flowers still the bud n & conspicuously are in stage (figs 12).

contains 8 The superior, slightly oblique, gynoecium —15 laterally concrescent, condupli- Unicellular cate carpels that are also adnate with the solid core of central tissue (fig. 10). trichomes the surface. ventral unbranched sparsely cover gynoecium Although sutures the remain closed the adnation of their the may open, carpels by ventral margins with

solid central core. Carpels never open ventrally into a central space as illustrated by

Swamy (1953). The region of lateral connation is histologically evident as a zone of remain antire all narrow, compressed cells; however, carpels fused along their length at

of are number and stages development. Stigmas equal in to carpels, sessile, conduplicately ovules folded. Each locule contains four rather small, anatropous, unitegmic that are The borne in a single row on axile placentas, again contradicting Swamy's interpretation. of thicker ovular integument is composed 3 —4 layers, the outer ofwhich is composed of

walled cells that are more The mother cell is densely staining (Plate III, 1). megaspore of within the locule embedded in 2—3 layers nucellus. The position of the ovules varies dorsal dorsal from epitropous to hypotropous (sensu Bjornstad, 1970). The ovules

completely fill the width of the rather narrow locule chamber.

Vascular of discrete bundles at the floral anatomy: A ring many is present base. Each perianth member receives three traces that undergo limited branching. Stamens from and which receive a single trace that originates directly the central cylinder in

filaments of stamens from bisexual flowers remains undivided along its entire length

rich vasculature (fig. 13). The enlarged filaments of stamens from male flowers have a from basal dimensional of the resulting a dichotomy and subsequent three branching

stamen trace (figs 11a, b, 12). Stamen traces are surrounded by darkly staining (tanni- ferous?) cells.

the central venation The vascular tissue in cylinder becomes organized into a gynoecial ventral pattern consisting of one prominent dorsal bundle per carpel and a less obvious

supply characterized by the fusionof the two ventral bundles of adjacent carpels (fig. 10).

The fused ventrals and at the lie in a plane between the locules of adjacent carpels, same lateral wall time constitute aninterrupted vascular cylinder ofthe central core. Additional bundles absent. from the are Ovules are vascularized by traces (or procambium) departing ventral bundles. distal of the the fused in In regions gynoecium ventrals separate resulting

dorsal and two unfused ventral bundles The three bundles one per carpel. carpellary the terminate in massive of extend to style apex where they a aggregation tracheary elements.

Pollen (Plate IV)

Solitary, radially symmetrical, small, tricolporate, semi-angular to more commonly

circular in outline in median polar view (Plate IV, 1), spherical to oblate-spheroidal in

IV, X m. shape (Plate 1—5), (22.4 —)24(—26.5) (21.5—)22.4(—25) fi Ektoaperture elongated and meridionally covered by a granular membrane. Polar index 0.29. Endoaperture

slightly elongated meridionally (3x5 fim), closed with granular material (Plate IV, 2, 6, 7),

Intine in the and forming a bridge. very prominent under the colpae and forming a plug

endoaperture. Exine2— thick in the center of mesocolpium. The exine is stratified 2.5 (im

into endexine, prominent foot-layer, columellae, and tectum (Plate IV, 6). Endexine

is thin in the {s.s.) electron dense with the staining method used and forms a very layer of the but rather thick the of the The center mesocolpium is along margins apertures.

boundary between endexine and ektexine is well defined. The foot-layer composes about

two thirds of the total exine thickness although it decreases in thickness along the margins W. C. Dickison & P. Baas: Paracryphia 425

of the ektoapertures. Total exine thickness along the margins of the ektoapertures,

however, remains uniform as a result of the corresponding increase in endexine deposition

unknown the surface (Plate IV, 6). In our preparations a deposit of origin covers external of Columellae are short and distributed a thefoot-layer. (c. 0.3 fim in height), thick, are in reticulate pattern in uniscriatc rows (Plate IV, 6 & 7). The tectum comprises about one fourth of the total exine thickness. Sculpturing is predominantly reticulate although

occasionally it is transitional between reticulate and rugulate (retirugate) in the meso-

colpium, becoming finer at the poles and along the colpae. Muri are about 0.4 —0.7 um wide.

Fruit (Plate III, 2)

The fruit is a glabrous, loculicidal capsule that dehisces longitudinally along the ventral

order for this each from the suture of each carpel. In to occur carpel at maturity separates

central column along its entire ventral margin, although some lateral concrescence among adjacent carpels remains. Although each carpel eventually becomes completely unsealed from the central axis, a distal connection is maintained by two strands apparently rep- resenting the lignified ventral carpellary vascular bundles. cells the Anatomically the fruit is distinguished by a massive sclerification of lining

locules (Plate III, 2). This is accompanied by the uniform sclerification of the central axis

cells and those ensheathing the vascular bundles. Sclerified cells are extensively pitted, and

more or less isodiametric in shape. Remaining ground tissue of the fruit is composed of

thin-walled parenchyma. There is no proliferation of vascular tissue in the fruit.

Seed (Plate III, 3)

seeds the Seeds are The number of per fruit locule equals number of ovules per carpel. thin small, 2.0—2.5 mm in length, compressed, exarillate, and albuminous. The seed coat

consists of a single cell layer c. 12 thick and is composed of cells with thickened —14 fim inner surface have lignified walls as viewedin transection. In view, cells of the testa highly

undulating anticlinal walls. At maturity the testa becomes extended to form small wings.

Endosperm abundant, cellular at maturity, and composed of thin-walled polyhedral cells.

A single vascular bundle extends into the dorsal wing of the seed coat for approximately

half the length of the seed. In Corner's terminology (1976) the seed of Paracryphia is exotestal. in radicle unitegmic Embryo dicotyledonous, straight, c. 0.5.—0.7 mm length, longer than the cotyledons.

DISCUSSION

General

Paracryphia is distinguished by the following unusual combination of characters: wood primitive structure with extremely long vessel members and scalariform perforation plates containing over 100 bars, scalariform intervessel pits, fiber-tracheids, heterogeneous three-trace rays; trilacunar, nodal anatomy; complex petiole vasculation; simple, ex- stipulate, pseudoverticillate leaves; anomocytic stomata; large styloid crystals; secondary phloem with primitive sieve tube elements and abundant fibres; bisexual and less fre-

staminate flowers in the quently same complex inflorescence; undifferentiated, 4-merous decussate 8 with basifixed of perianth; c. stamens anthers; a gynoecium numerous com- pletely concrescent carpels accompanied by the fusion of ventral traces ofadjacent carpels; 426 BLUMEA VOL. 23, No. 2, 1977

unitegmic ovules and axile placentation; tricolporate, retirugate pollen; and capsular fruits leaving a central column at dehiscence, with small, winged, albuminous, exotegmic seeds. is remarkablethat from It most characters vegetative anatomy are distinctly primitive least or at unspecialized, whereas those of the reproductive organs can be regarded as

advanced or specialized.

The fact that Paracryphia produces both bisexual and male flowers within the same Bausch inflorescence was notedby (1938). Swamy (1953), however, apparently concluded flowers The that only bisexual were formed. genus clearly produces inflorescences con- bisexual small number of taining predominantly flowers although a entirely staminate

flowers also of male floral buds has shown the stamen filaments are present. Examination

to be expanded laterally prior to anthesis. Associated with the enlarged condition of the

filamentsis the of branched vascular The functional presence athree-dimensionally system.

significance of these features with regard to the reproductive biology of the plant is

unclear. Pollen produced in anthers from male flowers appears normal. Another of floral about which aspect morphology some disagreement exists concerns the number and interpretation of the floral envelopes. Both Baker (1921) and Bausch the of (1938) referred to presence two imbricate, caducous perianth segments. Airy Shaw described bract (1965) Paracryphia flowers as having no perianth but one and two bracteol-

the androecium and have examined es subtending gynoecium. Preserved flowers we have than the two opposing pairs of accessory segments, one of which is distinctly larger

others. We see no compelling reason not to refer to these structures as perianth members.

Placement of Paracryphia in the General System

examined In this discussion the putative, previously suggested, allies of Paracryphia are

in the light of available evidence. This is followed by comparisons between Paracryphia

and additional families that for various reasons we feel deserve consideration as possible

relatives. Comparative data were gleamed from the literature or obtained from original observation.

1. The of ovules in severely —. presence unitegmic Paracryphia weakens that the should be included the families of the arguments genus among woody

subclass Magnoliidae (sensu Cronquist, 1968). Ovules in the Magnoliales are apparently families of the uniformly bitegmic. Unitegmic ovules do occur sporadically in a few

Ranunculales (Ranunculaceae, Menispermaceae, Circeasteraceae) and Saururaceaeand Piperaceae of the Piperales (Philipson, 1974). All of these unitegmic Magnoliidae, however, have no relationship to Paracryphia as demonstrated both morphologically and anatomically. Salient features of Winteraceae such as vesselless xylem; bitegmic ovules; pollen grains

always anaulcerate (i.e., aperture located distally), semitectate, reticulate, and consistently

shed in and ofleafvenation permanent tetrads (rarely monads); etherial oil cells; a pattern

in which secondary veins have an irregular course and intercostal areas lack uniform size

rank and shape (primitive first leaves, sensu Hickey, 1971) negate a close alliance with the well Paracryphia. The regular course of the major veins in addition to developed

of areolation in leaves advanced features to pattern Paracryphia are structurally according the and interpretations of Hickey (1971). Illicium, despite a somewhat similar appearance with vessel having xylem exceedingly primitive members, is clearly distinguished mor- phologically and anatomically from Paracryphia by unilacunar nodes; specialized stomatal

ethereal oil vein structure; cells; leaf margins with 'Chloranthoid teeth' having a medial W. C. Dickison & P. Baas: Paracryphia 427

'braced' by two prominent laterals which join it distally (see Hickey & Wolfe, 1975); unusual tricolpate pollen in whichthe three colpi join atthe poles resulting in an syncolpate

condition; fruit a single whorl of follicles; and a simple carpellary vasculation consisting of only a single dorsal and ventral strand that is more comparable to members of the

Lauraceous assemblage (Keng, 1965). Paracryphia diverges from the Chloranthaceae by the

of possession trilacunar nodes, tricolporate pollen, and anatropous ovules, as opposed to the unilacunar nodes, monosulcate and monosulcate derived pollen, and orthotropous ovules of The of tooth is absent in Chloranthaceae. very distinctive type Chloranthoid Paracryphia leaves.

Trochodendraceae. and Trochoden- 2. Suggestions of a close affinity between Paracryphia the draceae, e.g. by Airy Shaw (1965), appear equally unlikely even though two taxa have rather similar in a pollen morphology and a very superficial resemblance gross morpholo- derive gy. Although it would be theoretically possible to most Paracryphia character

from states those present in Trochodendron, in our opinion the differences outlined below make close a relationship between these taxa improbable.

Paracryphia Trochodendraceae

Vessels present Vessels absent Node trilacunar Node multilacunar (mature)

Petiole vasculation complex Petiole vasculation simple Chloranthoid teeth absent Chloranthoid teeth present Astrosclereids absent Astrosclereids present

Styloid crystals present Styloid crystals absent

Inflorescence paniculate Inflorescence cymose

Perianthpresent Perianth absent

Stamens few Stamens numerous Axial absent tissue present in gynoecium Axial tissue in gynoecium Simple floral vasculation Complex floral vasculation

Styles sessile Styles elongate Ovules unitegmic Ovules bitegmic Ovules few per carpel Ovules numerous per carpel

Fruit a capsule Fruit a follicetum

Inclusion of in the done in a 3. Eucryphiaceae. Paracryphia or near Eucryphiaceae, as number of tentative manner by Bausch (1938), also cannot be supported. The large differences significant are compared below.

Paracryphia Eucryphiaceae

Vessel elements extremely long Vessel elements moderately long Vessel Vessel perforations scalariform, many- perforations simple or scalariform, barred usually few-barred

Xylem parenchyma diffuse, scanty Xylem parenchyma terminal, scanty diffuse

Stipules absent Stipules present Stomata anomocytic Stomata paracytic

Styloids present Styloids absent Inflorescence paniculate Flowers solitary 428 I3LUMEA VOL. 23, No. 2, 1977

Perianth undifferentiated, caducous Perianth differentiated, persistent

Ovules unitegmic Ovules bitegmic

Floral vasculature simple Floral vasculature ± complex Pollen small, tricolporate Pollen minute, dicolporate

In the is addition, although the fruits initially appear similar, manner of dehiscence

differentwith regards to the attachment of the valves to the central axis. As pointed out by

Bausch, the valves of the mature fruit ofEucryphia are attached basally whereas in Para-

cryphia attachment of the valves to the persistent central axis is distal. related The suggestion that Paracryphia is to Eucryphia led us to an examinationof other Rosidae.

than Data on wood anatomy indicate a lower level of evolutionary advancement any

extant Rosalean and of before the oldest Cunonia- supports anearly divergence Paracryphia has Rosalean ceae, Brunelliaceae, or Rosaceae. Cronquist (1968) characterized a hypothetical

plant combining all the primitive features found in various members of the order to be

woody, with alternate, stipulate leaves, regular, hypogynous flowers with five separate

, numerous centripetal stamens, several separate carpels, and numerous crassinucel- late, bitegmic ovules, die ovaries ripening into follicles with endospermous seeds. Para- cryphia combines few of the above features. Furthermore, the presumably primitive absent Rosalean five-trace carpellary vascular pattern (Sterling, 1969) is in Paracryphia.

with other search for close relatives of 4. Comparisons families. A possible Paracryphia resulted concentrated those in comparisons with numerous families. Attention was on

which retained while advanced to groups primitive xylem traits possessing moderately floral ovules. The advanced characters such as unitegmic combination of morphological and families anatomical features present in Paracryphia is reminiscent in varying degrees to

compared in Table 1. from Members of the Theaceae, a family generally thought to be derived ,

have very primitive woods (subfamily Ternstroemioideae) that are comparable to Para-

cryphia although pores are never in radial multiples. Theaceous also approach

Paracryphia in foliar venation, having secondary veins brochidodromous often with

veins that Placentation is ascending arches and tertiary are random to weakly percurrent. axile Theaceae and be the of in up to ten carpels can present in syncarpous gynoecium

Laplacea (Keng, 1962). Small, unspecialized, tricolporate pollen of the Paracryphia type is

widespread in several groups of the Theaceous complex (subfamily Ternstroemioideae of

Theaceae). Other similarities are evident in the capsular fruit of the Camellioideae in which

the from valves at maturity separate a persistent central columella, fibrous endothecium, and sclereids in the perianth parts. A relationship to Theaceae is weakened, however, since

their nodes are strictly unilacunar, petioles have a simple vasculature, ovules are bitegmic, the and as far as is known mechanical cells are poorly developed or entirely lacking in bark.

Any similarity between gynoecia of Paracryphia and Medusagyne is quickly negated by the in the latter vessel occurrence genus of a highly advanced xylem, characterized by elements with with vascular bundles simple perforations, petioles numerous separate

arranged in an arc, opposite leaves, and flowers with well differentiated perianth parts and numerous stamens (Beauvisage, 1920). Table shares of with the As seen in 1, Paracryphia a number features sensu

lato. Included among these resemblances are primitive vessel elements, styloids, and uniteg- axile because of mic ovules borne on placentas. Styloids are ofparticular interest they are W. C. Dickison & P. Baas: Paracryphia 429

of characters in Table 1. Occurrence some of Paracryphia

selected families of

Q)

nj

id Q)

•H o Q) 2 fd

0) O 0 c c Characters of Pharacryphia Sai d) 4J (A 0 G

0) Actinidiac Ericaceae Aquifoliaceae x: Theaceae (incl. 04 CO

trilacunar nodes - ( + ) - + +

simple, exstipulate leaves + + + + +

leaf venation as in Paracryphia + - ( + ) ( + ) +

- complex petiole vasculation - ( + ) ( + ) (±)

anomocytic stomata ( + ) + + ( + ) +

- styloids - ( + ) + +

extremely long vessel elements, ( + ) ( + ) < + ) + + seal. 100 bars p.p.

fiber-tracheids + + + + +

undifferentiated perianth - - - - +

compound gynoecium + + + + +

sessile stigmas - - - + +

axile placentation + + + + +

anatropous, unitegmic ovules - + + + +

small, retirugate, tricolporate ( + ) ( + ) < + > + - pollen

loculicidal, capsular fruit ( + ) ( + ) ( + ) - -

albuminous seeds + + + + +

bark fibrous - - - - +

= = Legends: - character absent; + character present in most

of = character representatives of the family; (+) character present in a few

representatives only; + = tendencytendency to bebe similar inin this charac-

ter.

The wood of of restricted distribution in Dicotyledons. anatomy some species Saurauia is distinguished by vessel elements with highly oblique scalariform perforation plates intervessel containing over 100 bars and scalariform to opposite pitting, fiber-tracheids,

of sizes that contain a of cells and often heterogeneous rays two high percentage upright Vessels sheath cells, and diffuse and scanty axial parenchyma. are solitary, never in long radial multiples. The secondary xylem of has become modified in structure of have typical of lianas. A few Saurauias have trilacunar nodes but the majority species a bundles found of unilacunarnode more typical ofEricales. Medullary are in petioles some and surrounded modified cells. Loculicidal, species stomata are not by subsidiary capsular 430 BLUMEA VOL. 23, No. 2, 1977

fruits are known in Saurauia and seeds are small, unitegmic exotestal, with cellular endo- starch. have seeds that arillate. sperm lacking Many species, however, are The similarities between the Actinidiaceae and has been are impressive as pre- several of viously pointed out by workers including one us (Dickison, 1972). Among these resemblances the sclerified of are unitegmic-tenuinucellate ovules, syncarpy, nature cells

lining the fruit locules, and seeds that have the outer cells of the testa with thickened inner

all features that also in in walls; occur Paracryphia and when considered conjunction with

wood lead to the that the anatomy suggestion genus may represent an early divergent offshoot of the Dillenialean-Thealean-Ericalean line. Fibrous endothecia and spurless

stamens, as foundin Paracryphia, are not common in Ericales but do occur in Clethraceae.

material did not characterization of the ovule of Our permit an accurate Paracryphia as

crassinucellate or tenuinucellate, although this is clearly an important consideration (see that needs clarification. Philipson, 1974; Palser, 1975) Paracryphia differs significantly from the above assemblage, and particularly Actinidia-

undifferentiated ceae, in possessing an perianth, sessile stigmas, and in having a com- paratively simple floral vasculation. Paracryphia also lacks the complex trichome morpho- and leaf with that logy, raphides, craspedodromous venation rigidly percurrent tertiaries

are so typical of Actinidiaceae. The large number of carpels in Paracryphia is also non- ericalean this condition is in Actinidia and in although seen up to ten carpels may occur

some Ericaceae (e.g., Arctostaphylos, Gaylussacia). Although the floral vasculature of Para-

offers few obvious clues the of the flowers of cryphia regarding relationships genus,

Saurauia are richly vascularized and characterized by numerous fusions of floral bundles in

The the contrast to the pattern in Paracryphia. occurrence of septal bundles in gynoecium of Saurauia is similar to many members of the Ericales and unlike Paracryphia. Actinidia, of however, shows anabsence septal bundles accompanied by the fusion of the two ventral bundles of each carpel. Pollen of Actinidiaceae is small, tricolporate, and not highly

sculptured.

Few similarities are evident between Paracryphia and primitive Dilleniaceae which have

flowers with a well differentiated basically pentamerous perianth, an apocarpous gynoe-

cium containing bitegmic ovules, raphide crystals, and never produce capsular fruits. lacks the characteristic of all of Paracryphia rigidly percurrent secondary veins so genera Dilleniaceae of Hibbertia. except various species

In the course of reviewing the salient morphological and anatomical features of Para- became focused ofsimilarities between cryphia our attention again on the large number it and the genus , also from New Caledonia, and New Guineaand Queensland.

This a chaotic has been studied genus, having very taxonomic history, recently thoroughly

of and concluded that deserves familial status by one us (Baas, 1975) it was Sphenostemon and has affinities with Aquifoliaceae and Icacinaceae. Sphenostemon is anatomically circum- scribed by the following features: hairs if present unicellular, stomata anomocytic, petiole with closed vascular connected with the trilacunar, a simple open or system stem through a three-trace node, cork arising in the subepidermis, secondary phloem with diffuse to tangentially arranged fibers and sieve tubes with compound sieve plates, styloid crystals in the ground tissue of leaf and stem and in the secondary phloem, wood with scalari-

vessels scalariform intervessel fibre- formly perforated (with 32—185 bars), pits, long of tracheids, diffuse-in-aggregates parenchyma, heterogeneous II rays (Kribs, 1968) two All these distinct sizes. characters from vegetative anatomy agree remarkably well with of The differences and wood those Paracryphia. in petiole vasculation rays is not regarded

external the decussate of also as significant. In morphology 4 perianth parts Sphenostemon recall well the leaf Leaf Paracryphia, as as occasionally pseudo-verticillate arrangement. W. C. Dickison & P. Baas: Paracryphia 431

since veins venation harmonizes with this pattern both genera have secondary brochido- dromous with marginal arches developed apically; tertiary veins are weakly transverse. The detailed morphology of the reproductive structures: stamens, ovary, and pollen grains different. The ovules of are, however, quite pendulous anatropous Sphenostemon are, nevertheless, also unitegmic (Bernardi, 1964). The indehiscent fruit with one or two stony seeds is quite unlike the capsular fruit of Paracryphia*. Vasculation of the bilocular and gynoecium of Sphenostemon consists of8—12 bundles, two ofwhich are fused extend the locules. the of the locules the central vertically in the tissue separating two At apex strand bifurcates with each branch vascularizing one ovule (Bailey & Swamy, 1953).

In spite of the significant differences in reproductive morphology, we cannot ignore the most striking similarities between Sphenostemon and Paracryphia in all aspects of vegetative features such the anatomy, supported by general reproductive as 4-merous, decussate, with undifferentiated perianth; superior ovary anatropous, unitegmic ovules; basifixed the of the anthers; and to a lesser extent sculpturing pollen exine (cf. Lobreau-Callen, of evolution I975)- To explain all these similarities in terms convergent seems unreason-

it is in related. able and our beliefthat Sphenostemon and Paracryphia are some way Although

of concluded that should be treated a member of the one us (Baas, 1975) Sphenostemon as

Celastrales, this was only based on similarities with Aquifoliaceae, Phellinaceae, and Icacina-

the inclusion of and Icacinaceae in the order ceae. However, Aquifoliaceae appears

to be more and more doubtful. Thome (1968) referred Aquifoliaceae to the Theales, and although the second author (1975) strongly criticized the placement of this family in an that the order different from of apparently related Icacinaceae, there are a fairly large number of anatomical similarities between Theaceae and Aquifoliaceae, and accordingly, with Icacinaceae. According to Corner (1976) the seed structure (unitegmic, exotestal) of

Icacinaceae and Aquifoliaceae is incompatible with a Celastralean alliance and could be more in line with Theales, where (exclusive of Theaceae) there is a unitegmic tendency.

Recently, Dahlgren (1975) included Icacinaceae in , and earlier Huber (1963) favored inclusion of Aquifoliaceae in thatorder. IfParacryphia is related to these two families

'through' Sphenostemon, thenall families included in Cornales deserve attention as putative allies, as well as all families ofCelastrales. Likewise, the similaritiesbetweenParacryphia and

Actinidiaceae reported above should invite comparisons with all families of the orders have been Theales and Ericales to which that family has been assigned. Such comparisons all in widest that all made, taking four orders the possible sense; is, including families that have been assigned to these orders by different workers (Melchior in Engler, 1964; Hut-

et and chinson, 1959,1974; Lanjouw al., 1968; Cronquist, 1968; Takhtajan, 1973; Dahlgren, one-hundred families well all families listed I975)- Just over were considered, as as in Metcalfe Chalk scalariform their and (1950) as possessing perforations in wood. Following this a comprehensive survey of the anatomical and morphological characters of large sample of dicotyledons, only the following families emerged, sharing enough characters with Paracryphia to merit further considerations as a close relative: Sphenostemonaceae,

Actinidiaceae, Aquifoliaceae, Icacinaceae, Theaceae, Pentaphylacaceae, Clethraceae, Cyrillaceae, and . Of these Sphenostemonaceae, Theaceae, and Actinidiaceae were discussed above. Aquifoliaceae and Icacinaceae (p.p.) share a fairly large number ofanatomical features with Paracryphia; the floral morphology is, however, quite different and deviates to a from greater degree Paracryphia than either Sphenostemon or Actinidiaceae. Pentaphylacaceae,

In the with *) most descriptions fruits of Sphenostemonare referred to as drupes oneor two bony pyrenes.

This is, however, erroneous (cf. Bailey, 1956): the pericarp remains more or less soft throughout but the seed develops a very thick testa of many layers ofsclerified cells. 13LUMEA VOL. No. 432 23, 2, 1977

Clethraceae, Cyrillaceae, and Escalloniaceaeall share a rather primitive wood structure with has also been Floral Paracryphia. Escallonia reported to possess styloids. morphological

characters of these four families still deviate considerably from Paracryphia, however, and

affinities with if be rather their Paracryphia, any, must remote. Pollenmorphologically the small, unspecialized, tricolporate, reticulated pollen of

in Actinidia- Paracryphia is consistent with the type ofpollen grains found many Theaceae,

transitional between Theales and Ericales ceae, Clethraceae, and Cyrillaceae, i.e., groups

(Walker & Doyle, 1975). families above have time It is noteworthy that many of the mentioned at one or

another been transferred from the Celastrales to Theales or Ericales or vice versa. Cyrilla-

ceae have thus been treated in Ericales as well as in Celastrales, and although the former

affinity is favored by most contemporary systematists, there are striking anatomical Actinidiaceae ranked similaritieswith Aquifoliaceae (cf. Baas, 1975). (including Saurauia) are Theales. Clethraceae moved at the base of the Ericales or included in Similarly, have been

Theales Thorne also face uncertain future: from Ericales to by (1968). Aquifoliaceae now an under retention in Celastrales or transferal to Theales. Pentaphylacaceae can be found both

ordinal headings. much We believe the Cornales should be excluded from this discussion in as as the

of affinities rather haphazard suggestions Cornalean of Icacinaceae and Aquifoliaceae seem anatomical tendencies in that to be too much in conflict with the morphological and order. The fact that for the orders Ericales, Theales, and Celastrales several families seem ordinal to be exchangeable makes the problem of assigning Paracryphia to an existing

difficult. shows affinities, close ones, to concept very Paracryphia although never very

ofall threeorders. This well be taken an to some representatives may as argument support and from the Thea- the view that Paracryphia represents an independent early divergence With the dis- lean, Ericalean, and/or Celastralean (pro parte?) lines of evolution. present

about the definition of orders it is not which of the three agreements readily apparent of the families alternatives best reflects the evolutionary relationships genus. The Actini- and closest relatives of diaceae, Theaceae, Sphenostemonaceae appear to us to be the extant that familial Paracryphia. The present study fully confirms the view Paracryphia merits

status.

of between the The emerging synthesis is of a reticulate pattern relationships Theales, and Ericales families like Celastrales pro parte, through Paracryphiaceae, Sphenostemonaceae, should Actinidiaceae, Cyrillaceae, and Pentaphylacaceae. These relationships present an

interesting challenge to contemporary system makers.

ACKNOWLEDGEMENTS

from the Netherlands for the Advancement of This research was supported by a grant Organization

to acknowledge his appreciationfor Pure Research (Z. W. O.) awarded to W. C. Dickison. He would like

this and thank the staff of the for the considerations shown him his support Rijksherbarium many during due S. McKee New Caledonia) for collecting ex- stay in Leiden. Special thanks are to Dr. H. (Noumea, cellent materials used in this study; to Dr. Sherwin Carlquist (Claremont, California) for making available de Phanerogamie, the materials of Paracryphia in his collection; and to the Director of the Laboratoire of all herbarium of Museum National d'Histoire Naturelle (Paris) for the loan specimens Paracryphia. the Mr. J. Muller, palynologist,Prof. Dr. R. Hegnauer, and several members of the tropical staffat Rijks-

for their discussions. herbarium arc acknowledged stimulating W. C. Dickison & P. Baas: Paracryphia 433\I

x Plate of alticola. — of leaf I. Vegetative anatomy Paracryphia 1. Transverse section (MacKee 4993), 220.

— Paradermal 2. section of abaxial leaf epidermis showing anomocytic stomata (McKee 31070), x 220. —

3. Transverse section ofbark showing interrupted bands of fibrous cells ( McKee 31070), x 530. — 4. Longi- tudinalxsection of bark showing styloid crystals ( Schlechter 15326), 530. — 5. Longitudinalsection through

sieve a tube element from the secondary phloem showing a compoundsieve plate (sp) ( McKee 31070), x 870. BLUMEA 434\II VOL. 23, No. 2, 1977

1 2

Plate Wood of alticola — Transverse section, — 2. II. anatomy Paracryphia (McKee31070). 1. x 85. Tangential

x — — section of vessel elements scalariform 530. section, x 85. 3. Longitudinal showing perforation plates, — of vessel element scalariform x 5. 4. SEM perforationplate showing partial reticulation, 430. Longitudi-

nal section of vessel element showing scalariform intervascular pitting, x 530. W. C. Dickison & P. Baas: Paracryphia 435\III

— Plate III. Reproductive morphology of Paracryphia alticola. 1. Longitudinalsection of ovule illustrating

— of fruit anatropous, unitegmic condition ( McKee 26820), x 670. 3 (top). Transverse section mature showing prominent sclerification of cells lining the locules (McKee 31070), X 110. — 3 (bottom). SEM of mature seed ofthe anticlinal walls of cells showing undulating pattern epidermal (McKee 31070), x 50. VOL. No. 436\IV BLUMEA 23, 2, 1977 W. C. Dickison & P. Baas: Paracryphia 437

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EXPLANATION OF PLATE IV

Pollen of — view Plate IV. morphology Paracryphia alticola (McKee 26820). 1. Median polar showing _

circular outline, X1330. — 2. Equatorial view, high focal level showing endoaperture and reticulate

sculpture, X1330. — 3. Same grain, equatorial view at lower focal level showing pattern of columellae

distribution, x 1330. — 4. SEM of acetolyzed pollen grain as seenin equatorial view. Note the retirugate

— in sculpturing in the mesocolpium, X3500. 5. SEM of acetolyzed grain equatorial view, X1700. — 6. TEM of FAA preserved, nonacetolyzed grain sectioned obliquely along the equatorial axis. Note the

intine under the and the thick endexine the ofthe prominent (in) colpae (e) along margins apertures, x 4400.

— section exine is into 7. Same grain showing a median throughthe endoaperture(p). The stratified en-

dexine (e), foot-layer (f), columellae (c), and tectum (arrow), X9900.