BLUMEA 23 (197?) 289—300

Notes on ()

R.W.J.M. van der Ham

Rijksherbarium, Leiden, The Netherlands

Summary

revision been made Consequently upon a of Mischocarpus (Van der Ham, 1977a), a detailed study has of several in in of the characters Arytera order to get a better understandingof the delimitation two genera mutually.

The differences between and Radlkofer his and Mischocarpus Arytera given by (1931) in key are vague can only be used when fruiting material is available. However, several species in Arytera were described by him and others material of could be connected with on flowering only. Two species Mischocarpus species

One M. of Arytera. species of Arytera appeared to be the same as Mischocarpus exangulatus. exangulatus has, besides typical Mischocarpus characters, several features uniquein Mischocarpus but more regularly occurring in Arytera.

but Radlkofer distinguished 4 sections with 24 species. A few more species were described afterwards were not placed in a section. When searching for new characters on which to base a better delimitation against Mischocarpus, variation in several characters of Arytera turned out to be rather wide for a within the Cupanieae.Moreover, these variations appeared to be discontinuous. With help ofa few characters, of could be formed natural and better based than the made groups species which are more sections by

Radlkofer. Several of turned be least dubious in species or groups species out to wrongly placed or at the naturalness of here be Arytera. Even Arytera as provisionally accepted can questioned.

material L M all The studied mainly comes fom and (types of Radlkofer). Material of nearly species was

available.

I. Species excluded from Arytera

concolor Arb. — concolor Arytera (Gillespie) A. C. Smith, J. Am. 31 (1950) 298. Guioa

Gillespie, Bull. Bish. Mus. 83 (1931) 17, f. 19. — Type: Islands, Taveuni, near

Gillespie along streams in the coconut belt, alt. 200 Waiyevo, 4794 , 3-3-1928, m,

fl. (BISH, n.v.; iso in A, DS, GH, K, US).

Mischocarpus guillauminii Kanehira, Bot. Mag. Tokyo 46 (1932) 672. — Type: Caro-

in lines. Truk Islands, Tol, Kanehira 1368 alt. c. m, fr. (FU, n.v.; iso , -6-1931, 450 with Kanehira A, US 1268 on label).

Mischocarpus paradoxus auct. non Radlk.: Kanehira & Hatusima, Bot. Mag. Tokyo 57

(i943) 79-

CAROLINES. Truk Islands, Kanehira 1368; Hosokawa 8323 (US); Wong *77 (A).

FIJI ISLANDS. V i t i L e v U. Gillespie 3380 (A, BISH, DS, GH, K, US); Smith 4490 (A, L, US), 4837

L, L, — TAVEUNI. (A, US), 5033, 3287, 3580 (A, US); Degener 13398 (A, US), 13308 (A). Gillespie

4763, 4765, 4793, 4794, 4793 (A, BISH, DS, GH, K, US).

Arytera concolor from Fiji appeared to be almost identical with Mischocarpus guillauminii from the Carolines. Collections from the Carolines have fruits with a glabrous endocarp while those from Fiji have fruits which are densely hairy inside. When revising Mischo- the in carpus, M. guillauminii had to be excluded from that genus. In view of variability BLUMEA VOL. No. 290 23, 2, 1977

indumentum of the endocarp in various species of Mischocarpus, the collections from and from Fiji the Carolines may well belong to the same species. This unknown has been three until but species, to Radlkofer, placed in genera now, the of neither of these is satisfactory. It was described by Gillespie in Guioa on strength a

doubtful observation of two crested scales on each petal. Neither in his figure nor in the

material there is a trace of a crest on the scales, however. Guillaumin considered the

material from the Carolines to belong in Mischocarpus, probably because ofthe superficial

resemblance of the fruit and of the reticulationof the leaflets. Also the reasons to place it the he in Arytera are doubtful. Smith mentionedthe presence of hairs on endocarp which

said to be characteristic of Arytera brackenridgei (A. Gray) Radlk. However, collections

from the Carolines have a glabrous endocarp and such hairs as mentioned do occur in

other and Also the minute scales many genera (e.g. Mischocarpus Gongrodiscus). glandular in the said be feature of A. be found in present very young parts, to a brackenridgei, can other genera ( Cupaniopsis, Sisyrolepis, Dictyoneura, Lepiderema). Other characters present

in the not mentioned unlike those of Arytera: material, by Smith, are very

membranaceous at the mar- Calyx: opening late, sepals not connate, unequal, imbricate, this gins; is a typical 'Cupania-calyx' (Radlkofer, 1879, 1931), noticed by Gillespie but about not by Smith. Arytera is characterized by a 'Matayba-calyx': opening early, sepals

equal, not imbricate, small, not membranaceous at the margins. Fruit: neither Gillespie, nor Smith, nor Kanehira observed the presence of incomplete

i-locular fruit. This character is further met in Gon- septa leading to a 3-merous, only

grodiscus as far as I know. The seed is completely enveloped by an arilloid attached with arilloid a broad ring (pseudohilum) around hilum and micropyle. The in Arytera is either completely different, or attached with a narrow ring. in When On account of these characters, the species is not correctly placed Arytera. we follow Radlkofer's key (1931): fruit opening with three valves, embryo notorrhizal,

Pacific with scales without we distribution, petals two crest, 'Cupania-calyx', come to the Cupaniopsis. All ofthe characters mentioned above, except for the reduced septa, fit in

description of that genus. Also one of the sections ofCupaniopsis contains species provided scales. make combinationhere: with glandular With good reasons we can a new Cupa-

niopsis concolor (Gillespie) Van der Ham.

N The has B Muller o t e s. species pollen type (see: & Leenhouts, 1976). worth When revising Cupaniopsis it is remembering Gongrodiscus as a genus much

alike C. concolor.

sordida Arytera Radlk., Bot. Jahrb. 56 (1920) 301; Hartley c.s., Lloydia 36 (1973)

269. — Type: East , East Sepik Dist., Felsspitze, Kaiserin Augusta- Ledermann alt. fl. Fluss-Exped., 12492, —8—1913, 1400—1500 m, (B, lost; M, iso).

NEW GUINEA. alt. East. Morobe Dist., Edie Creek near Wau, Hartley 11676, 23-4-1963, c. 1950 m,

fl. Central fl. (L); Mt. Shungol, near Wagau, Hartley 12366, 17-12-1963, alt. c. 2100M, (L); Dist., 9.9 S,

147-43 E, NGF 34723, fr. (L).

fruits said be This species was described on flowering material only, while are to characteristic described it with minute scales for Arytera. Radlkofer as having petals two he did the scale but probably not see whole variation: from one big scale via a retuse to

the reduced two smaller, almost separated auricles. Possibly Radlkofer observed more

of scale. When a the condition or maybe he saw the remnants one big pulling out petal, of scale scale is very easily detached because its dense indumentum which makes the

the disk and Such is rather cling to densely hairy stamens. a condition of the scale strange R. W.J. M. van der Ham: Notes on Arytera (Sapindaceae) 291

for Arytera. When fruits became available to me, the species could be placed with cer- in Characteristic also the tainty Sarcopteryx. are very small club-shaped glands on the leaflets of the material. combination made: sordida in part A new is Sarcopteryx

(Radlk.) Van der Ham.

The has Notes. species pollen type B (see: Muller & Leenhouts, 1976). when be reduced In due time, Sarcopteryx is revised, this species may possibly to some species already known.

C. T. - Arytera subnitida White, Proc. R. Soc. Queensl. 47 (1936) 56 Mischocarpus

exangulatus (F. v. M.) Radlk., see Blumea 23 (1977) 266.

II. DUBIOUS SPECIES PROVISIONALLY KEPT IN ARYTERA

lautereriana Arytera (F. M. Bailey) Radlk., Fedde Rep. 20 (1924) 37, from Australia, is in section the only species Radlkofer's . It is a very strange species in fruit Arytera, having characters even unique in the Cupanieae. removed In Mischocarpus macrobotrys Merr. & Perry, a species which had to be from

Mischocarpus and which appeared to be closely related to A. lautereriana, these characters

turned out to be also and still known from present more extreme. M. macrobotrys is

several collections made in E. New Guinea (Brass 7464, 7618, 8057, and Hartley 12077).

The fruit is irregularly lobed and one or two out of three locules are abortive. Ripe fruits be are irregularly dehiscent with hardly a trace of valves; in young ones vague sutures can noticed. This character is not in accordance with the description of the fruit of Cupanieae which is loculicidally dehiscent by valves. The sclerenchymatic endocarp in a locule

radiates from the funicle and is in two the between them separated halves, gap more less the or indicating place of a suture in the pericarp. The sclerenchymatic endocarp is

only lining the proximal hemisphere of the locule and in ripe fruits (i.s.) it is completely detached from the seed and the outer fruitwall, forming a cupule-like structure around the arilloid, attached only around the funicule. The situationin fresh material is not known. The seed cover is said to be a translucent acid pulp. In the herbarium it is rather variable: the whole a subcoriaceous to somewhat fleshy to spongy brittle mass surrounding seed;

this the of probably, depends on the stage of maturity of fruit and the way drying. the On closer examination seed cover appears to be an arilloid attached around the hilum,

the leaving micropyle free (see: Van der Pijl, 1955, f. ic).

In lautereriana the characters A. mentionedare somewhat different but essentially the same.

there is a of at least two Concluding: relationship consisting (maybe_ _ Hartley x species v , Australia and demon- 12077represents a third one), distributed in E. New Guinea, which characters strates that are rather unique in Cupanieae. Till further knowledge is gathered, maintained dubious this relationship is as a group within Arytera: the mischarytera- group.

Consequently, one new combination must be made: Arytera macrobotrys (Merr. &

Perry) Van der Ham (Basionym: Mischocarpus macrobotrys Merr. & Perry, J. Arn. Arb. 21

(1940) 524. — Type: E. New Guinea, Western Dist., Middle Fly R., Lake Daviumbu,

Brass 7618, -8-1936, fl. (A; iso in BRI, L)). The of all collections of and rather with the Note. pollen are type B uniform, seen lightmicroscope.

from Arytera leichhardtii (Benth.) Radlk., Sapind. Holl.-Ind. (1879) 44, Australia, The of . position this species is doubtful: fruits are not known and with 292 BLUMEA VOL. 23, No. 2, 1977

connections in vegetative parts or flowers only no can be laid to other species Arytera.

Until complete material is available, this species is kept as dubious in Arytera. The pollen

but transitional to Pi. type is B, sometimes pollen type A occurs (see 2: 3—6).

III. UNKNOWN SPECIES

material available of No was two species: foveolata related Arytera F. v. M., probably to A. xerocarpa (Bl.) Adelb. Radlkofer related divaricata Arytera exostemonea Domin, according to to A. F. v. M.

IV. SUBDIVISION OF ARYTERA

be into With help of a few characters, the remaining species of Arytera can arranged

that to be more natural and better characterized than the sections made by groups seem

Radlkofer. The characters studied are:

leaflet: scales domatia glandular present or not; present or not.

pollen type: A or B.

fruit: habit; type of arilloid.

Glandularscales

have which less all Some species glandular scales, are more or densely spread over parts of somewhat Sometimes the . They give young parts a sticky shining appearance. they base of the leaflet are very scarce and only found in sunken parts near the or on very

leaflets. young

Domatia

Some species are provided with domatia, though mostly not all collections of one domatia species or all leaflets of one collection have them. In other species never occur.

Pollen

Muller Muller Leenhouts several in & (1976) distinguished pollen types in Sapindaceae.

Two of them, type A (spherical, tricolporate) and type B (triangular, syntricolporate),

in A the are found Arytera. According to Muller & Leenhouts, pollen type represents basic all tribes. and found type in Sapindaceae and occurs in Type B is more specialized is in Melicoccae, Schleichereae, and Cupanieae only. On the photoplates the two pollen types of the are shown as well as a range of sculptural development ektexine.

Fruit

Habit. The description of the fruit made by Radlkofer (1931) in his key to the genera: 'Frucht einander gewdhnlich knopfig-gelappt mit aus gespreitzten Lappen (fruit mostly lobed, be lobes spreading) is, as it already says, only partially true. Actually, two basic types can distinguished: accordance with the of Radlkofer: the axial of the fruit 1. In description parts are limited in and is growth 2 or 3 lobes are present.When an ovule abortive, the correspond-

is between locules This is ing lobe not expanding. Septa the are hardly present. type when mostly glabrous inside; only along the sutures hairs are sometimes present ripe.

the Mostly only lobes are opening; the axial part is not teared up (see fig. 1). R. W. J. M. van der Ham : Notes on Arytera (Sapindaceae) 293

cross-section fruit of lobe with fertile ovule, b: lobe with abortive ovule. Fig. 1. through type 1. x¾ (a:

line — cross-section c: pointof attachment of ovule. d: indicatingplace ofopening ofthe lobe.) fig. 2. through lobe, of attachment of ovule, line of of the fruit. fruit of type 2. x 1 (a: b: point c: indicating place opening

section x fold of arilloid. c: d: septum.) — fig. 3. through arilloid of type 1. 3 (a: arilloid,b. micropylar the slit in arilloid. d: testa of seed, e: internal fold of testa, f: micropyle. g: radicula of the embryo. h:

i: — arilloid of cotyledons of the embryo, funicle. j: bundle.) fig. 4. section through type 2. x 3 (a—j:

ofarilloid the — of the hilar of the seed see under fig. 3. k: cap overlying micropyle.) fig. 5. aspect region the arilloid attached hilum. (schematically) (a: testa free from arilloid. b: testa to which is (pseudohilum). c: d: bundle trace, e: micropylar region, f: micropyle. g: line indicating plane of section in fig. 3 and 4.) —

arilloid. b: of arilloid attached between fig. 6. arilloid of type 1 seen from below (hilar aspect), x 2 (a: part in hilum and micropyle. c: micropylar slit arilloid. d: bundle trace, e: hilum. f: line indicating plane of section in fig. 3). BLUMEA No. 294 VOL. 23, 2, 1977

axial is in in 2. Here, the part not much limited growth, which results a hardly lobed

fruit. There are locules and between them. The is two a septum endocarp always hairy. Seeds when the axial the fruit teared can only come free also part of is up (see fig. 2). Arilloid. For all species the seed cover is an arilloid originating from and attached to the

testa around hilum and der f. of arilloid micropyle (Van Pijl, 1955, ib). Two types are observed:

folded 1. Mostly inside near the base; thick, subcoriaceous-fleshy; some layers can be

distinguished with low magnification: an inner and outer one which are thin, soft,

light-coloured, and a middle one which is thick, firm, dark-coloured (see fig. 3).

2. Not folded inside; membranaceous, mostly somewhat fleshy near the base; only one

layer can be distinguished which is yellowish (see fig. 4).

is almost like the arilloid Type 1 an arillus: the descending parts of seem to be fused with the funicle; the funicle is nearly completely surrounded by the arilloid. The middle

the alike the the funicle with low layer of arilloid is very much tissue of as seen magni- fication.

The of the hilar of the and aspect region seed is about the same in type 1 2: the place

where the arilloid is attached is seen as a lightbrown testa (pseudohilum) around hilumand

micropyle (see fig. 5).

slit observed there a A micropylar is only in type 1 (see fig. 6). In type 2, is cap, over-

lying the micropyle and the part of the arilloid which is attached between micropyle and hilum such would be the situation in (when a cap lengthened, same as Mischocarpus reached: arilloid would be an with an appendix arising abaxially of the micropyle). table the characters mentioned above. In 1 the species are grouped by Arytera xero-

Adelb. = Bl. cited Radlkofer. of the carpa (Bl.) as by Some species were incompletely available.

Four these of groups are characterized. However, are not just phenetic groups species

which have some characters in common; during several months of careful study they

revealed themselves coherent gradually as groups of mutually distinctly related species.

xerocarpa-group (sect. Euarytera p.p.)

Pollen well characterized the type A is rare in Asian Cupanieae. The group is by arilloid It has SE. Asia via New Guinea E. Australia. type 1. mainly Malesian distribution from to

azarytera-group (cf. sect. Azarytera)

is in section of Noteworthy the presence of glandular scales also met aPacific Cupaniopsis other the and in a few genera in Cupanieae (see also chapter I. Arytera concolor). The calyx the is not always a pure 'Matayba-calyx': there are transitions to 'Cupania-calyx' (see also The distribution of A. concolor). the group is predominantly a Pacific one.

Arytera o’shanesiana (cf. sect. Xylarytera)

This is rather isolated in Distribution: Australia a species Arytera. (Queensland)

chartacea-group (sect. Euarytera p.p.) Remarkable also and is a character occurring in Cupaniopsis (C. fruticosa Radlk.)

Gongrodiscus: hairs onthe hypocotyl of the embryo. The 5-lobed disk, which sometimes

also known from Both of this well occurs, is Gongrodiscus. species group (as as Cupaniopsis fruticosa and Gongrodiscus) come from .

The to the sections distinguished by Radlkofer. Radlkofer groups roughly correspond these grouped sections (and sect. Mischarytera) in one genus because of a resemblance in R. W. J. M. van der Ham: Notes on Arylera (Sapindaceae) 295

TABLE 1

species of glandular domatia pollen fruit type of arilloid Arytera scales type type

1 1 gigantosperma describ ed on fruj.ts only

- X A 1 1 xerocarpa

- 1 1 brachyphylla X

densiflora - X A 1

- 1 divaricata X A 1

- A 1 distylis X

- 2 2 microphylla X

- B 2 xanthoneura X 2

2 arcuata X - 2

lepidota X - 2 2

- B gracilipes X ?

- 2 2 brackenridgei X B

- 2 samoensis X 2

- 2 setosa X 2

livida X -

o'shanesiana - X A resembling 2 2

chartacea - - B resembling 1 2

collina - - B resembling 1 2

lautereriana - X B ) unlike junlike

macrobotrys - X B j 1 or 2 Jl or 2

leichhardtii - X B not known 296 BLUMEA VOL. 23, No. 2, 1977

least fruit. Actually, however, at two fruit types can be distinguished: the xerocarpa-type and the The fruit the other classified (i) azarytera-type (2). types in groups are as: resem- is bling type 1 or 2, or as: totally different (mischarytera-group). Besides, there a correlation differences and arilloid The with in vegetative characters, pollen types, types. flowers, not discussed here, also seem to be different.

Conclusion: in fruit noticed Arytera proves to be very heterogenous; the resemblance Radlkofer be differences. Three have by appeares to superficial, covering many species to be be in other Three further excluded and can placed genera. species (inch the mischarytera- dubious The be group) are in Arytera. remaining species can arranged into 4 natural groups, which mutually, however, have not much in common. Future research may learn

other the For time whether they could better be added to some genera of Cupanieae. the

the are the The then being, however, groups kept in one genus Arytera. genus comprises:

and A. xerocarpa- group (8 species; moreover, probably including A. foveolata exostemonea) azarytera-group (10 species) chartacea-group (2 species) Arytera o’shanesiana and provisionally also: mischarytera-group (2 species)

Arytera leichhardtii

V. THE DELIMITATION OF ARYTERA AGAINST MISCHOCARPUS

A simple differentiation between Mischocarpus and Arytera in a key is not well possible of relations of because of the heterogeneous nature the latter. As long as the true hips best do with each Arytera are not clear, the one can is to compare Mischocarpus group mentioned for separately. With regard to the characters studied the following can be

Mischocarpus:

glandular scales: absent

domatia: present in most species der pollen type: B (exceptionally A; see Van Ham, 1977c). arilloid type: 2 fruit type: not lobed, 3-locular.

of be Mischo- On account these characters, Mischocarpus exangulatus appears to clearly a carpus. The resemblances with some species of the xerocarpa- group (style with stigmatic nervation in lines, pattern of the leaflets) are not based on true relationship.

REFERENCES

Blumea —288. VAN DER HAM, R. W. J. M. 1977a. A revision of Mischocarpus (Sapindaceae). 23: 251

— of Blumea 1977 c. Pollenmorphology the genus Mischocarpus (Sapindaceae. 23: 301—335.

of in relation to MULLER, J., & P. W. LEENHOUTS. 1976. A general survey pollen types in Sapindaceae

. In: I. K. Ferguson & J. Muller, eds.: The evolutionary significance of the exine. Linn. Soc.

Symp. Ser. 1: 407—445. VAN DER PIJI, L. 1955. Sarcotesta, aril, pulpa, and the evolution of the angiosperm fruit. Proc. Kon. Ned.

AK. Wetensch. C, 58, 2 and 3.

— of in 1957. On the Arilloids of Nephelium, Euphoria, Litchi, and Aesculus, and the seeds Sapindaceae

general. Acta Bot. Neerl. 6: 618—641. RADLKOFER, L. 1879. Ober Cupania und damit verwandte Pflanzen. Sitz. Ber. K. Bayer. Ak. Wiss. M.-Ph.

Kl. Munch. 9: 457—503.

— Pfl. R. 1268—1288. 1931. Sapindaceae. Heft 98a: 15, key; 1933. R. W. J. M. van der Ham: Notes on Arytera (Sapindaceae) 297

EXPLANATION OF PLATES

Plate I. Pollen A. — Adelb. Backer s.n. 1904, herb. Pollen type i—3. Arytera xerocarpa (Bl.) ( y Java, L). outline rounded grains tricolporate, subspheroidal [P/E i.oo (1.04) 1.05]. Equatorial or slightly rounded

Size: P E distinct, in the triangular. 21—23 /rm, 22—24 fim. Ektoapertures fairly long, narrow equatorial about Tectum plane. Endoapcrtures elliptic, 3 jum. striate-rugulate (i) to reticulate-rugulate (3), muri

in — — view anastomosing three dimensions. 1. oblique polar view, x 3700. 2. equatorial of apertural

view region, x 3700. — 3. equatorial of mesocolpium, x 3700.

Pollen 4—6- Arytera o’shanesiana (F. v. M.) Radlk. (Brass 19157» Queensland). grains tricolporate, sub- oblate outline rounded Size: P E [P/E 0.80 (0.82) 0.87]. Equatorial triangular. 21—22/um, 25—28 jum.

about The tectum to be built Ektoapertures distinct, fairly long. Endoapertures rounded, 3 jum. appears of basal membrane in which round On this membrane fused a to elliptic perforations are present. are more in reticulate-striate or less curving ridges anastomosing three dimensions, causing a appearance ofthe outer of surface ($). — 4. polar view, x 3400. — 5. oblique polar view, x 3400. — 6. detail 5, x 13600.

Plate II. Pollen B. — 1—2. Arytera collina (Pancher & Sebert) Radlk. ( McMillan New Cale- type 5049 , donia). Pollen grains parasyntricolporate, oblate to suboblatc [P/E 0.73 (0.77) 0.81]. Equatorial outline rounded Size: P E about triangularto triangular. 24—25 fim, 30—33 //m. Apocolpium 9—13 jum, triangular.

distinct. 2 Tectum striae Ektoapertures Endoapertures elliptic, —3 jum. striate, subparallel, anastomosing mostly in two dimensions. — 1. oblique polar view, x 2550. — 2. detail of 1, x 6300.

3—6. Arytera leichhardtii (Benth.) Radlk. (Lam 7631, Queensland). Pollen grains parasyntricolporate, suboblate outline rounded Size: P E [P/E 0.78 (0.81) 0.88]. Equatorial triangular. 21—22 jum, 23—28 jum. rounded continuous Apocolpium 4—6 fim, about triangular,sometimes with one mesocolpium. Ektoaper-

distinct. about Tectum as a membrane tures Endoapertures elliptic, 3 fim. reticulate-substriate, appearing

with which of muri is The lumina perforations upon a pattern threedimcnsionallyanastomosing lying. are

on superposed the perforations. — 3. polar view, x 2150. — 4. detail of 3, x 8300. — 5. equatorial view,

x 3800. — 6. detail of 5, X 7600. R. W. M. Ham: Notes J. van der on Arytera (Sapindaceae) 299\I

PLATE I No. 300\II BLUMEA VOL. 23, 2, 197

PLATE II