Studies in Fitchia (Compositae): Novelties from the Society Islands; Anatomical Studies

Reprinted from PACIFIC SCIENCE, vol. XVJI, no. 3, July, 1963

SHERWIN CARLQUIST and MARTIN L. GRANT1

THE GENUS Fitchia has attracted interest be TAXONOMIC DESCRIPTIONS cause of its arborescent habit, its distribution in southeastern Polynesia, the endemism of its 1. Fitchia cordata M. L. Grant et S. Cariquist, species, and its phylogenetic relationships. In sp. nov. a monograph of this genus, six species occur Fig. 31 ring on five islands were recognized (Carlquist, 1957). Following the appearance of that mono Frutex arboreus 2 m altus glaber, caule basi graph, Grant notified Cariquist of his collec liter 4 cm crasso. Folia simplicia petiolata petio tion, while on a Bishop Museum Fellowship lis 25—65 mm longis, longitudine media in 1931, of specimens of Fitchia from two ad (N= 42) 47 mm. Laminae late obovatae vel subor ditional islands (Bora Bora and Tahaa) in the biculares luteovirides 40—72 mm longae Society Group. The genus had not been pre 36—65 mm latae, magnitudine media foliarum grandi viously reported from these islands, and manu orum circa 58 >< 54 mm, basi cordata vel script names, as a new species and subspecies, ro tundata, apice obtuso acutove vel parum acu respectively, had been applied to the taxa. On minato, margine crenulato ye! subintegro, Grant’s invitation, Carlquist made the anatomi venu lis lateralibus 5—10 (numero medio 8.5) pro cal studies presented here which, it is felt, puts unico latere. Pedunculi terminales 20—40 mm the recognition of the two new entities on a longi, longitudine media (N=7) mm. Capi much firmer basis than gross morphology alone 33 tula solitaria circa 40-florata. Involucra could have provided. In addition, descriptions 25 mm longa 25 mm lata, bracteis 3—4 seriatis viride are given of one additional specimen each of lutei coriaceis marginibus scariosis, exterioribus two previously known species of Fitchia, one late triangularis 3 mm longis mm latis, from Tahiti, and one from Rapa. The occur 9 medi oribus semiorbicularibus, interioribus lanceo rence of the two novelties is of more than or laris 13—17 mm longis 5—7 mm latis. Bracteae dinary interest because of the nature of this receptaculares 12—17 mm longae 2—3 genus, and permits more extended remarks on (—5) mm latae. Corollae 15—20 mm longae, the nature of speciation in Fitchia. Flowering tubis circa 10 mm longis, sinibus subaequalis. Antherae material of F. rapensis, not available for the saccis 5 mm longis, apicibus staminum monograph, has been studied here. This paper, cuneato-lance olatis 1—1.5 mm longis. Ovaria ad anthesin therefore, may be considered an addendum to 7—8 mm longa, stylis 20—22 mm longis. the monograph. The taxonomic descriptions be Achaenia matura 13—16mm longa, aristis low have been prepared by Grant. The anatomi 8—10mm longis plerumque triangulatis cal account which follows is the in sectione transversa work of Carl subdeciduis. quist, and has been prepared to conform with Tree-like shrub, 2 the types of data offered in the monograph. m tall, 4 cm in diameter at the base, glabrous. Bark 1 mm thick, brown, Claremont Graduate School, Claremont, California; almost smooth, with fine longitudinal ridges and Iowa State Teachers College, Cedar Falls, Iowa. 0.5 mm deep. One-year twigs on flowering Manuscript received December 28, 1961. shoots greenish, 2.5—3 mm in diameter; two-

282 Studies in Fitchia—CARLQUIST and GRANT 283

FIGS. 1—6. Secondary xylem. Figs. 1—4, Fitchia cun eata ssp. tahaaensis. Figs. 5—6, F. cordata. Figs. 1, 2: Transections; note ryloses in vessels; Fig. 2, a scierified rylosis in which the lumen is nearly occluded. Figs. 3, 4: Tangential sections; Fig. 3 is a portion showing long, chick-walled fibers; Fig. 4 shows storied apotracheal parenchyma cells. Fig. 5: Transection; note band of apotracheal patenchyma above center of photograph. Fig. 6: Tangential section. Fig. 2, X 300; others, X 117. year twigs pale brown, 4—5 mm in diameter; broadly ovate to suborbicular, yellowish-green, internodes 4—12 mm long, on vegetative shoots with a narrow (0.3 mm) stramineous margin, to 20 mm long. Leaves simple. Petioles 2 5—65 and with a dark callous at the apex; 40—72 mm mm long, the average being 47 mm (N=42); long, 36—65 mm wide, the average (N=8) the stipular sheaths 0.4—0.6 mm high. Blades size of the largest blade on the flowering shoots 284 PACIFIC SCIENCE, Vol. XVII, July 1963 being 62 X 55 mm, the second largest 58 X 54 Otemanu (Mt. Temanu of the maps), eleva mm, and the smallest of the fully-developed tion 725 m, a peak which apparently has never leaves 64 >< 43 mm; the base cordate to a been climbed by anyone other than Polynesians. depth of 1—2 (—3) mm or rounded; the apex obtuse to acute, with the margins near the tip 2. Fitchia cuneata W. Moore, Bishop Mus. slightly rounded to straight, or, if barely acu J. Bull. 102:48. 1933 ssp. tahaaensis M. L. minate, departing at most 1 mm from a straight Grant et S. Carlquist, subsp. nov. line; the margin crenulate, with 2—3 indenta tions (up to 0.5 mm deep) per cm, to suben Fig. 32 tire, the base entire for 20 mm adjacent to the petiole; with 5—10 (average 8.5) pairs of major Frutex arboreus 2.5 m altus glaber, caule lateral veins, usually curving somewhat towards basaliter 2.5 cm crasso. Folia simplicia petiolata the tip of the leaf. Peduncles terminal, 20—40 petiolis 10—37 mm longis, longitudine media mm long, the average (N=7) 33 mm, often (N30) 20 mm. Laminae plerumque ellipti arcuate and reflexed. Head solitary, with about cae vel parum ovatae obovataeque vel thorn 40 flowers. Involucres 25 mm long and 25 mm boidales subatrovirides 40—110 mm longae 25— wide; the bracts in 3—4 series, greenish-yellow, 58 mm latae, magnitudine media foliarurn coriaceous, scarious-margined, the outermost grandiorum circa acuminato, margine crenu broadly deltoid (3 mm long, 9 mm wide), the lato serratove vel subintegro, venulis lateribus middle ones semicircular, and the inner ones 5—10 (numero medio 7.5) pro unico latere. lanceolate (13—17 mm long, 5—7 mm wide). Pedunculi terminales 2 5—50 mm longi, longi Receptacular bracts 12—17 mm long, 2—3(—5) tudine media (N=10) 33 mm. Capitula soli mm wide. Corollas 15—20 mm long, the tube taria circa 75-florata. Involuc 20—40 mm longa, about 10 mm long, the shallower sinuses nearly 15—25 mm lata, bracteis 3—4 seriatis, viride lutei coriaceis equalling the ventral cleft, and the lobes with marginibus chartaceis, exteriori - terminal hairs 0.1—0.6 mm long. Anther sacs bus breviter ovatis reniformibusque 2—5 mm 5 mm long, the stamen tips cuneate-lanceolate, longis 6—15 mm laris, medioribus semiorbiculari 1—1.5 mm long. Ovary at anthesis 7—8 mm long; bus 8—12 mm longis 9—13 mm lads, interioribus the styles 20—22 mm long. Ripe achenes 13—16 cuneatis 12—20 mm longis, 4—8 mm latis. Brac mm long, with awns 8—10 mm long; awns gen teae receptaculares lanceolatae 12—17 mm longae erally triangular in transection and somewhat 2—4 mm latae. Corollae ad anthesin 15—22 mm deciduous. longae, tubis 4—10 mm longis, dentibus 5—7 mm longis. DISTRIBUTION: Society Islands, endemic to Antherae saccis, 4.5—5 mm longis, apici Bora Bora. bus staminum cuneato-lanceolatis 1.5 mm longis. Ovaria ad anthesin 8—10 mm longa, SPECIMEN EXAMINED: Bora Bora, in dwarf 3—4 mm lara, stylis 24—30 mm longis. Achaenia rain-forest on the summit of Mt. Tarapaia, alti matura 10—20 mm longa, 4—6 mm lata, aristis tude 645 m, Jan. 3, 1931, M. L. Grant 4968 10—13 mm longis plerumque teretibus sub (BISH, type; RSA, ISTC, isotypes). persistentibus. This species was found in several places on Treelike shrub, 2.5 m tall, 2.5 cm in diameter Mt. Tarapaia, none less than 25 m in altitude at the base, glabrous. Bark 1 mm thick, brown, below the summit, but only four inflorescences roughened with irregular scaly ridges. One-year- were discovered, two in bud, one at anthesis, old twigs on flowering shoots greenish, 2 mm and one in fruit. Associated woody plants were in diameter; two-year twigs pale brown, 3 mm Aistonia, Mct’rosideros, Freycinetia, and Glochi in diameter; internodes 4—8 mm long, on vege dio’n. Native name: anei. tative shoots to 12 mm long. Leaves simple. Mt. Tarapaia appears on U. S. Hydrographic Petioles 10—37 mm long, the average (N= 30) maps as Mt. Pahia, elevation 2165 ft (== 660 20 mm; the stipular sheaths 0.5—0.8 mm high. m), the name and elevation being taken from Blades mostly elliptical, varying to slightly French maps. It has been recorded as “Tarao ovate, obovate, or rhomboid, dull green, with pai’a.” The only higher point on Bora Bora is a narrow stramineous margin, and with a dark Studies in Fitchia—CARLQUIsT and GRANT 285 callous at the apex; 40—110 mm long, 25—58 3. Fitchia nutans Hook f., London Jour. Bot. mm wide, the average size (N= 10) of the 4:640, t.23—24. 1845. largest leaf on flowering shoots 84 X 45 mm, the second largest 75 )< 42 mm, and the small Since so few specimens of this species are est of the fully developed leaves 59 )< 30 mm; available in the United States, and the several the base cuneate (usually) to barely rounded; Kew and Paris sheets are without detailed habi the apex acute, occasionally slightly acuminate; tat data, the citation of an additional collection the margin crenulate to toothed or subentire, may be of interest: Tahiti iti, district of Teahu with 2—4 indentations (up to 1 mm deep) per p00, on Mt. Ronui, in Metrosideros- W/einman cm; with 5—10 (average 7.5) pairs of major nia forest, altitude 890 m, July 2, 1930, M. L. Grant 3925 (BISH, ISTC). lateral veins, mostly curving slightly toward the The following de scription is from this collection alone. tip of the leaf. Peduncles terminal, 2 5—50 mm Treeshaped glabrous long, the average (N=10) 39 mm long, al shrub, 2.5 m tall, with a basal diameter of cm; ways reflexed and usually arcuate at anthesis. 5 bark 2 mm thick, brown, with longitudinal Heads solitary, with about 75 flowers. Involucres ridges about 1 mm deep; the wood very sweet-smelling. 15—25 mm long, 20—40 mm wide; the bracts in One-year twigs mm 3—4 series, greenish-yellow, coriaceous, charta 2—3 in diameter; two-year twigs 3.5— 4.5 mm thick; ceous-margined, the outermost short ovate or internodes 7—10 mm long. Peti .reniform (2—5 mm long, 6—15 mm wide), the oles 30—60 mm long; stipular sheaths 2—3 mm middle ones semi-circular (8—12 mm long, 9—13 high. Blades ovate, yellowish-green, 60—130 mm long, mm wide), and the inner cuneate (12—20 mm 45—80 mm wide, the average of 12 leaves long, 4—8 mm wide) Receptacular bracts lanceo being 96 X 61 mm; truncate or cordate at late, 12—17 mm long, 2—4 mm wide. Corollas the base, occasionally barely rounded, often at anthesis 15—22 mm long, the tube 7—10 mm somewhat oblique; the apex slightly acuminate; the margin irregularly long, and the teeth 5—7 mm long, the shorter and shallowly crenulate to entire; with teeth about half the length of the limb, or 8—10 major lateral veins. Pedun des two in each of the four eventually equalling it; lobes with hairs about inflorescences pres ent, long, 0.75—1 mm long. Anther sacs 4.5—5 mm long, 60—65 mm reflexed. Heads shattered with the stamen tips cuneate-lanceolate, 1—1.5 mm age, the involucral bracts and corollas hav ing long. Ovary at anthesis 8—10 mm long, 3—4 mm dropped. Receptacular bracts 13—14 mm long, wide. wide; styles 22—30 mm long, the stigmatic 3—5 mm Ripe achenes 9—10 mm long, branches 0.8 mm long, barely separating at an- 2—2.5 mm wide, with persistent awns 7—8 long. thesis. Mature achenes 10—20 mm long, 4—6 mm mm wide; the awns 10—13 mm long, generally This collection fits the general description of the species rounded in transection, subpersistent. (Carlquist, 1957: 63), except as follows, with Cariquists measurements and DISTRIBUTION: Society Islands, endemic to notes in parentheses: plant smaller (4.5—7.5 m Tahaa. tall), blades often crenulate (entire), maximum SPECIMEN EXAMINED: Tahaa, district of leaf size greater (115 >< 70 mm), base often Ruutia, slopes of Mt. Ohiri, in rain-forest of truncate to cordate (acute to obtuse), heads Crossostylis, Aistonia, and Morinda, altitude paired (solitary), receptacular bracts shorter 465 m, Jan. 25, 1931. M. L. Grant 5161 (BIsH, (20—22 mm long), achenes much shorter (16— type; RSA, ISTC, isotypes). 17 mm long at anthesis) and narrower (3—4 This subspecies was observed in three other mm). These differences, however, do not seem nearby localities, all within an altitudinal range significant enough to warrant separation, and of 15 m, and about 70 m below the top of the material matches Hookers plate closely. Ohiri, the highest point on the island. Other Although the type of the species was sup associated woody plants were Xylosma, Meryta, posedly from Elizabeth Island,” all the evi Metrosideros, Wikstroemia, Pagara, Astronia, dence (Carlquist, 1957: 63) suggests that it is and Hernandia. confined to Tahiti. The present specimen comes FIGS. 7—12. Figs. 7, 8, 11, Fitchia cordata. Figs. 9, 10, 12, F. cuneata ssp. tahaaensis. tion, adaxiat face above. Secretory Fig. 7: Leaf transec canal is above vein at right. Fig. 8: Transection awn adaxial face at right. Note of from an achene, lignification of hypoderrnal layers, presence of a single vascular Leaf transection, adaxial bundle. Fig. 9: face above. In addition to sec retory canals adjacent to vein, left, secretory may be seen, center and right. cavities Fig. 10: Transection of awn from an achene, adaxial face upper right. chomes in section, above and Tn below. Fig. 11: Transection of pith. Secretory canal with adjacent fibrosclereids, above; inner margins of vascular cylinder, below; note thin-walled cells at top of figure. Fig. 12: Longitudinal section of pith. All cells have secondary walls, some such are thicker walled. Figs. 7, 9, X 145. Figs. 8, 10, X 255. Figs. 11, 12, X 135. ______

Studies in Fitchia—CARLQUIsT and GRANT 287 from the smaller mountain (Tahiti-iti) of the because no other collections have been made of two which make up the island. Fitchia in the Fiji group. Also, Hemsley (1885: Hoffmann (1890: 353) reported F. nutans 20) reports that there is at Kew a specimen from Tabuai (= Tubuai), as well as from collected by Moseley in Tahiti at an altitude of Tahiti and Elizabeth (now Henderson) I., but 4,000 ft, and this is probably the specimen as Hemsley (1885: 20) had earlier shown, cited by Erdtman. this supposed locality is based on the type sheet Papy (1955: 325) mentions F. nutans as (Cuming 1424) which Hooker had earlier at growing on “Tahiti, Moorea, Raiatea, Rapa.” tributed to Elizabeth Island. The record from Rapa is probably based on the Erdtman (1952: 124) cites a specimen of report of Riley (1926: 55), which was made F. nutans as “Fiji, Moseley, anno 1875.” Carl before F. ra1bensis had been described as a quist (1957:63) suggested that this is in error, separate species (Brown, 1935: 366), although

13 I

FIGS. 13—16. Fig. 13: Fachia cordata. Transection of involucral bract taken about midway along length of bract; adaxial face at left. Fig. 14: F. cordata. Transection receptacular bract, adaxial face at left. In Figs. 13, 14, tracheary elements shown in bold outline, fibers and sciereids stippled. Fig. 15: “Typical” leaf of F. cordata, showing main veins. Fig. 16: “Typical” leaf of F. cu’neata ssp. tahaaensis. Figs. 13, 14, X 105. Figs. 15, 16, X 3. 288 PACIFiC SCIENCE, Vol. XVII, july 1963

Papy was aware of this. Neither Moore nor ANATOMICAL DESCRIPTIONS Grant found F. nutans on Raiatea, and Papy was aware of the description of F. cuneata from Anatomical studies were based on the her that island. Moorea is certainly a likely locality barium specimens cited above. Liquid-preserved for F. nutans, but for none of these localities heads of F. cuneata ssp. tahaaensis and wood does Papy cite specimens. samples of both F. cuneata ssp. tahaaenris and Thus, the geographic distribution offered by F. cordata, for which the herbarium specimens Carlquist (1957:2) must be broadened to in serve as vouchers, were available. The methods clude the occurrence of F. nutans on Tahiti-iti, of preparation of these materials are the same F. cordata on Bora Bora, and F. cu’neata ssp. as those employed for such materials in the tahaaensis on Tahaa. two papers (Cariquist, 1957, 1958) dealing with the anatomy of this genus. 4. Fitchia rapensis F. Brown, Bishop Mus. Bull. SECONDARY XYLEM: With respect to quali 130:366. 1935. tative features, the species of Fitchia other than F. speciosa are much alike in wood anatomy Since the ten previously recorded collections (Carlquist, 1958). Fitchia cuneata ssp. tahaaen species of this are all without flowers (Carl sis and P. cordata share the qualitative features quist, 1957: 62), the description of a flowering of those species. The bands of apotracheal specimen should be of interest: Rapa, moist parenchyma are clearly shown by the two new zone, altitude 520 m, Jan. 3, 1922, W. B. Jones taxa (Figs. 4, 5). Thick-walled fibers are readily 372 (Whitney Expedition) (BKL). apparent in F. cuneata ssp. tahaaensis (Figs. 1, “Tree,” glabrous. Petioles 40—7 5 mm long. 3); libriform fibers are less thick-walled in Blades ovate, the larger ones averaging 90 mm F. cordata (Figs. 5, 6). Scletosed tyloses have long and 84 mm wide, rounded or scarcely been reported in F. nutans and F. speciosa cuneate at the base, the apex acute; entire; with (Carlquist, 1957, 1958); they are abundant in about 12 pairs of major lateral veins. The sin F. cuneata ssp. tahaaensis (Figs. 1, 2), although gle inflorescence is terminal, with two heads, none were observed in F. cordata. the peduncles 50 and 60 mm long, reflexed. Quantitative features of the two new taxa Heads 45 mm long and 45 mm wide. Bracts in are as follows: F. cordata: vessels per group: abotrt. 4 series, those of the second layer semi 1.37; average vessel diameter: 61.5 ; diameter circular, 10 mm long and 22 mm wide, and of widest vessel: 88 c; average length vessel those of the inner layer 20 mm long and 12 elements: 325 s; average length libriform fibers: mm wide. Receptacular bracts cuneate, 16—17 502 ; average length apotracheal parenchyma mm long, 3—4 mm wide. Mature corollas 24—26 cells: 356 r; average height multiseriate rays: mm long, the ventral cleft reaching to 6—8 mm 1.17 mm. Fitchia cuneata ssp. tahaaensis: ves of the base, the segments eventually separating sels per group: 1.66; average vessel diameter: to within 10 mm of the base; the tips of the 62.8 c; diameter of widest vessel: 92 r; average lobes often densely covered with sclerified cells, length vessel elements: 326 r; average length which are up to 0.75 mm long. Anther sacs libriform fibers: 507 c; average length apo 5.5—6 mm long, caudate, the tail 0.3—0.4 mm tracheal parenchyma cells: 380 s; average long; the stamen tips cuneate, 1.5 mm long. height multiseriate rays: 1.51 mm. Mature style (broken off) at least 32 mm Quantitative as well as qualitative data point long. Achenes, at and just after anthesis, 10—12 up a close similarity between F. cuneata ssp. mm long, 4 mm wide, the awns 12—15 mm tahaaensis and the typical F. cuneata. Quanti long, strongly bent and twisted at the base. tative and qualitative data for F. cordata re

FIGS. 17—19. Fitchia cuneata ssp. tahaaensis. Fig. 17: Transection of involucral bract taken about midway along length; adaxial face below. Fig. 18: Transection of recepracular bract; adaxial face and adjacent left. Fig. achene at 19: Transection of style. At center of style, stigmatoid tissue. Exterior to each bundle is a canal, secretory except for bundle at upper left, which is flanked by a pair of canals. At right, anthers in transection. Note secretory canal in each of the two anthers. Fig. 17, X 100; Fig. 18, X 80; Fig. 19, X 120.

290 PACIFIC SCIENCE, Vol. XVII, July 1963

semble the figures given (Carlquist, 1958: 6-7) but they are lacking in sheaths of finer veinlets. for F. czmnea’a and F. rnaans. No secretory cavities are present in the meso PITH: Pith anatomy proved important in dis phyll, although secretory canals occur adaxially tinguishing s p e c i e s of Fi.tchia (Cariquist, or abaxially to the veins, or both (Fig. 7). 1957). Likewise, the two new taxa possess Mesophyll is about 10 layers thick. These char characteristics useful for taxonomic purposes. acteristics are cot matched by any other Fi.tchia Pith of F. co:data (Fig. 11) consists of both collection. Presence of a few fibers would ally thick-walled, lignified, and thin-walled, non F. cordata to F. ?zutans or F. tahitensis, but lignified, cells. Secretory canals are present; each these two species possess secretory cavities. Ab is surrounded by an eccentric zone of fibro sence of secretory cavities does characterize F. sciereids. Although secretory canals surrounded speciosa and the typical F. cuneata, but these by small lignifled thick-walled cells occur in F. taxa lack sclerenchyma in bundle sheaths. iraans and F. tahitensis, the presence of these Leaves of F. cuneata ssp. tahaaensis (Fig. 9) combined with occurrence of thin-walled non lack sclerenchyma along all but the largest lignifled cells in pith is characteristic only of veins. In addition to the secretory canals above F. corda;a. Pith of F. cuneaa ssp. tahaaensis and below veins (Fig. 9, left), secretory cavi (Fig. 12) lacks secretory canals and consists ties are present in the mesophyll (Fig. 9, right). of thin-walled and thick-walled lignified cells. The mesophyll is about 10 cells thick. This The only pith pattern which matches this is description agrees closely with that of typical that described in the monograph for typical F. F. cuneata except for the presence of secretory cuneata. cavities. Although such a dierence might con LEAF: Leaf dimensions are mentioned in the ceivably arise from a difference in maturity of taxonomic description above. However, aver a plant, it seems more likely a valid difference, ages of dimensions of leaves from a collection for specimens of both subspecies were of flow were used in the monograph, and these may be ering age. compared with those of the new taxa. Mature INVOLUCRE: The heads of F. cordata are leaves of the collection of F. ccrdata (Fig. 15) small, and the involucre of a living specimen show an average lamina width of 52.5 mm, an would probably measure about 2 cm in diam average lamina length of 53 mm, and a petiole eter—the smallest involucre size in the genus length of 49.7 mm. The only Fitchia which re except for F, tahüensis and F. cuneai’a, Involu sembles F. cordata both in form and dimensions cral bracts of F. cordata (Fig. 13) have a thick is the extreme population of F. rapensis from ness comparable to some bracts of F. nutans, but the summit of Rapa. Leaves of F. cordata have differ in their lack of the abundant scleren longer petioles than that collection, and there chyma which characterizes bracts of F. nu/ans. is certainly no close relationship between F. The mature state of the bracts of F. corja:a cordata and F. rapensis. Leaves of F. cuneata illustrated is certain, because they came from ssp. tahaaensis (Fig. 16) which could be termed a head in fruit which was in the process of mature show the following average dimensions: shattering. Occasional lignifled thick-walled lamina width, 48.0 mm; lamina width, 87.5 cells may be seen along the inner face of the mm; petiole length, 26 mm. When compared bract and around the larger veins. In presence with the chart in the monograph (Carlquist, of sclerenchyma and other histological charac 1957: 50) the lamina dimensions fall in the teristics, the involucral bracts of F. cordata are vicinity of F. nutans and some collections of midway between those of F. nutans and F. cu F. rap ensis. The leaves are wider than those of neata. The receptacular bracts of F. cordata F. tahitensis but longer than those of the typical (Fig. 14) show similar characteristics. As might F. cuneata. Thus, leaf shape distinguishes both be expected, lignifled thick-walled cells are more F. cordata and F. cuneata ssp. tahaaensis. frequent than in the involucral bracts. Histological features of leaves of the new Fitchia cuneata ssp. tahaaensis also has very taxa also distinguish them. Fitchia cordata pos narrow involucres, like those of the typical F. sesses fibers in bundle sheaths of many veins, cuneata or F. tahitensis. Histologically, involu Studies in Fitchia—CARLQu1ST and GRANT 291

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FIGS. 20—24. Figs. 20, 23, Fitchia cordata. Figs. 21, 22, 24, F. cuneata ssp. tahaaensis. Figs. 20, 21: Tips of corolla lobes, showing exterior surface. Scierenchymatous 14 cells are shown in black. Figs. 22, 23: Stamen tips, showing inner surface. Anther sacs shown in outline, below. Fig. 24: Base of anther and adjacent filament. Note caudate anther sac, right. All, X 65. 292 PACIFIC SCIENCE, Vol. XVII, July 1963 cral bracts of F. cuneata ssp. .tahaaensis (Fig. ravages of the living plant by insects), the vena 17) resemble closely the description given for tion pattern of this species is substantially like typical F. cuneata in the monograph. This re that figured for F. cuneata or F. tahitensis. There semblance is especially evident in the lack of is some variation in the levels at which adja scierenchyma. Only an occasional lignifled cell cent lateral veins in each lobe join beneath (usually near a vein) can be found. Collenchy sinuses, but differences in these levels are not matic thickening may occur near outer or inner nearly so extreme as the condition figured for faces of the bract. In the section shown, more F. nutans (Carlquist, 1957: 10). In their rela than a single series of bundles may be seen; tively short length (ca. 18 mm) the corollas some of these bundles have distorted orienta of F. cordata approximate those of F. tahitensis tion of xylem and phloem, a condition men also. tioned in the monograph for F. .tpeciosa. This Corollas of F. cuneata ssp. tahaaensis have condition is probably occasional among larger similar size and venation features. Typically, bracts of several species. The thickness of the the patterns are the same as mentioned for F. involucral bract, as well as its lack of scieren cordata. However, occasionally a pair of adjacent chyma, identifies the two subspecies of F. cu laterals do not join beneath a sinus, but con neata closely with each other. The receptacular tinue into the tube and achene without joining. bract (Fig. 18) of a head anthesis at certainly This condition does occur occasionally in the lacks sclereids, although a greater degree of other species, such as F. speciosa. A feature of lignificarion might apparent be in bracts of a interest in the corolla of F. cuneata ssp. taha head in fruit. aensis is the presence of one or two supernu ACFIENES: Awn shape and anatomy proved merary veins in each lobe. In addition to the useful characteristics in distinguishing species median and lateral veins, these fine supernu of Fitchia. In comparing illustrations of these merary veins may be present for a short distance (Carlquist, 1957: plate 8) with these of the near the base of the lobe. Such veins were not new taxa (Figs. 8, 10) this also proves to be seen in the typical F. cuizeata, for which, how true. In F. cordata (Fig. 8), the awns tend to ever, mature corollas were not available. Super be rounded in transectional shape, like those of numerary veins are occasional in corollas of F. F. nutans. However, they show a tendency to speciosa, and characteristically abundant in those be wider than those of F. nutans. They have a of F. mangarevensis. single vascular bundle and, unlike at least some The occurrence of larger numbers of veins awns of F. nutans, they lack a secretory canal. in corollas of F. mangarevensis is particularly Awns of F. cuneata ssp. tahaaensis (Fig. 10) are interesting, since that species is close to F. like those figured for the typical F. cuneata in rapensis. Corollas, now available, of F. rapensis all respects. Presence of three longitudinal zones confirm this relationship in the similarly elabo of trichomes tends make to them triangular as rate venation pattern (Fig. 26). This similarity seen in transectional view. Where such tn is shown in the presence of five veins per lobe, chomes are absent, awns may have a more rather than three. Such a condition is basic in rounded shape. the construction of the F. rapensis corolla, al COROLLA: In the monograph, considerable though additional veins or vein fragments may attention was focused on the importance of be present. The outermost veins of each lobe venation patterns of Fitchia corollas, both for fuse at the tips of many corolla lobes. The co their phylogenetic importance within Composi rolla venation of F. mangarevensis (Carlquist, tae and for their usefulness in distinguishing 1957:14) is somewhat more complicated than species. Corolla venation patterns are not illus this, because although the five-vein condition trated here for the new taxa because they con may be observed in some lobes in that species, tribute relatively few novel features and can additional veins are more frequent. The pres be referred to patterns illustrated in the mono ence of complex corolla venation in both F. graph. Although few flowers of F. cordata could rapensis and F. mangarevensis is interesting in be studied in this respect (on account of the that it raises the question of how complex Studies in Fitchia—CARLQUIST and GRANT 293 venation may have been present ancestrally in tips than the other species of the genus, with the Fitchia. Is the more complex condition primitive exception of F. .cpeciosa. or a specialization? The question cannot be A feature of some interest not previously answered in terms of the data available, and observed is the occurrence of somewhat caudate cytological information would be very desirable anther sacs in F. cuneata ssp. tahaaensis (Fig. as a line of supporting evidence. 24) and the other taxa of Fitchia. In view of Distribution of thin-walled and thick-walled Cronquist’s (1955) dictum that anthers of Coin trichomes and occurrence of sclereids on corolla- positae are primitively tailless, this would seem lobe tips have proved useful features in the curious, because Fitchia possesses so many ana systematics of Fitchia. In the two new taxa these tomical and morphological characteristics which features are also of interest. Fitchia cordata (Fig. appear primitive for the family. Cronquist’s 21) possesses lobe tips not identical to those of dictum, for which no appreciable evidence is any species previously figured. The frequency adduced, seems highly questionable. of sciereids at the lobe tip is notable. Long In transection the anthers of F. cuneata ssp. multiseriate trichomes, all the cells of which are tahaaensis (Fig. 19) exhibit a feature of interest. sclereids, are present. No trichomes composed of One or two secretory canals may be observed in nonlignified cells were observed. This condition the connective of each anther. This condition is most closely matched in the genus only in occurs sparingly in F. speciosa. but was not the new subspecies of F. cuneata (Fig. 20). In observed in other species of Fitchia. The oc F. cuneata ssp. taha4ensis sclereids are rare or currence of anther secretory canals has been absent at the lobe tips. The trichomes are long, noted in other genera which may be related to multiseriate (rarely uniseriate), and composed Fitchia, such as Petrobium and Oparanthus wholly of sclereids. This condition is not unlike (Carlquist, 1957). the condition in typical F. cuneata, except that STYLn: Like the anthers, the style of F. cuneata the trichomes in the Tahaa plants are few and ssp..tahaaensis (Fig. 19) seems to exhibit more long. numerous secretory canals than those of other The corolla-lobe tips of F. raensis (Fig. species of Fitchia. One canal (or occasionally a 25) are, as might be expected, rather similar pair) is present exterior to each of the four to those of F. manga’revensis (Carlquist, 1957: style bundles. Liquid-preserved material is re 27). They are different in that in F. iajbensis the quired for accurate demonstration of this phe sclerified trichomes are slightly longer, in gen nomenon. In the only other species for which eral, and extend farther down the lobe. Oc such material was available, F. speciosa, canals casional thin-walled hairs may be seen on the were much less abundant in styles (Carlquist, terminal portion of lobes of F. rapensis, but 1957). However, as indicated in the monograph, the extremely dense coating of thick-walled hg many other Heliantheae do show abundance of nified trichomes clearly marks this pattern as stylar secretory canals. closest to that of F. mangarevensis. POLLEN GRAINS: The drawings of pollen STAMENS: Stamen tips likewise offer taxo grains of Fitchia (Figs. 28—30) show some dif nomic criteria. The long stamen tips of F. co, ferences in representation compared with those data (Fig. 23) exceed those of F. nutans in size, of the monograph. Although the outer sculp and are thus the longest of the Society Islands tured layer (ectosexine) is composed of fine species of Fitchia. The stamen tips of F. cuneata rods and spaces, as illustrated earlier, a lacunose ssp. tahaaensis (Fig. 22) are longer than those inner sculptured layer (endosexine) has now of the typical F. cuneata, but do not differ mark been observed. The endosexine consists of rods edly from those of F. tahitensis or F. nutans. interspersed in large spaces. In addition, an The stamen tips of F. rapensis (Fig. 27) are inner and outer layer of nexine (below in each 1.5 mm long, thus matching those of F. man figure) may be distinguished. Size and spine garevensis. This is particularly interesting be shape are of especial interest, however, in com cause the two species are so much alike in this parison of the species. The markedly blunt respect and because they have longer stamen spines of F. cuneata ssp. tahaaensis (Fig. 29) 294 PACiFIC SCiENCE, Vol. XVII, July 1963

j25 27 FIGS. 25—27. Fitchia rapensis. Fig. 25: Tip of corolla lobe, showing exterior in black. surface; scierenchymarous cells Fig. 26: Corolla, showing venation. Fig. 27: Stamen tip, showing inner surface. Fig. 26, >< 8. Figs. 25, 27, X 65 Studies in Fitchia—CARLQuIsT and GRANT 295 match those of the typical F. cuneata (Cariquist, mangarevensis, although the difference is not 1957:32). In F. cordata (Fig. 28) the spines significant. The spines of pollen grains of F. are even more markedly blunt. Pollen grains rapensis have an inverted-funnelform shape, like of F. cordata (range, 39—5 1 ; average, 42 r) those of F. mangarevensis, but appear to be are the smallest in the genus. The pollen grains more sharply pointed. In addition, lacunae in of F. cuneata spp. tahaaensis (range, 45—5 1 ; spine tips were observed for the first time in average, 48 r) closely match those of ssp. cu the genus in F. rapensis. This feature, however, neata in size. The slightly smaller size of grains is not unexpected in Fitchia, since other Helian in the latter may be caused by the fact that theae have lacunae in spine tips (Carlquist, flowers of F. cuneata ssp. cuneata from which 1957:33). grains were taken were not quite mature. Pollen SPECIES CHARACTERISTICS: In the mono grains of F. rapensis (Fig. 30) average 60 graph three species groups were recognized: in diameter, and have a range from 51 to 66 . (1) F. speciosa; (2) F. nutans, F. tahitensis, They are thus slightly larger than those of F. and F. cuneata; and (3) F. mangarevensis and

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FIGs. 28—30. Optical sections of equator of pollen grains, showing a third of each grain. Germ pores indi cated at left and right in each. Fig. 28, F. cordata. Fig. 29, F. cuneata ssp. tahaaensis. Fig. 30, F. rapensis. 296 PACIFIC SCIENCE, Vol. XVII, July 1963

FIGs. 31—32. Type specimens of Fitchia. Fig. 31, F.cordata. Fig. 32, F. cuneata ssp. tahaaensis, Studies in Fitchia—CARLQUIST and GRANT 297

F. rapensis. The new taxa clearly belong to the PHYLOGENETIC CONSIDERATIONS: The new second group. The present study also emphasizes taxa described here, although they contribute the distinctness of the third group and the close no anomalous features to the genus, do enlarge relationship between the two species it contains. our picture of it and permit more detailed ob Fitchia caneata ssp. tahaaensis agrees with servations in this regard. The monograph of the typical F. cuneata in most respects. Those by genus provided some notions on the relation which it differs include: presence of scierified ships of Fitchia. These ideas do not seem to tyloses in wood (not yet observed in wood of require alteration. The new taxa do enhance the typical F. caneata); leaves with wider and longer diversity of the Society Islands Fitchias. Indeed, laminae; secretory cavities present in leaf meso Papy (1955) notes that the Society Islands form phyll; involucral bracts thicker; scierified tn a center of diversity for Fitchia, and that the chomes on corolla-lobe tips longer and fewer; variability of Fitchia on Tahiti is suggestive of anther tips longer. These differences are rela this. The latter phenomenon remains a problem tively minor. Close identity of the two sub worthy of study when more material from Tahiti species is revealed by such critical features as becomes available. general leaf shape, pith structure, bract anatomy, Postulation of phylogenetic trends within corolla vascularization, awn shape and anatomy, each species group seems inadvisable, both and pollen size and ornamentation. The two because of the limited quantity of material taxa are therefore best regarded as subspecifically known and the fact that the species seem to different. This situation is precisely what one be variations on a basic theme rather than stages would expect on account of the closeness of along a phylogenetic pathway. However, some Tahaa to Raiatea, sister-islands (within a com interpretations of relationship among the species mon fringing reef). groups may be offered here. The nearest island to the sister-islands of The larger size—in all parts of the plant— Raiatea is Bora Bora. One might expect, there of F. speciosa, as compared with the remaining fore, a similarity between F. cuneata and F. species, seems specialized. The larger pollen size cordata. These two do appear similar in their suggests the possibility of a greater chromosome small heads, relative lack of sclerenchyma in number, and the morphology of the awn base involucral bracts, venation and size of corollas, seems clearly a specialized feature. The fact that lack of secretory cavities in leaves (excepting F. speciosa has such an isolated geographical ssp. tahaaensis) and blunt shape of spines on position, the westernmost species of Fitchia, in pollen grains. They are dissimilar in that F. contrast with the likelihood of an American cordata possesses secretory canals and thin-walled ancestry of this helianthoid (e.g., Brown, 1935; nonlignified cells in the pith, has cordate leaves, Papy, 1955), reinforces this supposition. The and some sclerenchyma along veins of leaves. extremely large seeds of F. speciosa seem less These characteristics are reminiscent of F. nu adapted to long-distance dispersal than the tans. From F. nutans, however, F. cordata differs smaller seeds of the Society Islands species. The by such respects as those in which it resembles most likely interpretation, seemingly, is that F. cuneata. Features of F, cordata which do not seed size, and the peculiar adnation of achene match those of any species in the F. nutans—F. summit to awn base in F. speciosa are charac cuneata—F. tahitensis group include the presence teristics acquired subsequent to arrival of its of nonlignified cells in pith, the small leaf size ancestors in Rarotonga. Fitchia speciosa may be and cordate shape combined with relatively long regarded as a highly distinct derivative of the petiole length, the presence of both long mul Society Islands stock. Although corollas of F. tiseriate sclerified tnichomes and sclerified epi speciosa are the largest in the genus, their vena dermal cells on corolla-lobe tips, long stamen tion is like that of the Society Islands species, tips, small pollen size, and blunt spines on not that of F. rapensis and F. mangarevensis, pollen grains. These characteristics, combined which have complex corolla venation. with the distinction furnished by its geographi If pollen-grain size is an indication of chro cal isolation on Bora Bora, mark F. cordata as mosome number, then one would expect the worthy of recognition as a new species. Society Islands species to have the lowest num 298 PACIFIC SCIENCE, Vol. XVII, July 1963 ber, and progressively higher chromosome num CRONQUIST, A. 1955. Phylogeny and taxonomy bers would be indicated for the rapensis—F. F. of the Compositae. Amer. MidI. Nat. 53:478— mangarevensis group and F. speciosa. If this 511. were true, and it were a criterion of phylogeny within the genus, the complex floral venation ERDTMAN, G. 1952. Pollen morphology and of F. rapensis and F. mangarevensis would be plant taxonomy. Chronica Botanica, Waltham, a specialization. Information on comparative Mass. cytology within the genus, in any case, is highly desirable. HEMsLEY, W. B. 1885. Report on present state of knowledge of various insular floras. Bot., Challenger 1(1):1—75. REFERENCES HOFFMANN, 0. 1890. Compositae. In: Engler and Prantl, Die natürlichen Pflanzenfamilien BROWN, F. B. H. 1935. Flora of southeastern 4(5)87—391. Polynesia, III. Dicotyledons. Bishop Mus. Bull. 130: 1—386. PAPY, H. R. 1955. Tahiti et les lies voisines. La végétation des lies de la Société et de Makatéa. CARLQuIsT, S. 1957. The genus Fitchia (Com 2 partie. Tray. Lab. Forest. Toulouse 5(2/1: positae). Univ. Calif. Pubi. Bot. 29:1—144. 3): 162—386.

— 1958. Wood anatomy of Heliantheae RILEY, L. A. M. 1926. Notes on the flora (Compositae). of Trop. Woods 108:1—30. Rapa. Kew Bull. 1926:51—56.