CYCADACEAE UPDATED (1999)
 D. J. DE LAUBENFELS

2. SUBGENUS TRUNCATA
Cycas L. subgenus Truncata de Laub., Blumea 44 (1998) 367.  Type species: Cycas rumphii Miq. 

Trunk cylindrical.  Petioles with or without thorns.  Pinnules straight or curving towards the base of the leaf, flat or slightly revolute, 5 mm or more wide, midrib generally more prominent below.  Microsporophyll truncate (abruptly narrowed) beyond the fertile part so that it becomes more or less peltate, the flat end being diamond shaped with a short to long apical spine sharply bent nearly perpendicular to the microsporophyll and pointing towards the apex of the cone.  Megasporophyll with the apical portion with undulate and entire or more often dentate margins, the teeth more or less obscure basally becoming more prominent towards the apex, less than 5 mm long and usually much less (rarely slightly more), acuminate with the acumen often an elongated spike. 

    Distribution -- Six species from southern Thailand through the Philippines and Indonesia to New Guinea, Fiji, and various Pacific islands as well as outposts in northern Australia. 

    Habitat -- Consistently wet equatorial regions, mostly in the understory of rainforest, but one species in open grassy situations. 

    Notes -- The species of this section are all very closely related to one another, differing in minor ways.  The majority have a spongy layer inside the stony layer of the seed which suggests that the whole subgenus may have had its origin from a plant much like thouarsii but which had a truncate rather than an elongated apical part of the microsporophyll.  Thus several widespread species retain the strand habitat with buoyant seeds while others have moved inland sometimes loosing the buoyant seed character.  One species, schumanniana, grows in grasslands within the rainforest zone and resembles in some ways nearby species of the type section and may spring from some kind of genetic interaction with them. 

KEY TO THE SPECIES

9. Cycas rumphii Miquel, Bull. Sc. Phys. et Nat. Néer. 2 (1839) 45; Comment Phytogr. (1840) 120; Monogr. Cycad. (1842) 29; Linnea 17 (1843) 688 (in part); Tydschr. Wis. en Natuurk. Wet. 2 (1849) 286 (in part); Anelecta Bot. Indica 2 (1851) 32 (in part); Linnea 25 (1852) 589, t. 2 (in part); Prodr. Cycad. (1861) 7 (in part); Archives Néer. Sci. Exact. et Nat. 3 (1868) 231 (in part); De Candolle, Prodr. 16 (2) (1868) 527 (in part); Warburg, Monsunia 1 (1900) 181, t. 7f; Stapf, Kew Bull. (1916) 7 (in part); Schuster in Engler, Pflanzenr. 99 (1932) 74 (in part); Hill, Aust. Syst. Bot. 7 (1994) 549, f. 6.  Holotype: Rumphius in Herb. Amboin., I (1741) 86, t. 22--3 (Olus Calappoides or “Sajor Calappa”), Amboina.

C. glauca Miq., Comment. Phytogr. (1840) 127; Monogr. Cycad. (1842) 30; Linnea 17 (1843) 692; Lamarck ex Hort. Brit. (1830) 403, nomen; hort.  Zamia corsoniana G. Don, Gard. Mag. 18 (1842) 371 (Schuster, 1932, gives this as C. corsoniana D. Don in syn. with rumphii).  -- Type: Corson s.n., “South Seas” (holo LINN, not seen). 

C. circinalis (auct.) non L., Jones, Cycads of the World (1993) photos on p. 137. 

C. bougainvilleana Hill, Aust. Syst. Bot. 7 (1994) 557, f. 11.  -- Type: Sayers NGF 19739, Tonalu Harbour, Papua New Guinea (holo NSW, iso K). 

Palm-like trees to 12 m high, trunk diam. 20--30 (40) cm, often branched.  Leaves 150--310 cm long including a petiole of 30--65 cm with thorns for most of its length, 95--160 pinnules on each side of the rachis.  Pinnules 18--30 cm x (12) 14--18 mm, margins slightly bent, midrib more prominent below, apex usually distinctly acuminate, dark green and shiny above, lighter below.  Cataphyll 6--8 x 2.5 cm, pungent.  Pollen cone ovoid, tapering slightly, 35--45 x 14--20 cm, with an up to 13 cm peduncle; microsporophyll 15--20 mm wide, the apical part with a narrow c5 mm acumen.  Megasporophyll with 2--10 ovules; apical part 30--60 x 20--30 mm, triangular and acuminate, marginal teeth 8--16 per side, 2--3 mm basally and 4--5 mm towards the apex, acumen a spine 20--25 mm long.  Seed 43--60 x 32--44 mm, the apical part of the stony layer with a prominent crest not visible before the fleshy covering dries, buoyant due to an inner spongy layer, orange. 

    Distribution -- Near the coast from the Moluccas and Sulawesi, along the northern coast of New Guinea through the Solomon Islands as well as outposts in the Marshall and Caroline Islands and on Christmas Island (south of Java).  Specimens studied: LAE 52094, NGF 15538, Anderson 3719, Atasrip 60, Barclay 3540, Berry 79, Carroll 80, Darbyshire & Hoogland 7855, Docters v. Leeuwen 1531, Fosberg 26192, 32009, 36632, 46237, Gibbs 6278, Gillison NGF 25380, Guppy 356, Kajewski 1619, Hill s.n., Kanehira in 1937, Maire s.n., Mosely 88, Powell 412, Ridsdale & Katik NGF 38078, Robinson 563, Sands 3025, Sayer NGF 19739, Streimann LAE 53828, Vandenburg & Mann NGF 42232, Van Royen 3300, Vogel 3136, Whitmore BSIP 1347, 3932, Womersley NGF 24825, Yen BSIP 19795. 

    Habitat -- Mostly in the strand vegetation in rainforests environments.  Although clearly spread at least locally by flotation, it is known to be spread by people and this probably explains the more remote outposts of its distribution. 

    Notes -- The confusion with other taxa started by Miquel himself has meant that most purported descriptions of rumphii are mixed with that of other species or are completely of something else.  The ecology of rumphii corresponds with that of thouarsii and edentata but rumphii can be distinguished by the crested seed and the very short apical spine on the microsporophyll, as well as the wide acuminate pinnules.  Specimens with pinnules 20 and 21 mm wide, distinctly wider than any other Cycas have been collected in Palau.  Kanehira described a “forma palauica” from there (Jour. Jap. Bot. 14, 1938, 33, t. 4F, 5F) without a type specimen, based on a globular seed with a sharp (but low) apical ridge.  He did not mention the pinnules at all and it remains to be seen if his form corresponds to the wide pinnules and if a distinct taxon is involved.  Hill distinguished bougainvilleana on the basis of irregular or indistinct lateral teeth (spines) on the apex of the megasporophyll and he also found an interrupted upper hypodermis in the pinnules.  The presence or absence of a small gap in the upper hypodermis in a few samples is probably unreliable as a character and hardly the basis for a species.  Hill admits that the teeth are clearly visible on the type (plate 22 of Rumphius which he reproduces) so how he concludes that no more than irregular teeth characterize rumphii is not clear.  In fact, experience with these cycads shows that teeth tend to erode away during the more than a year that it takes the attached seed to mature.  Furthermore, different examples from the same collection show variations in surviving teeth.  Thus, three of the four female specimens cited by Hill show clearly developed teeth on the examples I have examined.  Miquel described glauca strictly from living plants cultivated at Potsdam and imported from Java by H. Roterod where it undoubtedly also was cultivated, there being no wild plants there which fit the description.  In fact, no wild plants with a similar form are known to be glaucous, a character favored by horticulturalists.  The form described best fits rumphii, which could easily have been carried to Java from Amboina.  Without fertile material, which was not available for this taxon to Miquel, not much more can be said.  Note that Miquel reported that immature pinnules of glauca had undulate margins.  In 1868 Miquel listed glauca as of uncertain status in that no wild material known to him was glaucous. 

10. Cycas scratchleyana Mueller, Victorian Naturalist 2 (1885) 18; Hill, Austr. Syst. Bot. 7 (1994) 562, f. 13.  -- C. circinalis L. var. scratchleyana (Muell.) Schuster in Engler, Pflanzenr. 99 (1932) 70. -- Type: Armit in 1883, Mt. Bedford, Terr. New Guinea (holo MEL, iso K).

C. apoa Hill, Aust. Syst. Bot. 7 (1944) 553, f. 9.  -- Type: Iwanggin BW 5245, Lake Sentani, Irian Jaya (holo CANB, iso A, L). 

Palm-like trees to 10 m high, trunk diam. 15--30 or more cm, rarely branched.  Leaves 150--270 cm long including a petiole of 33--73 cm with thorns for most of its length, 100--160 pinnules on each side of the rachis.  Pinnules 15--32 cm x (9) 10--15 mm, margins slightly bent and sometimes undulate, midrib more prominent below and sometimes yellow above, apex sometimes slightly acuminate, shiny dark green above, brighter green below.  Cataphyll 7--9 x 2,5 cm.  Pollen cone ovoid, 20--45 (67) x 10--13 cm, with a short peduncle; microsporophyll 14--20 mm wide, apical part with a narrow 4--8 mm acumen.  Megasporophyll with 4--8 ovules; apical part 30--50 x 20-35 mm, triangular and acuminate, marginal teeth 14--20 per side, 2--3 mm basally and 4--5 mm towards the apex, acumen a 5--15 mm spine.  Seed 38--60 x 30--44 mm, the apex of the stony layer with a weak crest, buoyant due to a spongy layer under the stony layer, yellow to yellow-orange. 

    Distribution -- From the Moluccas across New Guinea as far as New Ireland.Specimens studied: LAE 52546, Armit in 1883, Brass 994, 5643, 6752, Clemens 751, 1277, 8104, Craven & Schodde 750, 933, Croft et al LAE 61174, 68785A, Darbyshire 907, de Haan 1798, Docters v. Leewen 9623, Floyd 6423, Forman & Vinas LAE 60256, Frodin UPNG 4031, Hinds in 1841, Iwanggin BW 5243, Kanehira 3950, Larivita & Mann NGF 70598, Lauterbach 475, 1345, McGruder in 1889, Ridsdale & Katik NGF 36761, Schiefenhoevel 142, Van Royen 4790, 5297, Versteeg 1470, Warburg 21129, Womersley NGF 46475. 

    Habitat -- Rainforest environments, especially along streams to 900 m or occasionally to 1500 m.

    Notes -- The two species, scratchleyana and celebica form a closely related group with rumphii with which they share the form of the microsporophyll, a spongy layer inside the seed, and a crest on the stony part of the seed, although the crest is generally weaker than for rumphii which also has a longer spine at the apex of the megasporophyll and distinctly wider pinnules.  The pinnules of scratchleyana are further scarcely acuminate and the seed somewhat smaller, while celebica has few or no spines on the petioles.  In habitat by contrast scratchleyana appears to be strictly a forest species where it appears to be less subject to human inter-island dispersal even though use as human food does occur.  The retention of the buoyant seed character clearly illustrates its relationships.  Hill distinguishes apoa on the basis of irregular or indistinct lateral teeth (spines) on the apical part of the megasporophyll.  Two sheets of the type collection which I have seen have well developed teeth (see note concerning teeth under rumphii). 

11. Cycas celebica Miquel, Bull. Sc. Phys. et Nat. Néer. 2 (1839) 45; Comment. Phytogr. (1840) 126; Monogr. Cycad. (1842) 31; Linnea 17 (1843) 701; Tydschr. Wis. en Natuurk. Wet. 2 (1849) 288; Prodr. Cycad. (1861) 7; De Candolle, Prodr. 16 (2) (1868) 528.  -- Holotype: Rumphius in Herb. Amboin., I (1741) 87, t. 20--21 (Olus Calappoides), Sulawesi (Celebes).

C. circinalis (auct.) non L., Seeman, Fl. Viti. (1865) 268; Schumann & Lauterb., Fl. Deutch. Schutzgeb. Südsee (1901) 153; Yuncker, Bul. Bishop Mus. 120 (1959) 45; Stone, Am. Sci. 59 (1971) 313, f. 9; de Laub., Flore Nouv. Caléd. et Dépend. 4 (1972) 8, t. 1; Fosberg & Sachet, Smithsonian Contrib. Bot. 1, 20 (1975) 6.

C. seemannii Braun, Sitzungsber. Ges. Naturf. Freunde Berlin (1876) ll4; Carruthers, Jour. Bot. 31 (1893) 2, t. 330 (in part); Hill, Austr. Syst. Bot. 7 (1994) 560, f. 12.   -- C. circinalis L. subsp. seemanii (Braun) Schuster in Engler, Pflanzenr. 99 (1932) 71; Smith, Flora Viti. Nova I (1979) 90.  -- C. rumphii Miq. var. seemannii (Braun) Parham, Agr. Jour. Dept. Agr. Fiji 19 (1948) 94, f. 7.  -- Type: Seemann 572, Fiji (lecto K, Smith, Flora Viti. Nova I, 1979, 90, iso BM, GH, P).

C. neocaledonica Linden, Illust. Hortic. 28 (1881) 32, nomen.

C. micronesica Hill, Austr. Syst. Bot. 7 (1994) 554, f. 10. -- Type: Turner in 1992, Anderson Air Force Base, Guam (holo NSW, not seen).

Palm-like trees to 15 m high, trunk diam. 14--30 cm, often branched.  Leaves 100--300 cm long including a petiole of 25--60 cm with few or no thorns, 65--150 pinnules on each side of the rachis.  Pinnules 15--32 cm x 10--16 mm, margins somewhat bent, midrib more prominent below, the wider examples acuminate, green above, paler below.  Cataphyll 8--11 x 2--3 cm.  Pollen cone ovoid with a slight taper, 32--67 x 8--15 cm, with a 7--10 cm peduncle; microsporophyll 13--18 mm wide, the apical part with a narrow 4--5 mm acumen.  Megasporophyll with (2) 4--6 ovules; apical part 35--60 x 25--40 mm, triangular or elongated and acuminate, marginal teeth 7--14 per side, 2--3 mm basally and 4--5 (7) mm towards the apex, occasionally more, acumen a 7--30 mm spine, occasionally with a few reduced basal teeth.  Seed 55--70 x 40--60 mm, the apical part of the stony layer with a small crest, buoyant due to an inner spongy layer, tawny orange. 

    Distribution -- From the Marianas south to Borneo and the Moluccas and from Bougainville  to Tonga.  Specimens studied: Aet 704, Aston S17927, Baker 199, 301, Baumann 13896, Bernier 12029, Brass 2852, Brongiart in 1864, Bryan 1240, Clemens 11277, 21175, Degener 15233, 15262, 15286, Degener & Ordonez 14262, Evans 2148, 2149, Fosberg 25300, 31830, 32644, 46256, Fosberg & Evans 46228, 46237, Foster RLP 78-37, Greenwood 644, Gressitt 21, Guam Experiment Sta. 406, Guppy 133, Halle RSNH 6362, 6363, Hodel et al 514, 551, Horne 309, 316, Hurlimann 272, Kajewski 400, Kanehira in 1932, 233, 1189, 2408, 3743, McKee 4038, 7884, 10048, McPaul 13198, Moran 4375, Morrison in 1896, Neker R-87, 95, Nooteboom 5269, Pancher s.n., Parham 137, Parks 16336, 16373, Pleyte 978, Rinehart 16143, Rodin 503, 563, Seemann 572, Sirate’e BSIP 9635, Smith 4486, Steere 16, 17, 19, Stone 4110, Takamatsu 1849, Thompson 406, Thurston in 1882, Waterhouse 356B, Whiting C25, 320, Yuncker 15485. 

    Habitat -- In rainforest environments along the seacoast and inland, particularly on limestone, up to 360 m elevation.

    Notes -- Particularly close to rumphii because of the strand habitat (in part) and the buoyant seed but differing in the narrower pinnules and thornless petioles in particular.  Both Rumphius and Miquel were rather uncertain of the identity of this taxon but both emphasized the lack of thorns, which certainly distinguishes celebica.  The somewhat disjunct distribution including many islands where the plant is emphatically cultivated, suggests, as for rumphii, that its distribution has been extended by human introductions.  Hill, who is content to leave celebica in synonymy with rumphii, supplies micronesica for the plants from the Marianas and distinguishes it from seemannii by an elliptical (rather than triangular) apical part of the megasporophyll and fewer lateral teeth (spines), I have e examined a wide range of material from the Marianas, New Caledonia, Fiji, and Tonga and find both extremes in all four areas.  The circinalis described by Safford (1905) from the Marianas appears to refer to cultivated material of that species as it differs quite sharply from any known wild specimens collected there. 

12. Cycas silvestris Hill, Telopea 5 (1992) 181, t. 1 (figures 1d--e taken from the type). -- Type:  Clarkson & Neldener 8813, Cook Dist., Queensland (holo NSW, not seen).

C. circinalis (auct.) non L., Roxburgh, Fl. Ind. 3 (1832) 744 (excl. pl. masc.); Blanco, Fl. Filip. (1837) 745; Foxworthy, Philip. Jour. Sci. 6 (1911) Bot., 151 (in part); Smitinand, Nat. Hist. Bull. Siam Soc. 24 (1971) 169 (in part); Flora Thailand (1972) 186 (in part); Amoroso Philip. Jour. Sci. 115 (1986) Bot., 182, t. 5, 7--9. 27--30.

C. circinalis L. var. curranii Schuster in Engler, Pflanzenr. 99 (1932) 69.  -- C. sp. Foxworthy, Philip. Jour. Sci. 6 (1911) Bot., 152, t. 27.  -- Type: Curran 3842, Palawan (holo PNH, destroyed, lecto K, here designated, iso P). 

C. rumphii (auct.) non Miq., Merrill, Sp. Blancoanae (1918) 52; Enum. Philip. Flower. Plants 1 (1923) 1 (in part). 

C. media (auct.) non R. Br., Gardner, For. Dept. Bull. Perth 32 (1923) 30.

C. pranburiensis (no Latin description) Tang et al., Cycads of Thailand (1997)  28, figs.

Palm-like trees to 4.8 m high, trunk diam. 20--25 cm, occasionally branching.  Leaves 110--210 cm long including a petiole of 24--60 cm with thorns at least half way to the base, 70--110 (120) pinnules on each side of the rachis.  Pinnules 17--35 cm x 9--15 mm, margins slightly bent, midrib more prominent below, green above, lighter below.  Cataphyll 6--8 x 2--3.5 cm, not pungent.  Pollen cone ovoid and tapering, 28--50 x 8.5--15 cm, with a c3.5 cm peduncle; microsporophyll 15--22 mm wide, the apical part with a spine 12--23 mm long (longest towards the center of the cone).  Megasporophyll with 4--8 ovules; apical part 25--35 x 20--23 mm, triangular and acuminate, marginal teeth 7--12 per side, 2--3 mm basally and up to 5 mm towards the apex, acumen a subulate spine 25--30 mm or longer.  Seed 42--55 x 25--38 (50) mm, yellowish. 

    Distribution -- From Viet Nam and southern Thailand through the Philippines to Palau and south to Java, the Sunda islands, and one or two outposts on the northern coast of Australia.  Specimens studied: Backer 50, Borssun Waalkes 523, Cardona 23870, Chai SAN 29392, Chai et al S38561, Clemens 3972, Conklin 18682, Curran 3842, 7381, 7513, Docters v. Leeuwen 1881, Edaño 76373. Eyma 3727, Fosberg 32376, Gaudichaud in 1836, 97, Gressitt 21, Jaag 667, 893, Kondo & Edaño 36768, Lee S41866, Merrill 855, 3257, Meyer 2577, Murthey 9576, Noerkas 481, Podzorski SMHI 2119, Pulle in 1906, Mamos 3281, Ramos & Edaño 48953, Whitford 1235, 

    Habitat -- Lowland rainforest sometimes along the coast. 

    Notes -- Hill described this species form Queensland, well beyond the main area of distribution, based on female material only.  The microsporophyll is identical with that of edentata whose megasporophyll lacks teeth and whose seed has a spongy layer inside the stony layer.  Both these species and riuminiana (chamberlainii) are thoroughly mixed up in the literature, as in Merril’s 1923 paper or in his collection numbered 855 meant to illustrate Blanco’s species.  The variety curranii was given for an unusually small pollen cone but this may be no more than the lower limit of natural size variation. 

13. Cycas edentata de Laub., Blumea 44 (1998) 373, f. 1.  Type: Kondo & Edaño 38877, Mt. Cabucan, Sulu Archipelago (holo L, iso A, BM). 

C. rumphii Miq. var. timorensis Miq., Comment. Phytogr. (1840) 125.  -- Type: Serchrusult s.n, Timor (holo P). 

C. rumphii (auct.) non Miq., Dyer in Hooker f., Flor. Brit. India 5 (1888) 657; Ridley, Fl. Malay Pen. (1925) 186; Smitinand, Nat. Hist. Bull. Siam Soc. 24 (1971) 173. t. 3b, 4d, plate 26; Flora Thailand (1972) 186; Pant, Cycas & the Cycadales, ed. 2 (1973) 36, t. 13; Amoroso, Philipp. Jour. Sci. 115 (1986) 182, t. 2, 6, 17--19, 23--26.

C. javana (auct.) non (Miq.) de Laub., Cycads in China (1996) 65. 

C. literalis (no Latin description) Tang et al., Cycads of Tailand (1997) 25, figs.

Palm-like trees to 10 m high, trunk diam. 15--45 cm, base sometimes enlarged up to 100 cm, occasionally branching.  Leaves 100--260 cm long including a petiole of 45--90 cm with thorns for most of its length, 60--108 or more pinnules on each side of the rachis.  Pinnules 15--34 (38) cm x 10--17 (18) mm, margins slightly bent, midrib more prominent below, dark green above, lighter below.  Cataphyll 6--8 x 2--3 cm, not pungent.  Pollen cone ovoid and tapering, 30--59 x 11--17 cm, with a 4--6 cm peduncle; microsporophyll 14--25 mm wide, the apical part with a spine 12--24 mm long.  Megasporophyll with 2--8 ovules; apical part 20--30 x 18--23 mm, triangular or almost diamond-shaped, acuminate, the margins toothless and straight or undulating but sometimes with a few very weak teeth, the acumen a spike 30--40 (46) mm long.  Seed 42--60 x 38--50 mm, buoyant due to a spongy layer inside the stony layer, orange. 

    Distribution -- From the Andaman Islands through peninsular Thailand and the Philippines south to Indonesia and Timor.  Specimens studied: Abbe et al 9684, Backer 27471, Bartlett 13509, Buysmann 16, Cabiling 3707, Charoenphol et al 4064, Cockburn FRI 7565, Congdon 42, Conklin 37914, Copeland in 1905, Curran 10319, DeVriese & Teijsmann s.n., Fenix 15660, Fernando 7181, Forman 176, Gibbs 4337, Henderson 20338, 29160, Hoogerwerf 235, Hutchinson 3441 Iboet 237, Keith 11355, 12768, 16087, 16569, Klemme 15219, Kondo 32862, Kondo & Edaño 38877, Koorders 16648, Korthals s,n., Kurz s.n., Lörzing 12206, 16881, Lutjeharms 4691, Matiwang 253, Merrill 11638, Mondi 136, no name 16345, Prain in 1889, Ramos 80613, Ramos & Edaño 48310, Raap 7, Ram 3806, Ridley 7143, Robinson in 1916, Serchrusult s.n., Smitinand 12242, Steven & Justina SAN 86704, Stolk s.n., Stone & Anderson SAN 86714, Weber 3, Wilkes s.n., Williams 2890, 3058, Wyatt-Smith KEP 93185. 

    Habitat -- In strand vegetation in rainforest environments rarely far from the coast. 

    Notes -- In spite of the buoyant seed, its distribution suggests long distance dispersal by human introduction in part, inasmuch as this species, like its relatives, is widely used as a food crop.  Sterile plants strongly resemble thouarsii in form and habitat, albeit somewhat less robust. 

14. Cycas schumanniana Lauterbach in Schumann & Lauterbach, Fl. Deutsch. Schutzgeb. Sudsee (1901) 154; Hill, Aust. Syst. Bot. 7 (1994) 536, f. 3B, 8.  -- Type: Lauterbach 2745, Bismarck Mts. (holo B, destroyed).

C. campestris Hill (in part), Aust. Syst. Bot. 7 (1994) f. 9a--e, g--h.

Palm-like trees to 2.5 m, trunk diam. 20--25 cm, occasionally branched.  Leaves 75--150 cm long including a petiole of 24--55 cm with several thorns or none, 50--105 pinnules on each side of the rachis.  Pinnules 9--22 cm x 5--8 (9) mm, margins slightly bent, midrib more prominent below, shiny dark green above, paler below.  Cataphyll 5--8 x 2--2.5 cm, not pungent.  Pollen cone ovoid, 17--20 x 8--10 cm (probably immature), with a 2.5--4 cm peduncle; microsporophyll up to 20 mm wide, apical part with a narrow acumen c3 mm long.  Megasporophyll with 2--6 ovules; apical part 25--40 x 19--29 mm, triangular and acuminate, marginal teeth 10--20 per side, 1--2 mm near the base and 2--4 mm towards the apex, acumen 10--15 mm long with a few reduced teeth basally.  Seed 28--42 x 23--34 mm, yellow to orange. 

    Distribution -- Along either side of the major mountain chain in eastern Papua-New Guinea.  Specimens studied: Anderson MVNG-1, Brass 877, Carr 11236, 11555, Clemens 10751, 11369, Darbyshire 662, Henty & Lelean NGF 49272, 49273, Hoogland 5120, Katik LAE 70805, Lam in 1954, McGregor in 1889, Pullen 956, Sands 1864, Saunders 583, Schodde 2582, Streimann NGF 27783, 39487, Vandenberg NGF 42149, Vaughan in 1900, White et al NGF 1626, Womersley & Hoogland 5158, Womesley & Simmonds 5091. 

    Habitat -- Open grassland form sea level to 1350 m. 

    Notes -- Generally identified by collectors as papuana or media, both of which, however, have elongated tapering apices to their microsporophylls with only the tip bent towards the apex of the cone, not a blunt apex with a sharply bent spike.  The former has generally narrower pinnules and the latter generally longer apical parts to the megasporophyll.  They all have similar habitats, contrasting with the other New Guinea species, which also have wider pinnules and a spike for an acumen on the megasporophyll.  One could indeed say that schumanniana is distinctly intermediate between media and scratchleyana and there is a clear possibility of some sort of hybrid origin.  A neotype collected in the Bismarck Mts. is needed. 

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