Boianical Journal Ofthe Linnean Sociely (1988), 97: 33-48. With 9 figures

Notes on the genus Pandanus (Pandanaceae) in Tahiti

BENJAMIN C. STONE, F.L.S.

Department of Botany, Academy of Natural Sciences, 19th 623 Parkway, Philadelphia, PA 19103, U.S.A.

Received October 1986, accepted for publication January I987

STONE, BENJAMIN C., 1988. Notes on the genus Pandanus (Pandanaceae) in Tahiti. Pandanus tectorius Park. ex Z. is reinstated with choice o f a representative pistillate specimen, following the example o f St. John who selected a representative pistillate specimen for P . odoratissirnus L. fil., thus making possible the comparison and contrast of other taxa distinguished on carpological characters. Most other putative endemic species are reduced to synonymy under P . teclorius, but the interior, upland species P . papenooensis St. John is shown to be a well-marked endemic (but with P . aoraiewis St. John as a synonym). One new combination is made under P . iectorius for a variety from Raiatea.

ADDITIONAL KEY WORDS:-Endemic species - new combination - Pandanus papenooensis - pandanus tectorius - representative specimen - typification.

CONTENTS

Introduction . . . . . . . . . . .

Reinstatement, synonymy, typification, and description of Pandanus tectorius

Tahitian varieties ofpandanus teciorius . . . . . . . A variety of Pandanus tectoriur from Raiatea . . . . . .

Introduced species . . . . . . . . . . References . . . . . . . . . . . .

Diversity and species recognition in Tahitian Pandanus . . . .

A representative pistillate specimen for Pandanus tectorius

Endemic species . . . . . . . . . . .

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33 35 36 38 43 44 44 46 48

INTRODUCTION

The significance of Tahiti in the taxonomy of the genus Pandunus is considerable, since it was in Tahiti, on the first voyage of Captain James Cook, that European botany first encountered the pandan which today still bears the name that Joseph Banks and Daniel Solander assigned to it a t the time: Pandanus tectorius. However, by a strange quirk of fate the authorship (at least in strict bibliographic terms) of this binomial must be attributed to S. Parkinson, who was the young illustrator employed by the aristocratic Banks, and, unlike Solander (who was a pupil of Linnaeus), was not a trained botanist. As ex- plained elsewhere (St. John, 1972, 1979a, b; Stone, 1976, 1982), Parkinson's diary

33 0024-4074/88/050033 + 16 $03.00/0 0 1988 The Linnean Society of London

34 B. C. STONE

(in the form of a German translation attributed to “Z”, which probably refers to J. P. du Roi) is actually the source of the binomial in its first acceptable (i.e. botanically valid) appearance, dated 1774 (Parkinson, 1774), although it must certainly have been coined by Banks and Solander. Moreover, this publication is the first appearance (according to some authors) of the generic name Pandanus in a botanically valid form. However, the name can logically be argued to have first appeared in 0. Stickman’s thesis as published by Linnaeus (1754); arguments pro and con have been considered (Airy Shaw, 1962; St. John, 1960; Stone, 1983). But in any case, Pandanus tectorius Parkinson ex “Z” is clearly the first published species name of its genus, and i t is based clearly on a plant from Tahiti. The type of P. tectorius, recently discussed by St. John (1979a, b), is a pistillate specimen from Tahiti preserved, with paintings of the male, in the British Museum (Natural History). The type species of the genus Pandanus is in fact Pandanus tectorius, and not, as earlier supposed, P. odoratissimus L. fil. The understanding of P. tectorius is accordingly crucial to knowledge of the genus Pandanus; and the importance of Tahiti, therefore, in the taxonomy of Pandanus can be readily appreciated.

The species of Pandanus found in Tahiti have been the subject of an important paper by Martelli (1933a) and a recent paper by St. John (1979b). A botanist, knowing that Tahiti is the type locality of P. tectorius, would expect to find this species well accounted for in both these publications; surprisingly, he would be disappointed in this expectation. Martelli’s paper enumerates nine species of Pandanus, several as new species: P. menziesii Gaudich., P. drolletianus Martelli, P. macfarlanei Martelli, P. tahitensis Martelli, P. prismaticus Martelli, P. distinctus Martelli, P. papeariensis Martelli, P. odoratissimus var. savaiensis Martelli, and P. tessellatus Martelli. O n more thorough reading, one finds in the introductory part of Martelli’s account (Martelli, 1933a: 150), a reference to P. tectorius, which states: “To the flora of this island belongs also the form called by Solander, Pandanus tectorius, about which I have previously published a note (Pandanus tectorius o Pandanus odoratissimus?, Nuovo Giornale Botanic0 Italiano, 1929).” This shows that at this stage, Martelli regarded P. tectorius ‘of Solander’ as synonymous with P. odoratissimus (of Linnaeus the younger), and considered the latter name as the correct one. For this reason, it can be deduced that the plant named P. odoratissimus var. savaiensis in the account (though not numbered, it comes at the eighth place) is actually P. tectorius; however, i t is not the ‘form’ described by ‘Solander’ but a variety, originally described from Samoa by Martelli himself, reported as a cultivated plant in Tahiti,

Further surprises await the botanist perusing the recent review by St. John (1979b). In his study, 13 endemic species are accepted in the Tahitian flora, as well as two introduced cultivars. St. John provides a key in which there is no mention of P. tectorius. The explanation, given in the opening paragraph, is that since the holotype is a very young pistillate inflorescence, “P. tectorius must rest as a species dubia”. This might seem to lay the problem of P. tectorius to a final rest. However, this is not the case; the problem is neither so simple, nor so complex, that the name P. tectorius can be thus ‘abandoned’. In the following pages it is my intention to show that the name can and must be applied, and with understanding, to plants found presently in Tahiti. To do this, two basic requirements are here met: first, a new taxonomic study of the Tahitian pandans; and second, the choice of a ‘representative pistillate specimen’ for

PANDAJUS (PANDANACEAE) IN TAHITI 35

P. tectorius , which, at long last, provides a basis for the recognition of species and varieties considered to differ from it on the basis of pistillate characters.

DIVERSITY AND SPECIES RECOGNITION IN TAHITIAN PANDANUS

In Tahiti, Pandanus is highly diverse; moreover, there is certainly more than one species present. An attempt is made below to show that the number of distinct species accepted by Martelli (nine) and, more recently, by St. John (13) is too high. In this context, it may be pointed out that diversity in the Tahitian pandans can be traced to three different causes: (a) the presence of both endemic and non-endemic wild species; (b) the presence of introduced species; and (c) the infraspecific variation of some species.

Endemic species. According to Martelli, of the nine Tahitian species he accepted, eight are endemic. According to St. John, there are 13 endemic, and no non-endemic indigenous, species. The present review accepts one species as indubitably endemic, P. papenooensis.

Indigenous species, non-endemic. Pandanus tectorius, while its type collection is from Tahiti, is not limited to the island but has a broad distribution in the Pacific, extending north to the Hawaiian Islands and west to Australia and New Guinea, into the Moluccas and Philippines. It varies greatly and has evolved in to many local forms and varieties (Martelli called them “microspecies”) which are identifiable chiefly or only by carpological characters. It has also been subject to some human selection and has become a cultivated plant, existing in several cultigen forms, of which P. tectorius var. laevis (P. spurius CV. Putat) an unarmed form, and several horticultural forms with variegated leaves (e.g. P. sanderi and P. veitchii), are notable. In Tahiti, there are several carpological forms of P. tectorius. It is argued here that these forms, discriminated by Martelli as binomial species (though he called them “microspecies”) and by St. John as endemic species, should be considered as infraspecific variants. Some of these may deserve taxonomic status at varietal rank.

Introduced species. Both Polynesian and later European introductions have occurred. Such plants, often but not always in gardens, cannot readily be traced to their origin. Useful forms such as P. tectorius var. laevis may have originated in Polynesia but are now widespread, and no clear-cut history of domestication or transport is available. Variegated forms may be introductions, but they could also occur as spontaneous mutants among indigenous native populations. Indeed mutation and back-mutation with regard to leaf variegation and to the armed versus unarmed leaf margins occurs readily, and chimaeras are known (cf. P. baptistii; Stone, 1973).

Martelli’s taxonomic understanding of the variation encountered in the Tahitian (and similar) pandan-floras deserves further comment. In the paper of 1933a, and in a preceding paper (1932), Martelli several times indicates that the taxa he describes are “microspecies”. In some places he uses phrases such as “forms or microspecies” or “varieties or microspecies” indicating that, although allocated binomials, these taxa are not truly equivalent to species. Indeed, in his key (“Quadro sinoptico delle Specie”) he divides the Tahitian pandans into two

36 B. C. STONE

groups: Pandanus odoratissimus, and Pandanus pedunculatus ( 1932: 1 16- 1 1 7). Under these headings he enumerates the eight binomials in this manner:

Pandanus odoratissimus L. (subspecies). [The synopsis then leads to the binomials P. menziesii, P. drolletianus, P. macfarlanei, P. tahitensis, P. prismaticus, P. distinctus, and P. papeariensis] .

. . . P: tessellatus. Pandanus pedunculatus R. Br. (subspecies).

From this peculiar arrangement it would appear that Martelli intended the binomials to refer to subspecies, but he never uses the trinomial convention for this rank. Yet in other instances he actually refers to some of the binomials as species (e.g. P. mentiesii, P. distinctus, P. papeariensis, P . tessellatus). In other words, his language suggests that he meant to subordinate these taxa to P. odoratissimus (or, in the case of P. tessellatus, to P. pedunculuatus), while his usage (and sometimes his wording also) indicates that he accepted them as species. This inconsistency is only of importance in an interpretation of his understanding of the variation; from a nomenclatural point of view, the taxa accorded binomials are species without qualification.

The allocation of some of the “microspecies” to P. odoratissimus and one other to P. pedunculatus might suggest either a supraspecific classification (such as a ‘grex’, complex or superspecies), or a typological clustering with the aim of assigning geographical affinities.

The impression made by Martelli’s treatment is that, for some reason, he could not employ a trinomial nomenclature (although he did use it in other cases) and yet did not really feel convinced that the species (binomials) were justified. This ambiguity is not seen in his companion work on the pandans of Rarotonga (Martelli, 1933b).

REINSTATEMENT, SYNONYMY, TYPIFICATION AND DESCRIPTION OF PANDANUS TECTORIUS

Pandanus tectorjus Parkins., Journal of a Voyage to the South Seas in . . . H.M.S. Endeavour (1773), nomen jugatum; idem. ed. ‘Z’ (trans]. = J.P. du Roi?) in Der Naturforscher, 4: 250 ( 1774), nomen normale non-jugatum descriptioque.

TYPE: Tahiti, 1769, Banks and Solander (BM). St. John, Botanical Journal of the Linnean Society, 4: 309 (1972); Pacijic Science, 33: 395 (1980). Stone, Kew Bulletin, 31: 60 ff. (1976); Botanical Journal of the Linnean Sociely, 85: 136 (1982).

SYNONYMS: P. bothreus St. John, Pacijic Science, 33: 407, f. 41 1 (1979). TYPE: Tahiti, near Papeete, “in M r Brander’s yard” viii 1929, Wilder 7 (BISH).

P. dicheres St. John, Pacijic Science, 33: 409, f. 412 (1979). TYPE: Tahiti, Hitiaa District, Motu Puuru, xi 1930, Grant 4448.2 (BISH).

P. distinctus Martelli, Uniuersity of California Publications in Botany, 17: 153, pl. 19-20 (1933); St. John, Pacijic Science, 33: 410, f. 413 (1979). TYPE: Tahiti, Papeari, near the seashore, on Harrison Smith Property, viii 1929, Wilder 5

P. drolletianus Martelli, Webbia, 2: 427 (1907); 4 ( 1 ) : 12, 48, 94, t. 3, f. 6-8 (1913-14); St. John, Paczjic Science, 33: 413, f. 414 (1979). TYPE: Tahiti, Drollet in 1907 (No. X in Herb. Martelli, FI).

(UC) *

PANDANUS (PANDANACEAE) IN TAHITI 37

Figure I . Pandanus tectorius Park. ex Z. Phalanges of various collections (all from Tahiti) representing carpological variability as 1 I synonymous binomials. A. P. distinctus Martelli. B. P. drolletianus Martelli. C. P. rnacfarlanei Martelli. D. P. dicheres St. John. E. P. prismaticus Martelli. F. P. papeariensis Martelli. G. P. subradialus St. John. H. P. tahitenszs Martelli. I . P. kamptos St. John. J. P. parksii St. John. K . P. bothreus St. John. Drawn or redrawn from the type specimens or original protologue illustrations.

P. kamptos St. John, Pacific Science, 33: 416, f. 415 (1979). TYPE: Tahiti, Hitiaa District, Motu Puuru, xi 1930, Grant 4448.2 (BISH).

P. rnacfarlanei Martelli, Webbia, 2: 426 (1907); 4(2): 22, 54, 94, 97, t. 2, f. 2-4 (1913); St. John, Pacific Science, 33: 418, f. 416 (1979). TYPE: Tahiti, “presso Papeete nell’Isola Tahiti” in 1904 (No. XI in Herb. Martelli, FI).

P. papeariensis Martelli, Societa Toscana Scienta Naturali, Atti, Memoria, 42: 1 17 (1932); University of California Publications in Botany, 17: 153, pl. 21 (1933); St. John, Pacific Science, 33: 422, f. 417 (1979). TYPE: Tahiti, Papeari, near the seashore, Harrison Smith Property, ix 1929, Wilder 2 (UC).

P. parksii St. John, Pacific Science, 33: 426, f. 419 (1979). TYPE: Tahiti, Arue Point, seashore, vi 1922, W. A. Setchell & H. E. Parkes 367 (BISH).

38 B. C. STONE

P . prismaticus Martelli, Societa Toscana Scienza Naturali, Atti, Memoria, 42: 1 17 (1932) University of California Publicationss in Botany, 17: 152, pl. 18 (1933); St. John, Pac& Science, 33: 428, f. 420 (1979). TYPE: Tahiti, Papeari, near the seashore, on Harrison Smith Property, viii 1929, Wilder 2 (UC).

P . subradiatus St. John, Pacijic Science, 33: 430, f. 421 (1979). TYPE: Tahiti, Pueu District, Motu Nono, beach, coral soil, ii 1931, Grant 5430 (BISH).

P . tahitensis Martelli, Webbia, 2: 428 (1907); 4 ( 1 ) : 33, 90, 94, t. 7, f. 1-3 (1913); St. John, Pacijic Science, 33: 432, f. 422 (1979). TYPE: Tahiti, 1905, MacFarlane (no. 2, in Herb. Martelli, FI).

P . tessellatus Martelli, University of California Publications in Botany, 17: 174, pl. 25 (1933). TYPE: Cook Islands, Rarotonga, viii 1929, Wilder 13 (UC).

var. tectorius. (Figs 1-5, 9)

TYPE: as above.

REPRESENTATIVE PISTILLATE SPECIMEN: Tahiti, Point Venus, head of beach,

Pyramidal branched tree to 8 m tall, proproots to c. 1 m long, slightly prickly. Trunk with brownish to greyish bark. Leaves mostly 1.0-1.7 m long, 60-80 mm wide near the base, gradually attenuate to the subflagellate apex. Undersurface proximally somewhat glaucous on each side of the midrib. Margins near base with antrorse greenish brown-tipped prickles 2.0-2.5 mm long, mostly 2-8 mm apart; near the middle, the prickles more appressed but otherwise similar or slightly more crowded; near the apex, prickles smaller, 1.0-1.5 mm long, about 2 mm apart. Midrib beneath near base unarmed, beyond the lower fifth or fourth of its length with antrorse prickles 1.5-2.0 mm long, rather irregularly spaced, distant, mostly 15-25 mm apart; near the middle, the prickles similar but more regularly spaced, mostly 10-15 mm apart; near apex, the prickles shorter, mostly 1.0-1.5 mm long, 2-5 mm apart. Longitudinal veins near leaf base c. 0.5 mm apart, but near apex c.0.3 mm apart, typically about 120-150 per leaf. Pistillate head solitary, pendulous, subglobose-ellipsoid, about 170 x 140 mm, obscurely trigonous, composed of about 108 phalanges, these (2-)4- to 10-celled mostly 5- to 7-celled (mean carpel number: 6.5). Phalanges 50-55 mm long, mostly 25-40 mm wide, usually slightly compressed, 5-6-angled in general outline in cross-section; apical 15 mm free, low convex with outer carpel tips larger than inner, somewhat incurved; stigmas subapical plane to oblique or erect, irregularly reniform to subovate or deltoid, mostly 1.5-2.0 mm wide, at inner edge of small or obscure pilear platform. Upper mesocarp cavernose-medullose-fibrous, in outer carpel tips c. 12 mm high; lower mesocarp fibrous-fleshy, c. 22 mm long. Endocarp approximately central, 20 mm high in the middle, shallower peripherally, osseous reddish-brown, traversed by a few paler fibres; locules as many as carpels, inner (fertile) ones obovoid-subfusiform, 15 mm long or less; endocarp walls to 2-3 mm thick.

30 viii 1981, Stone 24712 (BISH, K, KLU, L, P, PH, US, etc.).

A REPRESENTATIVE PISTILLATE COLLECTION FOR PANDANUS TECTORIUS

In his discussion of P. tectorius, St. John (1979a, b) quotes the description published by Parkinson. This is not a technical description, it is certainly inadequate by modern standards but does contain several clear indications of

P A N D A N U S (PANDANACEAE) IN TAHITI 39

Figure 2. Pandanus tectorius Park. ex 2. Entire cephalium representing the standard pistillate specimen (Stone 14712, Pt. Venus, Tahiti).

0 2 cm

Figure 3. Pandanus tectorius Park. ex Z. Phalanges from the standard pistillate specimen, Stone 14712, in profile (A, B), top view (C, with enlarged view of one stigma shown in D), longisection (E) and cross-section views (F).

40 B. C . STONE

morphology. Summarizing these one obtains the following brief diagnosis: tree; leaves long, sedge-like, serrate; plants dioecious; bracts white; male flowers fragrant, small; fruit the size of a (human) head, orange, formed of units (“cones”); bases of these when ripe sweet, edible, juicy. St. John states that “Nothing in the notes by Solander or in the publication by Parkinson is of any diagnostic specific value. I t is clear that they were written about a species in the section Pundunus”. It is obvious that Solander, Banks and Parkinson (whose judgement is scarcely at stake here) had seen numerous individual pandans while in Tahiti; their judicious conclusion was that one species only was represented. If in the light of modern knowledge we know more, that is not to say that Parkinson’s and Solander’s descriptions lack any diagnostic specific value in their own terms. If we look at the other sources, i.e. Solander’s notes, the lectotype specimen (selected by St. John and preserved in the British Museum of Natural History) and Parkinson’s drawings, we have a reasonable basis for recognizing Pandunus tectorius. The chief gap is the particular form and nature of the ripe fruit. This is significant a t present because of the variability in carpological features in Pundanus, and because fruit characters have been used ever since as the main taxonomic basis of species discrimination.

What is lacking can be supplied by an appropriate choice of a fruiting collection that can be associated with the species. I designate this as the ‘representative pistillate specimen’. This is not an innovation, but relies on the precisely similar action by St. John himself, who designates and describes a “standard specimen” for Pundanus odoratissirnus L. fil. (St. John, 1979a). St. John remarks: “Of the Thunberg collection, there exists now only a staminate flowering specimen and this, in the Linnean Herbarium, is here designated as the lectotype.. .”. If this is a proper nomenclatural action (and I agree that i t is), then precisely the same action can be taken to establish P. tectorius; the two cases are in fact exactly alike. The reason why St. John has chosen a “standard specimen” for P. odoratissirnus is to document the ripe fruit, and he explained that since “Only one species occurs in this littoral zone on Ceylon; consequently it is considered definite that the lectotype and the standard specimen both represent the same species, P. odorutissimus. The species is restricted to Ceylon”. The “standard specimen” designated by St. John is his collection 26222 from the beach south of Colombo. Fruits of this specimen are illustrated by St. John (1979a) as fig. 409. However, St. John does not mention another of his own collections, no. 26228, also from Ceylon, which I previously illustrated (Stone, 1967: fig. 6). I t will be evident that the variation in form between these two collections is as great (carpologically) as that between some of the Tahitian pandans; for example, between Drollet X (the type of P. drolletiunus Martelli) and Mucfurlune X I (the type of P. mucfarlunei Martelli). Other Ceylon collections which I unhesitatingly refer to P. odoratissirnus (Stone, 1981) show that a comparable, though somewhat less extreme, variability is to be found among Ceylon pandans, as is seen in the Tahitian pandans. The conclusion that P . odoratissimus is the only maritime pandan species in Ceylon is not founded on fixed, invariable carpology. The implication that variation is lacking is neither correct nor necessary as a justification for the establishment of a “standard specimen”. The justification must really be based on a different consideration- the need to have a morphological reference against which other, and subsequent, collections of fruits can be tested. It is primarily to serve as a

P A N D A N U S (PANDANACEAE) IN TAHITI 41

5 Figure 4. Pandanus tectorius Park. ex Z. Cephalium of H . Harms s.n. (identical to the form named “P. papeariensis”) . Figure 5 . Pandanus leclorius Park. ex Z. Cephalium of type collection (Wilder I ) of “ P . papeariensis Martelli”.

standard for the classification of infraspecific variants. In previous studies (e.g. Stone, 1976) the recognition of varieties within P. tectorius has been sustained as a practical and biologically realistic basis for recording some of the inherent variability in morphology and geographical patterns in this difficult species. This method was used by Warburg, by Martelli, and even by St. John, and is used here.

By using the representative specimen designated here, it is now possible to discriminate other taxa on carpological characters. If other botanists wish to recognize more than one species or more than one subspecies or variety, the representative specimen permits this on a factual basis.

42 B. C. STONE

Figures 6 and 7. Pundunus Papenooenris St. John. Cephalium (Fig. 6) and phalanges (Fig. 7 ) from Sfone, Fosberg @ Gunin 14701, from Teravao plateau, Tahiti.

0. & I. Degener (1981, 1982) basing their statement on a cited work by H. Manitz, have made the claim that the person identified in Der Nuturforscher only as “Z” is actually J. P. du Roi. Irrespective of the validity of the claim (which may well be true), the name du Roi does not appear in the publication and is thus of little use in bibliographic search. The Degeners published the name “Pundunus tectorius (J. P. du Roi) Deg. & Deg., comb. nov.”, which has no function, as this is neither a transfer from another genus nor the change in status of the taxon.

Of the 12 synonyms listed above, all but one are based on Tahitian collections. The exception, P. tessellutus, is a peculiar case. Martelli described this species twice, based on two different collections, one (designated lectotype by St. John) from Rarotonga (Wi lder 13, cited above), the other from Tahiti (Wi lder 3,

PANDANUS (PANDANACEAE) IN TAHITI 43

in part). He did not designate a type as such. The Rarotongan collection included an entire cephalium, notable by its possession of a reduced lateral cephalium on the same peduncle (a rare but not unknown occurrence in the species). Martelli mentions this and concludes that the “racemose” inflorescence is not of taxonomic importance (in which he is certainly correct in this instance). St. John omits P. tessellatus in his account of the Tahitian species, so it is clear that he regards the Rarotongan collection as the lectotype. This is also preferable because, as Martelli states, the phalanges from Tahiti which he identifies as belonging to P. tessellatus were in a lot (Wilder 3) which was a mixed batch, containing phalanges of the taxa P. macfarlanei Martelli, and P. menziesii Gaudichaud. The possibility of mingling of specimens makes inadvisable the selection of the Tahitian material as lectotype for P. tessellatus.

The phalanges identified as P. menriesii by Martelli have been re-identified by St. John as his P. subradiatus (based on a different type collection, Grant 5430). Still other material (phalanges) from the Wilder 3 collection were identified by Martelli as P. macfarlanei. St. John excludes them from that species and re- identifies them as belonging to his P. dicheres (through this is not clear from his discussion of P. dicheres, rather it is implied in his remarks under P. macfarlanei).

It is likely, therefore, that Wilder 3 represented a collection of phalanges from the ground, picked up and collated under the assumption that they belonged to one species. In segregating the collection into three parts, Martelli and St. John are probably correct in that several parent trees were probably represented, so the collection may be regarded as a mixture. Despite this, I conclude that all are carpological forms of P. tectorius.

The discussion of P. distinctus Martelli by St. John is of interest also, as he remarks that it “is very similar to the slightly older species P. papeariensis Martelli, which grew in the same coastal grove at Papeari. The only actual differences seem to be that P. distinctus has the phalanges 2.8-4cm wide and mostly obovoid to broadly obovoid, while P. pafieariensis has them 2-2.7 cm wide and oblanceoloid. However, by careful picking one can find phalanges of each that seem inseparable”. Yet both species are maintained in St. John’s account. I conclude that both are carpological forms of P. tectorius.

Pandanus parksii, cited above, must not be confused with P. tectorius var. parksii (Martelli) Moore, which is based on a Tongan type specimen.

TAHITIAN VARIETIES OF PANDANUS TECTORIUS

Pandanus tectorius var. tectorius. The typical variety is represented by the type collection; by the standard

pistillate specimen cited above; and by other collections which match these.

P. tectorius var. australiensis Martelli, Webbia 4 ( I ) : 33 (1913); 4(2): 409, t. 18, f.

This variety, described from Australian specimens, is also represented in the New Hebrides and Fiji (Stone, 1976).

P. tectorius var. drolletianus (Martelli) Stone, Kew Bulletin, 31: 66 (1976).

The phalanges of this variety are slightly broader and longer than in var. tectorius.

1-3 (1914). SYNONYM: P. tahitensis Martelli, Webbia 4(1 ) : 33 (1913).

SYNONYM: P. drolletianus Martelli, Webbia, 4( I ) : 33 ( 19 1 3).

44 B. C. STONE

Some other varieties may deserve nomenclatural recognition, whether based on existing described taxa or not, but for the moment, no further discrimination is pursued pending a more intensive study of the variation and causes thereof.

A VARIETY OF PANDANUS TECTORIUS FROM RAIATEA

Further support for the view that P. tectorius is a variable species in regard to fruit form but comparatively invariant (apart from striking mutations) in other ways is provided by Moore’s study of Raiatean plants (Moore, 1933), wherein he recognizes a variety of P. tectorius (var. parksii (Martelli) Moore), and another of P. tahitensis (var. exiguus Moore). The latter is also clearly another carpological variant of P. tectorius, viz.:

Pandanus tectorius var. exiguus (Moore) B. C . Stone, comb. et stat. nov.

BASIONYM: P. tahitensis var. exiguus Moore, Bishop Museum Bulletin, 202: 1 7 ( 1933). TYPE: Raiatea, in valley at Tioo, 26 ii 1927, Moore 632 (BISH).

ENDEMIC SPECIES

There is one very distinctive endemic Pandanus in Tahiti; it has already received two names, P. aoraiensis and P. papenooensis. (The species was discovered by Martin L. Grant, who left manuscript names for the collections he made.) Because this species is an upland species, it was probably not seen by earlier collectors, or if seen was mistaken for the coastal species. Its distinctness resides not only in carpological characters; it is not, therefore, merely another more extreme ‘variant’ of the P. tectorius complex, although it is of course related, and certainly a member of section Pandanus. Of the two binomials available, I have selected P. papenooensis and include P. aoraiensis within it conceptually. St. John, who published both names, has provided good full descriptions and illustrations. A comparison will readily demonstrate their similarity. St. John discusses the slight differences between them and indicates the pair as being very closely related. In concurrence I go farther and consider them conspecific.

Because of the opportunity of seeing this species in the living state, I can add here what is a hitherto unnoticed, but very interesting and, I believe, important character. This character is vegetative, and immediately distinguishes P. papenooensis from all forms of P. tectorius. There is at the base of each mature leaf a large, purplish area, several centimetres in length and breadth, irregular in form. This area is of course less noticeable in dried specimens (though it may be the basis of St. John’s descriptive remark “base unarmed, reddish” in the diagnosis of P. aoraiensis), and its coloration is never present in P. tectorius. Additionally, there is a palpable difference in texture; the living leaves of P. Papenooensis are brittle, even near the base, quite unlike those of P. tectorius. The upper surface of the leaf in P. papenooensis is somewhat glaucous, and the lower surface is paler and glaucous.

These additional vegetative characters support the taxonomic distinctness of P. papenooensis.

Some additional descriptive notes, and a description of seedlings, are provided below.

Pandanus papenooensis St. John, Pacijic Science, 33: 424, f. 418 (1979). TYPE: Tahiti, Papenoo District, Ana Pua, 1220 ft alt, ix 1930, M. L. Grant 4079 (BISH) (Figs 5-8, 9).

PANDANUS (PANDANACEAE) IN TAHITI 45

mrn

Figure 8. Pandanus papenooensis St. John. Phalanges (from Stone, Fosberg, tY Guerin 14701). A-In profile; B-in longisection; C-in top view; D-in cross-section through endocarp; E-detail of apex of one carpel with stigma.

SYNONYM: P. aoraiensis St. John, Pacijic Science, 33: 404, f. 410 (1979). TYPE:

Tahiti, Mahina District, Aorai, 2530 ft alt, vi 1930, M . L. Grant 3734 (BISH). Syn. nov.

NEW COLLECTION: TAHITI: Teravao plateau, above Cattle Experiment Station, route to Vaiufaufa, in badly disturbed forest with Miconia calvescens among native trees, 650 m alt., 28 vii 1981, B. C. Stone, F. R. Fosberg, €9 M . Guerin 14701 (BISH, K, KLU, P, PH, US, etc.).

Tree 5-6 m tall, with 7 or 8 main branches; trunk c. 140 mm diameter, sparsely tuberculate; proproots few, basal, stout. Leaves rather dark green and somewhat glaucous above, paler subglaucous to green beneath, dorsally at base with a large brownish-purplish patch; midrib unarmed near base; it and the margins armed beyond with very small antrorse prickles; blades about 1.4 m long, 60 mm wide.

Description of seedlings (from plants grown from seed in Rimba Ilmu, Botanic Garden of the University of Malaya, derived from Stone et al. 14701) at age 19 months. Stem unbranched, 40 mm tall to base of oldest leaf, 17 mm diameter. Proproots several, up to 25 x 5 mm. Crown of leaves open, the blades

46 B. C. STONE

arched outward, uniformly light or medium green, the undersurface scarcely paler, the base not perceptibly darkened, not glaucous; leaves about 850 mm long, 32 mm wide, or less; margins near base and middle virtually unarmed, only a very few widely scattered and very minute prickles present, these at most 0.2 mm long; leaf apex with similar small regular antrorse prickles c. 0.2 mm long and up to 2.5 mm apart. Midrib near base and along the middle unarmed, rounded or more distally carinate, near the apex with minute prickles similar to those of the adjacent margins; longitudinal veins 36-38, c. 0.9 mm apart, darker green than the intervein areas, these latter dotted with stomata. Leaf apex acute-acuminate, somewhat drawn out but not flagellate, the tip less than 1 mm wide only on the ultimate 20 mm. Apical ventral pleats rounded, unarmed.

Pandanus pabenooensis must be critically compared with P. tamaruensis J. W. Moore and P. ternehaniensis J. W. Moore, both of Raiatea Island, in the Society Islands group. These have moderately high carpel number (to 16 and to 18, respectively) but smaller phalanges. They are interior, upland species.

INTRODUCED SPECIES

If it were feasible, it would be useful to make a discrimination between aboriginal introductions (plants known to have been brought to Tahiti by Polynesians) and European (or European-era) introductions. However, this is a matter requiring more information than is available.

Martelli (1933a) suggested two possible introduced pandans in his account of the Tahitian species. Of these, one (which he identified as P. menziesii) has been re-identified as P. subradiatus St. John (1979b), however, I regard this as merely a form of P. tectorius. Nothing about the collection suggests strongly whether or not this pandan is introduced; it is probably a native, though it may have been in a garden.

The other tentative introduction mentioned by Martelli definitely occurred in a garden (“Yard of Dr. Williams’ home at Mateia, Wilder no. 8”). This was identified as P. odoratissimus var. savaiensis Martelli. St. John does not mention this specimen, but cites another plant, a cultivar Farkoi in his key (it is not further mentioned).

St. John (1979b) adds Pandanus spurius cv. Putat in his key to Tahitian pandans. This is the same as P. tectorius var. laevis Warb. I t is a widespread unarmed cultigen whose origin is unknown; it could have arisen in Polynesia, indeed it is possible that the ‘variety’ is simply a congeries of plants, all of P. tectorius, but representing mutants which have arisen separately and spontaneously in a number of different provenances, have been noticed by local human groups, and have been brought into cultivation as useful and obviously more convenient for such tasks as making thatch, weaving mats and baskets, etc. This cultivar has been found in many localities, and both pistillate and staminate flowering specimens are preserved in various herbaria.

The other cultivar mentioned briefly by St. John is identified as F . odoratissimus var. novi-caledonicus cv. Farkoi without a reference. This was published in PaciJic Science, 29: 403, f. 363 (1975), and is based on a collection from Rotuma Island (Fiji), St. John 29452 (BISH). Although from Rotuma, it is

PANDANUS (PANDANACEAE) IN TAHITI 47

5

Carpels/phalange

Figure 9. Carpel numbers/phalange/cephalium in Pandanus papenooensis (0 ) and P. tectorius (0). Whole-cephalium samples (1470I and I4702 respectively). The 31-carpellate and 34-carpellate phalanges shown for P . papenooensis probably represent ‘doubled’ (fused) pairs of 16, respectively 17- carpellate phalanges.

evidently one of the edible-pulp cultivars typical of the Gilbert Islands and St. John specifically states that the plant had been introduced to Rotuma by a Gilbertese man from his home island. No specimen from Tahiti is cited. Whatever the case, this is not a member of P. odoratissimus. This cultivar, and many others well-developed in the Gilbert and Marshall Islands, is undoubtedly the result of selection from P. tectorius in the most inclusive sense; for convenience the large-fruited, pulpy cultivars are treated as P. Jischerianus Martelli (synonym P. pulposus (Warb.) Martelli). The cultivar Farkoi most emphatically does not belong to var. nouo-caledonicus (not noui-caledonicus) , which is a wild form originally obtained in New Caledonia and now known in several other Pacific Islands, but not known certainly to be involved in the domestication process which led to the appearance of cultivars such as Farkoi.

‘There are also in Tahiti various variegated-leaf mutants of P. tectorius, known in horticulture variously as P. sanderi and P. veitchii, and sometimes with fancy names. These mutant forms (which can back-mutate to all green leaf forms) were almost certainly present in Polynesia before Europeans arrived, and may perhaps have been deliberately planted and transported by Polynesian voyagers. Their reproduction is often, but not exclusively, vegetative. They are popular and are now cultivated in many places, including greenhouses in the temperate zone, hotel lobbies, etc., and outdoors in many tropical lands, in all continents. No reliable history of their origin or dissemination is available. I know of no voucher specimens for their occurrence in Tahiti, but have personal sight-records in, for example, the Botanic Garden at Papeari.

One other plant, mentioned by Martelli, represented by the collection Wilder 7, was left unidentified by Martelli with the remark that “on account of its incompleteness I am unable to establish its specific position”. St. John, however, described it as P. bothreus. The type came from a “Single tree in Mr. Brander’s yard near Papeete. . .” which might suggest i t was in cultivation. Whether it was or not, i t seems plainly to be another carpological form of P. tectorius, and thus surely represents an indigenous plant.

48 B. C. STONE

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MARTELLI, U., 1933b. Pandanaceae of Rarotonga. University o f California Publications in Botany, 17: 171-185. MOORE, J. W., 1933. New and critical plants from Raiatea. Bernice P . Bishop Museum Bulletin, 102: 1-53. PARKINSON, S., 1773. Journal o f a V y a g e to the South Seas in his Majesty’s ship, the Endeavour. London: Stanfield

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