CYCADACEAE UPDATED (1999)
 D. J. DE LAUBENFELS

1. SUBGENUS CYCAS
Cycas L. section Lemuricae Schuster in Engler, Pflanzenr. 99 (1932) 65.  -- Type: C. circinalis L. 

Trunks sometimes somewhat enlarged at the base (not subterranean).  Petioles with or without thorns.  Pinnules straight or curving towards the base of the leaf or, in four species, at least some curving towards the apex; flat or slightly revolute margins, at least 4 mm wide, the midrib usually more prominent below.   Microsporophyll lanceolate or acuminate in the apical part beyond the fertile part and this part further gradually curved upwards claw-like or its tip hooked abruptly upwards towards the apex of the cone.  Megasporophyll with the apical portion bearing marginal teeth which are smaller or more or less obscure basally and more prominent apically, a maximum of 6 mm long but usually much less (rarely slightly more), briefly or strongly acuminate. 

    Distribution -- Eight species from peninsular India, Madagascar, and Australia, reaching to the east coast of Africa and the southern coast of New Guinea. 

    Habitat -- Mostly in open places in tropical forests and savannas, usually under distinctly seasonal conditions. 

    Notes -- The species of India have much in common with those of Australia, this subgenus being well represented in both areas.  In this subgenus only C. thouarsii, occurring in Madagascar and nearby areas and apparently also in Sri Lanka, has buoyant seeds, otherwise conforming well to the subgenus.  Several Australian species share various distinctive characters, such as apically bending pinnules and pinnules raised to form a trough, that otherwise occur in subgenus Revoluta.  It is considered likely that this represents the result of some form of hybridization. 

KEY TO THE SPECIES

1.  Cycas circinalis L., Spec. pl. 2 (1753) 1188; Lamarck, Ill. de generes 5 (1799) t. 891; Richard, Comment. bot. Conif. & Cycad. (1826) 187 (pt.), t. 24; Hooker, Bot. Mag. 55 (1828) t. 2826--2827; Blume, Rumphia 4 (1849) 13, t. 176; Parlatore in DC, Prodr. 16, 2 (1868) 526; Dyer in Hooker f., Flor. Brit. India 5 (1888) 656; Schuster in Engler, Pflanzenr. 99 (1932) 66, t. 10A, F--G, 11F; Pant, Cycas & the Cycads, ed. 2 (1973) 34, t. 8--12; Wijnands, Acta Hort. 182 (l986) 70; Hill, Taxon 44 (1995) 25.  -- Lectotype: “Todda-pana s. Monta-panna,” Rheede, Hort. Malab. 3 (l682) t. 19 (Stevenson in Jarvis et al., 1993: 40), India.

C. undulata Desf. ex Gaudich. in Freyc., Voy. Autour du Monde 1817--1820, Bot. 19 (1826) 434, hort.

C. squarrosa Lodd. ex Hort. Brit. (1830) 403, hort.

C. circinalis L. var. angustifolia Miq., Comment. Phytogr. (1840) 119.  -- C. wallichii Miq., Monog. Cycad. (1842) 32.  -- Type: Gaudichaud s.n. (“given to Wallich”), cult. Calcutta (lect, here designated, P).

C. squamosa (sic) Lodd. ex Dyer, Gard. Chron. 16 (1881) 270, hort.

Palm-like trees to at least 8 m high. trunk diam. 27--43 cm, tapering, often branched.  Leaves 150--270 cm long including a petiole of 40--60 cm with thorns for most of its length, 80--180 pinnules on each side of the rachis.  Pinnules 14--27 cm x 7--10 mm, margins distinctly but not strongly bent, midrib more prominent below, green above, lighter below.  Cataphylls 5--8 x 2--2.5 cm, not pungent.  Pollen cone cylindric, tapering slightly, 30--45 x 8--11 cm, with a c5 mm peduncle; microsporophyll c12 mm wide, the apical part lanceolate or slightly acuminate and 20--45 mm long, gradually bent towards the apex of the cone.  Megasporophyll  with 3--10 ovules; apical part 40--60 x 20--25 mm, triangular and strongly acuminate, marginal teeth 12--19 per side, 1--2 mm basally and 2--4 mm towards the apex, acumen 10--30 mm and generally bearing a few reduced rather widely spreading teeth basally.  Seed  38--42 x 31--38  mm, dull orange or reddish yellow.

    Distribution -- Along the Malabar Coast of India (Kerala State) and in Shri Lanka.  Specimens studied: Bondam s.n., Buchanan s.n., Debeaux s.n., Fischer 2613, Erlanson 5398, Gamble 13668, 14597, 17052, Greenway 1686, Griffith 88, Gaudichaud s.n., Herold (Milne) 489, Hooker & Tomson 10, 11, Leschenault 814, Mark 714, Metz l69a, Ramamoorthy & Gandhi 2687, Read 2186, 2187, Rottler in 1799, Saldanha 15197, Sauliere 827, Wallich 8587B, Wright 2756.

    Habitat -- Dry deciduous forests and sometimes in wet hill forests to 1070 m.

    Notes -- Although Linnaeus (1753) referred to several places, he specified India as the habit for his species while later other species have been erected for the other places he mentioned leaving circinalis to be lectotypified by “Todda pana.”  According to Desfontaine, undulata was of unknown origin but very much like circinalis.  So far as I can tell, no cultivar with undulate leaves has ever produced reproductive material and furthermore it appears that undulate horticultural forms have sprung from a variety of species.  It remains uncertain which one of these was the plant that Desfontaine had in mind.  Gaudichaud, however, left various specimens in Paris and the only one without a specific wild origin is the type of wallichii, a synonym of circinalis.  Miquel (1840), being influenced by the description of Indonesian material by Roxburgh under the name circinalis and by his own variety javana, understood circinalis to have a wider pinnule so, when he saw a true example in Paris, he named it var. angustifolia which he later raised to a species as wallichii.  He was subsequently uncertain of the relationships of walichii, placing it in synonymy first with rumphii ln 1849 and then with pectinata in l868.  The Gaudichaud specimen “given to Wallich” in the Paris herbarium is evidently the specimen he had in mind and is here lectotypified.  Clearly the squarrosa of Loddiges became mutated accidentally to squamosa and when Dyer mentioned the latter in 1881 he gave no reference or description but did comment that Wendland said it was introduced from Travencore in 1824.  As was mentioned above, the name circinalis has been applied to so many things that most citations of it without provenance have to be disregarded.

2. Cycas sphaerica Roxb., Fl. Ind. 3 (1832) 147; Miquel, Linnea 17 (1843) 693; Tydschr. Wis. & Natuurk. Wet. 2 (l849) 287; Archives Néer. Sci. Exact. & Nat. 3 (1868) 230;  -- Type: Roxburgh in 1808, cult Calcutta [lect (Hill, Taxon 44, 1995, 25) BM].

C. circinalis L. var. orixensis Haines, Bot. Bihar & Orissa 6 (1924) 1228; Hill, Taxon 44 (1995) 25.  -- Type: Haines 5876, India, Mals of Puri (lect, here designated, K).

C. nathorstii Schuster in Engler, Pflanzenr. 99 (1932) 76, t. 10E; Hill, Taxon 44 (1995) 27.  -- Type: Thwaites 3689, Shri Lanka (holo G; iso A, K, P).

Palm-like trees to 4.5 m high, trunk diam. 25--40 cm.  Leaves 150--270 cm long including  a petiole of 27--60 cm with thorns for most of its length, 90--180 pinnules on each side of the rachis.  Pinnules 19--32 cm x 9--12 mm, margins more or less flat, midrib more prominent below, green above, lighter below.  Pollen cone cylindric, tapering slightly, up to 45 x 10 cm, peduncle unknown; microsporophyll 15--22 mm wide, the apical part acuminate with a narrow spine 20--45 mm long that first bends basally and then curves sharply upward.  Megasporophyll with 3--12 ovules; apical part 25--45 x 20--30 mm, triangular, acuminate, marginal teeth 11--18 per side, 2--4 mm basally and 4--6 mm or more towards the apex (shorter on immature specimens), the teeth rather widely spreading, acumen 10--25 mm and generally bearing several reduced teeth basally.  Seed  c38 x 30 mm, orange. 

    Distribution -- Southwest of Calcutta (Orissa Province) and Shri Lanka.  Specimens studied: Fosberg & Sachet 53148, Haines 4033, 5876, 5877, Mooney 2867, Roxburgh in 1808, Thwaites 423, 3689. 

    Habitat -- In disturbed areas and open forest with substantial rainfall at low to moderate elevation. 

    Notes -- Generally similar to circinalis differing in the shape of the microsporophyll, the larger teeth on the megasporophyll and the somewhat larger pinnules.  Roxburgh’s specimen no doubt originated in nearby Orissa but he was under the impression that it derived from the Moluccas.  Haines claimed for the var. orixensis that the apical spine of the microsporophyll towards the apex of the pollen cone is bifid or trifid but no specimens are available to confirm this. 

3. Cycas thouarsii Desf. ex Gaudich. in Freyc., Voy. Autour du Monde 1817--1820, Bot. 19 (l829) 434; R. Brown, Prodr. (1810) 347, nomen; Eichler, Nat. Pflanzenfam. 2,1 (1889) 21; Stapf, Kew Bull. Misc. Inf. 2 (1915) 1; Melville, Fl. Trop.East Africa (l958) 1, t. 1; de Laubenfels, Flore de Madagascar et des Comores: Gymnospermes (1972) 4, t. 1, photo p. 2; Jones, Cycads of the World (1993) 159.  -- C. circinalis L. subsp. thouarsii (Gaudich.) Engler, Pflanzenw. Africa 2 (1908) 82.  -- Type: du Petit-Thouars s.n., Madagascar (holo P).

C. circinalis (auct.) non L., du Petit-Thouars, Hist. Veg. (1804) 2, t. I--II; Richard, Comment. Bot. de Conif. & Cycad. (1826) 187 (in part), t. 25--26; Engler, Pflanzenw. Ost-Africas (1895) 422.

C. madagascariensis Miquel, Comment. Phytogr. (1840) 127.  -- C. circinalis L. subsp. madagascariensis (Miquel) Schuster in Engler, Pflanzenr. 99 (1932) 73.  -- Lectotype: du Petit-Thouars, Hist. Veg. (1804) t. I, Madagascar (here designated).

C. comorensis Bruant, Bull. Soc. Bot. France 35 (1888) 246.  -- Type: Boivin 2891, Comores (holo P).

C. rumphii Miq. subsp. zeylanica Schuster in Engler, Pflanzenr. 99 (1932) 75, t. 10C--D, 11K--M.  -- Type: Thwaites 3862, Shri Lanka (holo G, iso BM, GH, K).

Palm-like trees to 10 m high, trunk diam. to 45 cm, tapering, often branched.  Leaves 150--300 cm long including a petiole 30--50 cm with thorns for most of its length, 60--120 pinnules on each side of the rachis.  Pinnules 17--31 (35) cm x 8--13 (15) mm, margins slightly bent, midrib more prominent below, green above, lighter below.  Cataphylls 4--6 x 2-2,5 cm, not pungent.  Pollen cone cylindric, tapering slightly, 30--60 x 10--15 cm, with a peduncle to 5 cm; microsporophyll 15--17 mm wide, the apical part slightly acuminate and 12--15 mm long with a sharply hooked tip 5--8 mm long.  Megasporophyll with 4--10 ovules; apical part 60--80 x 20--25 mm, lanceolate and strongly acuminate, marginal teeth 5--13 per side, weakly developed basally and 2--3 mm towards the apex, acumen 1--3 cm and bearing reduced teeth basally.  Seed 54--60 x 42--50 mm, buoyant due to a spongy layer within the stony part of the seed, red.

    Distribution -- Along the east coast of Madagascar, in the Comores, delta of the Zambesi, and possibly elsewhere along the east coast of Africa, and Shri Lanca. Specimens studied: Baron 2163, Beck 108, Benson GC50, Boivin 2891, s.n., Commerson s.n., Davy 22603, De Carey 5219, 6524, Faden 74/317, Geay 7616, Gentry 11341, Greenway 9635, Hooker & Thomson s.n., Isaac EA1329, Jayaswriya 12378, Jussieu 17208, Kaslend 14783, Kirks.n., Kitching in 1880, Lam & Meeuse 6182, Livingston in 1858, Meller in 1861, Perrier de la Bathie 11784, 15980, 18801, du Petit-Thouars s.n., Stadman 12, Thwaites 3862, Viguier & Humberet 2000, Williams 148.

    Habitat -- In strand vegetation and in lowland forests in strictly rainforest environments to over 500 m. 

    Notes -- Generally similar to circinalis but differing in the fertile structures, the somewhat wider pinnules, and especially the large buoyant seed.  Certain material from Sri Lanka appears to be the same as that of Madagascar, which is remote from where all the rest of the genus occurs, leading to the suspicion that an origin in Sri Lanka is possible.  The strand habitat and buoyant seed characters have not been confirmed in Sri Lanka.  There are two possible mechanisms for transport across the Indian Ocean, should this hypothesis be correct.  Seeds could have floated across the intervening ocean, but details of the distribution of thouarsii indicate that many appropriate islands and coasts lack the species and therefore long-range dispersal seems not to be within its capacity.  The other possibility is human transport as an edible crop, a process that certainly occurs for thouarsii and continues locally into nearby areas at the present time. 

4.  Cycas media R. Br., Prodr. Fl. Nov. Holland (1810) 348; Miquel, Linnea 17 (1843) 687; Arch. Née. des Sci. Exact. & Nat. 3 (1868) 233; Parlatore in DC, Prodr. 16, 2 (1868) 527; Hill, Telopea 5 (1992) 184, t. 3; Jones, Cycads of the World (1993) 145, photos pp. 145--6.  -- Type:  R. Brown 3106 p. p., probably from Calder I. off Mackay (holo BM). 

C. gracilis Miq., Versl. en Meded. Akad. Wetensch. Amster. 15 (1863) 366.  -- Type: Mueller s.n., Cape Upstart (holo U). 

C. armstrongii Miq., Archiv. Néer. Sci. Exact. et Natur. 3 (1868) 235; Jones, Cycads of the World (1993) 130, photos pp. 130--131.  -- Type: Armstrong 380. Port Essington, N. T. (holo U, iso K, P). 

C. papuana Muell., Papuana Pl. 4 (l876) 71; Warburg, Monsunia 1 (1900) 180, t. 7, f. e; Jones, Cycads of the World (1993) 151; Hill, Austral. Syst. Bot. 7 (1994) 532, f. 3A, 5.  -- C. circinalis L. subsp. papuana (Muell.) Schuster in Engler, Pflanzenr. 99 (1932) 70.  -- Type: D’Albertis in 1876, Fly River, Papua (lecto, Hill, MEL, iso K).

C. kennedyana Muell. ex Warb., Monsunia 1 (1900) 181. t. 7: f. a (Melbourne Chemist and Druggist 4: 47, 1882, 85).  -- Type: Fitzalen in 1881, Normanby Ranges (SW of Bowen) (holo MEL, not seen).

C. megacarpa Hill, Telopea 5 (1992) 188, t. 5 (figures 5a--c taken from the type); Jones, Cycads of the World (1993) 147, photos.  -- Type: Hill & Stanberg 4142, Port Curtis Dist. (holo NSW, not seen).

Palm-like trees to 5 m high, trunk diam. 10--24 cm, base sometimes becoming enlarged, sometimes branched.  Leaves 90--180 cm long including a petiole of (15) 30--60 cm with thorns up to half way to the base or none, 80--150 pinnules on each side of the rachis, the two rows sometimes forming a trough.  Pinnules (6) 11--28 cm x (5) 6--10 mm, margins flat to distinctly bent, midrib usually more prominent below, mid to dark green above, lighter below, sometimes glaucous.  Cataphylls 5--12 x 1.5--2 cm, pungent.  Pollen cone ovoid, 15--25 x 8--13 cm, with a c5 cm peduncle; microsporophyll 14--18 (20) mm wide, the apical part narrowed and extending 8--18 mm, including a sharply upturned spine 6--10 mm long.  Megasporophyll with 2--10 ovules; apical part 4--7 x 2--3 cm, triangular and distinctly acuminate, marginal teeth (7) 10--20 per side, 2--3 mm basally and 4--6 mm towards the apex, acumen 1--3 cm and bearing a few reduced teeth basally.  Seed 30--40 x 25--36 mm, orange-yellow to orange-red. 

    Distribution -- Along the coast of Queensland and in the northern Territory to the southwestern part of Papua, rarely much over 100 km inland.  Specimens studied: Banks & Solander in 1770, Bidwell s.n., Brown 3106, Craven 3211, Byrnes 2406, 2606, Dansie in 1958. Hoogland 8548, Kajewski 1308, Mackee 9610, Maconochie B, 1722, 2690, 2718, 2725, 2726, 2762, McDonald 1539, Melville et al 3673, Mueller s.n., Podenzana s.n., Thare s.n., White 3406, 10190, Armstrong 379, 380. Blake 16330, Brass 6541, Byrnes 2465, D’Albertis in 1876, Dunlop 2955, Foreman & Stocker LAE 59094, Foreman et al LAE 60454, 60493, Jacobs 1774, Levitt in 1922, Mackee 8375, Maconochie 1302, 1305, 1306. 

    Habitat -- Open seasonal forest to sometimes reaching wet forest. 

    Notes -- The two species, circinalis and media, are quite similar but the latter is less robust, has a hooked not gradually bent apex to the microsporophyll, has fewer thorns on the petiole, and has shorter pollen cones. Although typical media may have glabrous flat pinnules and flat leaves, these characteristics show variation throughout its range with, by contrast, glaucous pinnules, bent pinnule margins, and pinnule rows forming a trough commonly seen.  For example, the type specimen of gracilis has distinctly bent pinnule margins.  Hill (1992) reports these characters as “intermediate” populations with respect to platyphylla (angulata) near Atherton and the common occurrence of a trough in plants from the northern part of Cape York Peninsula.  These traits could well represent genetic infiltration from cairnsiana or something else.  Hill further reports that the southernmost populations of media have larger seeds (megacarpa, 40--60 x 35--45 mm) and this might merit treatment as a variety if it proves consistent. 

5.  Cycas angulata R. Br., Prodr. Fl. Nov. Holland (1810) 348; Miquel, Linnea 17 (1843) 686; Arch, ,  . des Sci. Exact. & Nat. 3 (1868) 233; Hill, Telopea 5 (1992) 202, t. 17; Jones, Cycads of the World (1993) 128, photos pp. 128--129.  -- Type: Brown 3106 p. p., prob. Bountiful Island in the Gulf of Carpenteria (holo BM, iso K).  

C. ophiolitica Hill, Telopea 5 (1992) 190, t. 7; Jones, Cycads of the World (1993) 150, photos on p. 149.  -- Type: Hill & Sternberg 4140, Port Curtis Dist., N. S. W. (holo NSW, not seen).

C. platyphylla Hill, Telopea 5 (1992) 193, t. 9 (figure 9e taken from the type); Jones, Cycads of the World (1993) 152, photos.  -- Type: Hill & Sternberg 3764, near Petford, Cook Dist., Queensland (holo NSW, not seen).

C. couttsiana Hill, Telopea 5 (1992) 197, t. 13; Jones, Cycads of the World (1993) 140, photo.  -- Type: Hill & Sternberg 3738, Upper Stawell R., Burke Dist., Queensland (holo NSW, iso K).

C. brunnea Hill, Telopea 5 (1992) 200, t. 15; Jones, Cycads of the World (1993) 133, photos on p. 134.  -- Type: Maconochie 1661, Lawn Hill Creek, Burke Dist.,Queensland (holo DNA, iso K).

C. pruinosa (auct.) non Maconochie, Jones, Cycads of the World (1993) photo on pp. 102--103.

C. arnhemica Hill, Telopea 5 (1994) 693, t. 1.  -- Type: Maconochie 1477, Goyder R. Crossing, N. T. (holo NSW, iso K). 

C. orientis Hill, Telopea 5 (1994) 696. t. 3.  -- Type: Hill & Sternberg 3936, east of Badalngarrmirri Creek, N. T. (holo NSW, not seen).

C. canalis Hill subsp. carinata Hill, Telopea 5 (1994) 700. t. 4e--g (figures taken from the type).  -- Type: Hill & Sternberg 4063, 90 km from Stuart Highway on Dorisville Road, N. T. (holo NSW, not seen).

C. desolata Forst.,Austrobaileya 4 (1995) 345, f. 1-5.  -- Type: Forster 14671B (holo BRI, not seen), Australia, Queensland, NW of Charters Towers.

C. yorkiana Hill, Telopea 7 (1996) 18, f. 8 (f. 8a, b, d-f taken from the type),  -- Type: Hill & Stanberg 4711 (holo NSW, not seen).  Australia, Queensland, 20.5 km N of Wenloch R. crossing, Bamaga Rd.

C. badensis Hill, Telopea 7 (1996) 20, f. 9 (f. 9b, d, g taken from the type).  -- Type: Clarlson 4007 (holo BRI, not seen), Australia, Queensland, Badre Island.

C. tuckeri Hill, Telopea 7 (1996) 20, f. 10 (f. 10b taken from the type).  -- Type: Hill & Stanberg 4683 (holo NSW, not seen).  Australia, Queensland, Coen district.

C. semota Hill, Telopea 7 (1996) 23, f. 11 (f. 11 taken from the type).  -- Type: Hill & stanberg 4708 (holo NSW, not seen), Australia, Queensland, Banaga district.

C. maconoche Hill, Telopea 7 (1996) 48, f. 22.  -- Type: Hill & Anserson 4461 (holo NSW, not seen), Australia, Northern Territory, Mandorah Rd. at Bynoe Hoarbour Turnoff.

Palm-like trees to 12 m but often much shorter, base of trunk expanded to 25--40 cm diam., occasionally branching.  Leaves 90--200 cm long including a petiole of 15--60 cm with thorns up to half way to the base, 80--160 pinnules on each side of the rachis often distinctly forming a trough. Pinnules 6--19 (23) cm x 4--6 (7) mm, margins flat to slightly bent, midrib more prominent below, apex often bent towards the apex of the leaf, generally forming an acute angle of less than 60 degrees with the rachis towards the apex of the leaf, green above, lighter below, usually glaucous.  Cataphyll 6--9 x 2 cm.  Pollen cone ovoid, 30--45 x 12 cm with a c5 cm peduncle; microsporophyll 10--15 (20) mm wide, the apical part distinctly acuminate and 10--17 mm long with a sharply bent subulate tip 8--20 mm long. Megasporophyll with 4--12 ovules; apical part 30--55 x 15--27 mm, triangular and acuminate, marginal teeth 5--13 per side, 2--3 mm basally and c4 mm apically, acumen a 10--30 mm spike.  Seed 30--46 x 25--39 mm, yellow to orange. 

    Distribution -- from southern Queensland all the way across Northern Territory generally within 100 km or so of the coast. Specimens studied: Blake 7898, Brown 3106 p. p., Byrnes 2408, 2782, Craven 4155, Dovey AAG, Dunlop 2955C, Everist 7612, Fell 1798, Fryxell & Craven 4252, Gittons in 1962, Hill & Stanberg 3738, King s.n. Levitt in 1972, Maconochie 608, 962, 1464--6, 1471, 1477--8, 1508--9, 1511, 1514, 1516, 1556, 1620, 1622, 1658, 1661, 1724, 2133--5, 2495, 2498, 2667, 2669. 2735, 2756, McKean 1336, Mueller in 1872, Must 1088, s.n., Nelson 1099, Specht 1003, Thozet in 1872, Wickham & Stokes s.n., Woodman in 1976.

    Habitat -- Sparse grassy woodland, often in sandy soil. 

    Notes -- The usually glaucous pinnules often curving apically and often forming a trough contrast with media but more reliable is the pinnule midrib more prominent below and the pinnules placed at an angle in the horizontal plane, not perpendicular, to the rachis, as well as the enlargement of the trunk base.  These characteristics are entirely intermediate between media and cairnsiana suggesting some sort of hybrid origin, also suggested by the lack of constancy of these characters.  It would appear that angulata is more widespread than has been realized.  In this connection, Hill has recently proposed several new species and varieties for populations more or less within the range of media that correspond closely to angulata.  He relies on distinctions that tend to be unreliable, but are often selected for horticultural purposes, such as color or density variations in the tomentum on emerging leaves, degree of glaucousness, sharpness of the trough formed by opposing rows of pinnules, amount of revoluteness, and color of leaves.  Characteristically, angulata has white and orange-brown hairs (trichomes) mixed in a tomentum on new leaf growth, but this is variable (the tomentum that persists on cataphylls and megasporophylls is much less variable).  On some populations (brunea) the white is little evident on new growth, while in others (couttsiana, orientis, arnhemica) it is predominant.  Glaucousness is a notoriously variable trait and tends to correlate with habitat.  The same can be true of leaf color.  After reviewing a large number of specimens, I find the amount of revoluteness if any and the amount of a trough if any is everywhere variable.  For the time being I must place these species in synonymy pending further testing of their validity.  Hill (1992) reports a larger seed for angulata s.s., but all of the seeds I have seen are not so large and all are of similar size.  Two traits could separate the various taxa into three varieties.  The cataphylls of angulata sen. str. are pungent as are those of brunnea, desolata, and orientis.  The remainder are not pungent.  Of these couttsiana, arnhemica, tuckeri, and semota have densely wooly apices.  Finally, platyphylla, ophiolitica, maconochiei, badensis, and yorkiana have neither pungent cataphylls nor woolly apices. 

6. Cycas lane-poolei Gardner, For. Dept. Bull. Perth 32 (1923) 30, t. E; Jones, Cycads of the World (1993) 143, photos on p. 144.  -- C. media R. Br. var. lane-poolei (Gardner) Schuster in Engler, Pflanzenr. 99 (1932) 78.  -- Type: Gardner 1444, NE of Mt. Hann, near source of Moran R., Kimberly Dist., Western Australia (holo W Aust. Forests Dept. Herb., photo seen).

C. sp. (“Pine Creek”) Krempin, Palms and Cycads around the World (1990) photo p. 226; Jones, Cycads of the World (1993) 162.

C. conferta Chirgwin & Wigston, Journ. Adelaide Bot. Gard. 15 (1993)147; Jones, Cycads of the World (1993) photo on p. 125.  -- Type: G. Brown in 1987, N. Terr. (holo DNA, not seen).

C. canalis Hill, Telopea 5 (1944) 698, t. 4 a--d (figure 4c taken from the type).  -- Type: Hill & Stanberg 4034, 31.4 km from Labelle Downs homestead on track to Channel Point, N. Terr. (holo NSW, not seen).

C. xipholepis Hill, Telopea 7 (1996) 32, f. 15 (f. 15 a-d, f, g taken from the type).  -- Type: Hill & Stamberg 4712 (holo NSW, not seen), Australia, Queensland, Cook district, 10.5 W of Telegraph Line Rd on Batavia Downs to Indley Rd.

Palm-like trees to 6m high, trunk diam. 7--23 cm.  Leaves 60--120 cm long including a petiole of 14--30 cm mostly with thorns the entire length, 80--180 pinnules on each side of the rachis.  Pinnules 10--20 cm x (5) 6--8 (11) mm, margins flat, midrib weak and equal above and  below, straight or the apex bent towards the apex of the leaf, mid to deep green and glaucous when young.  Cataphyll 8--14 x 2, pungent.  Pollen cone ovoid, 22--33 x 12--15 cm with a c3.5 cm peduncle; microsporophyll c20 mm wide, the apical part acuminate and 8--14 mm long with a sharply bent subulate tip 11--18 mm long.  Megasporophyll with 2-4 ovules; apical part 40--65 x 22--32 mm, triangular and acuminate, marginal teeth 6--10 per side, 2--3 mm basally and up to 4mm towards the apex, acumen to 3 cm.  Seed 32--57 x 30--50 mm, yellowish, glaucous when young. 

    Distribution  -- Northeastern corner of Western Australia and northern parts of Northern Territory.  Specimens studied: Cunningham 286, Hinz A42-73, Maconochie 1238-9, 1242, 1264--5, 1308, 1320-3, 1316, 1319, Wilson 10888. 

    Habitat -- Dry stunted woodland on gravelly or sandy soil.

    Notes -- The short crowded relatively broad and flat apically bent pinnules of lane-poolei are unusual among the species where the pinnules may bend apically.  This species otherwise generally resembles angulata.  The seed in conferta is reported to be pear-shaped.  Overall the original description of conferta, involving unusual variation in pinnule width and number of thorns on the petiole, suggests that more than one taxon may have been included (one of which may be angulata).  Teeth on the megasporophyll are sometimes reported to be absent (see discussion under normanbyana). 

7. Cycas normanbyana Mueller, Fragm. Phytogr. Austr. 8 (1874) 169; Wieland, Amer. Fossil Cycads 1 (1906) f. 1260 (after Mueller).  -- C. rumphii Miq. subsp. normanbyana (Muell.) Schuster in Engler, Pfalnzenr. 99 (1932) 75.  -- Type: Fitzalajn in 1874 (MEL 68048), Burdekin R. (holo MEL, iso K).

C. basaltica Gardner, For. Dep. Bull. W. Austr. 32 (1923) 31; Jones, Cycads of the World (1993) 132, photo on p. 131.  -- C. media R. Br. var. basaltica (Gardner) Schuster in Engler, Pflanzenr. 99 (1932) 78.  -- Type; Gardner 1490, Kimberly Dist., W. Aust. (holo W. Austr. Forests Dept. Herb., photo seen). 

Palm-like trees to 6 m high, trunk base conspicuously enlarged and 25--40 cm diam. (to 110 cm in W. Australia).  Leaves 65--129 cm long including a petiole of (9) 21--28 cm with few thorns or none, c70--120 pinnules on each side of the rachis.  Pinnules 10--20 cm x 6--8 mm, straight or very slightly bent towards the apex of the leaf, margins flat or slightly bent, midrib weak, light to mid green.  Cataphylls 7--13 x 2.  Pollen cone connical-ellipsoid, 20--30 x 9--13 cm with a short peduncle; microsporophyll 12--18 mm wide, the apical part acuminate and 7--9 mm long with a sharply curved claw-like tip 5--7 mm long.  Megasporophyll with 2--8 ovules; apical part 50--80 x c18 mm, lanceolate and acuminate, marginal teeth 8--22 per side, 1--2 mm long, laterally spreading, acumen c10 mm long.  Seed 25--37 x 20--27 mm, glaucous when young. 

    Distribution -- Scattered and rare along the northern Queensland coast and along the northernmost coast of Western Australia.  Specimens studied: Brass 18924, Fitzalen in 1874, Maconochie 1196--7, 1269, 1272--3, 1283, 1285--6, 2692, 2694, Mueller in 1872. 

    Habitat -- Open woodland, sometimes on basaltic soils. 

    Notes -- The striking laterally spreading teeth on the elongated apical part of the megasporophyll together with sometimes upturned pinnules suggests a relationship with cairnsiana (pruinosa), a near neighbor in Western Australia, while in other ways the resemblance is more with media, a Queensland neighbor.  Some kind of hybrid origin is possible.  Hill (1992) reports searching the type area for examples without success.  It may, of course, have disappeared there (Mueller said it was rare).  Many specimens in Northern Australia lack teeth on the megasporophyll.  It appears that insect damage during early development often produces these toothless margins, which have been reported in several species in northern Australia.

8.  Cycas furfuracea Fitzgerald, Journ. & Proc. R. Soc. W. Austr. 3 (1918) 108; Jones, Cycads of the World (1993) 141, photo.  -- C. media R. Br. var. furfuracea (Fitzgerald) Schuster in Engler, Pflanzenr. 99 (1932) 78.  -- Type: Fitzgerald s.n., Kimberly Dist., W. Australia (holo NSW).

C. arenicola Hill, Telopea 5 (1993) 419, t. 1 (figures 1b--c, e taken from the type).  Type: Russell-Smith & Brock 8502, upper E. Alligator R., N. Terr. (holo NSW, not seen).

Palm-like trees to at least 2 m high, trunk diam. 15--22 cm.  Leaves 60--160 cm long including a petiole of 14--35 cm with thorns on the upper half, c60--140 pinnules on each side of the rachis.  Pinnules 9--19 cm x 4.5--7 mm, straight or slightly curved towards the apex of the leaf, margins distinctly bent, midrib more prominent below, glossy dark green with a rusty tomentum below.  Cataphylls 4--8 x 1.5, not pungent.  Pollen cone narrowly ovoid with a short peduncle, 25--38 x 5--9 cm; microsporophyll up to 20 mm wide, the apical part acuminate and 12--18 mm long with a sharply bent tip 5--12 mm long.  Megasporophyll with 2--6 ovules; apical part 25--40 x 20--32 mm, triangular and acuminate, marginal teeth c10 per side, 2--3 mm long, acumen 8--14 mm long.  Seed 28--32 x 25--29 mm, yellow to orange, glaucous when young. 

    Distribution -- Localized in the Leopold Ranges and Mitchell Plateau of W. Australia and on the upper reach of the E. Alligator R. in N. Terr. of Australia.  Specimens studied: Lazarides 7650, Lullfitz 57, Maconochie 1201. 

    Habitat -- Open woodland on sandy soil. 

    Notes -- This rare species is more or less intermediate between calcicola and media and could be of hybrid origin.

Click here to return to the Outline

This site is copyrighted © 1998-2006, Palm & Cycad Societies of Florida, Inc.
For questions or comments, e-mail the webmaster.
Internet hosting provided by Zone 10, Inc.