A QUARTERLY PUBLICATION FOR MEMBERS OF THE INTERNATIONAL AROID

The Araceae of Indomalaya & TropicalAustralasiaby Wong Sin Yeng & Peter C Boyce 1Scouting the Jama-Coaqueby Jason P. Hernandez 2The Infructescences of Hapalineappendiculata Ridl.by Wong Sin Yeng & Peter C Boyce 3Bernard Jadin, Husband of AroiderFrieda Billiet Dies in Belgiumby Thomas B. Croat 4

IAS Officers :

President : Zach DuFranVice President : Peter BoyceCorresponding Secretary : Kathy UptonRecording Secretary : Jason SarineTreasurer : Denis Rotolante

Newsletter :

Editor : Carla KostelacLayout : Albert Huntington

This edition of the IAS Newsletter is Copyright ©2012 by the International Aroid Society, Inc.

A QUARTERLY PUBLICATION FOR MEMBERS OF THE INTERNATIONAL AROID SOCIETY

by Peter C. Boyce & Wong Sin Yeng

Peter C. BoycePusat Pengajian Sains Kajihayat [School of Biological Sciences]Universiti Sains Malaysia 11800 USMPulau Pinang, [email protected]

Wong Sin YengDepartment of Plant Science & Environmental EcologyFaculty of Resource Science & TechnologyUniversiti Malaysia Sarawak94300 Kota SamarahanSarawak, [email protected]

ABSTRACTA key to the higher taxa of aroids of Indomalaya and tropical Australasia is offered. Example images are provided for mosttaxa.

INTRODUCTION

Work on the revising the aroids of Indomalaya and Australasia has progressed considerably in the last 15years. Nonetheless, much remains to be done before there is a comprehensive body of published data forall genera to species level. Indeed, in recent years increased fieldwork activities in some parts of theseecozones (notably on Borneo and more recently in NE Peninsular Malaysia) have revealed a wealth ofnovel species such that for even recently revised genera (e.g., Alocasia and Schismatoglottis, etc.), it isapparent that a great deal more awaits discovery, and that the aroid biota (at least those of Indomalaya) isconsiderably richer and more diverse than hitherto suspected.

Additional to the discovery in the wild of much that is taxonomically new, molecular analyses continue toprovide insights into pre-existing taxa, in many instances forcing re-examination of long-held generic

concepts.

Against this background of much new data published in numerous different journals, changing higher taxon concepts, and the high probability that ‘complete’monographs for most genera are still many years away, we thought it useful to provide a key to the higher taxa of aroids in Indomalaya and Australasia thatreflects current taxonomic conditions.

The key presented here is unorthodox in that it not only keys out all the currently accepted genera but also keys out separately major generic subordinal units,and those species that no longer fit comfortably in their former generic placement.

IAS Newsletter Vol 34, No 1 http://www.aroid.org/member/newsletters/nl34-01/

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Figure 1. Lemna minor L. Although radicallydifferent in appearance to the rest of the Araceae,molecular analyses and study of theinflorescences of the duckweeds (which arebisexual) have established their positionembedded in the family and sister to thePothoideae.

Figure 2. Anaphyllumwightii Schott. A genusof two species restrictedto SW India.Relationships ofAnaphyllum to the otherextant lasioids imply thatthey are vestiges of anancient lineage ofaroids. Image © RenéStalder, used withpermission.

Figure 3. Pycnospathaarietina Gagnep.Pycnospatha may beseen as the Asian‘equivalent’ ofNeotropical Dracontium.Although superficiallyvery similar in flora andvegetative appearance,Pycnospatha isimmediatelydistinguished by theflowers lacking tepals.The dracontioid leaf ofPycnospatha is sharedwith Dracontium andalso occurs, certainlyindependently, in theAfrotropics genusAnchomanes. Image ©René Stalder, used withpermission.

Figure 4. Lasia spinosa(L.) Thwaites. One of themost wide-spread speciesof lasioid, often persisting incleared land owing to theuse of the emerging leavesand inflorescences as avegetable. Here thedistinctive spinulous fruitsare shown.

Figure 5. Cyrtospermaferox N.E.Br. Anotherstriking species of theLasioideae. Here shown infruit. Unlike Lasia theinfructescence is declinateand the fruits smooth.

Figure 6. Podolasiastipitata N.E.Br. Whilerather similar in appearanceto Crytosperma, thepeastswamp-dwelling genusPodolasia produces erectspikes of red berries.

Figure 7. Pedicellarum paieiM.Hotta. Pedicallarum is veryclose in appearance to severalPothos species, notably those ofthe Goniurus complex withscattered flowers. Molecularwork is underway to resolve therelationships between thevarious species groups.

Figure 8. Pothoidium lobbianumSchott.Pothoidium closely resembles somespecies of Pothos, notably P. repens (Lour.) Druceand P. longipes Schott, but differs in the uniquearchitecture of the inflorescences, and in beingfunctionally dioecious. Pictured is a pistillate plantin young fruit.

Figure 9. Pothosovatifolius Merr. Ripeinfructescence. Note thatthe individual fruits arepointed (compare with thetruncate-topped fruits ofAnadendrum in Fig. 11).


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Figure 10. Spathiphyllum commutatum Schott.Spathiphyllum is currently the only aroid genusthat is definitely represented in the Asian andNeotropics. Formerly Schismatoglottis andHomalomena were both considered to bedistributed in the manner, but recent and on-goingresearch has established that the Neotropicalgenera are different. Spathiphyllum commutatumis one of three species in the Asian tropics.

Figure 11. Anadendrum. Isperhaps the least-wellunderstood genus ofclimbing aroids inIndomalaya. Most of thespecies are yet to bedescribed, including the onefigured here. Note thattruncate-topped fruits.

Figure 12. Amydriummedium (Zoll. & Moritzi)Nicolson. Amydrium fruitsare berries (here shownripe) and do not shed thestylar plate to reveal theseeds in the manner ofother Monstereae (see Fig.12).

Figure 13. Scindapsusgeniculatus Engl. Unlikethe Neotropics, epiphyticaroids are rare in the Asiantropics. Exceptions occur inRemusatia (most), a fewArisaema, and in the genusScindapsus, where the ca20 species (most yet to bedescribed) related to S.geniculatus are obligatetrash-basket epiphytes. Theplant here is growing onTristaniopsis beccarii(Myrtaceae).

Figure 14. Rhaphidophoralobbii Schott. Here shownwith a ripe infructescence(monsterocarp) sheddingstylar plates to reveal seedsembedded in pulp. Thestylar plates are full ofrhaphides and causeintense irritation if bitten orhandled incautiously.However, the pulp cavity isfree of these structures and,indeed, the pulp is sweet-tasting.

Figure 15. Epipremnumpinnatum (L.) Engl. This isa large growing species thatis frequently limestone-associated in habitat. Thespecies is widespread intropical Asia, with numerouslocalized and oftendistinctive populations,notably on the PhilippineIslands. The plant featuredhere originates from easternSabah, the only place onBorneo that this speciesoccurs.

Figure 16. Pistia stratiotes L. The ubiquitouswater lettuce, Pistia can become pestilential onslow-moving waterbodies.

Figure 17.Amorphophallusranchanensis Ipor,A.Simon & Meekiong.Amorphophallus continuesto surprise with newspecies. Amorphophallusranchanensis, describedfrom basalt close to Serianin western Sarawak seemsto be closest in appearanceto species from Sabah.Borneo’s Amorphophallusflora now stands at 19described species, anincrease of 15 since 2000.In Thailand half of all aroidsare either Amorphophallusor Typhonium.

Figure 18. Arisaemafiliforme Blume.Arisaema is richest inwarm-temperate andsubtropical Asia.However, there areseveral species in theAsian wet tropics. Image© Eric Hunt, used withpermission.

Figure 19. ‘Lazarum’, isa group of speciesrestricted to Australiaformerly included inTyphonium. Here an asyet undescribed speciesis shown, with thespathe artificially openedto show the thickened‘ring’ separating thelower spathe and spathelimb. Althoughwell-supported bymolecular data, themorphologicalseparation of Lazarumfrom Typhonium remainsproblematic.


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Figure 20. Typhonium trilobatum (L.) Schott.Typhonium is richest and most diverse in stronglyseasonal tropical habitats, notably Thailand andIndochina. Image © John Mood, used with permission.

Figure 21. Sauromatumhorsfieldii Miq.Sauromatum hasrecently beenonce-again removedfrom Typhonium andexpanded to includeseveral additionalspecies. Sauromatumhorsfieldii is awidespread species,occurring from SWChina, though Jawa, toBali.

Figure 22. Cryptocorynelongicauda Becc. ex Engl.Photographed in deepsaturated peat layersoverlaying limestone inMulu N.P. Cryptocorynespecies are importantenvironmental indicators asthey are highly dependenton unpolluted water andgood forest cover.

Figure 23. Aglaodorum griffithii (Schott)Schott. Superficially similar to (and closely relatedto) Aglaonema (Fig.24 & 25), Aglaodorum is ahelophyte with large green spongy fruits. Whenmature they fall and float away.

Figure 24. Aglaonemasimplex (Blume) Blume.One of several widespreadspecies in the genus. Herephotographed at the NiahCaves in N Sarawak.

Figure 25. Aglaonema nitidum (Jack) Kunth.Here fruiting in forest not far from Kuching. Thered fruits are eaten by birds such as spiderhunters(Arachnothera).

Figure 26. Nephthytisbintuluensis A.Hay,Bogner & P.C.Boyce.Plants in the type locality.Note that the inflorescencesare some distance from theaccompanying (preceding)foliage leaf. The locality,photographed here in thedry season, is inundated to1 metre (ca 3 ft) or more fordays at a time during thewet season.

Figure 27. Nephthytisbintuluensis A.Hay,Bogner & P.C.Boyce.Although vegetatively quitedissimilar to the relatedAglaonema, the similarity ofthe inflorescence ofNephthytis to that ofAglaonema in striking.


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Figure 28. Furtadoa mixta (Ridl.) M.Hotta. Althoughthe plants look very similar to Homalomena belonging tothe Chamaecladon Supergroup, the arrangement offlowers is quite different (see Fig. 31). Image ©Zulhazman bin Hamzah, used with permission.

Figure 29. Furtadoamixta (Ridl.) M.Hotta.Note that each staminateflower accompanied byand is overtopped by aflask-shaped pistillode.Image © Zulhazman binHamzah, used withpermission.

KEY TO HIGHER TAXA OF INDOMALAYAN AND TROPICAL AUSTRALASIAN ARACEAE

1. Plants comprising small to minute few-rooted or rootless free-floating thallus-like leafless bodies . . Lemnoideae (>2)2. Roots 1 per plant body; plant bodies with 1–3 veins, without a scale at the base . . .Lemna

3. Roots absent; plant body without veins; the daughter plant bodies originating from a single terminal pouch or cavity at the base of the plantbody; inflorescence originating in a cavity on the plant body upper surface Wolffia

2. Roots 1–21 per plant body; plant body with 1–21 veins; the daughter plant bodies and inflorescences originating from 2 lateral pouches at the base ofthe plant body

3. Roots (1–)2–21 per plant body; plant bodies with (3–)5–21 veins, surrounded at its base by a small scale (prophyll) covering the point ofattachment of the roots

4. Plant bodies 1–1.5 times as long as wide, with 7–12 roots of which 1(–2) perforate the scale . . Spirodela4. Plant bodies 1.5–2 times as long as wide, with (3–)5–7 veins and (1–)2–7(– 12) roots all of which perforate the scale . . Landoltia

1. Plants not as above. If free-floating then leaves forming a conspicuous rosette with copious roots hanging beneath5. Flowers bisexual, mostly with a perigone of conspicuous tepals

6. Plants terrestrial, mostly armed with prickles; leaf development acroscopic Lasioideae (>7)7. Geophytes with ‘dracontioid’ leaf blades; petioles somewhat asperate but never prickly; flowers without tepals . . Pycnospatha7. Armoured or unarmoured helophytes (rarely peatswamp mesophytes, or rheophytes) with hastate to 4×pinnatifid leaves; flowers with aperigone of conspicuous tepals

8. Unarmoured helophytes; spathe open, often twisted; seed testa membranous Anaphyllum8. Armoured helophytes, rarely peatswamp mesophytes, or rheophytes

9. Stems suffruticose, erect to decumbent, usually with prickly conspicuous internodes; leaves hastate to (4×)pinnatifid;spathe caducous or marcescent; placentation apical; fruit usually spinulous . . Lasia9. Stem a condensed rhizome, rarely with distinct internodes, and these then unarmed; leaves entire, sagittate to hastate;spathe persistent to marcescent; placentation not apical; fruit smooth

10. Infructescence nodding, fruits barely emerging from between the tepals, ripening dull purple; seeds crested orwarty; plant with spines mixed straight and up-turned . . Cyrtosperma10. Infructescence erect, fruits emerging fully from between the tepals, ripening bright red; seeds smooth; plant withspines mixed straight and downturned . . . Podolasia

6. Plants mostly climbing hemiepiphytes; flowers naked or perigoniate; if terrestrial helophytes then tepals fused into a thickened ring-likestructure; leaf development basiscopic

11. Climbing hemiepiphytes; tepals free or, if fused, then not forming a thickened ring; ripe fruits red, well-extruded from spadix . . Pothoideae: Potheae (>12)

12. Flowers pedicellate, with a distinct receptacle; tepals of perigone connate forming a conspicuous ‘cup’ . . . Pedicellarum12. Flowers sessile, without a receptacle; tepals of perigone free, very rarely united and then not forming a ‘cup’

13. Flowering shoot terminating in a (leafless) branching system of spadices; flowers functionally unisexual . . . Pothoidium13. Flowering shoot with inflorescences mostly axillary or infra-axillary, if terminal then inflorescence system not branching,and leafy; flowers bisexual . . . . Pothos

11. Terrestrial helophytes; tepals fused into a thickened ring-like structure; ripe fruits green, encased in persistent green tepals . . . Monsteroideae: Spathiphylleae (>14)

14. Inflorescences held down among the leaf bases; spathe limb enclosing the spadix throughout anthesis, not expanding;


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marcescent into fruiting . . Holochlamys14. Inflorescences held above the leaves; spathe limb spreading expanding; persistent and turning green during fruiting, laterfalling . . Spathiphyllum

5. Flowers bisexual, mostly without a perigone of tepals; if tepals present then membranous and very inconspicuous; or flowers unisexual15. Flowers bisexual, perigone (if present) membranous and inconspicuous; mostly climbing hemiepiphytes, rarely rheophyticMonsteroideae: Monstereae (>16)

16. Flowers with a perigon of fused membranous tepals; inflorescences long-slender-pedunculate, numerous in a distichous fan;spathe in bud conspicuously long-beaked (beak to 1/3 length of entire spathe); infructescence with discrete, truncate berries;trichosclereids absent Anadendrum16. Flowers naked; inflorescences very short to long-stout-pedunculate, solitary or few, not carried in a fan; spathe is notconspicuously long-beaked (beak if present less than 1/6 length of entire spathe); infructescence mostly a monsterocarp, if withdiscrete berries, then these not truncate; trichosclereids present (very sparse in Amydrium)

17. Venation reticulate; infructescence with discrete berries; trichosclereids very sparse . . . Amydrium17. Primary venation striate; higher order venation reticulate; infructescence a monsterocarp; trichosclereids abundant

18. Ovules solitary, placentation basal; fruits each with a solitary large seed . . . Scindapsus 18. Ovules 2–6 or more, placentation parietal; fruits with more than one seed

19. Ovules 8 or more, superposed on 2 (rarely 3) parietal placentas; seeds many, ellipsoid, straight, 1.3–3.2mm long, 0.6–1 mm wide; testa brittle, smooth . . . Rhaphidophora19. Ovules 2–4(–6) at base of a single intrusive placenta; seeds few, curved, 3–7 mm long, 1.5– 4 mm wide;testa bony and ornamented . . . Epipremnum

15. Flowers unisexual . . . . Aroideae (>20)20. Free-floating plants with leaves forming a conspicuous rosette with copious roots hanging beneath . . . . Pistieae(Pistia)20. Not free-floating and otherwise not as above

21. Tuberous-stemmed geophytes; leaf blades mostly variously divided22. Leaf blade decompound; inflorescences usually appearing before leaf emergence, with flowers of both sexes(i.e., monoecious) . . . Thomsonieae (Amorphophallus)22. Leaf blade often trifoliolate to pentafoliolate, rarely hastate or simple; inflorescences produced with leaves

23. Inflorescences usually single-sexed, paradioecious; ripe fruits red . . . Arisaemateae (Arisaema)23. Inflorescences monoecious; ripe fruits never red . . Areae (>24)

24. Spathe internally septate into two chambers or at least the constriction between the lower spatheand spathe limb internally thickened. Australia Lazarum24. Spathe not internally septate

25. Sterile zone between pistillate and staminate zones with smooth naked distal part, base withvarious numbers of staminodes, or whole zone covered with staminodes but then leavesalways entire and proximal staminodes spatulate (in T. flagelliforme) . . Typhonium25. Sterile zone between pistillate and staminate zones fully covered with staminodes, orgrooved and with staminodes only at base; leaves usually pedate, rarely entire; staminodesnever spatulate . . Sauromatum

21. Not geophytes; leaf blade simple, ranging from linear-lanceolate to hastate26. Aquatic plants; infructescences composed of carpels connate into a syncarp . . Cryptocoryneae (>27)

27. Carpels opening from base; leaf ptyxis involute Lagenandra27. Carpels opening from the apex; leaf ptyxis convolute. . Cryptocoryne

26. Plants not aquatic; infructescences comprised of berries28. Plants suffruticose, or with a creeping rhizome-like stem; fruits conspicuous red or pink or green berriesnot surrounded by a persistent spathe . . Aglaonematineae (>29)

29. Colonial helophytes with creeping spongy rhizomes and erect,spongy petioles; fruits large, ripeninggreen and functioning as floating dispersal units . . Aglaodorum29. Terrestrial forest herbs with erect (rarely creeping) firm stems, and erect solid petioles; fruitsmedium-sized, ripening red or rarely pink, not water-dispersed . . . Aglaonema

28. Plants not suffruticose; fruits various, mostly green, if red or orange-red then berries surrounded by apersistent spathe

30. Entire spathe closing after anthesis and then persisting until fruit maturity31. Rhizomatous geophytes; leaves solitary or rarely 2 together, distant from each other;inflorescences arising directly from the ground; spathe forced open by developing fruits but notactively splitting; fruits a large conspicuous red berry containing a large solitary green seed . . . .Nephthytidineae (Nephthytis)31. Clumping mesophytes, rarely colonial helophytes; leaves many together; inflorescencesarising from leafy shoots; spathe actively splitting from the base and peeling upwards to revealfruits; fruits many, whitish to very pale pink, each containing many tiny brown or black seeds . .Philodendrinae (>32)

32. Staminate flower consisting of solitary stamen overtopped by flask-shaped pistillode;ovary 1-locular, placenta basal . . . Furtadoa32. Staminate flower consisting of 2–6 stamens, pistillodes absent; ovary incompletely


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2–5 locular, placentas parietal and axile . . Homalomena30. Upper spathe mostly shedding during or soon after anthesis, and lower spathe persisting until fruitmaturity; if upper spathe persisting then attached portion of petiolar sheath very short and the remainderfree-ligular or absent and the protective role taken on by prophyll/cataphyll

33. Staminate flowers forming synandria . . (Colocasieae [& Caladieae] >34)34. Spathe differentiated into an upper limb and a tubular or convolute lower part separated byone or sometimes two pronounced constrictions

35. Plant with conspicuous erect aerial stolons bearing along their distal portionnumerous barbed bulbils . . Remusatia (incl. Gonatanthus)35. Plant without conspicuous erect aerial stolons with barbed bulbils; if stolons presentthen these decumbent and bearing tubercules at the tips

36. Leaf blades abaxially with waxy glands in the axils of the primary veins;mature infructescences erect; fruit red when ripe, odourless; seeds large, few perfruit . . . Alocasia36. Leaf blade glands absent

37. Seasonally deciduous lithophytes of Karst limestone; stolons simple,decumbent, bearing tubercules at the tips; mature infructescences erect;fruit red when ripe, odourless; seeds large, few per fruit . . . Alocasiahypnosa37. Evergreen plants of a variety of substrates; stolons if presentbranching, spreading. viviparous but without tuberules; matureinfructescences declinate to pendent or erect; fruit pale yellow to brownand fruit-smelling when ripe; seeds small, very numerous per fruit

38. Plants small to medium; leaf blades and petioles at most onlyweakly grey-glaucous; spathe usually yellow; matureinfructescences declinate or pendent; stolons mostly present . . Colocasia38. Plants usually gigantic; leaf blades and petioles pruinose;spathe pure white; mature infructescences erect; stolons absent . .Colocasia gigantea

34. Spathe not differentiated into an upper limb and lower part by one or more constrictions39. Synandria connate; thecae of adjacent synandria encircling pits in the spadix, eachpit with a somewhat prominent upper margin; leaf blade peltate . . . Ariopsis39. Synandria not so; leaf blade peltate or hastate

40. Spathes brightly coloured (internally commonly yellow or purple-red); pistillateflowers with staminodes; stem a repent or suberect epigeal rhizome . . . Steudnera40. Spathe white; pistillate flowers without staminodes; stem a hypogeal tuber orstolon . . . Caladieae (Hapaline)

33. Staminate flowers not forming synandria . . . Schismatoglottideae (>41)41. Modules monophyllous, congested in a distichous arrangement; petiolar sheath with along-persistent, long ligular free portion . . Pichinia41. Modules mostly polyphyllous, if monophyllous then never distichous; petiolar sheath fullyattached, persistent or marcescent; if free ligular then soon-marcescent

42. Wings of petiolar sheath fully or almost fully attached to the petiole; seeds never witha micropylar appendage

43. Inflorescences on very slender peduncles, nodding at anthesis, peduncle atspathe insertion flexing 180˚ from vertical axis; infructescences narrowlycampanulate, nodding; plants of podzols . . . Hestia43. Inflorescences erect to nodding at anthesis, if nodding, then plants massivepachycauls, and peduncle very stout; infructescences fusiform with a constrictedorifice, if campanulate, then thick-walled and erect, never nodding; plants ofvarious substrates but never on podzols

44. Shoot modules hapaxanthic . . Schismatoglottis [Calyptrata Group]44. Shoot modules pleionanthic

45. Petiole sheathing only at extreme base; each foliage leafalternating with a cataphyll . . . Schismatoglottis [TecturataGroup]45. Petiole usually sheathing for at least a third of its length (rarelyless); individual foliage leaves not alternating with cataphylls

46. Pistillate and staminate flower zones separated by aconspicuous partly naked somewhat swollen interstice;inflorescences erect; spathe limb semi-persistent and barelyopening during anthesis and then clasping the spadix, falling


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with spent parts of the spadix; petiolar sheath wings usually(but not always) deciduous . . Apoballis46. Pistillate and staminate flower zones not separated by anaked interstice; spathe not persistent or, if so, theinflorescence nodding; petiolar sheath wings persistent

47. Small to medium plants, often with asperatepetioles and sometime the whole plant coarsely hairy;inflorescence erect; spathe limb irregularly crumblingand breaking away at or after staminate anthesis .. Schismatoglottis [Asperata Group]47. Massive glabrous pachycauls; inflorescencenodding; spathe limb clasping the spadix andmore-or-less marcescent after anthesis, finally fallingwith spent parts of spadix . . Schismatoglottis[Corneri Group]

42. Wings of petiolar sheath always extended into a free ligular portion; seeds often witha micropylar appendage

48. Spathe not constricted; plants glabrous or at most minutely puberulent49. Thecae of anther never with horn- or needle-like projections

50. Spadix free; spathe limb caducous during anthesis;infructescence on erect peduncle; lower spathe forming asplash-cup . . . Piptospatha (>51)

51. Anther connective extended into a pronouncedelongated beak . . . Piptospatha insignis51. Anther connective not so, or if elevated then shortly soand obtuse

52. Spathe white; leaf blade with conspicuouslytessellate tertiary venation on both surfaces . .Piptospatha perakensis52. Spathe pink; leaf blade without tesselate venation,or if present then on very faint and only abaxially . .Piptospatha

50. Spadix partially to almost completely adnate to spathe; spathelimb not caducous during anthesis; infructescence on declinatepeduncle; lower spathe not forming a splash-cup

53. Staminate flowers mostly sterile with a narrow zone offertile flowers exposed by the spathe opening; spathepersistent into fruiting, at fruit maturity swiftly drying, reflexingand opening basally by tearing at peduncle insertion toexpose fruits but remaining distally convolute and while inthis situation clasping the spadix; ovules basal; seeds with ablunt micropyle . . Bakoa53. Staminate flowers all fertile; spathe persistent throughoutthe fruit dispersal; ovules parietal; seeds with a pronounced,hooked, micropylar appendage . . . Ooia

49. Thecae of anther each with a horn- or needle-like projection, althoughsometimes visible only after pistillate anthesis

54. Thecae with needle-like projection extending only after pistillateanthesis; projection tipped with a weakly peltate ovate-triangularflap; appendix composed of pistillodes . . . Schottariella54. Thecae with a horn- or needle-like projection present prior topistillate anthesis; with the projection pointed and never associatedwith a terminal flap; appendix, where present, composed ofstaminodes

55. Sterile interstice of spadix with flattened scale-likestaminodes; anthers not excavated . . . Bucephalandra55. Sterile interstice absent, or with truncate staminodes;anthers mostly with the top excavated

56. Anthers clearly excavated . . . Aridarum56. Anthers not excavated

57. Thecae horns short, robust, their basesoccupying the whole upper surface of theanther; stamens and staminodes smooth .


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. Aridarum incavatum57. Thecae horns very slender; stamens andstaminodes coarsely verriculate . . Aridarumrostratum

48. Spathe mostly constricted58. Thecae of anther each with horn- or needle-like projections; ovules onbasal placenta; seeds with a long, hooked micropylar appendage . . . Phymatarum58. Thecae of anther without horn- or needle-like projections; ovules onparietal or basal placenta; seeds without a micropylar appendage

59. Placentation basal . . . . Schottarum59. Placentation parietal . . . Schismatoglottis [Multiflora Group]


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Figure 30. Homalomena symplocarpifoliaP.C.Boyce, S.Y.Wong & Fasih. A recentlydescribed species from central Sarawak.Homalomena is by far the largest Indomalayangenus of Araceae, with probably in excess of 350species, the majority yet to be formally described.

Figure 31. HomalomenasymplocarpifoliaP.C.Boyce, S.Y.Wong &Fasih. Many Homalomenaare pollinated by largebeetles of the Scarabidae,however, H.symplocarpifolia and itsrelative appear to haveevolved pollination utilizingmuch smaller beetlesbelonging to the Nitidulidae,seen here on the spathelimb. Just visible behind thespadix is a Chrysomelidaebeetle; chrysomelids arefrequently pollen-robbers inAraceae, often chewingthrough the unopenedspathe to reach the spadix.

Figure 32. Remusatiapumila (D.Don) H.Li &A.Hay. Detail of thebulbil-bearing stolon.Each scale is a minuteleaf, the lower portionequivalent to the petiolarsheath, while the ‘hook’is a petiole and leafblade analogue. Image© John Mood, used withpermission.

Figure 33. Alocasia sp. Portion of the abaxialsurface of the leaf blade showing the genericallydiagnostic waxy glands.

Figure 34. Alocasiahypnosa J.T.Yin,Y.H.Wang & Z.F.Xu.Growing in habitat onKarst limestone inThailand. The colonyhas formed primarilythrough the extensiveproduction of stolons.Image © David Prehsler,used with permission.

Figure 35. Alocasiahypnosa J.T.Yin, Y.H.Wang& Z.F.Xu. Although currentlyplaced in Alocasia, thisspecies differs on a numberof key characters, includinglacking waxy glands on theabaxial surface of the leafblade. The inflation of thespathe limb is also notmatched in Alocasia.

Figure 36. Colocasiaoresbia A.Hay. The onlyunequivocally indigenousColocasia in Malesia. Herephotographed in the hillsabove Nanga Gaat, centralSarawak.

Figure 37. Colocasiagigantea Blume. Anultimately enormous plantwith glistening whitespathes carried in a fan.The peduncles remain erectinto fruiting, in contrast toother Colocasia species inwhich the peduncle isdeclinate. Note the pruinosepetiole and peduncles.


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Figure 38. Ariopsis protanthera N.E.Br. A genus oftwo species. One restricted to the Western Ghats of SWIndia and one (depicted here) occurring from NE India toNE Thailand. Image © Rachun Pooma, used withpermission.

Figure 39. Steudneradiscolor W.Bull. One ofperhaps nine speciesoccurring from NE India toSW China and south intoIndochina. Image © WilbertL.A.Hetterscheid, used withpermission.

Figure 40. Hapaline benthamiana Schott.Another small genus, with most species locallyrestricted and seemingly also rare. One exceptionis H. benthamiana, which occurs through much ofThailand on exposed limestone.

Figure 41. Apoballismutata (Hook.f.)S.Y.Wong & P.C.Boyce.Long included inSchismatoglottis despitesome strikingmorphologicaldifferences, recentmolecular andpalynological researchhas shown it to be highlydistinct. The spadix isshown here (with thespathe artificiallyremoved) to reveal theinflated, almost nakedinterstice separating thepistillate (below, palegreen) and staminate(above, salmon-orange)flower zones.

Figure 42. Pichinia disticha S.Y.Wong &P.C.Boyce. Endemic to an isolated limestonemassif in SW Sarawak, Pichinia has a uniquevegetative morphology in the tribeSchismatoglottideae; with each leaf in the fanrepresenting a single branch (similar morphologyoccurs in some Anthurium and Philodendron).Although reminiscent of Schismatoglottis MultifloraGroup (see Fig. 46) Pichinia falls outsideSchismatoglottis in molecular analyses, althoughits exact position is yet to be established.

Figure 43. Hestialongifolia (Ridl.) S.Y.Wong& P.C.Boyce. A recentlyrecognized segregate fromSchismatoglottis, Hestia isreadily distinguished by thefascicles of very slender-pedunculate inflorescenceswith a sharply noddingspathe.

Figure 44. Schismatoglottis viridissima A.Hay.A member of the Schismatoglottis CalyptrataGroup, and typically for this group formingcolonies by the production of numerous stolons.

Figure 45. An undescribedspecies in theSchismatoglottisTecturata Group. Thisphoto shows the very shortpetiolar sheath. Theprotective role of thepetiolar sheath is taken overby the large prophyll,remains of which aremarked by the dark brown,papery tissue at the base ofthe petiole.


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Figure 46. Schismatoglottis nicolsonii A.Hay,a typical member of the Schismatoglottis MultifloraGroup. The free-ligular portion of the petiolesheaths are clearly visible.

Figure 47.Schismatoglottis asperataEngl., photographed inhabitat on Matang,Sarawak, exemplifies someof the critical morphologiesof the SchismatoglottisAsperata Group. The fruitingspathe is held erect, withthe thick-walled persistentlower part half constricted atthe mouth. Note, too, theasperate petioles and (justvisible) the short, open,petiolar sheath.

Figure 48. Schismatoglottis corneri A.Hay, thesole member of the Schismatoglottis CorneriGroup, showing the diagnostic inflorescences.Plants can attain over 2 m in height. Image ©Kohei (Takenaka) Takano, used with permission.

Figure 49. Piptospathaviridistigma P.C.Boyce,S.Y.Wong & Bogner.The nodding pinkinflorescences and erectsplash-cup arediagnostic for the genus.

Figure 50. Piptospathaperakensis (Engl.) Ridl.A highly distinctivespecies, the onlyPiptospatha with a whitespathe, and a powerfullyfragrant inflorescence.Molecular analysisclearly separates P.perakensis from the‘core’ Piptospatha asrepresented by P.viridistigma (Fig. 49).

Figure 51. Bakoabrevipedunculata(H.Okada & Y.Mori)S.Y.Wong. The seconddescribed species of thisBornean-endemic genusdemonstrates well thedifficulties thatsometimes bestbotanists. Originallydescribed in 2000 in thegenus Hottarum, it wasmoved in the same yearto Piptospatha, althoughfitting neither genusconvincingly. Latercombined molecular andmorphological analysesrevealed it to fall outsideof both and, togetherwith Hottarum (laterPiptospatha) lucens,supported therecognition of a newgenus: Bakoa.

Figure 52. Ooia kinabaluensis (Bogner)S.Y.Wong & P.C.Boyce. Another species that hasbeen bounced from genus to genus before, withthe help of combined molecular and morphologicalanalyses, being found a taxonomically convincing‘home’. Image © Kohei (Takenaka) Takano, usedwith permission.

Figure 53. Schottariellamirifica P.C.Boyce &S.Y.Wong. One ofseveral newlyrecognized genera of theSchismatoglottideaeoccurring almostexclusively on Borneo.The damage to thespathe limb interior iscaused by the pollinator,a chrysomelid beetle(just visible behind thespadix).

Figure 54. Bucephalandramotleyana Schott.Inflorescence at pistillateanthesis (spathe limb andthe nearside part of thelower spathe artificiallyremoved). The shield-likestaminodes above thepistillate flowers are erect.


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Figure 55. Bucephalandramotleyana Schott.Inflorescence at staminateanthesis (also with thespathe limb and thenearside part of the lowerspathe artificially removed).Note that the shield-likestaminodes above thepistillate flowers are nowlowered, closing off thelower spathe (which ispersistent throughoutfruiting) and protecting thedeveloping infructescence.

Figure 56. Phymatarumborneense M.Hotta.Inflorescence at pistillateanthesis. The numeroussmall flies (Drosophilidae)are attracted by a powerfulesteric (model airplaneglue) smell.

Figure 57. Phymatarumborneense M.Hotta. Detailof the spadix showing thetuberculate staminateflowers with their need-likethecae horns, and theappendix comprised ofsimilarly tuberculatestaminodes.

Figure 58. Aridarum montanum Ridl. This plantwell matches the only known specimen of the typespecies, collected once, in 1905, and never sincerefound; confirmation of its identity throughflowering is eagerly awaited. Image © Mike Lo,used with permission.

Figure 59. Aridarumrostratum Bogner &A.Hay. One of severalrelatively recently describedAridarum speciesrecognition of which hasconsiderably stretched theboundaries of the genus. Itseems probable that as withPiptospatha molecularanalysis will result inchanges to the genericboundaries of Aridarum.

Figure 60. Schottarumsarikeense (Bogner &M.Hotta) P.C.Boyce &S.Y.Wong. Although verysimilar in overallappearance toSchismatoglottis of theMultiflora Group (see Fig.46), Schottarum is in factmuch more closely relatedto Aridarum.


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A QUARTERLY PUBLICATION FOR MEMBERS OF THE INTERNATIONAL AROID

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