CYCADACEAE UPDATED (1999)
 D. J. DE LAUBENFELS

3.  SUBGENUS PECTINATA
Cycas L. subgenus Pectinata de Laub., Blumea 44 (1998) 375.  Type: Cycas pectinata Hamilton. 

Trunk cylindrical.  Petioles more than 25 cm long, usually armed with thorns.  Pinnules straight or more commonly turning basally, narrowly acute, flat or slightly bent on the margins, 6--17 mm wide.  Microsporophyll with an acuminate to blunt apical part where the upper margin including the acumen is bent sharply towards the apex of the pollen cone.  Megasporophyll with the margin of the expanded apical part lacinate or pectinate, the comb-teeth at least 6 mm and usually more than one cm long, usually with the longest in the middle or all close to the same size, the apex acuminate with the acumen from narrowly lanceolate to an elongated spike, either usually with a few reduced teeth basally. 

    Distribution -- Seven species from eastern India and southern China through southeastern Asia to Luzon, Indonesia, and New Guinea. 

    Habitat -- In the understory or in open disturbed areas in rainforests or seasonal forests. 

    Notes -- The comb-teeth of the megasporophyll are obviously homologous with foliar pinnules and are probably more primitive than are the short marginal teeth in subgenera Cycas and Truncata, which otherwise strongly resemble this subgenus and could well have been derived from it.  Subgenus Pectinata more nearly occupies a central (and perhaps ancestral) area for the genus and is more diverse than the other subgenera.

 KEY TO THE SPECIES

15. Cycas pectinata Hamilton, Mem. Wern. Soc. 5, 2 (1826) 322; Hill, Taxon 44 (1995) 26; Wang et al, Cycads in China (1996) 97, pl. 19; Tang et al, Cycads of Thailand (1997) 10, figs.  -- Type: Hooker & Thomson s.n., Chittagong, E. Bengal (neotype K).

C. pectinata Griff. (non Hamilton), Notul. 4 (1854) 10, t. 360, f. 3; Dyer in Hooker, Fl. Brit. Ind. 5 (1888) 657; Leandri, Fl. Indo- Chine (1931) 1090; Smitinand, Nat. Hist. Bull. Siam Soc. 24 (1971) 169, f. 3d, 4a; Flor. Thai. (1972) 190, f. 13d, 14a; Pant, Cycas & the Cycadales, ed. 2 (1973) 37; Jones, Cycads of the World (1993) 151.  -- C. circinalis L. var. pectinata (Griff.) Schuster in Engler, Pflanzenr. 99 (1932) 68.  -- Holotype: the figure given by Griff.

C. jenkinsiana Griff., Notul. 4 (1854) 9, t. 360, f. 1--2, t. 362, f. 1.  -- Type: Jenkins s.n., Assam (holo BM).

C. dilatata Griff., Notul. 4 (1854) 15.  No type given (see notes). 

C. siamensis (auct.) non Miq., Dyer in Hooker, Fl. Brit. Ind. 5 (1888) 657; Ridley, Fl. Mal. Pen. 5 (1925) 285; Leandri, Fl. Indo-Chine (1931) 1088 (in part); Smitinand, Nat. Hist. Bull. Siam Soc. 24 (1971) 173 (in part), f. 3a, 4b; Flor. Thai. (1972) 190 (in part), f. 13a, 14b.

Palm-like trees to 15 m high, trunk diam. c14 cm, expanded base to 100 cm diam., sometimes branched.  Leaves 120--240 cm long including a petiole of 25--54 cm with thorns a t least half way down, 80--140 pinnules on each side of the rachis.  Pinnules 13--25 cm x 6--9 mm, margins distinctly bent, midrib prominent, dark green above, lighter below.  Cataphyll 4--10 x 1.5--2cm, not pungent.  Pollen cone cylindric, tapering, 35--69 x 11.5--15 cm or more, with a 6 cm peduncle; microsporophyll 17--23 mm wide, the apical part narrowed rather abruptly to an acuminate spike bent sharply upward, 3--4 mm wide at the base and 10--40 mm long.  Megasporophyll with 2--6 ovules; apical part 30--50 x 25--45 mm, triangular or somewhat rounded and acuminate, 12--18 (24) comb-teeth along each margin 12--25 mm long, acumen linear to lanceolate and 24--40 or more x 4--8 mm long usually with a few reduced basal teeth.  Seed 30--44 x 28--36 mm, yellow to orange. 

    Distribution -- From eastern Nepal and the southernmost edge of China across most of southeastern Asia.  Specimens studied: Abbe & Smitinand 9361A--B, Dickason 7226, 7623, Gamble 325B, 2675A, Gammie s.n., Geesink & Phengkhlai 6157, Grierson & Long 2357, Haines 3983, Henry 12637, Hooker & Thomson s.n. (Sikkim, Khasia, Chittagong), 595, Jenkins s.n., Kerr 8858, Kingdon-Ward 21772, Kurz 2322, 2622, Meebold 8166, Nicholson 3078, Parry 100, Stainton 5733, 6638, Wallich 8587D, Watt 6796. 

    Habitat -- In the understory of moist evergreen forests on clay, mostly above 600 m and to 1300 m. 

    Notes -- The incomplete notes of Griffith were published posthumously.  He certainly did not intend pectinata to be a new species as he did not append his name and no specimen can be associated with his figure.  Because no reference to Hamilton’s species of the same name was included, Griffith’s pectinata becomes a synonym and the species is most often attributed to him, quite inappropriately.  In fact, Griffith described jenkinsiana which is also a synonym.  Further, he had obviously laid the name dilatata aside in favor of jenkinsiana, but it was published anyway, creating yet another synonym.  Hamilton cited angulata with a question mark and also thought his species might be what Rumphius had in mind for Olus calappoides, neither of which proved correct.  Hamilton specified east Bengal for his species but no specimen. The early collection by Hooker & Thomson from that area can be taken as the neotype.  Later, Epicycas siamensis has been thoroughly confused with pectinata in the literature.  Smitinand, for example, (1971, 1972) illustrated pectinata fertile structures under siamensis while Leandri’s (1931) figures under pectinata are mostly siamemsis. 

16.  Cycas taiwaniana Carruth., Jour. Bot. 31 (1893) 2, t. 331; Wang et al, Cycads in China (1996) 127, pl. 28 & f. 17; .  -- C. revoluta Thunb. var. taiwaniana (Carruth.) Schuster in Engler, Pflanzenr. 99 (1932) 84.  -- Type: Swinhoe14119, Taiwan (holo BM, iso K).

C. hainanensis C. J. Chan ex C. Y. Cheng, W. C. Cheng, & L. K. Fu, Acta Phytotax. Sin. 13 (1975) 82; S. H. Fu et al, Flora Rep. Pop. Sin. 7 (1978) 16, f. 2, 5--6 (taken from the type); Jones, Cycads of the World (1993) 143.  --Type: Zhong 4706, Hainan (holo IBSC, not seen).

C. fairylakea D. Y. Wang in Wang et al, Cycads in China (1996) 54, pl. 7 & f. 2.  Type: Wang & Peng H2978, S China Bot. Gard. (holo SZG, not seen). 

C. parvulus S. L. Yang iin Wang et al, Cycads in China (1996) 93, pl. 18 & f. 12.  Type: Yang 317, Yunnan, Honghe co., Mengdian (holo HWA, not seen). 

C. multifida Chang & Zhong, Acta Sci. Nat. Univ. Sunyatseni 36 (1997) 70,  f.1 (5).  Type: Zhong 80196, Guangxi, Xiling (holo SYS, not seen). 

Palm-like trees to 3.5 m high, trunk diam. 13--35 cm.  Leaves 65--200 cm long including a petiole of 20--40 cm with thorns for most of its length, 66--170 pinnules on each side of the rachis.  Pinnules 12--32 cm x 6--12 mm, margins flat, midvein prominent above, sometimes weaker below, shining dark green above, paler below.  Cataphyll 6--13 x 1.5--2.5 cm, not pungent.  Pollen cone 30--70 x 9--13 mm with a 5 cm peduncle; microsporophyll 12--18 mm wide, the apical part narrowing abruptly to a triangular c4 mm beak on the upturned sometimes serrulate margin.  Megasporophyll with 2--4 ovules; the apical part 35--60 x 25--40 mm, triangular to rounded and acuminate, 10--24 comb-teeth along each margin 12--25 mm long, acumen triangular to lanceolate, 34--50 x 25--35 mm,  with reduced teeth or serrations along the entire length.  Seed 35--45 x 30--36 mm, yellow, becoming irregularly rugose, the stony layer verricose. 

    Distribution -- From Hainan, along the nearby coast, and apparently formerly on the southwest coast of Taiwan.  Specimens studied: Merrill 18752, Phillips in 1883, Swinhoe in 1867, 14119. 

    Habitat -- Scattered and endangered in moist forested locations from near the sea to 1200 m. 

    Notes -- Long confused with another species known from the interior of Taiwan recently named taitungensis.  The authors of this new species argue for an origin of the type material of taiwaniana from the mainland due to the fact that it is not known today from Taiwan and that Swinhoe did collect extensively on the mainland.  Although they quoted evidence from Dyer (1903), they ignored the comments quoted therein of Watters (who served under Swinhoe) that he had “had several conversations about a Cycas which grew abundantly on Saracen’s Head at the entrance of the harbour. That was in 1865, and since then the Chinese have altered the flora of the place very much by cutting away everything for the erection of a fort and the construction of roads.”  (original in K).  This location is on the southwestern coast of Taiwan in an area where Swinhoe is known to have collected.  These authors further included material of more than one species in their description of taiwaniana.  Schuster made Carruther’s species a variety of revoluta based on “living” (cultivated) material which was in fact taitungensis material, a close relative of revoluta.  By contrast, taiwaniana is very similar to pectinata which differs in the long apical spike on the microsporophyll and the smooth not vericose surface to the stony layer of the seed.  Hill & Chen (1994) distinguished taiwaniana from hainanensis by pinnules more than 13 mm wide.  However the type of taiwaniana has pinnules less than 12 mm wide.  Apparently Hill & Chen have included material of szechuanensis in their description of taiwaniana.  The same confusion occurs in "Cycads in China."  The acumen of the megasporophyll of taiwaniana is unusual in that it is normally booth wide and it bears numerous reduced lateral teeth or serrations.  When these are counted, as in multifida, an abnormally large number is achieved. 

17. Cycas szechuanensis Chen & Fu, Acta Phytotax. Sin. 13 (1975) 81; S. H. Fu et al, Flora Rep. Pop. Sin. 7 (1978) 12, f. 1: 7--8 (taken from the type); Wang et al, Cycads in China (1996) 121, pl. 26.  -- Type: Xiong et al 33221, Sichuan (holo PE, not seen). 

C. rumphii (auct.) non Miq., Acad. Sinica, Flora Rep. Pop. Sin. 7 (1978),16 (excl. pl. fem. ), f. 3: 4, 6--8. 

C. guizhouensis Lan & Zou, Acta Phytotax. Sin. 21 (1983) 209, f. 1; Wang et al, Cycads in China (1996) 57, pl. 8 & f. 3.  -- Type: Lan & Zou 81-8-0001, Xingyi, Guizhou (holo PE, photo seen). 

C. ferruginea Wei, Guihaia 15 (1994) 300, f. 1-6.  -- Type: -- Wei 1986, cult. Guangxi (holo IBK, not seen). 

C. diannanensis Guan & Tao, Sichun Forestry Survey & Design (1995) f. 1-7; Wang et al, Cycads in China (1996) 49, pl. 5.  -- Type: Tao 95014, Yunnan, Gejiu, Manhao (holo SFEDI, not seen).

C. pectinata Griff. subsp. manhaoensis Chen & Yun, Acta Bot. Yunnan. 17 (1995) 400.  -- Type: Min-chen & Po 22, Gejiu, Manhao (holo KUN, not seen). 

C. multiovula D. Y. Wang in Wang et al, Cycads in China (1996) 83, pl. 15 & f. 10.  -- Type: Wang 5574, cult. Gejiu, Yunnan (holo SCG, not seen). 

C. longlinensis Chang & Zhong, Acta Sci. Nat. Univ. Sunyatseni 36 (1997) 68, f. 8.  -- Type: Zhong 80848, Guangxi, Longlin, Jin Zhong Shan (holo SYS, not seen). 

C. xilingensis Chang & Zhong, Acta Sci. Nat. Univ. Sunyatseni 36 (1997) 69, f. 6-7.  -- Type: Zhong 80866, Guangxi, Xiling (holo SYS, not seen).                                                                                                              

Palm-like trees to 5 m high, trunk diam. 30--40 (50) cm.  Leaves 140--300 cm long including a petiole of 40--70 cm with thorns for most of its length, 67--138 pinnules on each side of the rachis.  Pinnules 17--40 cm x 11--16 mm, margins slightly bent, midrib prominent above, sometimes weaker below, dark green above, lighter below.  Cataphyll 5--7 x 1,5--2.5, pungent.  Pollen cone Cylindric-ovate, to 53 x 11 cm; microsporophyll 12--17 mm wide, the apical part narrowing abruptly to a small triangular 1--3 mm beak on the upturned margin.  Megasporophyll with 2--8 (14) ovules; apical part 40--50 x 20--40 mm, triangular or rounded and acuminate, 8--23 comb-teeth along each margin 20--45 mm long, acumen 30--45 x 7--17 mm usually with reduced teeth basally.  Seed 30--36 x 38 mm (wider then long), yellow to reddish. 

    Distribution -- From southern Sichuan Prov. to the south coast of China.  Specimens studied: Esquirol in 1909, Hu 7059, McClure 13315, Rock 2654, Wang 2737, 76684, Wang & Liu 274l, 2743, Wang & Peng 2673, 2977. 

Habitat -- In the understory of forested and in grassy areas to 1800 m. 

Notes -- The long comb-teeth resemble those of several species in subgenus Revoluta but the pinnules are much broader with a prominent upper midrib.  The megasporophyll of the type appears abnormal with its multiple apex and overlapping seeds and comb-teeth.  The type is known from a cultivated plant that exists in clones in several gardens.  Without the abnormalities, it corresponds morphologically with guizhouensis, which was published later.  Some forms appear to have narrower pinnules than do others. 

18. Cycas campestris Hill, Aust. Syst. Bot. 7 (1994) 538, f. 3c, 9f (figures taken from the type).  -- Type: Jacobs & Conn 5950, Lea Lea Lakes, Cent. Dist., Papua-New Guinea (holo NSW, photo seen).

Palm-like trees to 8 m high, trunk diam. 15--30 cm.  Leaves 180--250 cm long including a petiole of 33--69 cm with a few to several thorns, 140--180 pinnules on each side of the rachis.  Pinnules 18--35 cm x 9--13 mm, margins distinctly but not strongly bent, midvein prominent, green above, lighter below.  Pollen cone ovoid, 22 x 5.5 cm, with a 2--3 cm peduncle; microsporophyll to 20 mm wide, the apical part blunt with an upturned edge.  Megasporophyll with 4--8 ovules; the apical part 30--40 x 25--35 mm, triangular and acuminate, 8--15 short comb-teeth along each margin 6--8 mm long, acumen poorly developed to 35 mm long with reduced teeth basally.  Seed 38--48 x 30--38 mm, red. 

    Distribution -- Most of the length of the island of New Guinea.  Specimens studied: NGF 34119, Brass 23830, Eyma 5451, Gideon LAE 76964, Hoogland 4750, Streimann NGF 28556, Van Royen 4269, 4769, Womersley & Van Royen 5872. 

    Habitat -- Scattered in disturbed areas of lowland rainforest, often along streams to 300 m. 

    Notes -- Only the type collection in the original publication corresponds to this species, the remainder of the material cited represents perfectly typical schumaniana.  Both this latter and scratchleyana which are also found in New Guinea have shorter teeth on the apical part of the megasporophyll which are also smaller towards the base, not uniform in size, and microsporophylls with distinct apical spikes, while their seeds do not become red.  The seed of scratchleyana has a distinct ridge on the apical part of the stony layer and the pinnules of schumanniana are much narrower.  The relatively short comb-teeth of campestris are transitional in size to those of subgenera Cycas and Truncata. 

19. Cycas javana (Miq.) de Laub. in Wang et al, Cycads in China (1996) 65; Blumea 44 (1998) 381, t. 2.  -- C. circinalis L. var. javana Miq., Monogr. Cycad. (1842) 28, t. 2f, E.  -- Type: Blume 1089, Java (holo L).

C. rumphii (auct.) non Miq., Schuster in Engler, Pflanzenr. 99 (1932) 74, f. 11G; Backer & Bakhuizen v. d. Brink, Fl. Java 1 (1963) 87.

Palm-like trees to 6 m high, trunk diam. unknown.  Leaves 120--200 cm long including a petiole of 45--50 cm with thorns for most of its length, c90 or more pinnules on each side of the rachis.  Pinnules 23--36 cm x 11--14 (15) mm, margins slightly bent, midrib prominent, green above, lighter below.  Pollen cone ovoid, 30--70 x 12--17 cm, with a short peduncle; microsporophyll c15 mm wide, the apical part acuminate and extending 5--7 mm and then sharply bent upward in a beak 10--15 mm long.  Megasporophyll with 4--8 ovules; apical part 30--50 x 16--19 mm, expanding gradually and then lanceolate, 6--9 comb-teeth on each margin c4--6 mm basally and 6--9 mm towards the apex, a 35--50 mm apical spike with several reduced teeth basally.  Seed 45--60 x 35--50 mm, orange. 

    Distribution -- Java and nearby parts of Sumatra including Krakatoa.  Specimens studied: Backer in 1908, in 1918, 1476, 37088, Blume 1089, Borbey s.n., Burck 1904, DeVriese 11153, 11154, Horsfield 80, Junghuhn s.n., 1079, Koorders 961, 1563, 1564, 12727, 20106, 24998, Korthals 961, Kuripan s.n., Lang in 1927, Limbergang s.n., Luibungan s.n., Pottinga s.n., Pulla in 1906, Schouta in 1903, Stolk s.n., Van Steenis 3162, Zollinger 1161. 

    Habitat -- Understory of lowland rainforest. 

    Notes -- Sterile material strongly resembles sylvestris and edentata, both of which occur in or near Java, but the fertile material is quite different.   In "Cycads in China" it is actually edentata which is described. 

20. Cycas macrocarpa Griff., Notul. 4 (1854) 11, t. 362, f. II (see Ridley, 1925, 285: a “forest form”).  -- Type: Griffith 6361, Crew, near Tabong, Malaya (holo K).

C. circinalis (auct.) non L., Smitinand, Nat. Hist. Bull. Siam Soc. 24 (1971) 169 (in part); Flora Thai. (1972) 186 (in part). 

Palm-like trees to 9 m tall, trunk diam. 13--39 cm.  Leaves 100--250 cm long including a petiole of 40--68 cm with thorns up to half way to the base, c120 pinnules on each side of the rachis.  Pinnules 19--39 cm x 12--17 mm, flat, midrib prominent, often acuminate, glossy dark green.  Cataphyll 4--7 x 1.5--3 cm, pungent.  Pollen cone cylindric, 33-35 x 7.6 cm, with a short peduncle; microsporophyll c12 mm wide, apical part triangular or acuminate and extending c10 mm and then sharply bent upward with a 7--8 mm beak.  Megasporophyll with 2--8 ovules; apical part c20 x 15--30 mm, triangular to nearly blunt and acuminate, 6--7 comb-teeth along each margin 8--12 mm long, acumen a 25--35 mm spike.  Seed 50--75 x 35--45 mm, yellow. 

    Distribution -- Peninsular Thailand and peninsular Malaysia.  Specimens studied: Corner 30294, Field 2960, Griffith 6361, Kerr 10868, 10972, King 4871, Larsen 30984, Larsen et al 1263, Meijer & Yong KEP 94932, Ng FRI 5581, no name 15699, Ridley s.n., Sangkhachand & Smitinand 234, Sinclair 8554, Sinclair & Kiah SF 40769, Wallich 8587C, Whitmore FRI 4232. 

    Habitat -- Understory of lowland rainforest and on limestone hillsides. 

21. Cycas riuminiana Porte ex Regel, Gartenfl. 12 (1863) 16, f. on p. 17.  -- C. circinalis L. subsp. riuminiana (Porte) Schuster in Engler, Pflanzenr. 99 (1932) 68.  -- Type: Regel 6210, hort. (lecto K, here designated).

C. chamberlainii Brown & Kienholz, Philipp. Jour. Sci.  26 (1925) Bot., 47, t. 1--2; Zamora & Co, Guide Philipp. Flora & Fauna 2 (1986) 219; Amoroso, Philipp. Jour. Sci. 115 (1986 181, f. 4, 14--6, 39--41; Jones, Cycads of the World (1993) 136.  -- Type: Brown & Kienholz 42539, Mt. Arayat, Luzon (holo US, iso A, BM, K, NY, P).

Palm-like trees to 8 m high, trunk diam. 15--25 cm.  Leaves 115--220 cm long including a petiole of 26--57 cm with thorns at least half way to the base, 80--150 pinnules on each side of the rachis.  Pinnules 18--30 cm x 7--11 (12) mm, margins slightly bent, midrib prominent, green above, lighter below.  Cataphyll 6 x 1--1.5 cm, pungent.  Pollen cone ovoid, c45 x 12 cm, with a c 15 mm peduncle; microsporophyll 16--20 mm wide, apical part abruptly rounded with a lanceolate 12--20 mm beak spreading but strongly curved upward.  Megasporophyll with 4--8 ovules; apical part 15--20 x 20--30 mm 5(wider than long), rounded to nearly blunt, 4--10 comb teeth along each margin 8--18 mm long, an apical spike 2--4 cm long.  Seed 39--50 x 29--36 mm, yellowish. 

    Distribution -- Luzon to Sulawesi and the Moluccas.  Specimens studied: Balgooy 3930, Brown & Kienholz 42539, Buwalda 4622, Hennipman 6113, Loher 4153, 4844, 7133, Madulid et al 7232, McGregor 42080, Merrill 9795, Prawiroatmodjo & Soewoko 1757, Ramos 22391, Ramos & Edaño 48912, Vogel 5304, 5734, Worcester in 1907. 

    Habitat -- Understory of lowland rainforest or in disturbed areas and often on limestone, to 1030 m. 

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