CYCADACEAE UPDATED (1999)
 D. J. DE LAUBENFELS

EPICYCAS
Epicycas de Laub. Blumea 44 (1998) 388.  Type species: Cycas micholitzii Dyer [= Epicycas micholitzii (Dyer) de Laub.].

Cycas L. section Indosinenses Schuster in Engler, Pflanzenr. 99 (1932) 65.  -- Type: C. siamensis Miq. 

Cycas L. section Stangerioides Smitinand, Nat. Hist. Bull. Siam Soc. 24 (1971) 168.  -- Type: C. micholitzii Dyer.

Palm-like plants with a mostly underground bulbous base, one or more leafy apices on the surface of the bulb, each eventually developing a short usually slender trunk or a branching underground bulb, generally reaching the fertile stage before any trunk forms.  Leaves with numerous bilateral pinnules and with a well-developed petiole usually with numerous thorns.  Pinnules with flat and often undulate margins, linear, narrowed sharply at the base to produce a short petiolule or sessile, dichotomous in some species and then sufficiently dispersed so that adjacent pinnules do not overlap, otherwise spaced as in Cycas.  Pollen cone cylindrical and often tapering, with a short scaly peduncle; microsporophyll wedge-shaped, the apical part rounded and truncate with the margin bent sharply towards the apex of the cone and bearing a small apical spur or acuminate with an apical spur bent towards the apex of the cone.  Megasporophyll with several ovules, the apical part lacinate with the comb-teeth often dichotomous even when the pinnules are not.  Ovules 1--4 (10). 

     Distribution -- Seven species from the southern margins of China across southeastern Asia into northernmost Malaysia. 

     Habitat -- Understory of moist forests or more often in open disturbed areas. 

     Notes -- Most closely related to the genus Cycas but differing in the bulbous underground base with relatively slender short trunks or a branching bulb and, where present, by the dichotomous divisions of the pinnules and the comb-teeth of the megasporophylls.  No other cycads of any kind produce a combination of a slender stem emerging from a bulbous base, although bulbs without stems and all degrees of subterranean and semi-subterranean stems occur in various genera of other families of cycads.  Distinctly divided into two groups.  The type group has the entire width of the truncated apex of the microsporophyll sharply bent towards the apex of the cone the margin of which is serrate and bears a short acumen, the underground bulb is elongated, and the stem (when present) only a few cm in diam., while the petiole is very long with thorns the whole way.  The other group has the apex of the microsporophyll is sharply tapering with a narrow acumen sharply bent towards the apex of the cone, the underground bulb is globular and sometimes huge, while the trunk is like that of Cycas, the petiole tends to be rather short, and the number of thorns is few.  The former microsporophyll resembles those of subgenera Pectinata and Revoluta of the genus Cycas while the latter microsporophyll resembles those of subgenus Cycas.  One species of subgenus Pectinata, macrocarpa, has microsporophylls of the latter type (but no bulb).  It may be that the two groups in Epicycas are not closely related. 

KEY TO THE SPECIES

1. Epicycas micholitzii (Dyer) de Laub., Blumea 44 (1998) 389.  --  C. micholitzii Dyer, Gard. Chron. 38 (1905) 142, f. 48--49; Fedde Rep. Spec. Nov. 4 (1907) 171; Prain, Bot. Mag. 135 (1909) t. 8242; anon., Kew Bull. (1910) 163; (1912) 301, t.; Leandri, Flore Indo-Chine (1931) 1091; Schuster in Engler, Pflanzenr. 99 (1932) 81; Smitinand, Nat. Hist. Bull. Siam Soc. 24 (1971) 169, f. 1; Jones, Cycads of the World (1993) 148; Wang et al, Cycads in China (1996) 71, pl. 12 & f. 7.  -- Type: Micholitz s.n., hort. (holo K).

C. rumphii Miq. var. bifida Dyer, Jour. Linn. Soc. 26 (1903) 560.  -- Type: Morse 273, Lungchow, China (holo K). 

C. multifrondis D. Y. Wang in Wang et al, Cycads in China (1996) 80, pl. 14 & f. 9.  -- Type: Wang 5024, cult. (holo SZG, not seen).

A subterranean or protruding bulb 10--30 cm in diam. from which eventually grows a trunk 4--12 cm in diam. and to 60 cm tall.  Leaves solitary of few, 190--350 cm long including a petiole of 60--270 cm with widely spaced thorns along its length, 10--44 pinnules along each side of the extremely tomentose rachis 4--12 cm apart, apex with several pinnules merging at their bases rather like a single terminal dichotomous pinnule.  Pinnules dispersed, 4--12 cm apart, two to three times dichotomous, less so towards the apex and on juvenile leaves, the segments 20--30 cm x 20--25 mm, margins flat and slightly undulate, midrib prominent on both sides, narrowed at the base to a 2--7 mm petiolule which participates in the first dichotomy, acuminate, dark green above and paler below, glaucous at first.  Cataphyll 3--6 x 1.5--4 cm, not pungent.  Pollen cone cylindrical and tapering to both ends, to 35--42 x 5--8 cm, with a 3--12 cm peduncle; microsporophyll 12--17 mm wide, the apical part rounded truncate and sharply bent with a 1--5 mm acumen, the edge slightly serrulate.  Megasporophyll with 1--6 (8) ovules; apical part c26 x 26 mm, rounded, with 5--19 comb-teeth along each margin 12--40 mm long and sometimes dichotomous, the apex with a spike 4--7 cm long with one or two reduced teeth near the base.  Seed 26--31 x 23--28 mm, yellow. 

     Distribution -- Northeastern Viet-Nam near the China border and northeastward into China.  Specimens studied: Chevalier 37888, Condomines 16, Dournes in 1967, Micholitz s.n., Morse 273, Schmid 872, Van Du s.n. 

     Habitat -- Scattered in rainforest environments. 

     Note -- When the bulbous base protrudes, the trunk looks like a narrow-necked vase. 

2. Epicycas multipinnata (Chen & Yang) de Laub., Blumea 44 (1998) 391.  -- C. multipinnata Chen & Yang, Acta Phytotax. Sin. 32 (1994) 239; Wang et al, Cycads in China (1996) 86, pl. 16 & f. 11.  -- Type: Yang 9202, S. Jianshui Co., Yunnan (holo PE, photo seen).

C. longpetiolula D.Y. Wang in Wang et al, Cycads in China (1996) 68, pl. 11 & f. 6.  -- Type: Yang & Peng 5523, Yunnan, Pingbian, Yuanjiang valley (holo SZG, not seen).

A subterranean or protruding bulb 10--25 cm in diam.from which eventually grows a trunk c3 cm in diam. and to 40 cm tall.  Leaves one or two (3) at a time, 300--700 cm long including an 80--400 cm petiole with thorns along the entire length, twice pinnate with 6--11 (25) primary pinnae on either side of a rachis 25--90 cm long, longest in the middle, the longer pinnae with 7--11 dispersed pinnules, apices with several pinnules merging at their bases rather like a single dichotomous terminal pinnule.  Pinnules non-symmetrically spaced at 6--20 cm, two or five times dichotomous, less so towards the apex of the leaf, 20--40 cm long, the segments 8--12 cm x 10--25 mm and acuminate, narrowed at the base  to a 15--25 mm (up to 8 cm) petiolule which participates in the first dichotomy, margins flat and undulate, midrib prominent on both sides, dark green above, lighter below.  Cataphyll 8--10 x 2.5--3.5 cm, not pungent.  Pollen cone fusiform cylindric 36 x 6 cm with a 4 cm peduncle; microsporophyll 15--18 mm wide, the blunt upturned edge slightly serrulate, with a 2--4 mm acumen.  Megasporophyll with 2--8 ovules; apical part c30 x 30 mm, triangular, with c15 comb-teeth along each margin 20--35 mm long, the apex with a spike hardly longer than the comb-teeth.  Seed c35 x 30 mm, yellow. 

     Distribution -- Southern Yunnan near the Viet-Nam border.  Cultivated plants studied. 

     Habitat -- Somewhat open forest on limestone. 

3. Epicycas tonkinensis (Lind. & Rod.) de Laub., Blumea 44 (1998) 391.  -- Zamia tonkinensis Lind. & Rod., Ill. Hort. 32 (1885) 27, t. 547.  -- Cycas tonkinensis (Lind. & Rod.) Lind. & Rod., Ill. Hort. 33 (1886) 27.  -- Holotype: the plate (547).

Cycas bellefontii Lind. & Rod., Ill. Hort. 33 (1886) 27, t. 586.  -- Holotype: the plate (546). 

C. balansae Warb., Monsunia 1 (1900) 179.  -- C. siamensis Miq. subsp. balansae (Warb.) Schuster in Engler, Pflanzenr. 99 (1932) 68.  -- Type: Balansa 4084, Tonkin, near Hanoi (holo P, iso K).

C. chevalieri Leandri, Flore Indo-Chine (1931) 1092.  -- Type: Fleury 32612, N. Annam, Nghe-An Prov.  (holo P).

C. siamensis (auct.) non Miq., S. H. Fu et al, Flore Rep. Pop. Sin. (1978) 11, t. 3: 1--3.

C. simplicpinna (Smitinand) Hill, Proc. 3rd Intern. Conf. Cycad Biol. (1995) 39; Wang et al, Cycads in China (1996) 117, pl. 25 & f. 16; Tang et al, Cycads of Thailand (1997) 33. -- C. micholitzii Dyer var. simplicipinna Smitinand, Nat. Hist. Bull. Siam Soc. 24 (1971) 164, f. 2, 3e, 4f; Flora Thai. (1972) 192, f. 13e, 14f, 15; Jones, Cycads of the World (1993) 149. -- Type: Smitinand 4757, Doi Suthep, N. Thailand (lecto BFK, here designated, not seen, f. 2a--b taken from the type).

C. segmentifida Wang & Deng, Encephalartos 43 (1995) 11; Wang et al, Cycads in China (1996) 108, pl. 22.  --Type: Wang & Peng 2967, Guangzhou (holo SZG, not seen).

C. tanqingii D. Y. Wang in Wang et al, Cycads in China (1996) 134, pl. 29 & f. 18.  -- Type: Wang 5538, Yunnan (holo SZG, not seen).

C. shiwandashanica Chang & Zhong, Acta Sci. Nat. Univ. Sunyatseni 36 (1997) 67, f.1-4.  -- Type: Zhong 88015, Guangxi, Fangcheng, Shiwanda Shan (holo SYS, not seen). Wang et al, Cycads in China (1996) 111, pl. 23 & f. 14 (nomen).

A subterranean or protruding bulb 15--30 (50) cm in diam. from which may eventually grow a trunk to 100 (200) cm, trunk diam. c 10 cm.  Leaves few in each whorl, 65--250 (330) cm long including a petiole of 25--105 (164)  cm with thorns for most of its length, 18--81 (149)  pinnules on each side of the rachis.  Pinnules 17--40 (45) cm x 13--28 mm, margins flat and sometimes undulate, midrib prominent above, weaker below, narrowed at the base to a short petiolule or sessile, weakly to distinctly acuminate, glaucous at first, shiny dark green above paler below.  Cataphyll 4--9 x 1.5--3 cm, not pungent.  Pollen cone cylindrical and tapering to both ends, 30--60 x 7--12 cm, peduncle 3--5 cm long; microsporophyll c17 mm wide, apical part rounded and truncate with a distinct 1--3 mm acumen on the upturned slightly serrulate margin.  Megasporophyll with 1--4 (6) ovules, apical part c35 x 15 mm, oval, with 6--19 comb--teeth on each margin 20--35 mm long and often dichotomous, an apical spike 20--40 x 4--9 mm sometimes with basal reduced teeth.  Seed 25--40 x 23--35 mm, yellow. 

     Distribution -- From southern China and northern Thailand south through Viet-Nam.  Specimens studied: Abbe & Smitinand 9473, Balansa 563, 4084, Bon 2706=4146, Cavalerie 7106, de Loureiro s.n., Fleury 326122, Ford s.n., Kerr 3206, 5347, 5645, Nicholson 1688, Pierre 5517, Rock 2500, Smitinand & Abbe 6349, Sorenson et al 1320, Wang 75868, 78922, Wang & Peng 2964, Wilson 5420, Winit 1686, Yates s.n. 

     Habitat -- Understory of lowland rainforest. 

     Notes -- The original descriptions of tonkinensis and bellefontii were made from young plants shipped from Tonkin.  The authors distinguished the first by undulate pinnule margins from the latter.  The illustration of tonkinensis shows a barrel-shaped base narrowed below.  It is almost certain that this was naturally mostly underground, the bulbous base which is associated with leaves with such pinnules.  In fact, the pinnules of tonkinensis are given as a not very precise 2--3 cm wide, a little large for the species as it is generally known but nothing actually above 25 mm wide has ever been collected, and this should be accepted as exaggeration due to imprecision.  It is not at all unusual for the wider pinnules in this species to be undulate.  The growth form and wide pinnules resemble micholitzii without the pinnule dichotomy.

4. Epicycas miquelii (Warburg) de Laub., Blumea 44 (1998)  393.  -- C. miquelii Warburg, Monsunia 1 (1900) 179; Wang et al, Cycads in China (1996) 76, pl. 13 & f. 8.  --Type: Warburg s.n., S. China (holo B, destroyed).  -- Neotype: Kerr 11873, S. Tailand, K. Taco, Chumphon (holo K).

Cycas inermis Lour., Fl. Cochinch. (1790) 632, illeg. (given in synonymy with C. revoluta Thunb.).  -- C. revoluta Thunb. var. inermis (Lour.) Miq., Prodr. Cycad. (1861) 16.  -- C. siamensis Miq. subsp. inermis (Lour.) Schuster in Engler, Pflanzenr. 99 (1932) 80.

C. chamaoensis (no Latin description) Tang et al, Cycads of Tailand (1997) 11, figs.

C. clivicola (no Latin description) Tang et al, Cycads of Tailand (1997) 14, figs. 

C. tansacha (no Latin description) Tang et al, Cycads of Tailand (1997) 21, figs.   

A subterranean or slightly protruding bulb to 100 cm in diam. from which eventually grows a slender trunk to 8 m, sometimes several trunks, often branched.  Leaves 50--190 cm long including a 15--55 cm petiole with few or no thorns, 40--100 pairs of pinnules.  Pinnules 15--30 (48) cm x 6--12 (13) mm, margins slightly bent and sometimes undulate, midvein prominent on both sides, dark green above and paler below.  Cataphyll 4.5--8.5 x 1.5--2.5 cm, not pungent.  Pollen cone ovoid, 25--63 x 8--15 cm, peduncle unknown; microsporophyll c12 mm wide, apical part sharply narrowed with a c10 mm apical spur bent towards the apex of t'he cone.  Megasporophyll with 2--5 ovules; the apical part 45 x 40 mm, triangular and acuminate, 10--16 comb-teeth along each margin 15--25 mm long, sometimes forked, acumen 22--40 x 3--7 mm with a few reduced basal teeth.  Seed 18--22 x 18--25 mm, yellow to orange red. 

     Distribution -- Southern China, across Viet-Nam, and in peninsular Thailand.  Specimens studied:  Curtis 2424, 2427, Henderson SFN 23826, Kerr 11873, 14058, 16079 Kingdon-Ward SFN 37538, Machado in 1903, Ogata 10293, Scortechini 913,  Whitmore FRI 12993, Yang et al 649, 650.  

     Habitat -- Low elevation limestone. 

     Notes -- Although Cycas inermis is not a legitimate name, the description and nearby location fit Epicycas miquelii.  A Laureiro specimen in BM has been taken for the type of inermis (and therefore for the variety and the subspecies) but has pinnules 32 cm long while Laureiro specified 15 cm for his species.  The specimens from Saraburi (tansacha) have a distinctly wooly apex.  Specimens of miquelii from eastern Thailand and nearby Vietnam have often been taken to be siamensis or Cycas pectinata. 

5. Epicycas elongata (Leandri) S. L. Yang in de Laub. & Adema, Blumea 44 (1998) 393.  -- Cycas pectinata Griff. var. elongata Leandri, Flore Indo-Chine (1931) 1091; C. elongata (Leandri) D. Y. Wang in Wang et al, Cycads in China (1996) 51.  -- Type: Poilane9328, Ca Na, Annam (holo P). 

A large subterranean or slightly protruding bulbous base from which eventually grows a trunk up to 5m, diam. c15 cm.  Leaves 75--165 cm long including a petiole of 15--50 cm with thorns for at least half the distance to the base, 51--92 pinnules on each side of the rachis.  Pinnules 12--21 cm x 6--10 mm, midrib prominent, green above, paler below.  Cataphyll 3--5 x 1.5--2 cm, not pungent.  Pollen cone shape unknown, c11.5 cm in diam., peduncle unknown; microsporophyll c20 mm wide, the apical part erect and triangular, 15 mm long including an upturned acumen.  Megasporophyll with 1--4 ovules; apical part 25--45 x 20--30 mm, triangular and acuminate, 8--10 comb-teeth along each margin 12--20 mm long, the acumen 30--60 mm long usually with a few reduced basal teeth.  Seed 24--44 x 23--30 mm, color unknown. 

     Distribution -- Mostly near the coast from central Viet-Nam through Cochin China. Specimens studied: Abbe & Smitinand 9573,, Kiep s.n., Pierre 1722, 5516, s.n., Poilane 2746, 4656, 9095, 9328, Robinson 1304, Schmidt s.n., Thorel 374.

     Habitat -- In dry rocky and sandy hillsides near the sea to 400 m. 

     Notes -- Superficially like Cycas pectinata but distinguished by the narrow megasporophyll with a distinctly elongated apical spike and particularly by the bulbous base.  The description under Cycas elongata is of C. immersa.

6. Epicycas siamensis (Miq.) de Laub., Blumea 44 (1998) 393.  -- Cycas siamensis Miq., Bot. Zeit. 21 (1863) 334; DC, Prodr. 16, 2 (1868) 528; Leandri, Flore Indo-Chine (1931)1088 (in part); Schuster in Engler, Pflanzenr. 99 (1932) 80; Smitinand, Nat. Hist. Bull. Siam Soc. 24 (1971) 169 (in part), t. 27; Flora Thai. (1972) 189 (in part); Krempin, Palms & Cycads Around the World (1990) 228, photo lower left; Jones, Cycads of the World (1993) 157 (in part); Wang et al, Cycads in China (1996) 114, pl. 24 & f. 15; Tang et al, Cycads of Thailand (1997) 393, figs.  -- Type: Teysmann s.n., Kan-Boerie, Thailand (holo U, iso L).

C. pectinata (auct.) non Hamilt., Leandri, Flore Indo-Chine (1931) f. 121: 3--5, f. 122: 1, 3--4.

C. changjiangensis N. Liu, Acta Phytitax. Sin. 36 (1998) 552.  -- Type: Liu 97002, Haiinan, Changjiang (holo IBSC, not seen).

A subterranean or protruding bulb to 100 cm diam. from which eventually grows a trunk to 2.5 m high, trunk diam. 8--20 cm.  Leaves 45--120 cm long including a petiole of 8--18 cm usually covered with thorns or sometimes thornless, pubescent when young, 50--103 pinnules on each side of the rachis.  Pinnules 6--12 cm x 4--7 mm, margins flat, midrib prominent, apex acute to rounded, green above, paler below, gradually reducing tpwards the base of the leaf.  Cataphyll 2--4 x 1--1.5 cm, not pungent.  Pollen cone cylindrical or oblong, 20--30 x 6--8 cm, peduncle 1.4--4 cm; microsporophyll 17--20 mm wide, the erect apical part acuminate, 7--12 mm long with a sharply upturned 5--10 mm apical hook.  Megasporophyll with 2--4 ovules; apical part 22--35 x 17--30, triangular and acuminate, 7--16 comb-teeth along each margin 6--15 mm long, the acumen a 14--30 mm spike with several basal reduced teeth.  Seed 29--35 x 30--35 mm, yellow or orange. 

     Distribution -- From Myanmar across most of Thailand, into southern Vietnam, and to Hainan.  Specimens studied: Aplin in 1887, Chevalier 40474, 40601, Gamble (Robertson) 73, Hosseus 361, Kerr 5737, 10208, 16198, 19763, 19963, 20299, Lakshnakara 999, Larsen 8102, Marcan 891, 892, 2466, Martin 1374, Maxwell 75-215, 75-237, 76-264, 76-272, Murata et al 49760, 49779, Pierre 532, Put 2187, Schmid 857, Seidenfaden 11636, 51483, Shimizu T22450, Soejarto et al 6009, Sorenson et al 864, 6463, Teysmann s.n., Thorel 2189, Van Beusekom et al 4067, Van Steenis 17625. 

     Habitat -- Dry deciduous forest on poor rocky soil to 1200 m. 

     Notes -- The small pinnules and short petiole of this and the following contrast with other members of the genus but the retain the flatness that prevails, nevertheless.  There is some overlap in leaf morphology with Cycas pectinata and this has led to confusion (see discussion under C. pectinata), in spite of substantial differences.  The shorter pinnules of siamensis have a mostly rounded apex, the microsporophyll is more elongated with an erect apical part, the pollen cone cylindrical, and the teeth on the megasporophyll are much shorter, while the base is bulbous.  Bifurcated pinnules occur sporadically in Larsen 8102.  Several apparent hybrids involving siamensis with miquelii have ben detected in southeastern Thailand.               

7. Epicycas lindstromii (Yang, Hill & Hiep) de Laub., Blumea 44 (1998) 395.  -- Cycas lindstromii Yang, Hill & Hiep, Novon 7 (1997) 213, fig.  -- Type: Yang 557, Viet Nam, Binh Thuan, Tuy Phong, Xa Chi Cong, near Doc Bao Da.

Often branched, semi-underground bulbs 13--25 cm in diam. without any trunk.  Leaves sometimes few in a whorl, 50--78 cm long including a petiole of 6--15 cm with thorns along at least half its length, 18--60 pinnules on each side of the rachis.  Pinnules 10--13 cm x 7--10 mm, diminishing towards the apex of the leaf, margins flat, midrib prominent, dark green above, lighter below.  Cataphyll 2--5 x 0.3--0.7 cm, not pungent.  Pollen cone cylindrical, 16--20 x 4--5 cm with a c3 cm peduncle; microsporophyll 11--15 mm wide, apical part triangular and slightly rounded, c 6 mm long with a sharply upturned 2--8 mm apical spine.  Megasporophyll with 2 ovules; apical part c60 x 20 mm, oval and acuminate, 6--13 comb-teeth along each margin 8--12 mm long, acumen c30 mm long with a few basal reduced teeth.  Seed 31--35 x 20--30 mm, reddish orange. 

     Distribution -- Along the east coast of southern Vietnam.  Specimens studied: Yang 557, 563, 729, 733. 

     Habitat -- Sandy grassland and open woodland at low elevation. 

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