BARRO COLORADO ISLAND
INTRODUCTION 63 INTRODUCTION 63
10. POLYPODIACEAE
84
PTEROPHYTAlong, bluntly acuminate apex, the lobes 2-3 mm wide, somewhat falcate, entire to minutely crenate; veins mostly 1-forked near base, the major veins sparsely pubescent beneath with purplish, mostly stellate scales. Sori round, usually 4-6, clustered in proximal half of lobes; indusium globose to urn-shaped, dark brown, completely enclosing sporangia, breaking into irregular segments at maturity, deciduous to persistent, scarious. Croat 6529.
Rare on the island, restricted to deep ravines. Seasonal behavior undetermined. Seen fertile in April, August, and September.
Nicaragua to Peru, Bolivia, Paraguay, and Brazil. In Panama, known only from tropical moist forest on BCI and in Darien.
See Fig. 11.
TRICHOPTERIS Presl
Trichopteris microdonta (Desv.) Tryon, Contr. Gray Herb. 200:46. 1970
Akophila microdonta Desv.
Tree fern, 1-5 m tall, slender, the trunk only a few cm diam. Leaves few, 2-pinnate-pinnatifid; petioles to ca 1 m long, lustrous, purplish near base, becoming light brown, glabrate, sparsely scaly near base and bearing numerous distant, narrowly conical spines, these curved and to 1 cm long; scales to ca 1.5 cm long and 2 mm wide, brown, very long-caudate-acuminate; leaves ovate-oblong, abruptly acuminate, to 1.5 m long and 1.2 m broad, the primary rachis brown or yellowish-brown, armed throughout;
pinnae petiolate, narrowly oblong, acuminate, 30-60 cm long, 10-25 cm wide; secondary rachis yellowish-strigose above, thinly scurfy-hirtellous below (glabrate in age), distantly aculeate with spines to ca 4 mm long; leaflets ± sessile, ± oblong, long-attenuate, 5-13 cm long, 1.5-3 cm wide, the costa densely yellowish-strigose above, scurfy- hirtellous and with a few minute, mostly caducous scales below, the segments of 19-25 pairs, linear, ± acute, 8-18 mm long, 2-4(5) mm wide, crenate-serrate, the costules distantly hispid above, thinly scurfy-hirtellous below. Sori in 6-11 pairs, the paraphyses very numerous, equaling sporangia. Kenoyer 7, Munch s.n.
Apparently rare, in ravines; more common elsewhere in the Canal Zone.
Mexico to Peru and Brazil, principally at low eleva- tions; West Indies (Isle of Pines). In Panama, known only from tropical moist forest in the Canal Zone.
Trichopteris trichiata (Max.) Tryon, Contr. Gray Herb. 200:44. 1970
Akophila trichiata Max.
Graceful tree fern, to 3.5 m tall. Leaves 2- or 3-pinnate- pinnatifid on lower parts of large leaves, to 3.5 m long and 2 m wide at the middle; petioles densely pubescent with moderately stifli' trichomes, sparsely armed throughout with short spines; rachis with short, soft, villous pubes- cence interspersed with long, stiff, jointed trichomes;
pinnae lanceolate to oblong-lanceolate, 50-90 cm long.
to 30 cm wide, abruptly acuminate and confluent toward apex; leaflets sessile, linear-lanceolate, attenuate-caudate, pinnatifid to beyond the middle, mostly to 9 (15) cm long, to 2 cm wide, both surfaces pubescent especially on veins, the segments 5-10 mm long, 1.5-3 mm wide, usually bi- dentate at apex. Sori exindusiate, round, on veinlets on proximal two-thirds of lobes, consisting of moderately few sporangia; paraphyses few, scarcely exceeding spo- rangia. Croat 11729.
Apparently rare; collected once on Gross Point Penin- sula {Croat 11729) and twice elsewhere by Shattuck {740,1149). Probably fertile throughout the year.
The generic name Trichipteris, used by some authors, is an orthographic error.
Costa Rica to Ecuador and Venezuela, from near sea level in Costa Rica to ca 1,000 m in Ecuador. In Panama, known from tropical moist forest in the Canal Zone, Panama, and Darien and from premontane wet forest in the Canal Zone and Panama.
10. POLYPODIACEAE/ADIANTUM 85
KEY TO THE SPECIES OF ACROSTICHUM
Leaf veins glabrous, scarcely if at all raised A. aureum L.
Leaf veins at least partly pubescent, standing up sharply A. danaifoUum Langsd. & Fisch.
Throughout tropical America. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Tore, San Bias, Panama, and Darien and from tropical wet forest in Colon (Miguel de la Borda).
See fig. on p. 12.
Acrostichum danaifolium Langsd. & Fisch., Icon. Fil.
5,t. 1.1810
Very similar to A. aureum, but the fertile leaves having more of their leaflets fertile; and the leaflets 1-4 cm apart near apex, pubescent at least on the veins, and the veins beneath raised. Croat 6165.
Found with A. aureum along the shore of the island.
Throughout tropical America. In Panama, known only from BCI and the Pacific slope of the Canal Zone and in Panama.
ADIANTUM L.
The genus Adiantum, distinguished by its oblique leaflets and indusiate marginal sori, provides the dominant terres- trial fern flora on the island.
Adiantum decoratum Max. & Weath., Amer. J. Hot.
19:165. 1932
Terrestrial, to 70 cm tall; rhizome ± erect, with dense linear scales. Leaves 2-pinnate; petiole, rachis, and pin- nular rachis with long, conspicuous, ± dense, filiform scales; petioles as long as or longer than blades; leaflets asymmetrical (the midrib nearly submarginal), ± oblong, blunt at apex, truncate and parallel to rachis at base, to 2.5 cm long and 8 mm wide, glabrous, usually glaucous beneath, the sterile margins irregularly, finely serrate.
Sori interrupted along upper and distal margins. Shattuck 282.
Rare, in the forest.
Mexico to Panama. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, Los Santos, Panama, and Darien and from premontane wet forest in Panama (Cerro Azul).
Adiantum fructuosum Spreng., Syst. Veg. 4:113. 1827 Terrestrial, rarely taller than 50(80) cm. Leaves usually several, closely clustered on a short-creeping, scaly rhi- zome, usually 1-pinnate (often 2-pinnate elsewhere);
KEY TO THE SPECIES OF ADIANTUM
Leaflets borne on slender stalks more than 7 mm long:
Leaflets ± reniform, rounded at apex, usually less than 4 cm long on stalks less than 2.5 cm long A. lunulatum Burm.
Leaflets ± ovate, acimiinate, usually more than 5 cm long on a stalk more than 3 cm long A. seemannii Hook.
Leaflets not borne on stalks more than 7 mm long:
Sori continuous along margins:
Leaflets more than 5 cm long, narrowly triangular; sori continuous along upper and lower margins A. lucidum (Cav.) Sw.
Leaflets less than 3 cm long, ± falcate; sori continuous on upper margin, sometimes with a shorter segment on the distal edge, lacking on lower margin A. pulverulentum L.
Sori interrupted along margins:
Rachis bearing long, conspicuously spreading, threadlike scales, clearly visible to the naked eye, these interspersed with shorter, fine scales:
Leaflets pubescent beneath A. humile Kunze Leaflets glabrous beneath A. decoratum Max. & Weath.
Rachis lacking clearly visible scales or bearing scales ± appressed, not spreading, and less conspicuous:
Sterile leaflets unevenly and coarsely serrate on upper and distal margins, the apex usually blunt to rounded A. fructuosum Spreng.
Sterile leaflets evenly and usually finely serrate or biserrate:
Leaflets not glaucous beneath, the sterile ones usually acute to acuminate A. ohliquum Willd.
Leaflets glaucous beneath (fertile leaflets sometimes pubescent on inner margin), the sterile ones usually blunt at apex:
Leaflets pubescent beneath (often nearly glabrous on juvenile or sterile leaflets), usu- ally not tapered much to apex A. humile Kunze Leaflets usually glabrous beneath, usually much broader at base than apex
A. petiolatum Desv.
86
PTEROPHYTApetiole and rachis dark brown, with fine, short threadHke scales persisting at least on rachis; leaflets nearly oblong, blunt to acute at apex, 1-3 cm long, glabrate beneath (elsewhere sometimes with filiform scales), midrib very near lower margin, the lower edge ± straight, the proxi- mal edge paralleling the rachis, the upper and distal edges of sterile leafiets unevenly and coarsely serrate.
Sori interrupted along upper and distal margins. Croat 8576.
Occasional, in the forest.
Distinguished by its small, coarsely serrate leaflets. The species is easily confused with and possibly not separable from A. tetraphyllum H. & B. ex Willd., which has a long-creeping rhizome with the petiole bases not closely crowded as in /I. fructuosum. Moreover, in A. tetra- phyllum the sterile apices of the leaflets are acute and turned toward the apex of the pinnae, whereas in A.
fructuosum they are mostly straight and obtuse.
Mexico to Brazil; West Indies. In Panama, known from tropical moist forest in the Canal Zone, San Bias, Panama, and Darien, from premontane wet forest in Panama and Code, and from tropical wet forest in Colon.
See Fig. 12.
Adiantum humile Kunze, Linnaea 9:80. 1834 A. killipii Max. & Weath.
Terrestrial, usually less than 50 cm tall. Leaves 2-pinnate;
petioles shiny and atropurpureous; rachis similar but usually covered with two types of scales•small, fine,
± appressed scales and longer, subulate-filiform scales;
leaflets asymmetrical, mostly 1-3 cm long, acute at base, the lateral leaflets ± oblong or sometimes broader at the base with apex usually rounded and the proximal edge nearly paralleling the rachis, the lower surface usually glaucous and usually bearing numerous to few simple trichomes and sometimes filiform scales; sterile leaflets finely and evenly serrate. Sori interrupted along inner, upper, and outer margins, frequently fertile at apex.
Croat 8553.
Occasional, along trails in the forest.
Belize to Peru, Brazil, and French Guiana; Trinidad.
In Panama, known from tropical moist forest on the Atlantic slope of the Canal Zone and in San Bias, and from premontane wet forest in Panama (Lago Cerro Azul).
See Fig. 13.
Adiantum lucidum (Cav.) Sw., Syn. Fil. 121. 1806 Terrestrial, to 70 cm tall; rhizome moderately short- creeping with slender spreading scales. Leaves usually few, 1-pinnate or rarely 2-pinnate; petiole and rachis dark brown or black, persistently pubescent-scaly (at least on rachis); leaflets asymmetrical, usually 5-10 cm long, ± narrowly triangular, acuminate at apex, markedly inequi- lateral at base, glabrous or sparsely scaly below; sterile leaflets with margins unevenly and coarsely serrate but the proximal third of lower margin as well as the proxi- mal margin entire; terminal leaflet ± ovate. Sori very
long and continuous on upper and lower margins (except near apex). Croat 7731.
Abundant in the forest.
Distinguished by its long pointed leaflets bearing sori along most of both margins. Tryon (1964) reported the species to be 2-pinnate in Peru, a condition that is rare in Panama.
Panama to Peru and the Guianas; Trinidad and To- bago. In Panama, known from tropical moist forest on both slopes of the Canal Zone and in Panama and Darien, from premontane moist forest in the Canal Zone (Anc6n Hill), and from tropical wet forest in Colon.
See Fig. 14.
Adiantum lunulatum Burm., Fl. Ind. 235. 1768 Terrestrial, to 55 cm tall; rhizome erect, with a few dark, lanceolate-linear scales. Leaves 1-pinnate; petiole and rachis dark reddish-brown, shiny; petiolules slender, 0.3-2.5 cm long; leaflets lunate to reniform, lobed on semicircular distal edge, truncate to obtuse on lower edge, to 4 cm long and 1.8 cm wide, glabrous. Sori inter- rupted along lobes on distal edge. Shattuck 1058.
Rare, in the forest.
This species has been confused with the name Adian- tum philippense L., which should probably be considered a nomen duhium.
Mexico to Colombia and Venezuela; West Indies. In Panama, known from tropical moist forest on both slopes of the Canal Zone and in Veraguas; known also from premontane moist forest in Panama (Panama City) and from premontane wet forest in Chiriqui, Code, and Panama.
Adiantum obUquum Willd., Sp. PL 5:429. 1810 Terrestrial, to 60 cm tall. Leaves mostly 2-pinnate, a few 1-pinnate; petiole atropurpuraceous, sparsely to moder- ately scaly at least when young, the threadlike, curly scales persisting on rachis; leaflets asymmetrical, mostly 3-6.5 cm long, seldom more than 1.5 cm wide, narrowly tapered, the apex acute to acuminate, the lower edge nearly straight, the proximal edge nearly paralleling the rachis, the underside not glaucous, the sterile margins unevenly and finely biserrate. Sori interrupted along upper and distal margins with the apex usually sterile. Croat 7815.
Occasional, along forest trails and no doubt also in other areas of the forest.
Often confused with A. petiolatum, but differing in having the leaflets more narrowly triangular and irregu- larly serrate and in having the apex of the fertile leaflets free of sori and tapered to a sharp point. The leaflets of A. petiolatum are glaucous beneath, with the inner edge frequently cordate.
Throughout the West Indies and Central America, as well as from Colombia to Bolivia, Brazil, and the Gui- anas. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, and Darien and from pre- montane wet forest in Panama (Chiman).
ff*^ ^^^-''W^
Fig. 12. Adiantum fructiwsum*..••
Fig. 13. Adiantum humile
Fig. 14. Adiantum lucidum
I
t
1-1
Fie. 16. Anetium citrifolium :• ""d "•
\ (ij.
w*.*
jff
Fig. \1. Aspkniuni delitescens
•^ ft:
^^^'
Adiantum petiolatum Desv., Ges. Naturf. Freunde Berl. Mag. 5:326. 1811
Terrestrial, to 40 cm tall; rhizome with spreading scales, the petiole bases closely spaced. Leaves 1- or 2-pinnate;
petioles dark, glabrate; leaflets asymmetrical, obscurely triangular to oblong, rounded to bluntly acuminate at apex, mostly 2-5 cm long, 1-3.5 cm wide, the proximal edge frequently semicordate and overlapping the rachis at base, usually glaucous and glabrous beneath, the ster- ile margins evenly and finely serrate. Sori interrupted along upper, lower, and sometimes proximal margins, frequently extending to very near the apex. Croat 6948.
Common on trails in the forest and at the edge of • clearings.
See the discussion oi A. obliquum for a comparison of the two species.
Mexico to Peru, Bolivia, and Brazil; West Indies. In Panama, known from tropical moist forest in the Canal Zone (BCI and the Pacific slope), Herrera, Panama, and Darien and from tropical dry forest in Panama (Taboga Island).
See Fig. 15.
Adiantum pulverulentum L., Sp. PL 1096. 1753 Terrestrial, to 1 m tall. Leaves 2-pinnate; petiole and rachis dark reddish-brown, densely pubescent-scaly, the scales fine and curly; leaflets asymmetrical, 1.5-3 cm long, usually of about equal width throughout but gradu- ally curved upward to the sharply toothed, ± acute apex, the lower edge entire and usually straight, the proximal edge ± paralleling rachis, glabrate to scaly beneath, the sterile leaflets and apex of the fertile leaflets coarsely serrate. Sorus 1 (rarely 2), long, on upper margin, with occasionally a shorter one on outer margin. Croat 4340.
Occasional, in the forest.
Distinguished by the long sorus on the upper edge of each leaflet.
Mexico to Colombia, Bolivia, Brazil, and the Guianas;
West Indies. In Panama, known from tropical moist forest in the Canal Zone, Panama, and Darien, from premontane wet forest in Colon and Panama, and from tropical wet forest in Colon.
Adiantum seemannii Hook., Species filicum 2(5):81A. 1851
Terrestrial, usually to about 50 cm tall; rhizome short- creeping, densely scaly. Leaves 1- or 2-pinnate; petiole, rachis, and petiolules blackish and shiny; leaflets ovate, acuminate at apex, oblique and obtuse to truncate or cordate at base, to 9 cm long and 7 cm wide, glabrous and glaucous beneath, the sterile margins coarsely and unevenly serrate; petiolules slender, 2-7 cm long. Sori interrupted along margin. Aviles 79.
Uncommon to rare, in the forest.
Not confused with any other species on the island.
Distinguished by its large, ovate, long-petiolate leaflets.
Mexico to Colombia. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, Pan-
ama, and Darien, from premontane moist forest in Pan- ama (Panama City), and from premontane wet forest in Chiriqui.
ANANTHACORUS Und. & Max.
Ananthacorus angustifolius (Sw.) Und. & Max., Contr. U.S. Nad. Herb. 10:487. 1908
Epiphyte; rhizome moderately short with many fine root- lets densely covered with threadlike, reddish-brown scales; rhizome scales linear-lanceolate, attenuate, clath- rate, iridescent. Leaves simple, entire, sessile, ± linear, tapering gradually at both ends, mostly 10-30 cm long, 1-1.5 cm wide (the sterile leaves shorter and relatively broader), ± thick, glabrous to minutely scaly. Sori exin- dusiate, in continuous or irregularly interrupted lines very near margins. Croat 4367.
Occasional, in the forest, usually on smaller trunks and branches moderately near the ground.
Throughout much of tropical America. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, Veraguas, and Darien, from premontane wet forest in Chiriqui and Panama, and from tropical wet forest in Panama.
ANETIUM (Kunze) Splitg.
Anetium citrifolium (L.) Splitg., Tijdschr. Natuurl.
Gesch. Physiol. 7:395. 1840
Epiphyte with a creeping, scaly rhizome; rhizome scales broadly ovate-attenuate, clathrate, iridescent with short- ciliate margins; petioles well spaced with many fine root- lets densely covered with threadlike scales. Leaves simple, entire, glabrous, sessile or nearly so, oblanceolate to ob- tuse, abruptly acuminate at apex (round on juveniles), gradually tapered to base, usually less than 15 cm long and 4 cm wide, thick, the midrib indistinct except at base, the veins anastomosing. Sporangia scattered chiefly on veins, forming a reticulum. Croat 11703.
Infrequent, in the forest, usually on tree trunks very near the ground; often on stilt roots of Scheelea zonensis (19. Palmae). Tryon (1964) reported seeing leaves to 100 cm long.
Throughout tropical America. In Panama, known from tropical moist forest on both slopes in the Canal Zone and from tropical wet forest in Veraguas (Atlantic slope)
and Colon (Portobelo).
See Fig. 16.
ASPLENIUM L.
The genus occurs in epiphytic habitats similar to those of Polypodium. It can be distinguished by its elongate indusiate sori along major veins on the underside of leaves. Most species have oblique leaflets.
90
PTEROPHYTAKEY TO THE SPECIES OF ASPLENIUM
Leaves simple A. serratum L.
Leaves 1-pinnate:
Leaflets usually more than 6 cm long:
Plants epiphytic; terminal leaflet similar to lateral leaflets, not pinnatifid A. falcinellum Max.
Plants terrestrial or epipatric (on rocks); terminal leaflet unlike lateral leaflets, pinnatifid A. delitescens (Max.) A. R. Smith Leaflets mostly less than 6 cm long:
Stipe and lower rachis dark purplish, shiny; proximal half of lower edge of leaflet frequently entire A. laetum Sw.
Stipe and lower rachis not dark purplish and shiny; proximal half of lower edge of leaflet usu- ally toothed:
Veins mostly 1- or 2-forked; blades not very thin A. auritum Sw. var. auriculatum (Hook.f.) Mort. & Lell.
Veins nearly all simple; blades thin A. pteropus Kaulf.
Asplenium auritum Sw. var. auriculatum (Hook.f.) Mort. & Lell., Mem. New York Bot. Gard. 15:19. 1966 Epiphyte, to 35 cm tall. Leaves I-pinnate, glabrous; leaf- lets oblong, reduced above and tapered to a narrowly rounded apex, usually with a small auricle on upper side near base, to 3 cm long, less than 1 cm wide, the margins finely and ± irregularly serrate or the proximal one- fourth of the lower edge entire, the veins mostly I- or 2- forked. Sori several, oblong, 2-5 mm long, along upper edge of lateral veins, sometimes also on auricle, sharply oblique to midrib; indusium ± lunate, thin. Croat 11293.
Uncommon, on trees (seen mostly near the ground) or on rocks, within the forest.
Throughout much of tropical America. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, Panama, and Darien, from premontane wet forest in Chiriqui, Code, and Panama, from tropical wet forest in Colon, Chiriqui, and Darien, and from lower montane wet and lower montane rain forests in Chiriqui.
Asplenium delitescens (Max.) A. R. Smith, Amer. Fern J. 66:120. 1976
Diplazium delitescens Max.
Terrestrial, to 35 (43) cm tall; rhizome short-creeping, with minute, dark, fibrous scales, the petiole bases closely spaced. Leaves few, 1-pinnate; petioles 4.5-20 cm long, with a few scales similar to rhizome scales near base;
lateral leaflets mostly narrowly lanceolate, gradually long- acuminate, inequilateral at base with the lower proximal edge acute and the upper proximal edge obtuse to trun- cate and nearly paralleling rachis, 3-12 cm long, 1-2 cm wide, irregularly serrate, sometimes with a small auricle on upper side; veins 3- or 4-forked; terminal leaflet pin- natifid. Sori linear along upper branch of lateral veins, slightly curved, nearer midrib than margin, usually on distal side of veins, much less frequently on both sides;
indusia thin, broad, ending abruptly. Croat 8643.
Occasional, in the forest, especially older forest.
Its terrestrial habitat, nonclathrate rhizome scales, and pinnatifid terminal leaflet (unlike the lateral ones) dis- tinguish it irora Asplenium falcinellum, which has a simi- lar leaflet shape. A. falcinellum, an epiphyte, has clathrate rhizome scales and a conform terminal leaflet.
Belize to Panama. In Panama, known from tropical moist forest in the Canal Zone (Pacific slope), Panama, and Darien and from premontane moist forest in Panama.
See Fig. 17.
Asplenium falcinellum Max., Contr. U.S. Natl. Herb.
13:14. 1909
Epiphyte, to 40 cm long, at least sometimes pendent;
rhizome scales clathrate. Leaves 1-pinnate, few; petioles mostly 8-10 cm long, ± glabrous; leaflets in 7-12 pairs, narrowly lanceolate, gradually long-acuminate, equally or inequilaterally acute at base (the lower side often more acute than the upper), 6-11 cm long, 9-20 mm wide, entire or irregularly and inconspicuously crenate, the terminal leaflets much like the lateral ones. Sori linear, subequal, on distal side of lateral veins. Croat 12562.
Infrequent, along trails in the old forest.
See the discussion oi Asplenium delitescens for a com- parison of the two species.
Guatemala to Panama; near sea level to reportedly 1,200 m. In Panama, known from tropical moist forest in the Canal Zone (on and adjacent to BCI), from premon- tane wet forest in Veraguas, and from tropical wet forest in Colon (Santa Rita Ridge) and Panama.
Asplenium laetum Sw., Syn. 79:271. 1806
Epiphyte, usually 25-50 cm tall. Leaves 1-pinnate; petiole and base of rachis atropurpureous and shiny; blades reduced toward apex and tapered to a slender tip; leaflets
± oblong, very unequal at base, the lower edge straight to curved, entire, the proximal margin nearly paralleling rachis, the upper and distal margins serrate, the upper margin infrequently auriculate, 2-6 cm long, 8-15 mm wide, the veins simple or 1-forked. Sori several, linear- oblong, on upper side of lateral veins oblique to mid- rib, mostly on distal half of leaflets; indusium thin.
Croat 8475.
Occasional, on rocks in stream beds of larger ravines;
locally abundant.
Mexico to Paraguay and Venezuela; the Antilles. In Panama, known from tropical moist forest in the Canal Zone (BCI), Bocas del Toro, and Panama and from trop- ical wet forest in Colon.