80 AROIDEANA, Vol. 24 A Revision of Schott (:Aionstereae)

J. Bogner Botanischer Garten Miinchen Menzinger Strde 63 D-80638, Miinchen, Germany P. C. Boyce , Royal Botanic Gardens, Kew Richmond, Surrey, TW9 3AB, u.K. C. M. Sakuragui Departamento de Biologia Universidade de Maringa Av. Colombo, 5790 CEP 87020-000 Maringa PR, Brazil

ABSTRACT premnum Schott, Adans., Rha­ phidophora Hassk., Poepp., The Alloschemone Schott is re­ Schott and vised. One new species, A. inopinata Schott. The genera are mostly separated Bogner & P. C. Boyce, is recognized, and on gynoecial characters, notably locule an expanded description of A. occidentalis number, (and thus ) number, (Poepp.) Eng!. & K. Krause is presented. placenta position, seed shape and endo­ The distinguishing characters of the genus sperm presence or absence. Due to their are discussed, especially with regard to the great plasticity, vegetative characters alone genus Scindapsus Schott, from which Al­ are of little use for generic determination. loschemone has been considered insepa­ In the past some botanists have sought to rable. Both genera have unilocular ovaries reduce several genera into one (e.g. Bak­ with basal placentation with one ovule; Al­ huizen van den Brink 1958; Hotta 1970). loschemone differs from Scindapsus by a However, recent work on vegetative anat­ pinnatifid lamina (entire in Scindap­ omy (e.g. French & Tomlinson, 1981) and sus), architecture, anatomical differ­ floral anatomy (e.g. Carvell, 1989) has ences, and Neotropical distribution (Scin­ shown that these 'weak' genera are prob­ dapsus Paleotropical). Our new species ably soundly based. has fused filaments, a unique character in Previously Alloschemone has been treat­ Monstereae, and further differs from A. oc­ ed as part of Scindapsus (e.g. Madison, cidentalis by the narrower leaf pinnate 1976) due to shared gynoecial characters. separated by an oblong sinus. , often However, Madison (1979), French & Tom­ diagnostically very useful in Monstereae, linson (1981) and Carvell (1989) have are still unknown in Alloschemone and are demonstrated convincingly that Allosche­ highly desirable. mone is not only distinct from Scindapsus but also probably not closely related. INTRODUCTION IDSTORY Generic delimitation within Araceae tribe Monstereae has long been subject to Schott (1858) established Alloschemone differing interpretations. Mayo et al. (1997) (as Alloschemone poeppigiana Schott, recognize eight genera for the tribe: Allos­ nom. illeg.) for Scindapsus occidentalis chemone Schott, Schott, Epi- Poepp. (Poeppig 1845) based on a collec- J. BOGNER, P. C. BOYCE, C. M. SAKURAGUI, 2001 81 tion made by Eduard Poeppig in 1831 at cluding Alloschemone. Unfortunately the Ega (modern Tefe) in Brazilian Amazonas. distribution of species with vestigial Poeppig produced a good description of is, in this instance, of little taxonomic sig­ his new species, which is fortunate since nificance. the single spadix collected was lost. In Alloschemone has been seldom collect­ separating Alloschemone from Scindapsus, ed. Following Poeppig's original gathering Schott (1858, 1860) cited the disjunct dis­ it was recollected by B. A. Krukoff in 1934 tribution of Poeppig's species (Scindapsus and then by M. Madison in 1978. The same is wholly Paleotropical) and the divided year it was also found by staff from the leaf lamina (Scindapsus leaf laminae are Instituto Nacional de Pesquisas de Ama­ always entire). zonia (INPA) while working on the Projeto Koch (1856) and Bentham & Hooker Epffita; in 1982 T. Plowman made (1883) included Alloschemone in the Neo­ what was then the best collection. In 1977 tropical genus Monstera but this was never sterile specimens were collected by Kirk­ universally followed. Engler and others bride et at. in Rondonia state, Brazil and (Engler, 1879, 1889; Engler & Krause, in 1987 Solomon made the only known 1908) recognized Alloschemone but with collection from Bolivia. Recently a fine the caveat 'genus dubium, imperJecte cog­ collection has been made by C.M. Sakur­ nitum'. Madison (1976) argued that the agui at Tefe. This collection, which in­ leaf shape and disjunct distribution of At­ cludes for the first time pre-adult loschemone were secondary to the gyn­ and in spirit, forms the basis oecial characters. Alloschemone and Scin­ for the expanded deSCription presented dapsus have a unilocular with a sin­ below. All collections seen by us originate gle ovule on a basal placenta and Madison from Brazilian Amazonia and Bolivia concluded that the genera should be unit­ (Dept. Pando). ed. Later, based on observations along the Alloschemone has been introduced into Rio Negro in Brazilian Amazonas, Madison cultivation only once, from the 1978 Mad­ reported that the growth of Allosche­ ison collection (Madison 6310, see Croat, mone was so different to that of Scindap­ 1986). However, there has been some con­ sus that he subsequently considered Allos­ fusion as to the origin of the in cul­ chemone to be a distinct genus (Madison, tivation under this number and it seems 1979). that it may have become mixed. A speci­ French and Tomlinson (1981) surveyed men of this Madison collection grown by vascular patterns in the stems of Monster­ one of us (J.B.) flowered after several oideae. They found that in Alloschemone years in cultivation and proved to be Mon­ the course of the vascular bundles in the stera spruceana (Schott) Eng!. stem central core differed from that of all other vining Monstereae while Scindapsus REIATIONSIllPS showed similarities to Monstera, Rhaphi­ dophora, and A mydrium. The maintenance of Alloschemone as a Carvell (1989) showed the floral vascu­ genus is well supported, but its generic re­ lature of Alloschemone to be substantially lationships are less clear. Gynoecial char­ different to that of Scindapsus but similar acters are important for the delimitation of to that of Liebm. Allos­ genera in Monstereae. Alloschemone and chemone also shares some features in its Scindapsus, the only genera with a single vasculature typical of Rhodospatha and basal ovule in a unilocular ovary, would Stenospermation. Much of Carvell's work appear to be closely allied. However, oth­ involved studying vestigial tepals at the er characters do not support this assump­ base on the . Monstereae has na­ tion. Alloschemone and Scindapsus differ ked flowers (i.e. lacking a perigon), al­ in features of vegetative and floral anato­ though Carvell found that many species my, shoot architecture, leaf shape and studied have traces of vestigial tepals, in- geographical distribution. Gynoecial char- 82 AROIDEANA, Vol. 24 acters aside, Alloschemone is most similar arrangement of characters is unknown to the Neotropical genera of Monstereae. elsewhere in . Alloschemone has a chromosome number After R. Keating's unpublished anatom­ of 2n = 84 and would be hexaploid if a ical data the following leaf data can be basic number of x = 14 is considered. summarized here (with permission). Two Neotropical genera, Stenospermation Leaf lamina: surface with a smooth, thin and Rhodospatha, have 2n = 28, Rhodos­ cuticle, cells undulate adaxially, patha also 2n = 56, while Monstera and but polygonal abaxially, with straight an­ the other genera of the Monstereae have a ticlinal walls at the cell base. Stomata ab­ certain number of 2n = 60; some other axial, level with the surface and brachy­ numbers have been published but must be paracytic to brachyparahexacytic; subsidi­ confirmed. The chromosome number of ary cells narrow and rounded. Palisade 2n = 84 for Alloschemone suggests a clos­ cells of the mesophyll short, with 6-8 er relationship to Rhodospatha and Sten­ spongy layers, air cavities accounting for ospermation. All counts cited are from up to 600Al of this layer. There are small work by Dr Gitte Petersen (Petersen 1989). cavities at the palisade/spongy boundary and larger cavities regularly spaced within VEGETATIVE ANATOMY the spongy layer. Collenchyma weakly de­ veloped abaxially at the mid-rib, with thin French & Tomlinson (1981) investigated wall layering on epidermal and subepi­ the vascular patterns in stems of subfamily dermal walls. Mid-rib with one to two vas­ Monsteroideae. Their findings mostly sup­ cular bundles. The xylem has small pro­ port the infrageneric classification pro­ toxylem cells and two to five medium­ posed by Engler & Krause (1908) and also sized metaxylem cells. The phloem is a Mayo et al. (1997). French & Tomlinson small irregular strand. Sclerenchyma fibers defined three groups; 'Group l' and '2' to­ ensheathing larger vascular bundles. Tri­ gether equating to Monstereae, 'Group 3' chosclereids are present in small clusters to tribe Spathiphylleae. Alloschemone has in air cavities just beneath the pallisade vascular patterns not fitting one of these layer, larger trichosclereids (up to 40-50 groups. j.Lm) in bigger air cavities deeper in the 'Group I' has simple vascular bundles spongy layer. Tannin cells with dark con­ with the axial bundles formed by the ag­ gregation of small traces branching from tents are scattered and very common in existing bundles. traces are promi­ the mesophyll. Raphide cells are large, nent, enter the central cylinder at the thin-walled and elongate, filled with mul­ node, are superficially attached and fre­ tiple crystals. Druses are very common in quently asymmetrically organized, five small cells in the palisade layer. genera: Amydrium, Epipremnum, Mon­ : sparse in the outer stera, , and Scindapsus. ground . is divided 'Group II' is essentially very similar, dif­ into and central cylinder, the cortex fering most notably in the presence of with eight to ten paranchyma layers and some compound vascular bundles, two central cylinder delimited by bands of fi­ genera: Rhodospatha, Stenospermation. bers. Air cavities are numerous throughout 'Group III' has amphivasal vascular bun­ ground tissue. The vascular bundles are dles forming a highly condensed, irregu­ scattered and numerous in the central cyl­ larly branching system. Bud traces are in­ inder. Sclerenchyma fibers ensheathing conspicuous and do not form nodal arcs, the vascular bundles. Trichosclereids are two genera: Holochlamys, Spath iphyllu m. numerous in all ground tissue. There are Alloschemone lacks basally aggregated also unusual crystals of small or large ag­ axial vascular bundles and has a 'Rhaphis­ gregations of fine or coarse prismatics like' (see Zimmerman & Tomlinson, 1972) throughout the ground tissue and these do upward branching of the leaf trace. This not resemble typical druses. J. BOGNER, P. C. BOYCE, C. M. SAKURAGUI, 2001 83

POLLEN MORPHOLOGY the condition in Monstera deliciosa al­ though M. deliciosa lacks the arching pat­ Grayum (1992) studied the grains tern of the outer vascular ring traces. Arch­ of Alloschemone occidentalis from the ing vascular traces are diagnostic for Rho­ Krukoff 7162 collection (NY). The pollen dospatha and Stenospermation but both grain of this collection is not fully zonate have only a single ring of traces. Scindap­ (=sulcate), ellipsoid and with a shallowly sus has a single ring of vascular traces ter­ foveolate exine. Grayum was not satisfied minating adjacent to the stylar canal but with the examined material and said " ... these do not arch. Carvell concluded that additional fertile collections of this little­ the flowers of Alloschemone were anatom­ known species are needed" (Grayum ically distinct from Scindapsus but showed 1992:15). some similarities to Rhodospatha, Stenos­ We studied pollen material of Plowman permation and Monstera. et al. 12207 for which there are several duplicates all with abundant pollen. How­ ever, it turned out much more problematic SHOOT ARCIllTECTIJRE than Grayum's study when examined with Alloschemone displays a mature (fertile) SEM. The Plowman collection also shows shoot architecture that is very unusual in ellipsoid pollen but with a very inconspic­ Monstereae (Madison, 1979; Sakuragui, uous sulcus best described as "hardly bor­ pers comm.). Mature fertile stems of Allos­ dered diffuse field." The ex­ chemone bear foliage interposed by ine is more-or-Iess rough (compare Figs. up to six nodes each bearing a cataphyll. 3-5 with Grayum 1992:55, Figs. 107-108). This type of shoot architecture is very rare We must agree with Grayum and say in Monstereae and occurs otherwise only that more and better material is needed to in shoots of, e.g. Rhaphidophora decursi­ give a clear interpretation of the pollen va (Roxb.) Schott, Amydrium humile grains. Schott and A. medium (Zoll. & MoritzO Nicolson. The seedling Alloschemone are FLORAL ANATOMY still unknown. Carvell (1989) reported that floral vas­ There are three climbing Neotropical cular anatomy supports the integrity of Monstereae genera: Alloschemone, Mon­ Monstereae (sensu Engler & Krause, 1908; stera and Rhodospatha. Monstera has the Mayo et al., 1997). Genera are generally most diverse shoot architecture. Madison supported by differences in floral vascu­ (1977) describes seven types of 'hetero­ lature. Carvell met with problems inter­ morphic shoots' for Monstera, defined pri­ preting the variability of vascular traces in marily by leaf development sequences. Rhaphidophora and Epipremnum. This, Madison's Monstera models include seed­ particularly for Epipremnum, cast some ling data and the occurrence of skototrop­ doubt on generic integrity. However, the ic shoots (see Strong & Ray, 1975), data percentage of variability appeared to part­ currently not available for Alloschemone. ly be an artifact of limited sampling and, However, comparison of mature and fer­ despite these problems, sufficient data tile shoot data available for Alloschemone were found to show that a close relation­ suggests that none of Madison's Monstera ship between Epipremnum, Amydrium models match Alloschemone. and Scindapsus was unlikely and that fur­ Little data are available for shoot archi­ thermore these genera have little to do tecture in Rhodospatha. Observations of with Alloschemone. herbarium specimens suggest that Rhodos­ Carvell (1989) found that Alloschemone patha species have fairly 'conventional' has two rings of gynoecial traces, the inner shoots with regularly produced foliage trace terminating along the stylar canal, leaves. A specimen of R. oblongata Poepp. the outer trace arching to a position adja­ (Harley et at. 18193 (K)) has a foraging cent to the stylar canal. This is similar to flagellum, suggesting a growth pattern 84 AROIDEANA, Vol. 24

similar to Madison's Fig. 27 (Madison 1977: ception of tribe Anthurieae (sole genus 23 Fig. 27, M. siltepecana Matuda) or Fig. Anthurium Schott) Monstereae displays 29 (Madison 1977:23 Fig. 29, M. acumi­ the greatest range of leaf shape in the Ar­ nata C. Koch). There is no evidence of aceae. Even individual genera have a be­ foraging shoot production in Alloschemo­ wildering range of leaf types (see, e.g. ne. Little data exist for seedlings of Rho­ Madison, 1977). However, it is interesting dospatha. The few juvenile specimens in to compare the distribution of divided Kew herbarium suggest that Rhodw,patha leaves among genera. Two genera of Neo­ produce conventional foliage leaves soon tropical Monstereae have divided leaves, after germination but it is not clear wheth­ namely Alloschemone and Monstera. Ster­ er pass through a 'stolon-like' (see ile specimens of some Monstera species Madison 1977:17-20) seedling stage. (especially M. spruceana) are superficially The fourth Neotropical monsteroid ge­ similar to Alloschemone, although M. spru­ nus, Stenospermation, is a genus of epi­ ceana is easily distinguished by having phytes or terrestrial herbs. However, sev­ more than one primary lateral vein per eral species produce elongated -climb­ pinna. Much has been written about the ing shoots but are not as -like as other significance of split and perforated leaves genera. The shoot architecture of Stenos­ in vining Araceae (e.g. King, 1962; Taylor permation will not be discussed further & Sexton, 1972) but there is no conclusive here. evidence as to the precise function of In the Paleotropics one genus of Mon­ these features. stereae (Epipremnum) is an obligate In the Paleotropics, two species of Epi­ climber and four genera have almost all premnum, four species of Amydrium and species climbing, namely Amydrium (5 nine species of Rhaphidophora have di­ spp., one non-climbing), Rhaphidophora vided leaves. Other features, especially (ca. 100 spp., two none-climbing), Scin­ gynoecial characters, separate Rhaphido­ dapsus (ca. 35 spp., five none-climbing). phora and Amydrium from Alloschemone Of these genera Scindapsus has the most and a close relationship between these diverse shoot architecture of any Paleo­ genera seems unlikely. Epipremnum is tropical monsteroid genus, with four dis­ overall more similar to Alloschemone. tinct patterns. One species, Scindapsus However, the species with divided leaves, certhios P. C. Boyce, has long root-climb­ E. pinnatum (1.) Eng!. and E. aureum ing sterile shoots bearing cataphylls and (Linden & Andre) Bunting are anomalolls scattered, somewhat depauperate, foliage in the genus by characters including stem leaves. These shoots later produce scat­ sculpturing and presentation tered fertile shoots with distichously ar­ and require further study. ranged fully developed foliage leaves functioning as a litter basket. The sterile As noted above, Scindapsus is indistin­ climbing stage of S. certhios is superficially guishable from Alloschemone on gynoecial similar to that of Alloschemone but and the characters but leaf shape and shoot archi­ similarity between the genera does not ex­ tecture are distinct. tend to the fertile shoots. This type of shoot function differentiation occurs else­ DISTRIBUTION where in Monstereae (e.g. Rhaphidophora schlechteri K. Krause) and in tribe Philo­ Alloschemone is known only from Brazil dendreae (e.g. Philodendron Iinnaei and Bolivia. Sterile collections from Vene­ Kunth) and is probably not of taxonomic zuela tentatively assigned to Alloschemone significance. have all proved to belong to Monstera. All genera of Monstereae recorded for the LEAF SHAPE Neotropics (Le. Alloschemone, Monstera, Leaf shape is of less use for determining Rhodospatha and Stenospermation) are generic affiliations. With the possible ex- not fOllnd outside of Mexico, Central and J. BOGNER, P. C. BOYCE, C. M. SAKURAGUI, 2001 85

South America and the Caribbean except Prantl, Die natiirl. Pflanzenfam. II, 3 as introductions. 120 (1889); Engl. & K. Krause in Engl., Scindapsus, as defined here, is strictly Das Pflanzenr. 37 (IV. 23B): 117-118 Paleo tropical, occurring from India to the (1908); Madison, Selbyana 1(4):325- western Pacific (Samoa). Except for Rha­ 327 (976) & Aroideana 2 (3):73 phidophora (ca. seven, mostly undescri­ (979); French & Tomlinson, Amer. J. bed, species in West Africa) all Paleotrop­ Bot. 68(8): 1123-1124, 1128 (981); ical Monstereae genera are restricted to the Carvell, FI. Anat. Pothoideae Monster­ area from the Ryukyu Islands and Okasa­ oideae 317-325, 525-528; (989); wara Guntu (Bonin Islands) to northern Mayo et al., Genera of Araceae 125- Australia and from India to Rarotonga 126, PI. 18. (Cook Islands). Trichosclereids abundant. Robust to TO WHAT IS ALLOSCHEMONE large evergreen root climber. Stem terete, epidermis smooth at first, becoming dis­ RElATED? tinctly corky and longitudinally furrowed The available evidence suggests that Al­ with age. Juvenile and pre-adult plants loschemone is not closely related to Scin­ with long flagelliform shoots and some­ dapsus. The similarities of the times forming sparse terrestrial colonies. are probably derived independently by densely arising along entire length the loss of a gynoecial septum. If the gy­ of climbing stem, terete, gray, tomentose, noecium prior to septal loss had a few adhering firmly on the trunk. Juvenile large seeds on an axial or deeply intrusive leaves spreading, slightly scattered, 1-3 ca­ placenta (i.e. as in Epipremnum or Amy­ taphyllary nodes between each foliage drium) then the loss of the septum would leaf, lamina entire, ovate to ovate-elliptic, most likely result in a basal placenta with apex acute, base slightly cordate to almost fewer and thus a single basal ovule truncate, margin entire to shallowly divid­ could have arisen independently in both ed (in more intermediate stage); petiole genera. The anatomical characters re­ long, geniculate apically, sheath short. viewed above suggest a link to Neotropi­ Adult leaves spreading, laxly recurved, cal rather than Paleotropical genera. Allos­ scattered, up to six cataphyllary nodes be­ chemone has unique stem anatomy but tween each foliage leaf; cataphylls long, shares gynoecial anatomical characters often reaching almost the length of the with all three genera of Neotropical Mon­ petiole, at first membranous, light green, stereae, basal placentation with Stenosper­ soon drying chartaceous and turning dark mation, vining habit but not shoot archi­ brown; petiole geniculate apically, thick­ tecture with Monstera and Rhodospatha, ened at base, sheath short, with margin in­ and divided leaves only with Monstera. curved and smooth on older leaves; leaf Much work on the phylogeny of the Ar­ lamina coriaceous, ovate-subcordate in aceae and related alismatid monocot fam­ outline, pinnatifid, mid-green, paler be­ ilies. is currently being undertaken using neath, pinnae 4-6 on each side of the mid­ molecular techniques. Preliminary work rib, subalternate, separated from each oth­ on the phylogeny of the Monstereae sup­ er by a rounded or oblong sinus and often ported a phylogenetic link between Allos­ with curtailed pinna development at the chemone, Stenospermation and Rhodos­ base, falcate, acute, each pinna with only patha (Tam, pers. comm.). one primary lateral vein, secondary lateral veins more-or-less parallel pinnate, higher Alloschemone Schott, Gen. Aroid. App. order venation reticulate. Inflorescence (1858) & Prodr. Syst. Aroid. 358 solitary, erect, (strongly sweet-fragrant on (1860); Engl. in Martius, FI. Bras. 3(2): opening in A. occidentalis); 115 (1878) & in A. & C., DC., Monog. shorter than petiole; spathe ovate-cymbi­ Phanerog. 2:267 (1879); Eng!. & form, coriaceous, deciduous at anthesis, 86 AROIDEANA, Vol. 24 convolute to half way, shallowly concave Gen. Spec. Pl. 3:88 (1845}-Monstera and gaping above at female receptivity, occidentalis (Poepp.) C. Koch ex En­ apex more-or-Iess obtuse, exterior creamy der, Ind. Aroid. 4, 54 (1864); Koch in white, glossy, interior flushed pink; spadix Bonplandia 4:4-10 (1856); Bentham & shorter than spathe, sessile to short-stipi­ Hook.f, Gen. Pl. 3:992 (1883). Type: tate, cylindric, apex obtuse, pale yellow to Brazil, Municipio de Tefe, Tefe ('Ega'), dull pink or orange, stipe terete. Flowers 1831, Poeppig s.n. ('Addenda 107') (ho­ bisexual, naked; stylar areas yellow. Sta­ lotype P; isotype LE). Figs. 1, 2(B-M), mens four, filaments flattened, free or par­ 3-5 & back cover C-D, F-J. tially to completely connate, shorter than gynoecium, thecae oblong, dehiscing by Robust to very large evergreen root oblique apical short slit; pollen not fully climber, at least 5 m high when flowering. zonate, ellipsoid, medium-sized (mean 46 Stem in adult plants adherent, unbranched /-Lm), exine shallowly foveolate. Gynoeci­ (unless apical damaged and then urn with prismatic ovary, I-locular, with reiterating from a node close to the dam­ abundant locular mucilage, ovule one, am­ aged apex), terete, (1-)1.5-2.5 cm diam., phitropous, funicle bearing , pla­ internodes ca. 1 cm long, epidermis centa basal; stylar region densely packed smooth at first, mid-green to pale gray or with trichosclereids, sessile, elliptic; brown, becoming conspicuously longitu­ often some gynoecia at base of spadix dinally corky-winged with age. Juvenile larger and apparently sterile and nectari­ and pre-adult plants with long, robust fla­ ferous. and seed unknown. Chro­ gelliform shoots and sometimes forming mosome number 2n = 84. sparse terrestrial colonies. Roots densely produced from entire climbing stem Type species-Alloschemone poeppigi­ length, adhering firmly to trunk, cylin­ ana Schott, nom. illeg. (=A. occidentalis drical, 2-6 cm X 2-3 mm diam., more-or­ (Poepp.) Engl. & K. Krause). less grayish, appearing tomentose, be­ Two species restricted to Brazil (Ama­ cause densely covered with 1-2 mm long zonas, Rondonia) and Bolivia (Dept. Pan­ root hairs. Juvenile leaves spreading, do). slightly scattered (one to three cataphyl­ lary nodes between each foliage leaf), Etymology: Greek atlos (=other), sche­ lamina lanceolate, ovate or ovate-elliptic, ma, schematos (=form) and one (=be­ up to 17 X 7 cm, apex acute, base cuneate ing); in allusion to the plant's habit, which to slightly cordate to almost truncate, mar­ looks different to other aroids. gins entire to shallowly divided in pre­ adult leaves, petiole up to 9 X 0.3 cm, ge­ Key to the species of AUoschemone niculum ca. 1 cm long; adult leaves laxly recurved, scattered (up to six cataphyllary 1. Pinnae of the leaf lamina broader (5-10 nodes between each foliage leaf); leaf em wide), sinus between the pinnae lamina thinly coriaceous, ovate-subcor­ round; filaments of the free ...... 1. A. occidentalis date in outline, pinnatifid, 45-95 X 48-66 1. Pinnae of the leaf lamina narrower (2-3 cm, shining mid-green, paler beneath, pin­ em wide), sinus between the pinnae ob­ nae subalternate, oblong-elliptic, 18-32 X long; filaments of the stamens connate .. 5-10 cm, falcate, apex acute, lacinae ex­ ...... 2. A. inopinata tending 1-4 cm from midrib, separated by 1. Alloschemone occidentaJis (Poepp.) a narrow to broad, rounded sinus, and of­ Engl. & K. Krause in Engl., Das Pflan­ ten with curtailed pinna development at zenr. 37 (N.23B):117 (1908}-Allosche­ the base, primary lateral veins 1 per pinna, mone poeppigiana Schott, Gen. Aroid. secondary lateral veins parallel-pinnate, App. (1858), nom. illeg. pro. A. occi­ 15-17 on each side of pinna, higher order dentalis (Poepp.) Engl. & K. Krause­ venation reticulate; petiole geniculate api­ Scindapsus occidentalis Poepp., Nov. cally, thickened at base, (26-)30-70 X J. BOGNER, P. C. BOYCE, C. M. SAKURAGUI, 2001 87

c A

Fig. 1. Alloschemone occidentalis. A leaf tip and base, mid-portion omitted, basal part with left hand lobe folded over to show abaxial surface X 1h; B inflorescence, spadix longitudinally sectioned and spathe damaged in nature X 1h; C detail of post-anthesis spadix X 2; D four flowers, three quater view X 5; E stylar region, plan view X 10; F gynoecium, longitudinal section with associated spadix axis tissue X 5. A-B Plowman et al. 12207, C-F Krukoif 7162. Drawn by Emmanuel Papadopoulos. 88 ARO IDEANA, Vol. 24

A

Fig. 2. A. ALloschemone inopinata. A leaf tip and base, mid-portio n o mitted V2. B-M ALloschemone occidentalis. B young stem showing clasping roots, cataphyllary node scars and two fo liar leaf peti ole bases X 213 ; C inflorescence, with portio n of spathe base X 73; D detail of pre-anthesis spadix X 1 liz; E three fl owers, sid e view X 4; F three fl owers, three quarter view X 4; G stylar regio n, plan view X 6; H gynoeciulll, lo ngitudinal secti on showing abundant trichomes X 6; J ovule X 10; K - M anthers in adaxial, three quarter and abaxial view X 4. A fro m Pro/eta Flora Epijita 3 10, B-M fro m Plowman el al. 12207. Drawn by Emmanue l Papado po ul os. J. BOGNER, P C. BOYCE, C. M. SAKURAGU1, 2001 89

Fig. 3. Alloschemone occidentaLis. Po ll en SEM from Plowman 12207. SEM micrographs by M. Hesse.

(0.7-)1-1.5 cm, petiolar sheath sho rt, 2-10 ing above on opening, apex obtuse, ex­ cm long, margins on older specime ns in­ terior creamy white, glossy, interior curved, smooth; geniculum 2- 2.5 cm long. flu shed pink, strongly sweet-fragrant; spa­ l1~f1.01'escence solitary, erect; peduncle dix sessil e to short-stipitate, cylindric, 10- erect, terete, 9-15 X ca. 2 cm diam. , red­ 12 X 2.5-3.5 cm, apex obtuse, stylar tissue dish; spathe deciduous, cori aceous, ovate­ pale yell ow to dull pink or orange, also cymbiform, 17-20 X 7-9 cm , convolute to very sweet-fragrant; stipe terete, (0-)-1.2 half way up, shallowly concave and gap- X ca. 3 cm. FLowers: bisexual, naked; sta- 90 AROlDEANA, Vol. 24

Fig. 4. Alloschemone occidentalis. Pollen SEM from Plowman 12207. The pollen is not acetolyzed; the re is a li ttle poll enkitt visible. SEM micrograph by M. Hesse.

m.ens 4, filaments fl attened, free (4-)5-8 X Distribution-Brazil (Amazonas, Ron­ 1-2 mm, thecae oblong, ca. 2111m lo ng, de­ donia), Bolivia (Dept. Pando). hiscing apically by an oblique, sho rt slit. Pollen not fu lly zonate, ellipsoid, 44--46 X Specimens seen-BRAZIL. Amazonas: 27-28 jJ..111 , exine shallowly foveolate. Gy­ No further data, Hutchinson 8687, 8699 noecium prismatic, 8- 9 X (2.5- )3.5-4 mm (UEC); Municfpio de Humayta (Humafta), at top, below 2.5-3 mm wide (gynoecia at Rio Madeira basin, plateau between Rio base of spadix larger, 8-9 X 4-6 mm, ap­ Livramento and Rio Ipix una, 7-8 Nov. parently sterile and nectariferous), stylar 1934, J(ru.l~o.fl 7162 (GH, NY); Municfpio regio n de nsely packed with trichoscler­ de Maraa , Rio Japura, near Maraa, La go eid s, sti gma prominent, e lli ptic in plan Maraa, 10 51'S, 65°36'\Xf, L't-5 Dec. 1982, view 1.3-1.5 mm X ca. 1 mm, stigmas o ri ­ Plowman e/ at. 12207 (INPA, K, MO, JY); entated longitudinally along the spadix Municfpio de Tefe, Tefe ('Ega'), 1831, axis; ovary I-locul ar, with abundant locu­ Poeppig s. n . ('Addenda lOT) (holotype P, lar mucilage; ou-u.te 1, amphitropous, 1-1.2 isotype LE); Mu ni cfpio de Tefe, Tefe, road mm tall , ca. 1.4 mm w ide, funicl e bearing to airport, ca. 5 km from Tefe, 17 Oct. trichomes, placenta basal. Fruit unknown. 1995 , Sakuragui 1050 (K, M; SPF). Ron­ Seed unknown. Chromosome number 2n donia: at km 11 on the road to Saldanha, = 84. close to Guajara-Mi rim, 12 Jan. 1977 Kirki­ Flowering-October to Novembe r; bride /I'., Lleras & Leite-Lleras 2712 (US). spathe and spadix strongly sweet-fra grant. BOLIVIA. De pt. Pando, Provo Gral. Fred- J. BOGNER, P. C. BOYCE, C. M. SAKURAGUI , 200 ] 9]

Fig. 5. Alloschemone occidentalis. Pollen SEM fro m Plowman 12207 showing ± ro ugh, but not foveolate, exine. SEM micrograph b y M. H esse.

erico Ho man, Hio Negro , 100 34'S, 66°06'w , 3--4 cm lo ng, 3 nodes bearing ca taphylis 17 June 1987, Solomon 17114 (MO). fo llowed b y foliage lea f (note by Madi­ son); clasping roots covered dense ly w ith Ecology-Caatinga, primary o r degrad­ root hairs. Juvenile leaves unknown; adull ed seasonal evergreen fo rest on terra firme lec!/Iamina p innatifid; petiole 17-25 X ca. of grayish clay o r sa ndy clay w ith deep 0.5 cm , geniculum 1-2 cm long, shea th litter. 10-120 m . sho rt (ca. 1 cm long); lamina pinnatifid, 45-50 X 35-40 cm , m ostl y 4 pinnae o n 2. Alloschemone inopinata Bogner & each side, midrib quite stro ng, ca. 3 mm P.e. B oyce sp. nov., differt ab A. oc­ diam. , 1 primary lateral vein running from cidentalis pinnis angustioribus (2-3 the midrib into each pinna, pinnae 13- 23 cm latis) , sinus inter pinnis oblongo, X 2-3(--4) cm, acute to som ewhat cuspi­ filamentis stamino rum connatis. Ty­ date, apex o f upper pinnae turning up­ pus: Brazil , A mazonas, H.io egro, wards, ca. 15 secondary lateral veins o n mo uth o f the Hi o Marie, 16 Oct. 1978, each side of the primary vein, these thin­ Madison et al. 6310 (I< , ho lotypus; ner, parallel-pinnate; higher o rder vena­ INPA, MO, SEL, isolyp i). Figs. 2(A) & tion reticulate; sinus between the p innae back cover A-B, E. oblong, lacinae extending 1- 1. 5 cm fro m Eobust evergreen root-climber, several midrib . J/~flO1'escence solitary; peduncle ca . meters tall; Stem terete, outside corky in 10 X 0.7 cm; spathe deciduo us; spadix cy­ adult stage, 0.8-1.5 cm diam. Internodes lindric, ca . 11 X 1.8 cm , ap ex o btuse . 92 AROIDEANA, Vol. 24

Flowers bisexual; gynoecium ca. 5 X 4 only. This can be concluded from the few mm, truncate, more-or-Iess rhombic in collections made within 170 years. plan view; stigma broadly elliptic; stamens The seed structure is very important in shorter than gynoecium, filaments con­ the Monstereae, but and seeds are nate, thecae oblong, dehiscing by a broad still unknown for Alloschemone. There­ apical slit. Fruit and seeds unknown. Chro­ fore, it would be very useful to get mature mosome number unknown. fruits. The living plants in cultivation grow only slowly, and it will take many years Flowering-October to December. for them to reach a size for flowering and Distribution-Brazil (Amazonas). perhaps fruiting. Nevertheless it can be predicted from the structure of the ovule Specimens seen-BRAZIL. Amazonas: that the seed cannot be straight, it must be Humaita, Cid 8781 (INPA); Rio Negro, somewhat curved. Further, it would be mouth of the Rio Marie, 16 Oct. 1978, very useful to know the structure of the Madison et al. 6310 (K, holotype; INPA, testa, if is present or absent, MO, SEL, isotypes); Rio Negro, mouth of and how the is formed. the Rio Marie, 16 Oct. 1978, Projeto Flora Mike Madison sent a color postcard pic­ Epifita 310 (INPA); Humaita, Teixeira et turing an Amerindian and a leaf in the up­ al. 96 (INPA); per left corner which is clearly an adult Ecology-Caatinga on terra firme. ca. leaf of Alloschemone occidentalis. It is not 10 m. known exactly where this photograph was taken. Croat (1986 '1985':80--82, Figs. 1-3) pic­ REMARKS tured juvenile leaves as Alloschemone oc­ The completely connate filaments of AI­ cidentalis, but these were later shown to loschemone inopinata are unique not only belong to Monstera spruceana, when the in this genus but also in the whole of Mon­ plant flowered. stereae. The Krukoif 7162 collection clear­ Plate 18 in Mayo et al. (1997), shows A­ ly shows free filaments which are never B, E, Alloschemone inopinata, and C-D, & connate partly at all. The Madison 6310 F-], A. occidentalis. collection has stamens where all filaments of a are connate, forming a ring. ACKNOWLEDGMENTS The Plowman et al. 12207 collection shows both free filaments and also sta­ The Keepers and Curators of the follow­ mens where two are connate and thus this ing herbaria are thanked for assistance character is variable. The leaf pinnae of during visits, and for the loan of material: the Plowman et al. 12207 collection are GH, INPA, K, LE, MO, NY, P, SEL, SPF, broader, and the sinus of the pinnae is UEG, US. Thanks are due to Emmanuel more-or-Iess round, as it is in typical A. Papadopoulos and Eleanor Catherine for occidentalis. From the available material it executing the illustrations, Gitte Petersen, is difficult to decide if Alloschemone com­ Copenhagen for counting the chromo­ prises one very variable species or wheth­ somes, Prof. M. Hesse, Wien, for the SEM er there are two, more narrowly defined micrographs of the pollen grains, Dr H. species. For the present we believe that RoeSler, Miinchen, for providing the Latin there are two species. We hope that this diagnosis and to Prof. R. Keating, Ed­ paper will stimulate people into trying to wardsville, Illinois, for additional anatom­ collect more, fertile, ideally fruiting, collec­ ical data. tions of this enigmatic genus. The genus Alloschemone has a wide dis­ UTERATURE CITED tribution On Amazonian Brazil and one re­ cord is known from Bolivia) but it seems Bakhuizen van den Brink, R. C. 1958. Are to be never common and occurs locally Epipremnum Schott, Rhaphidophora ]. BOGNER, P. C. BOYCE, C. M. SAKURAGUI, 2001 93

Hassk., and Monstera Adans. conge­ King, H. W. 1962. The physiology of Mon­ neric? Blumea suppl. 4:91-92. stera deliciosa. J. Queckett MiG. Club, Bentham, G. & J. D. Hooker, 1883. Ara­ Ser. 2, 32:125-136. ceae ('Aroideae'). Genera Plantarum, Koch, C. 1856. Monstera und Scindapsus. vol. 3:955-1000. Lovell Reeve & Co., Bonplandia 4:4-10. London. Madison, M. T. 1976. Alloschemone and Carvell, W. N. 1989. Floral anatomy of the Scindapsus (Araceae). Selbyana 1: Pothoideae and Monsteroideae (Ara­ 325-327. ceae) Unpublished Ph.D. Disserta­ ---. 1977. A revision of Monstera (Ar­ tion, Miami University, Oxford, Ohio. aceae). Contrib. Gray. Herb. 207:3- Croat, T. B. 1986 ('1985'). A new collection 100. of the rare Alloschemone occidentalis ---.1979. Notes on some aroids along (Poepp.) Engl. & Krause. Aroideana the Rio Negro in Brazil. Aroideana 8(3):80-82. 2(3):67-77. Engler, A. 1879. Araceae. In A. & c. de Mayo, S.]., J. Bogner & P. C. Boyce. 1997. Candolle, Monographiae Phaneroga­ The Genera oj Araceae. xii + 370 pp. marum, vol. 2:1-681. Masson, Paris. Royal Botanic Gardens, Kew. ---. 1889. Araceae. In A. Engler & K. Petersen, G. 1989. Cytology and system­ Prantl (eds.), Die naturlichen Pjlan­ atics of Araceae. Nord. J. Bot. 9:119- zenJamilien 2(3):102-153. W. Engel­ 166. mann, Leipzig. Poeppig, E. 1845. Nova Genera ad Species Engler, A. & K. Krause. 1908. Araceae­ Plantarum, vol. 3. Leipzig. Monsteroideae. In A. Engler (ed.), Schott, H. W. 1858. Genera Aroidearum Das Pjlanzenreich 37(IY.23B):4-139. Appendix. Ueberreuter, Vienna. W. Engelmann, Leipzig. ---. 1860. Prodromus Systematis Aro­ French, J. c. & P. B. Tomlinson. 1981. Vas­ cular patterns in stems of Araceae: idearum, p. 358. Typis congregationis subfamily Monsteroideae. Amer. J. mechitharisticae, Vienna. Bot. 68(8):1115-1129. Strong, D. R. & T. S. Ray. 1975. Host tree Grayum, M. H. 1992. Comparatve External location behavior of a tropical vine Pollen Ultrastructure of the Araceae (Monstera gigantea) by skototropism. and Putatively Related Taxa. Mono­ Science 190:804-806. graphs·Syst. Bot. Missouri Bot. Gard. Taylor, S. E. & O. J. Sexton. 1972. Some 43:1-167. implications of leaf tearing in Musa­ Hotta, M. 1970. A system of the family Ar­ ceae. Ecology 53:143-149. aceae in Japan and adjacent areas. Zimmerman, M. H. & P. Tomlinson. 1972. Mem. Fac. Sci. Kyoto Imp. Univ., Ser. The vascular system of monocotyle­ Bioi. 4:72-96. donous stems. Bot. Gaz.133:141-155.