FERN
_GAZET TE

_ Edited by
vd M.Camus & J.A. Crabbe

iE NEI SR aie ee
Volume 15 Part 7 — 1999

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FERN GAZ. 15(7) 1999 221

CHECKLIST OF THE PTERIDOPHYTES OF THE MBARACAYU
FOREST NATURE RESERVE, PARAGUAY

M. C. PENA-CHOCARRO’, B. JIMENEZ’, G. MARIN? & S. KNAPP*
I, *Department of Botany, The Natural History Museum, Cromwell Road,
London SW7 5BD, U.K.

3 Fundacion Moisés Bertoni, Casilla 714, Asuncion, Paraguay

Key words: checklist, new records, Paraguay pteridophytes, phytogeography.

ABSTRACT

A checklist of the pteridophytes of the Mbaracayu Forest Nature
Reserve, eastern Paraguay is presented. This is the first such detailed
list for the area and includes 115 taxa in 22 families and 46 genera.
Fifteen of these are new published records for Paraguay. Distribution
details are provided, and a brief analysis of the phytogeography of the
pteridophyte flora is presented. Habitat specialisation in the area is also
discussed. Collections for the taxa in the checklist are cited.

INTRODUCTION

The Mbaracayt Natural Forest Reserve (Reserva Natural del Bosque Mbaracayu) is
situated in the north-eastern part of the Oriental region of Paraguay in the department
of Canindeyu between 55°20-55°33’W latitude and 23°59-24°16’S longitude (Fig.
1). The Reserve covers an area of approximately 600 km”, largely covered with
Atlantic Forest (bosque Atldantico Interior: FMB, 1991; Selva Paranaénse, Laclau,
1994: Mata Atlantica, Davis et al., 1997) and is between 140-150 m above sea level.

In the 1980s the original area of the Reserve (then much smaller than at present)
was the property of the World Bank and during its tenure some parts of the forest
were selectively logged for tropical hardwoods. In 1991, thanks to the efforts of
many individuals and organisations (such as The Nature Conservancy) the area was
purchased and passed to the Fundacién Mbaracayu for management. At this time the
Fundacion Moisés Bertoni, a Paraguayan NGO involved in conservation throughout
the country, undertook the management and protection of the Reserve. With the help
of private donations and the ceding of government controlled lands, the Reserve has
grown considerably from its original size. Between 1992 and 1994 approximately
6000 hectares of grassland, the “campos cerrados’” known as Aguara Nu, were
incorporated into the area of the reserve.

The current area of the Reserve was previously occupied by nomadic hunter-
gatherers of the Aché people (Hill & Hurtado, 1996). At the time of contact in the
late twentieth century there were four independent populations of Aché. The largest
group, the northern Aché, roamed an area of about 18,500 km? in North-eastern

VOLUME 15 PART 7 (1999)

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PTERIDOPHYTES OF MBARACAYU FNR ZED

Paraguay. The northern limit of this group’s present territory is in the current
Reserve. This northern group numbered about 550 persons before contact (in the
1970s) and inhabited the Rio Jejui drainage, using the natural resources of the then
pristine forest. Later, many of the groups mixed and established themselves in
association with various missions. The northern Aché people today number about
600 (Hill & Hurtado, 1996). Although all the communities are outside the core
protected area of the reserve, the Aché have retained the right to hunt and gather
within its limits in their traditional ways.

Other indigenous groups, all members of the Guarani linguistic family, have more
recently migrated to the area and have established communities around the limits of
the Reserve. In the area of the cerrado the Ava Katuete or Ava Chiripa, who
traditionally inhabit the transition zone between cerrado and forest, have established
communities; near the south-eastern boundary are Mbya communities; and towards
the Rio Jejui Guazt are communities of Pa’i Tavytera (see Fig. 1).

Although all of the communities, Aché and Guarani, are more or less integrated
into Paraguayan society, they all retain much of their traditional knowledge and
many of their cultural practices, above all in the uses of the plant life around them.
This knowledge needs to be taken into account in planning any study in the region.

With funding from the UK Government’s Darwin Initiative for the Survival of
Species we have been carrying out an inventory of the plants and selected insect
groups in the Mbaracayt Reserve. While preparing a larger checklist of all vascular
plants of the Reserve collected during the course of the project, we found an
unexpected high diversity of pteridophytes. The checklist we present includes our
own collections and those of a past collector in the region, Emile Hassler, a Swiss
national who collected in Paraguay between 1885-1913 and in Mbaracayui between
1898-1899. Hassler enumerated 41 species of pteridophytes for the Mbaracayi area.

STUDY AREA

Approximately 80% of the area of the Mbaracayt Reserve is covered with Atlantic
Forest (Fig. 2). This forest is a subtropical semideciduous wet forest (Hueck, 1978;
Keel & Herrera-MacBryde, 1997). The extent of the forest cover has been reduced
severely in recent times due to deforestation, and now this forest type appears only as
fragmented and isolated patches (Bozzano S. & Weik, 1992). These continental
forests are similar to those found along the eastern coast of Brazil (see Mori et al.,
1983; Mori, 1989), but are distinct in that they harbour several endemic subtropical
genera and some tropical and cerrado species at the southern edge of their
distribution. The affinities of these forests are both with the tropical forests to the
north and also with the southern temperate forests, thus making them
biogeographically interesting (Keel & Herrera MacBryde, 1997). Atlantic Forest has
been designated one of the high priority habitats for conservation on a global scale
(Davis et al., 1997) and the Mbaracayu Reserve is the only example of this habitat
with effective protection in Paraguay.

The non-forested part of the reserve consists of savannah vegetation made up of
campos cerrados and other types of grasslands (see Table 1). Cerrado vegetation is
represented by a wide range of open woodland, open scrub and grassland forms
(Eiten, 1972), but is always found on uplands such that the soils are well-drained and
do not remain water-logged. In this way it is very different from other grass-
dominated vegetation types in South America (Eiten, 1972). Cerrado is a climax

224 FERN GAZETTE: VOLUME 15 PART 7 (1999)

Figure 2. Canopy of Atlantic forest, Salto de Karapa. (Photo S. Knapp).

Figure 3. Cerrado vegetation with Butia paraguayensis (Barb. Rodr.) L.H. Bailey,
Horqueta mi. (Photo S.Knapp).

PTERIDOPHYTES OF MBARACAYU FNR psp)

vegetation type and is not maintained by human interference such as annual burning
(Eiten, 1972). Paraguay is the southern limit for cerrado vegetation (Dinerstein et al.,
1995) which is largely found in Brazil and to a lesser extent in western Bolivia. In
Paraguay, cerrado vegetation is found mostly in the departments of Amambay,
Concepcion and Canindeyiu. In the Reserve the cerrado is a very biodiverse habitat,
with grasses, palms and legumes being prominent and an important part of the
vegetation (Fig. 3). Wet or semi-flooded grasslands also occur within the limits of
the Reserve; here sedges and grasses dominate and the shrubby component of the
vegetation is absent.

LIST OF SPECIES

The following enumeration lists 115 taxa, these belong to 22 families and 46 genera.
Many of these records are new published citations for Paraguay, these taxa are
indicated with an asterix in the checklist. We have listed as separate taxa species that
we have been unable to identify at present, but that are clearly different from other
species in the checklist (e.g. Thelypteris sp. #2). For these taxa, only habitat in the
Reserve and collections are cited - they are not included in the biogeographical
discussion. Names of authors of taxa are abbreviated in accordance with Pichi
Sermolli (1996). The taxonomic arrangement follows Flora Mesoamericana
(Davidse et al., 1995) and within families we use an alphabetical sequence for
genera, species and infraspecific taxa. Herbarium acronyms follow Holmgren et al.,
1990. In the list we have used the terms for forest types commonly in use in
Paraguay (FMB, 1991). A brief description of each forest type and the number of
fern species found therein is presented in Table 1.

* = new records for Paraguay
# = unidentified specimens

PSILOTACEAE

Psilotum nudum (L.) P. Beauv., Prodr. Aethéogam. 106, 112. 1805.

Lycopodium nudum L., Sp. Pl. 1100. 1753; Psilotum triquetum Sw., J. Bot. (Schrad.)
1800 (2):109. 1801., nom. superfl. for Lycopodium nudum L.; Psilotum triquetum
var. gracile Grev. & Hook., Enum. Fil. in Bot. Misc. 2:362. 1831.

Epiphyte. Bosque alto.
Distribution: New and Old World tropics and subtropics.

Material examined: in regione Yerbalium de Mbaracayi, Hassler 5805 (BM); Jejui
Mi, sendero principal, B. Jiménez 1254 (PY).

EQUISETACEAE

Equisetum giganteum L., Syst. Nat. ed. 10, 2:1318. 1759.
Equisetum schaffneri Milde, Verh. Zool.-Bot. Ges. Wien 11:345. 1861.

Terrestrial. Flooded grasslands.
Distribution: Mesoamerica, South America to Uruguay and Chile, Antilles.

Material examined: in dumeto ad marginem paludis pr. Igatimi, Hassler 5509 (BM);
Valinotti-cue, Jiménez & Marin 1295 (PY, BM, CTES, MO, RP)

226 FERN GAZETTE: VOLUME 15 PART 7 (1999)

LYCOPODIACEAE

Huperzia mandiocana (Raddi) Trev., Atti Soc. Ital. Sci. Nat. 17:248. 1874.
Urostachys mandiocanum (Raddi) Herter, Repert. Spec. Nov. Regni Veg. 19:164.
1923; Lycopodium mandiocanum Raddi, Opusc. Sci. 3: 280. 1819.

Epiphyte. Bosque medio, on Sorocea (Moraceae).
Distribution: Brazil, Paraguay.
Material examined: Carapa, Jiménez & Marin 0192.

Lycopodiella alopecuroides (L.) Cranfill, Amer. Fern J. 71:97.1981.

Lycopodium alopecuroides L., Sp. Pl. 1102. 1753; Lycopodium matthewsii Hook.,
Icon. Pl. 1: t 26. 1836; Lepidotis alopecuroides (L.) Rothm. Repert. Spec. Nov.
Regni Veg. 54:66. 1944; Plananthus alopecuroides (L.) P. Beauv., Prod. Aethéogam.
111. 1805; Lycopodiella matthewsii (Hook.) Holub, Folia Geobot. Phytotax. 18:441.
1983.

Terrestrial. Flooded grasslands.

Distribution: eastern temperate North America, Cuba, throughout continental tropical
America.

Material examined: in uliginosis pr. Ipé-hu, Hassler 5264 (BM); Carpa-cue, pastizal
sur, 1764 (BM, PY, CTES, RP); Valinotti-cue, Jiménez & Marin 2008 (BM, PY);
Valinotti-cue, Pena-Chocarro 336 (BM), Aguara Nu.

Lycopodiella caroliniana (L.) Pic. Serm., var. meridionalis (Underw. & F.E.
Lloyd) B. Ollg. & P.G. Windisch, Bradea 5:27. 1987.

Lycopodium meridionale Underw. & F.E. Lloyd, Bull. Torrey Bot. Club 33:121.
1906; Pseudolycopodiella meridionalis (Underw. & F.E. Lloyd) Holub, Folia
Geobot. Phytotax. 18:442. 1983.

Terrestrial. Flooded grasslands.
Distribution: widely distributed in tropical America, rare in the Andes.

Material examined: Aguara Nu, Jiménez 1465 (PY, BM, CTES); Carpa-cue, pastizal —
Sur, Jiménez 1772 (BM, PY).

Lycopodiella cernua (L.) Pic. Serm., Webbia 23:166. 1968.

Lycopodium cernuum L., Sp. Pl. 1103. 1753. Lycopodium cernuum var. capillaceum
Spring., Mém. Acad. Roy. Belg. 15 [Mon. Lyc. 1]:80. 1842; Lycopodium
capillaceum (Spring) Hieron., Bot. Jahrb. Syst. 34:573. 1905.; Palhinhaea cernua
(L.) Vasc. et Franco, Bol. Soc. Brot. 41:25. 1967.

Terrestrial. Flooded grasslands, river banks.
Distribution: pantropical.

Material examined: in palude pr. flumen Jejui-guazu, Hassler 4677 (BM); in palude
pr. Ipé-hu, Sierra de Maracayu, Hassler 5261 (BM); Fazenda Cerro Pora, Marin &
Jiménez 314 (PY, BM), Carpa-cue, pastizal sur, Jiménez 1763 (BM, PY, CTES,
MO); Lagunita, sendero arroyo Moroti, Jiménez & Marin, 1242 (BM); Valinotti-cue,
Jiménez & Marin 1287 (PY, BM, CTES, AA); Valinotti-cue, Pefia-Chocarro 337
(BM), Valinotti-cue, Pefia-Chocarro 342 (BM).

PTERIDOPHYTES OF MBARACAYU FNR 227

SELAGINELLACEAE
Selaginella marginata (Humb. & Bonpl. ex Willd.) Spring, Flora 21:194. 1838.

Lycopodium marginatum Humb. et Bonpl. ex Willd., Sp. Pl. ed. 4, 5:41. 1810;
Selaginella excurrens Spring in Martius, Fl. Bras. 1(2):128. 1840; Selaginella
marginata subsp. distorta Spring, Flora, Jena 21:196. 1838; Selaginella marginata
var. minor Spring I.c.; Selaginella distorta (Spring) Spring, Bull. Acad. r. Belg.
10:229. 1843; Selaginella distorta var. minor (Spring) Spring, Mém. Acad. r. Sci.
Lett. Belg. 24:213; Selaginella distorta var. major Baker, J. Bot., Lond. 21:335.
1883; Selaginella chromatophylla Silveira, Bolm Comm. Geogr. Geol. Minas Geraes
5:124. 1898; Selaginella urbani Hieron., in Engl. & Prantl, Nat. Pflanzenfam. 1
(4):709, no. 297. 1901 (1902); Selaginella moseni Hieron., in Engl. & Prantl, Nat.
Pflanzenfam. 1 (4):709, no. 400. 1901 (1902); Selaginella burchellii Hieron., in Engl.
& Prantl, Nat. Pflanzenfam. 1 (4):709, no. 403. 1901 (1902).

Terrestrial. Flooded grasslands.
Distribution: Mexico, Venezuela, Bolivia, Brazil, Uruguay, Paraguay and Argentina.

Material examined: Jejui Mi, pastizal sur, Jiménez & Marin 1261 (BM, PY), Jejui
Mi, pastizal sur, Jiménez & Marin 1278 (BM, PY).

Selaginella muscosa Spring in Martius, Fl. Bras. 1 (2):120. 1840.

Lycopodium brasiliense Raddi, Pl. Bras. Nov. Gen.:82. 1825; Lycopodium
crassinervium Desv., Mém. Soc. Linn. Paris 6:190. 1827; Lycopodium pallidum
Beyr. ex Gaudich. in Freyc., Voyage Autour Monde, Bot.:285. 1828. Nomen
abortivum; Lycopodium albidulum var. majus Hook. & Grev., Bot. Misc. 2:399.
1831, Selaginella crassinervia (Desv.) Spring in Martius, Fl. Bras. 1(2):119. 1840;
Selaginella apus var. tetragonostachya Spring in Martius, Fl. Bras. 1(2):119. 1840;
Selaginella polysperma Spring, Bull. Acad. R. Belg. 10:138. 1843; Selaginella
beyrichii A. Braun, Index Sem. Hort. Bot., Berl. Appendix 1857:16. 1857;
Selaginella anocardia A. Braun, Ann. Sci. Nat. (Bot.) V, 3:290. 1865, non (Desv.)
Spring. 1838; Selaginella deltoides A. Braun, Ann. Sci. Nat. (Bot.) V, 3:287. 1865;
Alston, Reprium Spec. nov. Regni veg. 45:312. 1936; Selaginella brevipes Fée,
Crypt. Brés.:226. 1869, non A. Braun. 1867; Selaginella trifurcata Baker, J. Bot.,
Lond. 21:98. 1887; Alston, J. Bot., Lond. 72:38. 1934; Knox, Trans. Bot. Soc. Edinb.
35:278. 1950; Selaginella niederleinii Hieron., Bot. Jb. 22:418. 1896; Selaginella feei
Hieron. in Engl. & Prantl, Nat. Pflanzenf. 1 (4):713. 1901; Selaginella brasiliensis
var. crassinervia Hieron. ex Bonap., Notes Ptérid. 2:137. 1915; Selaginella os
sensu Hieron., Hedwigia 58:321. 1917, non Jenman. 1897.

Terrestrial. Flooded grasslands.

Distribution: Tobago, Trinidad and Guyana, west to Colombia and south to northern
Brazil; also Argentina, Uruguay and Paraguay.

Material examined: Jejui Mi, 2 km antes del puesto principal, Jiménez & Marin 1965
(PY, BM).

228 FERN GAZETTE: VOLUME 15 PART 7 (1999)

MARATTIACEAE

* Danaea nodosa (L.) Sm., Mem. Acad. Roy. Sci. (Turin) 5:420. 1793.
Acrostichum nodosum L., Sp. Pl. 1070. 1753; Danaea longifolia Desv., Ges. Naturf.
Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5:307. 1811.

Terrestrial. Gallery forest.

Distribution: S. Mexico to Panama, West Indies, Colombia, Venezuela, Surinam,
Ecuador, Brazil, Peru, Paraguay.

Material examined: camino Jejui Mi a Lagunita, arroyo Pira-Cajon, Pefia-Chocarro
359 (BM).

OSMUNDACEAE

Osmunda cinnamomea L., Sp. Pl. 1066. 1753.

Osmundastrum cinnamomeum (L.) C. Presl, Abh. B6hm. Ges. Wiss. 5:326. 1848;
Osmunda imbricata G. Kunze, Farrnkr. 2:29. 1849; Osmunda bipinnata L., Sp. Pl.
LOGS: 1753:

Terrestrial. Swamps with Sphagnum.
Distribution: eastern and central United States and Canada, West Indies, S. Mexico,
Mesoamerica, South America to Paraguay and Brazil, South East Asia.

Material examined: Lagunita, sendero arroyo Moroti, Jiménez & Marin 1516 (BM,
PY); Lagunita, sendero Arroyo Moroti, Pefia-Chocarro 214 (BM, PY).

Osmunda regalis L., Sp. Pl. 1065. 1753.

Osmunda mexicana Fée, Mém. Foug. 9:43. 1857; Osmunda palmeri A.E. Bobrov,
Novesti Sist. Vyssh. Rast. 1968:8. 1968; Osmunda regalis L. var. spectabilis (Willd.)
A. Gray, Manual ed. 2:600. 1856; Osmunda spectabilis Willd., Sp. Pl. ed. 4, 5:98.
1810; Osmunda palustris Schrad., Gott. Geh. Anz. 866. 1824; Aphyllocarpa regalis
(L.) Cav., Anal. Cienc. 5:164. 1802.

Terrestrial. Swamps with Sphagnum.
Distribution: cosmopolitan.
Material examined: in sylva pr. flumen Tapiraguay, Hassler 4150 (BM).

SCHIZAEACEAE

Anemia phyllitidis (L.) Sw., Syn. Fil. 155. 1806.
Osmunda phyllitidis L., Sp. Pl. 1064. 1753; Anemia haenkei C. Presl, Relig. Haenk.
17451825,

Terrestrial. Bosque alto, disturbed bosque alto, bosque medio, gallery forest.

Distribution: Mexico to Panama, Greater Antilles, Trinidad, Venezuela and
Colombia to Argentina and Uruguay.

Material examined: in dumeto pr. flu. Jejui-guazu, Hassler 5704 (BM); Jejui Mi,
sendero Aguara’ i, Jiménez & Marin 133 (BM, PY, CTES); La Morena, Jiménez &
Marin 1348 (PY, BM, CTES, RP); Carapa, Pefia-Chocarro 227 (BM, PY); Lagunita,
sendero arroyo Moroti, Pefia-Chocarro 232 (BM, PY).

PTERIDOPHYTES OF MBARACAYU FNR 229

Anemia phyllitidis var. tweediana (Hook.) Hassl., Trab. Inst. Bot. Farm. Fac. C.
Méd. Bs. As. 45:85. 1928.
Anemia tweediana Hook., Icon. Pl. t. 906. 1854.

Terrestrial. River banks.
Distribution: E. Paraguay, S. Brazil, Uruguay, N.E. Argentina.
Material examined: Jejui Mi, 1500 m N caseta, Marin & Jiménez 192 (BM, PY).

Anemia tomentosa (Sav.) Sw. var. anthriscifolia (Schrad.) Mickel, Iowa State J.
Sci. 36 (4):424. t. 28. 1962.
Anemia anthriscifolia Schrad., Gott. Gel. Anz. 1824:625. 1824.

Terrestrial. Cerrado, pastures with Butia (Arecaceae).
Distribution: Brazil, Bolivia, Paraguay and Argentina.

Material examined: Lagunita, Marin & Jiménez 152 (PY); entrada a Aguara Nu,
Jiménez & Pena-Chocarro 1240 (BM, PY).

Anemia tomentosa (Sav.) Sw. var. tomentosa Syn. Fil. 157.1806; Mickel, Iowa
Siate J. Sci. 36 (4):423. 2. 27. f. B. 1962.
Osmunda tomentosa Savigny in Lam., Encycl. 4:652. 1797.

Terrestrial. Cerrado, pastures.
Distribution: S. Brazil, E. Paraguay, Uruguay and Argentina.
Material examined: Lagunita, sendero arroyo Moroti, Pefia-Chocarro 262 (BM).

Lygodium volubile Sw., J. Bot. (Schrad.) 1801 (2):304. 1803.

Lygodium micans J. W. Sturm in Mart., Fl. Bras. 1(2):178. 1859.

Terrestrial, scandent on shrubs and low trees. Along streams and river banks, bosque
alto.

Distribution: S. Mexico to Panama, Trinidad, Greater Antilles, Venezuela to the
Guianas, south to Argentina and Brazil.

Material examined: Jejui Mi, Rumbo Norte, Marin & Jiménez 150 (PY, BM, CTES);
Jejui Mi, Rumbo Norte, Pefia-Chocarro 255 (BM); Valinotti-cue, Pefia-Chocarro
287 (BM).

GLEICHENIACEAE

Dicranopteris flexuosa (Schrad.) Underw., Bull. Torrey Bot. Club 34:254. 1907.
Mertensia flexuosa Schrad., Gott. Gel. Anz. 863. 1824; Mertensia rigida Kuntze,
Linnaea 9:16. 1834; Gleichenia flexuosa (Schrad.) Mett., Ann. Mus. Bot. Lugduno-
Batavum 1:50. 1863; Gleichenia rigida (Kuntze) J. Bommer & Christ, Bull. Soc.
Roy. Bot. Belgique 35:174. 1896.

Terrestrial. Cerrado, river banks in cerrado habitat.

Distribution: pantropical.

Material examined: procumbens in argillosis pr. Igatimi, Hassler 4864 (BM);
Valinotti-cue, Jiménez & Marin 1285 (PY, BM, CTES, RP); Aguara Nu, camino a la

230 FERN GAZETTE: VOLUME 15 PART 7 (1999)

comunidad Tekoha Ryapu, Jiménez & Marin 2039 (BM, MO, CTES, G); Aguara Nu,
Marin & Jiménez 247 (PY, BM).

HYMENOPHYLLACEAE

Trichomanes cristatum Kaulf., Enum. Filic. 265. 1824.
Trichomanes sellowianum C. Presl, Hymenophyllaceae 15, 37. 1843.

Terrestrial. Swamps with Sphagnum.
Distribution: Venezuela and Colombia south to Brazil and Argentina.

Material examined: in sylvis Sierra de Maracayié, Hassler 5143 (BM); Lagunita,
sendero arroyo Moroti, Jiménez & Marin 1508 (BM, PY); Lagunita, sendero arroyo
Moroti, Pefia-Chocarro 216 (BM, PY).

* Trichomanes diaphanum Humb., Bonpl. & Kunth, Nov. Gen. Sp. 1:25. 1816.

Trichomanes leptophyllum Bosch, Ned. Kruidk. Arch.4:363. 1859, non A. Cunn.,
1836, nom illeg.; Trichomanes hymenophylloides Bosch, Ned. Kruidk. Arch. 5:209.
1863, nom. nov. for T. leptophyllum Bosch and type based on that name;
Vandesboschia diaphana (Humb., Bonpl. & Kunth) Copel., Philipp. J. Sci. 67:53.
1938; Vandesboschia hymenophylloides (Bosch) Copel., Philipp. J. Sci. 67:53. 1938.

Epiphyte. Gallery forest.

Distribution: S. Mexico to Panama, West Indies, Trinidad, Guianas to Colombia,
south to Paraguay and Brazil.

Material examined: Jejui Mi, sendero Aguara’i, arroyo Amambay, Pefia-Chocarro
325 (BM); camino Jejui Mi a Lagunita, arroyo Pira-Caj6n, Pefia-Chocarro 350
(BM); camino Jejui Mi a Lagunita arroyo Pira-Cajon, Pefia-Chocarro 360 (BM)

Trichomanes radicans Sw., J. Bot. (Schrad.) 1800 (2):97. 1802.

Trichomanes kunzeanum Hook., Sp. Filic. 1:127. 1884; Trichomanes brachyblastos
Kuhn, Linnaea 35:388. 1868; Vandenboschia radicans (Sw.) Copel., Philipp. J. Sci.
67:54. 1938; Trichomanes radicans var. kunzeanum (Hook.) Duek & Lellinger,
Amer. Fern J. 68:120. 1978; Crepidomanes radicans (Sw.) K. Iwats., J. Fac. Sci.
Univ. Tokyo, sect. 3, Bot. 13 (5):530. 1985.

Hemi-epiphyte. Gallery forest.
Distribution: cosmopolitan.
Material examined: Jejui Mi, Jiménez & Marin 155 (BM, PY, CTES, RP); Jejui Mi,

sendero Aguara’ i, arroyo Amambay, Pefia-Chocarro 324 (BM); camino Jejui Mi a
Lagunita, arroyo Pira-Cajon, Pefia-Chocarro 358 (BM).

Trichomanes rigidum Sw., Prodr. 137. 1788.

Cephalomanes rigidum (Sw.) K. Iwats. Acta Phytotax. Geobot. 35 (4-6):177. 1984;
Selenodesmium rigidum (Sw.) Copel., Philipp. J. Sci. 67:81. 1938; Trichomanes
daucoides C. Presl, Epim. Bot. 12, t. 7. 1849.

Terrestrial. River banks.

PTERIDOPHYTES OF MBARACAYU FNR 231

Distribution: S. Mexico to Panama, West Indies, South America to Brazil and
Paraguay, Old World tropics.

Material examined: Lagunita, arroyo Moroti, Marin & Jiménez 238 (PY, BM);

Lagunita, sendero arroyo Moroti, Pefia-Chocarro 270 (BM).

CYATHEACEAE

Cyathea atrovirens (Langsd. & Fisch.) Domin, Pterid. 262. 1929.

Alsophila atrovirens (Langsd. & Fisch.) C. Presl, Tent. Pterid. 61. 1836; Polypodium
atrovirens Langsd. & Fisch., Pl. Voy. Russes Monde (Icon. Filic.) 12, t. 14. 1810;
Trichipteris atrovirens (Langsd. & Fisch.) R. M. Tryon, Contr. Gray Herb. 200:45.
1970.

Terrestrial. Disturbed Bosque alto, bosque medio on saturated soil, bosque bajo on
saturated soils.

Distribution: S. Brazil, Paraguay.

Material examined: in dumetis humidis pr. San Estanislao, Hassler 4173 (BM); Jejui
Mi, Jiménez & Marin 113 (BM, PY, CTES); Jejui Mi, Ycua, Jiménez & Marin 123
(PY); Jejui Mi, Jiménez & Marin 125 (BM, PY); camino Lagunita-Horqueta-Mi,
entre Km 18 y entrada arroyo Moroti, Marin & Jiménez 231 (PY, RP); Jejui Mi,
sendero Aguara’i, Marin & Jiménez 510 ( BM, RP); Lagunita, sendero arroyo
Moroti, Peria-Chocarro 220 (BM, PY).

Alsophila cuspidata (Kuntze) D.S. Conant, J. Arnold Arbor. 64:371. 1983.
Nephelea cuspidata (Kuntze) R. M. Tryon, Contr. Gray Herb. 200:40. 1970; Cyathea
cuspidata Kuntze, Linnaea 9:101. 1834.

Terrestrial. Gallery forest.
Distribution: Nicaragua to Panama, northern South America to Bolivia and Paraguay.

Material examined: camino Lagunita Horqueta-Mi, arroyo Pira Caj6n, Marin &
Jiménez 217 (PY, BM, CTES, MO); Carapa, sendero Uru’i, Marin & Jiménez 332
(PY, BM, CTES, MO, RP).

PTERIDACEAE

Adiantopsis chlorophylla (Sw.) Fée, Mém. Foug. 5:145. 1852.
Cheilanthes chlorophylla Sw., Kongl. Vetensk. Acad. Handl. 1817:76. 1817.

Terrestrial. Disturbed and open areas in bosque alto, pastures.

Distribution: Mexico, Guatemala, Costa Rica, Colombia south to Argentina and
Brazil.

Material examined: camino Jejui Mi a Horqueta-Mi, entre Km 18 y sendero arroyo
Moroti, Marin & Jiménez, 232 (PY); Jejui Mi, sendero principal Km1-2, Pefa-
Chocarro 205 (BM, PY); Valinotti-cue, Pefia-Chocarro 285 (BM).

Adiantopsis radiata (L.) Fée, Mém. Foug. 5:145. 1852.
Adiantum radiatum L., Sp. Pl. 1904. 1753; Cheilanthes radiata (L.) J. Sm., J. Bot.
(Hook.) 4:159. 1841; Adiantopsis ternata Prantl, Gartenfl. 32:101. 1893.

Mele FERN GAZETTE: VOLUME 15 PART 7 (1999)

Terrestrial. Bosque medio, bosque medio on saturated soil.

Distribution: tropical America.

Material examined: Jejui Mi, sendero Aguara-i, Jiménez & Marin 128 (PY, BM,
CTES); Jejui Mi, sendero Jaku-apeti, Marin & Jiménez 303 (BM, PY, CTES);
Lagunita, sendero arroyo Moroti, Pefia-Chocarro 261 (BM).

Adiantum latifolium Lam., Encycl. 1:43. 1783.
Adiantum humile Kunze, Linnaea 9:80. 1834; Adiantum killipii Maxon & Weath.,
Amer. J. Bot. 19:166. 1932.

Terrestrial. Bosque alto, bosque medio, bosque bajo.

Distribution: Mexico, Mesoamerica, Antilles, Trinidad, Colombia to Venezuela south
to Brazil and Paraguay.

Material examined: Jejui Mi, Jiménez & Marin 137 (BM, PY, CTES); Jejui Mi,
Rumbo Norte, Pefia-Chocarro 259 (BM); Jejui Mi, sendero Aguara’i, Pefa-
Chocarro 327 (BM).

* Adiantum petiolatum Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Endeck.
Gesammten Naturk. 5:326. 1881.

Adiantum kaulfussiti Kunze, Linnaea 21:221. 1848; Adiantum obliquum Kaulf.,
Enum. Filic. 200. 1824, non Willd.; Adiantum oblongatum Schrad., Gott. Gel. Anz.
1824:872. 1824.

Terrestrial. River banks.
Distribution: from S. Mexico to Brasil and Paraguay.
Material examined: camino Lagunita a Horqueta-Mi, arroyo Moroti, Marin &

Jiménez 239 (PY); Lagunita, sendero Arroyo Moroti, Pefia-Chocarro 231 (BM, PY);
Jejui Mi, sendero Jaku-apeti, Peria-Chocarro 209 (BM, PY).

Adiantum pseudo-tinctum Hieron., Engl. Bot. Jahrb. 22. 394. 1896.

Terrestrial. Open areas in bosque alto, degraded bosque medio.

Distribution: Brazil, Argentina and Paraguay.

Material examined: Jejui Mi, sendero principal Km 1, Pena-Chocarro 200 (BM,

PY); Jejui Mi, Rumbo Norte, Pefia-Chocarro 244 (BM); camino Jejui Mi a Lagunita
Km. 4-5, Penia-Chocarro 323 (BM).

Adiantum raddianum C. Pres], Tent. Pterid. 158. 1836.

Adiantum cuneatum Langsd. & Fisch., Pl. Voy. Russes Monde (Icon. Filic.) 23, t. 26.
1810; Adiantum tinctum Moore, Gard. Chron. 1862:932; Adiantum amabile Moore,
Gard. Chron. 1868:1090. 1868, non Liebm. 1849; Adiantum decorum Moore, Gard.
Chron. 1869:582. 1869; Adiantum moorei Baker, Gard. Chron. 1873:811. 1873,
based on A. amabile Moore; Adiantum werckleanum Christ, Bull. Herb. Boisser, sér.
2, 4:1093. 1904.

Terrestrial. Bosque medio on saturated soil, gallery forest.

PTERIDOPHYTES OF MBARACAYU FNR 29

Distribution: S. Mexico, Mesoamerica, Antilles, Trinidad, Colombia, south to Chile
and Uruguay.

Material examined: salto Carapa, Marin & Jiménez 339 (PY); Carapa, camino al
salto, Penia-Chocarro 310 (BM).

Adiantum serratodentatum Humb. et Bonpl. ex Willd., Sp. Pl. ed. 4, 5:445. 1810.
Terrestrial. Bosque medio, pastures, flooded grasslands.
Distribution: tropical America.

Material examined: in sylva Sierra de Maracayu, Hassler 5399 (BM); Lagunita,
Jiménez & Marin 48 (BM, PY, CTES); pastizal Sur, Jejui Mi, Jiménez & Marin 1247
(PY, BM, CTES, MO, RP); entrada a Lagunita, Pefia-Chocarro 213 (BM, PY);
Lagunita, Pefia-Chocarro 233 (BM, PY); Valinotti-cue, Pefia-Chocarro 284 (BM);
Valinotti-cue, Pefia-Chocarro 338 (BM); rio Jejui-Guazu, Rojas 362] (BM).

Adiantum tetraphylium Humb. et Bonpl. ex Willd., Sp. Pl. ed. 4, 5:441. 1810.
Terrestrial. River banks, bosque medio, bosque bajo.
Distribution: from S. Mexico to Brazil and Paraguay.

Material examined: in sylva Sierra de Maracayu, Hassler 5149 (BM); Fazenda Cerro
Pora, Marin & Jiménez 313 (PY); Lagunita, sendero arroyo Moroti, Pefia-Chocarro
230 (BM, PY); camino Jejui Mi a Lagunita Km 11, Pefia-Chocarro 334 (BM).

Doryopteris concolor (Langsd. & Fisch.) Kuhn., Bot. Ost-Afrika 3(3):19. 1879.
Pteris concolor Langsd. & Fisch., Pl. Voy. Russes Monde (Icon. Filic.) 19. 1810;
Cheilanthes concolor (Langsd. & Fisch.) R.M. Tryon & A. F. Tryon, Rhodora
m1 35. 1981:

Terrestrial. Bosque alto, disturbed areas in bosque medio-alto.

Distribution: tropical America, Asia, Africa, Australia.

Material examined: La Morena, Jiménez & Marin 1341 (PY, BM, CTES, MO, RP);
Carapa, Pefia-Chocarro 226 (BM, PY); Jejui Mi, sendero Jaku-apeti, Pefia-Chocarro
234 (BM, PY).

Doryopteris lomariacea Klotzsch, Linnaea 20:343. 1847.
Terrestrial. Swamps with Sphagnum.
Distribution: Guyana, Peru, S. Brazil and Paraguay.

Material examined: in sylva Sierra de Mbaracayu, Hassler 5191(BM); in sylva pr. fl.
Jejui-guazu, Hassler 5680 (BM); Lagunita, sendero arroyo Moroti, Jiménez & Marin
1513 (BM, PY, CTES, MO, RP); Lagunita, sendero arroyo Moroti, Pefia-Chocarro
213 (SN, PY).

234 FERN GAZETTE: VOLUME 15 PART 7 (1999)

Doryopteris nobilis (Moore) C. Chr., Index Filic. 244. 1905.

Litobrochia nobilis Moore, Gard. Chron. Oct. 1862:932. 1862; Doryopteris raddiana
var. patula Fée, Crypt. Vasc. Brésil 1:45. 1869; Doryopteris patula (Fée) Fée, Crypt.
Vasc. Brésil 2:30. 1872-3.

Terrestrial. Bosque medio, gallery forest.
Distribution: Paraguay, Brazil, Argentina.

Material examined: in sylva pr. fl. Tapiraguay, Hassler 4372 (BM); in sylva Sierra de
Maracayu, in sylva pr. fl. Jejui-guazu, Hassler 570Ja (BM); in sylva pr. fl. Jejui-
guazu, Hassler 5701b (BM); Hassler 5388 (BM); Jejui Mi, sendero Jaku-apeti,
Jiménez & Marin 309 (BM, PY); salto de Carapa, Marin & Jiménez 334 (PY, BM,
RP); Lagunita, sendero arroyo Moroti, Pefia-Chocarro 268 (BM); Carapa, camino al
salto, Peria-Chocarro 309 (BM)

Doryopteris pedata (L.) Fée var. multipartita (Fée) R. M. Tryon, Contr. Gray
Herb. Harvard Univ. 143:38, t. 5b. 1942.

Doryopteris raddiana var. multipartita Fée, Crypt. Vasc. Brésil 1:45. 1869;
Doryopteris multipartita (Fée) Sehnem, FI. Ilustr. Catar. I, Pteridac., 176. 1972.
Terrestrial. Bosque alto, disturbed bosque alto, bosque medio, gallery forest.
Distribution: Bolivia, Paraguay, Brazil, Argentina.

Material examined: Jejui Mi, Jiménez & Marin 142 (BM, PY, CTES); Jeyui Mi,
sendero Jaku-apeti, Jiménez & Marin 308 (BM, PY, CTES); Jejui Mi, sendero Jaku-
apeti, Marin & Jiménez 310 (PY); salto de Carapa, Marin & Jiménez 338 (PY, BM,
RP); Jejui Mi, sendero principal, Marin & Jiménez 460 (PY, BM, CTES); Carapa,
Pena-Chocarro 225 (BM, PY); Jejui Mi, sendero Jaku-apeti, Pefia-Chocarro 235
(BM, PY); Carapa, camino al salto, Pefia-Chocarro 295 (BM).

* Pityrogramma calomelanos (L.) Link var. austroamericana (Domin) Farw.,
Amer. Midl. Naturalist 12:280.1931.

Pityrogramma austroamericana Domin, Publ. Fac. Sci. Univ. Charles 88:7. 1928;
also in Bull. Misc. Inform. 1929:221; Pityrogramma calomelanos var. aureoflava
auct., non (Hook.) Weath. ex Bailey, Man. Cult. Plants: 64. 1926.

Terrestrial. Open areas, in saturated soils.

Distribution: Mesoamerica, South America to N.E. Argentina, Brazil and Paraguay.

Material examined: Jejui Mi, Jiménez & Marin 1255 (BM, PY, CTES); Jejui Mi,
sendero principal Km1-2, Pefia-Chocarro 203 (BM, PY).

Pityrogramma calomelanos (L.) Link, var. calomelanos Handbuch 3:20.1883.
Acrostichum calomelanos L., Sp. Pl. 1072. 1753; Acrostichum ebeneum L., Sp. PI.
1071.1753; Gymnogramma calomelanos var. denudata Harr., J. Linn. Soc., Bot.
16:37. 1877; Pityrogramma ebenea (L.) Proctor, Brit. Fern Gaz. 9:219.1965.

Terrestrial. Open and disturbed areas, flooded grasslands.
Distribution: Mexico, Mesoamerica, South America to Paraguay and Argentina.

Material examined: in sylva apric apr. Igatimi, Hassler 4810 (BM); Jejui Mi, Jiménez
& Marin 75 (BM, PY); Valinotti cue, Jiménez & Marin 1291 (PY, BM, CTES, MO,

PTERIDOPHYTES OF MBARACAYU FNR 235

RP); ampliacion de la Reserva, Jiménez 1757 (BM, PY, CTES, MO); camino Jejui
Mi a Horqueta-Mi, Marin & Jiménez 233 (PY); Lagunita, Marin & Jiménez 271 (PY,
BM, CTES, MO, GENEVA, RP); Jejui Mi, sendero principal Km1-2, Pefa-
Chocarro 202 (BM, PY); Valinotti-cue, Pefia-Chocarro 339 (BM); Valinotti-cue,
Pena-Chocarro 340 (BM); Valinotti-cue, Pefia-Chocarro 344 (BM).

Pityrogramma trifoliata (L.) R. Tryon, Contr. Gray Herb. 189:68. 1962.
Acrostichum trifoliatum L., Sp. Pl. 1070. 1753; Trismeria microphylla Fée, Mém.
Foug. 5:165. 1852; Trismeria trifoliata (L.) Diels, Nat. Pflanzenfam. 1(4):265. 1899.
Terrestrial. Pastures, open areas.

Distribution: Mexico, Mesoamerica, South America to Chile, N. Argentina and
Uruguay.

Material examined: Carapa, camino al salto, Pefia-Chocarro 316 (PY, BM).

* Pteris altissima Poir. in Lam., Encycl. 5:722. 1804.

Litobrochia kunzeana (J. Agardh) Fée, Mém. Foug. 5:132. 1850-52; Pteris elata
J. Agardh, Recens. Spec. Pter. 63. 1839; Pteris kunzeana J. Agardh, Recens. Spec.
Pter. 62. 1839; Pteris protea Liebm., Kongel. Danske Vidensk. Selsk. Skr.
Naturvidensk. Afd. 5, 1:228 (seors 76). 1849.

Terrestrial. Gallery forest.

Distribution: from S. Mexico to S. Brazil and Paraguay.

Material examined: Carapa, sendero Uru’i, Marin & Jiménez 331 (PY, BM, CTES,
RP); Carapa, camino al Salto, Pefia-Chocarro 297 (PY, BM, RP).

Pteris deflexa Link, Hort. Berol. 2:30. 1833.
Pteris polita Link, Hort. Berol. 2:30. 1833.

Terrestrial. Bosque alto.
Distribution: tropical South America.

Material examined: Jejui Mi, Rumbo Norte, Pefia-Chocarro 250 (BM); Jejui Mi,
sendero principal, Km 4, Pefia-Chocarro 322 (BM).

Pteris denticulata Sw., Prodr. 129.1788.

Pteris brasiliensis Raddi, Opusc. Sci. 3:293. 1819; Litobrochia denticulata (Sw.)
C. Presl., Tent. Pterid. 149, t. 5, f. 20. 1836.

Terrestrial. Bosque alto, bosque alto on saturated soil, bosque medio, gallery forest.
Distribution: Paraguay, Brazil and Argentina.

Material examined: Jejui Mi, Jiménez & Marin 138 (BM, PY); Jejui Mi, Jiménez &
Marin 151 (PY, BM, CTES, RP); La Morena, Jiménez & Marin 1336 (PY, BM,
CTES); Jejui Mi, sendero Aguara’i, Jiménez 1784 (PY, BM, CTES, MO); Lagunita,
sendero arroyo Moroti, Marin & Jiménez 166 (PY); Jejui Mi, sendero Jaku apeti,
Marin & Jiménez 305 (PY, BM, CTES, MO, G).

236 FERN GAZETTE: VOLUME 15 PART 7 (1999)

Pteris quadriaurita Retz., Observ. Bot. 6:38. 1791.
Pteris plumula Desv., Mém. Soc. Linn. Paris 6:297. 1827; Pteris edentula Kunze,
Linnaea 9:75. 1834.

Terrestrial. River banks.
Distribution: tropical America, Old World tropics.

Material examined: camino Lagunita a Horqueta-Mi, arroyo Pira Caj6n, Pefa-
Chocarro 356 (PY, BM, RP).

Hemionitis tomentosa (Lam.) Raddi, Opusc. Sci. 284. 1819.
Asplenium tomentosum Lam., Encycl. 2:308. 1786; Gymnoptenris tomentosa (Lam.)
Underw., Bull. Torrey Bot. Club 29:627. 1902.

Terrestrial. Gallery forest.
Distribution: Peru to Argentina, Paraguay and Brazil.

Material examined: Carapa, camino al salto, Pefia-Chocarro 308 (BM); Carapa, el
salto, Peria-Chocarro 314 (BM).

VITTARIACEAE

Vittaria lineata (L.) Sm., Mém. Acad. Roy. Sci. (Turin) 5:421. 1793.
Pteris lineata L., Sp. Pl. 2:1073. 1753; Vittaria filiformis Cav., Descr. Pl. 270. 1802.

Epiphyte. Cerrado, bosque bajo on saturated soil, gallery forest, epiphytic on
Syagrus romanzoffiana (Arecaceae).

Distribution: tropical America.

Material examined: Jejui Mi, Rumbo norte, Jiménez & Marin 0176 (PY); Aguara Nu,
1996, Jiménez & Albert 1437 (PY, BM); Lagunita, sendero arroyo Moroti, Pefia-
Chocarro 269 (BM); Carapa, camino al salto, Pefia-Chocarro 30] (BM).

DENNSTAEDTIACEAE

Dennstaedtia globulifera (Poir.) Hieron., Bot. Jahrb. Syst. 34:455. 1904.
Polypodium globuliferum Poir. in Lam., Encycl. 5:554. 1804.

Terrestrial. Bosque alto, gallery forest.

Distribution: Mexico, Mesoamerica, Venezuela and Colombia, south to Argentina
and southern Brazil.

Material examined: in campo pr. fl. Jejui-guazu, Hassler 5702 (BM); Jejui Mi, Jiménez
& Marin 129 (BM, PY, CTES); Carapa, camino al salto, Pefia-Chocarro 298 (BM);
Jejui Mi, camino a Lagunita, Km1, Perfia-Chocarro 362 (BM).

Pteridium aquilinum (L.) Kuhn var. arachnoideum (Kaulf.) Brade, Zeitschrift
Deut. Var. Wissen. Kunst, Sao Paulo 1:56. 1920.

Terrestrial. Clearings in bosque bajo.
Distribution: from Mexico to South America.
Material examined: Lagunita, Marin & Jiménez 269 (PY).

PTERIDOPHYTES OF MBARACAYU FNR 257

Lindsaea guianensis (Aubl.) Dryand. subsp. lanceastrum K.U. Kramer, Acta Bot.
Neer: 6:217.:1957.

Terrestrial. Bosque alto on saturated soil, bosque bajo on saturated soil.
Distribution: C. E. S. Brazil and Paraguay.
Material examined: Jejui Mi, Rumbo Norte, Jiménez & Marin 172 (BM); Lagunita,

sendero arroyo Moroti, Jiménez & Marin 1514 (BM, PY, CTES, RP); Jejui Mi,
sendero Jaku-apeti, Marin & Jiménez 301 (PY, BM, MC, GENEVA, RP).

Lindsaea lancea (L.) Bedd., Suppl. Ferns Brit. Ind. 6. 1876.
Adiantum lanceum L., Sp. Pl. ed. 2, 1557. 1763.

Terrestrial. Bosque alto, bosque medio, bosque bajo, on saturated soil.
Distribution: tropical America.

Material examined: in sylva pr. Curuguaty, Hassler 4614 (BM); in sylva pr. Igatimi,
Hassler 4698 (BM); arroyo Moroti, Marin & Jiménez 167 (PY); Lagunita, sendero
Jaku-apeti, Pefia-Chocarro 210 (BM); sendero Jaku-apeti, Pefia-Chocarro 211
(BM).

Lindsaea portoricensis Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Endeck.
Gestammten Naturk. 5:326. 1811.

Terrestrial. Swamps with Sphagnum.

Distribution: Mexico to Guatemala, Greater Antilles, Guianas to Colombia, south to
Bolivia, Brazil and Paraguay.

Material examined: Lagunita, sendero arroyo Moroti, Pefia-Chocarro 218 (BM, PY);
Lagunita, sendero arroyo Moroti, Pefia-Chocarro 228 (BM, PY); Lagunita, sendero
arroyo Moroti, Pefia-Chocarro 229 (BM, PY).

Lindsaea quadrangularis Raddi, Opusc. Sci. 3:294. 1819.

Terrestrial. Bosque medio on saturated soils.

Distribution: Brazil, Paraguay.

Material examined: Jejui Mi, sendero Jaku-apeti, Marin & Jiménez 302 (PY, BM,
CTES, MO, G; RP).

THELYPTERIDACEAE

Macrothelypteris torresiana (Gaudich.) Ching., Acta Phytotax. Sinica 8:310. 1963.
Polystichum torresianum Gaudich. in Freyc., Voy. Uranie. 333. 1828; Aspidium
uliginosum Kunze, Linnaea 20:6. 1847; Dryopteris uliginosa (Kunze) C. Chr., Index
Filic. suppl. 3:100. 1934; Thelypteris torresiana (Gaudich.) Alston, Lilloa 30:111.
1960.

Terrestrial. Bosque alto, bosque medio, bosque medio on saturated soils, river banks,
bosque bajo.

238 FERN GAZETTE: VOLUME 15 PART 7 (1999)

Distribution: from S.E. United States to S. Brazil, Paraguay and N. Argentina,
tropical and subtropical Africa and Asia, Pacific Islands. Adventive in the New
World tropics and subtropics.

Material examined: Aguara Nu, Valinotti-cue, Jiménez & Marin 1288 (PY, BM);
Jejui Mi, sendero Jaku-apeti, Marin & Jiménez 31] (PY, BM, CTES, MO, G,RP);
Jejui Mi, sendero Aguara-i, arroyo Amambay, Marin & Jiménez 515 (PY); Jejui Mi,
Rumbo Norte, Pefia-Chocarro 258 (BM); camino Jejui Mi a Lagunita, Km 18,
arroyo Moroti, Pefria-Chocarro 289 (BM); Carapa, camino al salto, Pefia-Chocarro
299 (BM); camino Jejui Mi a Lagunita, Km 5, Pefia-Chocarro 318 (BM).

* Thelypteris (Amauropelta) abbiattii Reed, Phytologia 17 (4):257. 1968.
Goniopteris burkartii C. Chr. ex Abiatti, Darwiniana 13 (2-4):556. 1964, non
T. burkartii Abiatti, 1964.

Terrestrial. Gallery forest on saturated soils, river banks.
Distribution: Argentina, Paraguay.
Material examined: Jejui Mi, sendero Aguara-i, Jiménez 1783 (PY, BM, CTES, RP);

Jejui Mi, 1500 m N de la caseta, Marin & Jiménez 189 (PY, BM, RP); Jejui Mi,
sendero Aguara’i, Penia-Chocarro 346 (BM).

* Thelypteris (Goniopteris) gemmulifera (Hieron.) A.R. Sm., Opera Bot. 56:30.
1980.

Dryopteris gemmulifera Hieron., Hedwigia 46:326. 1907; Goniopteris gemmulifera
(Hieron.) Vareschi, Fl. Venez. 1 (1):450. 1969.

Terrestrial. Bosque alto in disturbed areas, river banks.
Distribution: Panama, Colombia, Venezuela, Guyana, Ecuador, Paraguay.

Material examined: camino Jejui Mi to Horqueta-Mi, arroyo Moroti, Marin &
Jiménez 261 (PY, BM); Jejui Mi, sendero Aguara’i, arroyo Amambay, Marin &
Jiménez 513 (BM, PY); Carapa, camino al salto, Pefia-Chocarro 291 (BM); Carapa,
camino al salto, Pefia-Chocarro 306 (BM).

Thelypteris (Cyclosorus) hispidula (Decne.) C.F. Reed, Phytologia 17:283. 1968.
Aspidium hispidulum Decne., Nouv. Ann. Mus. Hist. Nat. 3:346. 1834; Nephrodium
quadrangulare Fée, Mém Foug. 5:308. 1852; Dryopteris parasitica (L.) var.
glanduligera Rosenst., Repert. Spec. Nov. Regni Veg. 7:304. 1909; Dryopteris
quadrangularis (Fée) Alston, J. Bot. 75:253. 1937; Thelypteris quadrangularis (Fée)
Schelpe, J. S. African Bot. 30:196. 1964; Christella hispidula (Decne.) Holttum,
Kew Bull. 31:312. 1976; Dryopteris limonensis Christ, Repert. Spec. Nov. Regni
Veg. 8:18. 1910.

Terrestrial. Bosque alto, bosque medio, bosque bajo on saturated soil, gallery forest.
Distribution: New and Old World tropics and subtropics.

Material examined: Jejui Mi, sendero Aguara-i, arroyo Amambay, Marin & Jiménez
517 (PY); Jejui Mi, Km 2 sendero principal, Pefia- Chocarro 199 (PY, BM); Jejui
Mi, rumbo norte, Pefia-Chocarro 256 (BM, CTES); camino Jejui Mi a Lagunita, Km

PTERIDOPHYTES OF MBARACAYU FNR 239

7, Pena-Chocarro 260 (BM); camino Jejui Mi a Lagunita, Km 4-5, Pefia-Chocarro
332 (BM); Jejui Mi, sendero Aguara’ i, Pefia-Chocarro 348 (BM, PY, CTES).

* Thelypteris (Steiropteris) leprieurii (Hook.) R. M. Tryon, Rhodora 69:6. 1967.
Nephrodium leprieurii Hook., Sp. Filic. 4:106. 1862.

Terrestrial. Bosque alto.
Distribution: Mesoamerica, Colombia to Guianas, south te S. Brazil and Paraguay.
Material examined: Carapa, Yku’a, Jiménez & Marin 1628 (PY, BM, MO, CTES).

* Thelypteris (Amauropelta) pachyrhachis (Mett.) Ching, Bull. Fan Mem. Inst.
Biol., Bot. 10:253. 1941.

Nephrodium crassipes Sodiro, Anales Univ. Centr. Ecuador 9:323. 1893; Dryopteris
pachyrhachis (Kunze) Kuntze, Rev. Gen. Pl. 2:813. 1858; Aspidium pachyrhachis
Kunze ex Mett., Farngatt. 4: 83, n. 199. 1858.

Terrestrial. Gallery forest.

Distribution: Costa Rica, Panama, Antilles, Colombia, Venezuela, Ecuador, Peru,
Bolivia, S. Brazil and Paraguay.

Material examined: Carapa, sendero Uru’i, Marin & Jiménez 349 (PY, BM, CTES),
Carapa, camino al salto, Pefia-Chocarro 296 (BM).

Thelypteris (Amauropelta) rivularioides (Fée) Abiatti, Rev. Mus. La Plata, n.s.,
Bot. 9:19. 1958.

Aspidium rivularioides Fée, Crypt. Vasc. Brésil 1:145, t. 50, f. 1. 1869; Dryopteris
rivularioides (Fée) C. Chr. [apud. Rosenst.], Hedwigia 46:125. 1906.

Terrestrial. Bosque bajo on saturated soils, pastures on saturated soil.
Distribution: Brazil, Paraguay, Argentina.

Material examined: S Jejui Mi, Puente Carona, Jiménez & Marin 277 (PY),
Valinotti-cue, Peria-Chocarro 335 (PY, MO, CTES, BM).

Thelypteris (Goniopteris) scabra (C. Presl) Lellinger, Amer. Fern J. 74:60. 1984.
Polypodium scabrum C. Presl, Del. Prag. 1:169. 1822; Lastrea scabra (C. Presl)
C. Presl, Epim. Bot. 41. 1849; Polypodium tetragonum var. incompleta Lindm., Ark.
F. Bot. 1:229. 1903, Dryopteris pseudotetragona Urban var. gemmulifera Hieron. ex
Rosenst. forma major Rosenst., Hedwigia 46:119. 1906/07; Dryopteris
pseudotetragona Urban var. foecunda Rosenst, Hedwigia 46:199. 1906/07; Aspidium
caesarianum Christ, Denkschr. Akad. Wien 69:114. 1907; Dryopteris scabra
(C. Presl) C. Chr. var. incompleta (Lindm.) C. Chr., Kongel. Danske Vidensk.-Selsk.
Skr. 7 Naturvidensk. Math. Afd. 10 (2):238. 1913; Dryopteris scabra (C. Presl)
C. Chr. var. caesarina (Christ.) C. Chr., Kongel. Danske Vidensk.-Selsk. Skr. 7
Naturvidensk. Math. Afd. 10 (2):238. 1913.

Terrestrial. Gallery forest.
Distribution: Brazil, Paraguay and Argentina
Material examined: salto Carapa, Marin & Jiménez 335 (PY, CTES).

240 FERN GAZETTE: VOLUME 15 PART 7 (1999)

Thelypteris (Meniscium) serrata (Cav.) Alston, Bull. Misc. Inform. Kew 1932:309.
1937.

Meniscium serratum Cav., Descr. Pl. 548. 1802; Dryopteris serrata (Cav.) C. Chr.,
Index Filic. 291. 1905.

Terrestrial. Bosque bajo on saturated soil, pastures on saturated soil.

Distribution: from Florida, Antilles and Mexico, to Paraguay, Brazil and
N. Argentina.

Material examined: Jejui Mi , Rumbo Norte, Jiménez & Marin 177 (BM); Valinotti-
cue, Peria-Chocarro 286 (BM).

Thelypteris sp. #1 (Cyclosurus)
Terrestrial. Bosque medio.
Material examined: Jejui Mi, sendero Aguara’i, Pefia-Chocarro 347 (BM).

Thelypteris sp. #2 (Amauropelta)
Terrestrial. Bosque medio on saturated soil.

Material examined: Jejui Mi, sendero Jaku-apeti, Marin & Jiménez 300 (PY, BM,
MO).

TECTARIACEAE

Ctenitis eriocaulis (Fée) Alston, Lilloa 30:112. 1960.

Aspidium eriocaulon Fée, Crypt. Vasc. Brésil 1:136, t. 4, p. 1. 1869; Dryopteris
eriocaulis (Fée) Kuntze, Rev. Gen. Pl. 2:812. 1891.

Terrestrial. Bosque medio.

Distribution: Brazil, Paraguay.

Material examined: Jejui Mi, Rumbo norte, Pefia-Chocarro 243 (BM).

* Ctenitis cf. falciculata (Raddi) Ching, Sunyatsenia 5:250. 1940.

Aspidium falciculatum Raddi, Opusc. Sci. 3:289. 1819; Dryopteris falciculata
(Raddi) Kuntze, Rev. Gen. Pl. 3(2):378. 1898; Aspidium schomburgkii Klotzch,
Linnaea 20:369. 1847.

Terrestrial. Bosque alto.
Distribution: Guyana, Paraguay and Brazil.

Material examined: Jejui Mi, sendero Jaku-apeti, Marin & Jiménez 306 (PY, BM,
CTES, RP).

Ctenitis submarginalis (Langsd. & Fisch.) Ching, Sunyatsenia 5:250. 1940.

Polypodium submarginale Langsd. & Fisch., Pl. Voy. Russes Monde 12. 1810;
Aspidium caripense Mett., Fil. Hort. Bot. Lips. 90, t. 18, f. 9. 1856; Aspidium
karstenii A. Braun, Index Sem. Hort. Berol. 3. 1857; Aspidium microcarpon Fée,
Mém. Foug. 8:105. 1857; Aspidium obtusilobum Fée, Mém. Foug. 8:105. 1857;
Dryopteris collina Christ, Bull. Herb. Boissier, sér. 2, 7:922. 1907; Dryopteris

PTERIDOPHYTES OF MBARACAYU FNR 241

karstenii (A. Braun) C. Chr., Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk.
Afd. 7, 10:98, f. 6. 1913; Dryopteris submarginalis (Langsd. & Fisch.) C. Chr., Index
Fil. 296. 1905.; Polypodium caripense Humb. & Bonpl. ex Willd., Sp. Pl. ed. 4, 5.
1810.

Terrestrial. Bosque alto, bosque medio, bosque bajo on saturated soils.

Distribution: Florida, Hispaniola, S. Mexico to Panama, Venezuela, Colombia, south
to Argentina and Uruguay.

Material examined: Jejui Mi, Rumbo Norte, Pefa-Chocarro 246 (BM); Jejui Mi,
Rumbo Norte, Pefia-Chocarro 257 (BM); Lagunita, sendero arroyo Moroti, Pefia-
Chocarro 267 (BM); Lagunita, sendero arroyo Moroti, Penta-Chocarro 282 (BM);
Carapa, camino al salto, Pefia-Chocarro 290 (BM); Carapa, camino al salto, Pefia-
Chocarro 307 (BM); camino Jejui Mi a Lagunita, Km 5, Pefia-Chocarro 320 (BM);
Jejui Mi, sendero Aguara’i, Pefia-Chocarro 349 (BM); Jejui Mi, sendero Jaku-apeti,
Pena-Chocarro 361 (BM).

Ctenitis sp. #1
Terrestrial. Bosque alto.

Material examined: Jejui Mi, sendero principal, Jiménez & Marin 1587 (PY, BM,
CTES).

Lastreopsis amplissima (C. Presl) Tindale, Vict. Nat. 73:185. 1957.

Polystichum amplissimum C. Presl, Epim. Bot. 58.1851.

Terrestrial. River banks.

Distribution: from Venezuela south to S.E. Brazil and Paraguay.

Material examined: Fazenda Cerro Pora, Marin & Jiménez 312 (PY, BM); Lagunita,

arroyo Moroti, Pefia-Chocarro 274 (BM); Lagunita, arroyo Moroti, Pefia-Chocarro
276 (BM).

Lastreopsis effusa (Sw.) Tindale, Vict. Nat. 73:185. 1957.
Polystichum amplissimum C. Presl, Epim. Bot. 58. 1851; Polypodium effusum Sw.,
Prodr. 134. 1788.

Terrestrial. Bosque alto.
Distribution: from Venezuela to S.E. Brazil and Paraguay.

Material examined: Jejui Mi, sendero principal Km 1-2, Pefia-Chocarro 20] (BM,
PY); camino Jejui Mi a Lagunita, Km 5, Pefia-Chocarro 319 (BM).

Megalastrum connexum (Kaulf.) A.R. Sm. & R.C. Moran, Am. Fern. J. 77(4):127.
1987 (Publish. 1988)

Polypodium connexum Kaulf., Enum. Filic. 120. 1824; Ctenitis connexa (Kaulf.)
Copel., Gen. Fil. 124. 1947.

Terrestrial. Bosque bajo on saturated soil, bosque medio, gallery forest.
Distribution: S. Brazil and Paraguay.

242 FERN GAZETTE: VOLUME 15 PART 7 (1999)

Material examined: Lagunita, arroyo Moroti, Pefia-Chocarro 221 (BM, PY); Jejui
Mi, sendero Aguara-i, Jiménez & Marin 123 (BM); Jejui Mi, sendero Aguara-i,
arroyo Amambay, Marin & Jiménez 516 (PY, BM); Jejui Mi, sendero Aguara-i,
Marin & Jiménez 511 (PY, BM, CTES).

Tectaria incisa Cav., Descr. Pl. 249. 1802.

Aspidium macrophyllum Sw., Syn. Fil. 4:209. 1806; Aspidium martinicense Spreng..,
Anleit. Kenntn. Gew. 3:133. 1804; Tectarnia martinicensis (Spreng.) Copel., Phil.
Journ. Sci. Bot. 2:410. 1907.

Terrestrial. Bosque alto, bosque bajo.

Distribution: S. Florida, Mexico, Mesoamerica, S. America to N. Argentina and
Paraguay.

Material examined: Jejui Mi, Jiménez & Pefia-Chocarro 1210 (BM, PY, ); Jejui Mi,
1500m N de la caseta, Marin & Jiménez 190 (PY, BM, CTES, MO, RP).

DRY OPTERIDACEAE

Cyclodium meniscioides (Willd.) C. Pres] var. meniscioides, Tent. Pterid. 85. 1836.
Aspidium confertum Kaulf., Enum. Filic. 232. 1824; Cyclodium confertum (Kaulf.)
C. Presl, Tent. Pterid. 85. 1836; Dryopternis meniscioides (Willd.) Kuntze var.
conferta (Kaulf.) C.V. Morton, Bull. Torrey Bot. Club 66:50. 1939.

Terrestrial. Bosque bajo on saturated soil, gallery forest.

Distribution: Venezuela, Trinidad, the Guianas, Brazil, Colombia, Bolivia,
N.E. Argentina and Paraguay.

Material examined: in sylva pr. Igatimi, Hassler 4783 (BM); Valinotti-cue, Jiménez
& Marin 1286 (PY, BM, CTES, RP); Lagunita, sendero arroyo Moroti, Pefa-
Chocarro 219 (BM, PY).

Didymochlaena truncatula (Sw.) J. Sm., J. Bot. (Hook.) 4:196. 1841.
Aspidium truncatulum Sw., J. Bot. (Schrad.) 1800(2):36. 1801; Adiantum lunulatum
Houtt., Nat. Hist. 14:209. 1783, non Burm.f.

Terrestrial. Bosque medio.
Distribution: New and Old World tropics and subtropics.
Material examined: Lagunita, sendero arroyo Moroti, Pefia-Chocarro 272 (BM).

WOODSIACEAE

* Diplazium celtidifolium Kuntze var. puberulum Stolze, Fieldiana, Bot. n.s.
27:84. 1991.

Terrestrial. River banks.
Distribution: Peru, Paraguay.

Material examined: camino Jejui Mi a Horqueta-Mi, arroyo Moroti, Marin &
Jiménez 257 (PY, BM, RP).

PTERIDOPHYTES OF MBARACAYU FNR 243

Diplazium cristatum (Desr. in Lam.) Alston, J. Bot. 74:173.1936.

Meniscium cristatum Desr. in Lam, Encycl. 4:94. 1797.

Terrestrial. Bosque medio on saturated soil, bosque medio, river banks.

Distribution: Mexico, Mesoamerica, Antilles, Colombia to Venezuela, south to
N. Argentina and Paraguay.

Material examined: Jejui Mi, sendero Aguara’i, Jiménez & Marin 117 (PY, BM,
CTES); Lagunita, sendero arroyo Moroti, Pefia-Chocarro 281 (BM); Carapa, camino
al salto, Pefia-Chocarro 300 (BM); Carapa, camino al salto, Pefia-Chocarro 303
(BM); Jejui Mi, sendero Aguara’ i, Pefia-Chocarro 330 (BM).

* Diplazium hians Kunze ex Klotzsch, Linnaea 20:361. 1847.
Terrestrial. Bosque alto, on saturated soil.

Distribution: S. Mexico, Mesoamerica, Jamaica, Hispaniola, Colombia, Venezuela
south to Brazil and Paraguay.

Material examined: Jejui Mi, Jiménez & Marin 115 (BM, PY, CTES).

Diplazium plantaginifolium (L.) Urb., Symb. Antill. 4:31. 1903.

Asplenium plantaginifolium L., Sys. Nat. ed. 10, 2:1323. 1759; Asplenium
plantagineum L., Sp. Pl. ed. 2, 1537. 1759; Diplazium plantagineum (L.) Sw., J. Bot.
(Schrad.) 1800 (2): 2. 1801.

Terrestrial. River banks.

Distribution: Mexico to Panama, West Indies, Colombia, Venezuela, Ecuador, Peru,
Bolivia, Brazil, Paraguay.

Material examined: Jejui Mi, Jiménez & Marin 116 (BM, PY, CTES).

* Diplazium riedelianum (Bong. ex Kuhn) Kuhn ex C. Chr., Index Filic. 230.
1905.

Asplenium riedelianum Bong. ex Kuhn, Linnaea 36:102. 1869; Asplenium verapax
Donn.Sm., Bot. Gaz. (Crawfordsville) 13:77, t. 2. 1888; Diplazium verapax
(Donn.Sm.) Hieron., Hedwigia 59:322. 1917.

Terrestrial. River bank, gallery forest.

Distribution: Mesoamerica, Puerto Rico, Venezuela, Peru, Brazil, Paraguay.

Material examined: camino Lagunita a Horqueta-Mi, arroyo Pira Cajon, Marin &
Jiménez 226 (PY, BM, CTES); Lagunita, arroyo Moroti, Pefia-Chocarro 279 (BM);
camino Jejui Mi a Lagunita, arroyo Pira-Cajén, Pena-Chocarro 351 (BM).

* Diplazium urticifolium Christ, Prim. Fl. Costaric. 3:29. 1901.
Diplazium gemmiferum Christ, Bull. Herb. Boissier, sér. 2, 7:270. 1907.

Terrestrial. Bosque alto on saturated soil, gallery forest.

Distribution: Mexico, Mesoamerica, Colombia, Venezuela, Ecuador, Paraguay.
Material examined: Jejui Mi, Jiménez & Marin 114 (BM, PY, CTES, RP); camino
Jejui Mi a Lagunita, arroyo Pira-Cajén, Pefia-Chocarro 355 (BM).

244 FERN GAZETTE: VOLUME 15 PART 7 (1999)

LOMARIOPSIDACEAE

Bolbitis serratifolia (Mert. ex Kaulf.) Schott, Gen. Filic. t. 13. 1835.

Acrostichum serratifolium Mert. ex Kaulf., Enum. Filic. 66. 1824; Bolbitis mexicana
(Christ) C. Chr., Index Filic. suppl. 3:49. 1934; Leptochilus mexicanus Christ, Bull.
Herb. Boissier, sér. 2, 7:414. 1907.

Terrestrial. Disturbed bosque alto.

Distribution: S. Mexico, Mesoamerica, Colombia to Guianas south to Paraguay and
N.W. Argentina.

Material examined: camino Lagunita a Horqueta-Mi, sendero arroyo Moroti, Marin
& Jiménez 227 (PY, BM, CTES); Carapa, sendero Uru’1, Marin & Jiménez 330 (PY,
BM, CTES); Carapa, camino al salto, Pefia-Chocarro 304 (BM).

* Lomagramma guianensis (Aubl.) Ching, Amer. Fern. J. 22:17. 1932.

Polypodium guianense Aubl., Hist. Pl. Guiane 2:962. 1775; Leptochilus guianensis
(Aubl.) C. Chr., Bot. Tidsskr. 26:288. 1904; Bolbitis guianensis (Aubl.) Vareschi,
Flora Venezuela 1:376. 1969.

Terrestrial, scandent on tree trunks. Gallery forest.

Distribution: Greater Antilles, Colombia to the Guianas, south to Brazil, Paraguay
and Argentina.

Material examined: camino Jejui Mi a Lagunita, arroyo Pira-Cajén, Pefia-Chocarro
357 (BM).

Elaphoglossum balansae C. Chr., Index Filic. 303. 1905. nom. nov. pro
Acrostichum tenerum Baker.

Acrostichum tenerum Baker, J. Bot. 1878:302. 1878; Elaphoglossum tenerum
(Baker) Christ, Mon. 62. 1899.

Epiphyte.
Distribution: Paraguay.
Material examined: in sylva pr. Curuguaty, Hassler 4612 (BM)

Elaphoglossum hassleri Christ, Bull. Herb. Boissier, sér. 2, 7:425. 1907.

Epiphyte (?).

Distribution: Paraguay.

Material examined: in silva prope Igatimi, Hassler 5660 (G - n.v., fide Christ, 1907).

Elaphoglossum subcochleare Christ, Bull. Herb. Boissier, sér. 2, 7:425. 1907.
Epiphyte.

Distribution: Paraguay.

Material examined: in sylva pr. Igatimi, Hassler 566] (BM, type).

PTERIDOPHYTES OF MBARACAYU ENR 245

ASPLENIACEAE

Aspenium auriculatum Sw., Kong]. Vetensk. Acad. Handl. 1817:68. 1817.
Asplenium salicifolium L. var. auriculatum (Sw.) Proctor in R.A. Howard, Fl. Lesser
Antilles 2 (Pteridophyta):321. 1977.

Epiphyte on trunks and tree ferns. Bosque medio, gallery forest.

Distribution: S. Mexico, Mesoamerica, Antilles, Colombia, Venezuela, south to
Paraguay and Brazil.

Material examined: in sylva pr. Jejui-guazu, Hassler 5697 (BM); Lagunita, sendero
arroyo Moroti, Marin & Jiménez 163 (PY); Carapa, Marin & Jiménez 180 (PY);
camino Lagunita a Horqueta-Mi, arroyo Pira Cajon, Marin & Jiménez 22] (PY, BM,
MO) Lagunita, sendero arroyo Moroti, Lagunita, Pefia-Chocarro 278 (BM);

Asplenium claussenii Hieron., Hedwigia 60:241. 1919.

Terrestrial or epipetric. Bosque alto, gallery forest.

Distribution: northern South America, Greater Antilles and Brazil, Paraguay.
Material examined: camino Lagunita a Horqueta-Mi, arroyo Pira Cajon, Marin &
Jiménez 224 (PY, CTES); salto de Carapa, Marin & Jiménez 337 (PY, BM, RP),

Carapa, Pefia-Chocarro 224 (BM, PY); Carapa, camino al salto, Pefia-Chocarro 312
(BM).

Asplenium cuspidatum Lam., Encycl. 2:310. 1786.

Terrestrial. Bosque alto, bosque bajo on saturated soil, bosque medio, bosque medio
dominated by Myrtaceae.

Distribution: S. Mexico, Mesoamerica, Antilles, Colombia, Venezuela, south to
Paraguay and Argentina.

Material examined: in sylva pr. fl. Capibary, Hassler 589] (BM); Jejui Mi, Jiménez
& Marin 153 (BM, PY, CTES); Jejui Mi, Rumbo Norte, Jiménez & Pefia-Chocarro
195 (BM); Lagunita, sendero de arroyo Moroti, Marin & Jiménez 159 (PY); Carapa,
Pena-Chocarro 223 (BM, PY); Lagunita, sendero arroyo Moroti, Pefia-Chocarro
265 (BM); Carapa, camino al salto, Pefia-Chocarro 315 (BM); Camino Jejui Mi a
Lagunita Km 5, Pefia-Chocarro 321 (BM).

Asplenium formosum Willd., Sp. Pl. ed. 4. 5:329. 1810.

Epiphyte. Bosque alto.

Distribution: tropical America, Asia, tropical Africa.

Material examined: Jejui Mi, Jiménez & Penia-Chocarro 1202 (BM, PY, CTES).

Asplenium laetum Sw., Syn. Fil. 79, 271. 1806.

Asplenium lugubre Liebm., Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Afd.
5, 1:243 (seors 91). 1849; Asplenium schkuhrianum C. Presl, Tent. Pterid. 107. 1836;
Asplenium virens Desv., Mém. Soc. Linn. Paris 6:273. 1827, non C. Pres] 1825.

Terrestrial. Bosque alto on saturated soil, river banks.

246 FERN GAZETTE: VOLUME 15 PART 7 (1999)

Distribution: tropical America.

Material examined: Jejui Mi, Jiménez & Marin 150 (BM, PY, CTES); Carapa,
camino al salto, Pefia-Chocarro 302 (BM).

Asplenium mucronatum C. Pres], Delic. Prag. 1:178. 1822.

Epiphyte. Bosque medio, on tree ferns.

Distribution: Brazil, Paraguay.

Material examined: camino Lagunita a Horqueta-Mi, arroyo Pira Cajén, Jiménez

1382 (PY, BM, CTES, MO, RP); Carapa, Marin et al. 179 (PY); Carapa, sendero
Urui, después del Ycua Saraki Jhovy, Marin & Jiménez (BM, PY).

Asplenium serratum L., Sp. Pl. 1079. 1753.
Epiphyte. Bosque alto.

Distribution: S. Florida, Mexico, Mesoamerica, Antilles, Trinidad, Tobago,
Colombia, to Guianas, and south to Brazil, Paraguay, Argentina.

Material examined: Jejui Mi, Jiménez & Marin 134 (BM, PY, CTES).

Phyllitis balansae (Bak.) C. Chr., Ind. 492. 1906.
Scolopendnium balansae Baker, Icones Pl. 1653. 1886.

Terrestrial. Gallery forest, river banks.
Distribution: Paraguay.

Material examined: Salto de Carapa, Marin & Jiménez 336 (PY, BM, CTES); salto de
Carapa, Pefia-Chocarro 311 (PY, BM).

BLECHNACEAE

Blechnum asplenioides Sw., Kongl. Vetensk. Acad. Hand]. 1817: 72, t. 3. 1817.
Asplenium blechnoides Sw., Syn. Fil. 76. 1806; Blechnum unilaterale Sw., Ges. Naturf.
Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 4:79. 1810, also Willd. Sp.
Pl. ed. 4, 5:407. 1810; Blechnum polypodioides Raddi, Opusc. Sci. 3: 294. 1819, non
(Sw.) Kuhn. 1868; Blechnum blechnoides (Sw.) C. Chr.

Terrestrial. Gallery forest, bosque alto.
Distribution: tropical America.

Material examined: arroyo Moroti, Marin & Jiménez 161 (PY); La Morena, sendero
Guazu pyta, Jiménez & Marin 1334 (BM, PY, CTES, MO).

Blechnum binervatum (Poir.) C.V. Morton & Lellinger subsp. acutum (Desv.)
Tryon & Stolze, Fieldiana, Bot. n.s. 32:64. 1993.

Lomaria angustifolia Humb. Bonpl. & Kunth, Nov. Gen. Sp. 1:18. 1816; Lomaria acuta
Desv., Mém. Soc. Linn. Paris 6:290. 1827; Lomaria cuspidata Kunze, Linnaea 9:59.
1834; Lomaria meridensis Klotzsch, Linnaea 20:345. 1847; Lomaria ensiformis Liebm.,
Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Afd. 5, 1:234. 1849; Blechnum
meridense (Klotzsch) Mett., Fil. Hort. Bot. Lips. 61. 1856; Blechnum acutum (Desv.)

PTERIDOPHYTES OF MBARACAYU FNR 247

Mett., Ann. Sci. Nat. Bot. Ser.5, 2:225. 1864; Blechnum angustifolium (Humb. Bonpl. &
Kunth ) Hieron., Bot. Jahrb. Syst. 34:472. 1904, non Willd., 1810; Blechnum ensiforme
~ (Liebm.) C. Chr., Index Filic. 153. 1905; Blechnum kunthianum C. Chr., Index Filic. 16.
1913, nom. nov. for Lomaria angustifolia Humb. Bonpl. & Kunth, non Blechnum
angustifolium Willd., 1810.

Hemi-epiphyte. Gallery forest.

Distribution: Mesoamerica, Venezuela and Colombia, south to Bolivia and S.E. Brazil.
Material examined: La Morena, sendero Guazu Pyta, Jiménez & Marin 1337 (PY, BM,
CTES, MO, RP); camino Jejui Mi a Horqueta-Mi, arroyo Moroti, Marin & Jiménez 234
(PY, CTES); Lagunita, sendero arroyo Moroti, Pefia-Chocarro 273 (BM); Carapa,

camino al salto, Pefia-Chocarro 305 (BM); camino Lagunita a Horqueta-Mi, arroyo Pira
Cajon, Penia-Chocarro 354 (BM).

Blechnum brasiliense Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck.
Gesammten Naturk. 5:330. 1811.

Blechnum nigrosquamatum Gilbert, Bull. Torrey Bot. Club 24:258. 1897; Blechnum
corcovadense Raddi, Opusc. Sci. 3:294. 1819.

Terrestrial. Bosque alto, on saturated soils, bosque medio on saturated soil.

Distribution: Guatemala, Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil,
Paraguay, Uruguay and Argentina.

Material examined: in sylva pr. Igatimi, Hassler 4869 (BM); in palude pr. flumen Jejui-
guazu, Hassler 5706 (BM), Jejui Mi, Jiménez & Marin 126 (BM, PY); Jejui Mi, Rumbo
Norte, Marin & Jiménez 149 (PY, BM, CTES); Jejui Mi, sendero Aguara’i, Marin &
Jiménez 509 (BM, RP); Jejui Mi sendero Jaku-apeti, Pefia-Chocarro 207 (BM, PY).

Blechnum fraxineum Willd., Sp. Pl. ed. 4, 5:413. 1810.
Blechnum longifolium Willd., Sp. Pl. ed 4, 5:413. 1810, non Cav. 1802; Blechnum
gracile Kaulf., Enum. Filic. 158. 1824.

Terrestrial. River banks.
Distribution: tropical America.
Material examined: Lagunita, arroyo Moroti, Jiménez & Marin 1198 (PY); Lagunita,

arroyo Moroti, Pefia-Chocarro 277 (BM); camino Jejui Mi a Lagunita, arroyo Pira
Cajon, Peria-Chocarro 353 (BM, PY).

Blechnum lanceola Sw., Kong]. Vetensk. Acad. Handl. 1817:71, t. 3, f. 2. 1817.
Terrestrial. River banks.
Distribution: Peru, south to Argentina and east to Brazil.

Material examined: in sylva Sierra de Maracayu, Hassler 5237 (BM); Lagunita, sendero
arroyo Moroti, Marin & Jiménez 160 (PY); Lagunita, sendero arroyo Moroti, Pefia-
Chocarro 271 (BM); camino Lagunita-Horqueta-Mi, sendero arroyo Pira-Caj6n, Pena-
Chocarro 352 (BM, PY).

248 FERN GAZETTE: VOLUME 15 PART 7 (1999)

Blechnum occidentale L., Sp. Pl. 1077 [in error as orientale]. 1753.
Blechnum caudatum Cav., Descr. Pl. 262. 1802; Blechnum glandulosum Link., Enum.
Hort. Berol. Alt. 2:462. 1822; Blechnum pectinatum C. Presl, Relig. Haenk. 1:51. 1825.

Terrestrial. Gallery forest, river banks.
Distribution: United States, tropical America.

Material examined: in sylva pr. fl. Capibary, Hassler 4437; in sylva pr. Igatimi, Hassler
5656 (BM); Jejui Mi, sendero Jaku-apeti, Pefia-Chocarro 208 (BM, PY); Lagunita,
sendero arroyo Moroti, Pefia-Chocarro 280 (BM).

* Blechnum obtusifolium (C. Presl) Ettingsh., Denkschr. Kaiserl. Akad. Wiss. Wien
23259515 6, 1.4. 1604:
Lomaria obtusifolia C. Presl, Tent. Pterid. 143. 1836.

Terrestrial. Swamps with Sphagnum.
Distribution: Peru and N.W. Argentina, S.E. Brazil and Paraguay.
Material examined: Lagunita, sendero arroyo Moroti, Pefia-Chocarro 217 (BM, PY).

POLYPODIACEAE

Campyloneurum leuconeurum Fée, Crypt. Vasc. Brésil 1. 103, t. 35, f. 1. 1869.
Epiphyte or terrestrial. Gallery forest, bosque medio, disturbed bosque alto.

Distibution: S. Brazil, Uruguay, Paraguay.

Material examined: Carapa, sendero Uru’i, Marin & Jiménez 326 (PY, BM, CTES);

Carapa, camino al salto, Pefia-Chocarro 293 (BM); Carapa, camino al salto, Pefia-
Chocarro 313 (BM).

Campyloneurum nitidum C. Presl, Tent. Pterid. 190. 1836.
Epiphyte. Gallery forest, on tree-ferns.
Distribution: S. Brazil, Uruguay, Paraguay.

Material examined: Camino Lagunita a Horqueta-Mi, arroyo Pira Cajon, Jiménez 1381
(PY, BM, CTES, MO, RP).

Campyloneurum phyllitidis (L.) C. Pres], Tent. Pterid. 190. 1836.

Polypodium phyllitidis L., Sp. Pl. 1083. 1753; Cyrtophlebium phyllitidis (L.) J.G. Sm.,
J. Bot. (Hook.) 4: 58. 1841.

Terrestrial. Bosque medio.

Distribution: southern United States, Mesoamerica, West Indies, Colombia and
Venezuela to Bolivia and C. Brazil.

Material examined: Jejui Mi, sendero Aguara’i, Peria-Chocarro 331 (BM).

Marginaria hirsutissima (Raddi) Pic. Serm., Webbia 31(1):248. 1977.

Polypodium hirsutissimum Raddi, Opusc. Sci. 3:286. 1819; Polypodium rufulum
C. Presl, Delic. Prag. 1:164. 1822; Polypodium vexillare Chr., Bull. Herb. Boissier,
ser. 2, 2:373. 1902:

PTERIDOPHYTES OF MBARACAYU FNR 249

Epiphyte. Cerrado on Butia (Arecaceae), bosque bajo, bosque bajo on saturated
soils.

Distribution: Brazil, Paraguay, Uruguay, Argentina.

Material examined: Lagunita, sendero arroyo Moroti, Marin & Jiménez 156 (PY);
Aguara Nu, Jiménez & Marin 1318 (PY, BM, CTES); Jejui Mi, Jiménez & Marin
193 (BM, PY, CTES); Valinotti-cue, Pefia-Chocarro 288 (BM).

Microgramma lindbergii (Mett.) de la Sota, Opera Lilloana 5:56, f. 2, 5. 1961.
Polypodium lindbergii Mett., Linnaea 36: 136. 1869.

Epiphyte. Bosque alto, bosque medio, bosque medio on saurated soil.

Distribution: Brazil and Paraguay.

Material examined: in sylva Igatimi, Hassler 4892 (BM); Jejui Mi, Jiménez & Marin
130 (BM, PY, CTES); Jejui Mi, Jiménez & Marin 147 (BM, PY, CTES); Jejui Mi,
sendero Jaku-apeti, Marin & Jiménez 304 (PY, BM, CTES, G, RP); Jejui Mi, sendero
principal Km1-2, Pefia-Chocarro 204 (BM, PY); Jejui Mi, sendero Jaku-apeti, Pefia-
Chocarro 206 (BM, PY); Jejui Mi, sendero Aguara’i, Peria-Chocarro 329 (BM).

Microgramma persicariifolia (Schrad.) C. Presl, Tent. Pterid. 214. 1846.
Polypodium persicaritfolium Schrad., Gott. Gel. Anz. 1824:867. 1824.

Epiphyte. Disturbed bosque alto, bosque bajo on saturated soil. |
Distribution: Central America, Antilles, Colombia to Guyana south to C. Brazil and
Paraguay.

Material examined: in sylva pr. San Estanislao, Hassler 4145 (BM); Jejui Mi, Rumbo
Norte, Peria-Chocarro 253 (BM); Jejui Mi, Rumbo Sur, Jiménez & Marin 1283 (PY).

Microgramma squamulosa (Kaulf.) de la Sota, Opera Lilloana 5:59. 1961.
Polypodium squamulosum Kaulf., Enum. Filic. 89. 1824.

Epiphyte. Bosque alto, bosque medio, bosque bajo on saturated soils, gallery forest.
Distribution: Peru, Bolivia, S.E. Brazil, Argentina, Paraguay and Uruguay.

Material examined: in sylva pr. Sierra de Maracayu, Hassler 4887 (BM); in sylva pr.
Sierra de Maracayu, Hassler 5372 (BM); Jejui Mi, rumbo Norte, Jiménez & Marin 1267
(PY); Arroyo Guyra-Keha, Marin & Jiménez 254 (PY, BM); camino Jejui Mi-Lagunita,
Km 2, Marin 500 (PY, BM, CTES); Jejui Mi, sendero Jaku-apeti, Pefia-Chocarro 236
(BM, PY); Jejui Mi, Rumbo Norte, Pefia-Chocarro 251 (BM); Jejui Mi, sendero
Aguara’i, Pefia-Chocarro 328 (BM).

Microgramma vaccinifolia (Langsd. & Fisch.) Copel., Gen. Fil. 185. 1947.
Polypodium vaccinifolium Langsd. & Fisch., Pl. Voy. Russes Monde (Icon. Filic.) 8.
1810.

Epiphyte. Bosque alto, bosque medio.

Distribution: West Indies, Venezuela and Colombia, south to Paraguay and Argentina.
Material examined: Jejui Mi, sendero principal, Jiménez 174] (BM, PY, CTES, MO,
RP); Jejui Mi, sendero Aguara’i, Jiménez & Marin 0136 (PY); Jejui Mi, sendero

250 FERN GAZETTE: VOLUME 15 PART 7 (1999)

Aguara’i, Peria-Chocarro 326 (BM); Jejui Mi, sendero principal Km 1, Pefia-Chocarro
343 (BM).

* Pecluma camptophyllaria (Fée) M.G. Price var. lachnifera (Hieron.) Lellinger,
Amer. Fern J. 74:59. 1984.

Polypodium camptophyllarium Fée var. lachniferum (Hieron.) Evans, Ann. Missouri
Bot. Gard. 55: 254. 1969; Polypodium lachniferum Hieron., Bot. Jahrb. Syst. 34: 515.
1904.

Epiphyte. Bosque alto.

Distribution: Mesoamerica, Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil and
Paraguay.

Material examined: Carapa, Pena-Chocarro 222 (BM, PY).

Pecluma filicula (Kaulf.) M.G. Price, Amer. Fern. J. 73:114. 1983.

Polypodium filiculum Kaulf., Enum. Filic. 275. 1824.

Epiphyte. Bosque alto, bosque medio, bosque bajo on saturated soil.

Distribution: Colombia to Argentina and S. Brazil.

Material examined: Jejui Mi, camino principal Km 8, Jiménez & Marin 168 (BM);
Lagunita, sendero arroyo Moroti, Pefia-Chocarro 266 (BM).

Pecluma ptilodon (Kunze) M. G. Price var. robusta (Fée) Lell., Am. Fern. J. 74:60.
1984.

Polypodium ptilodon Kunze, Linnaea 9:42. 1834. Polypodium robustum Fée, Crypt.
Vasc. Brésil 1:92, t. 28, f. 1. 1869.

Terrestrial. Bosque alto, gallery forest, bosque bajo on saturated soils.
Distribution: Bolivia and E. Brazil to N. Argentina.

Material examined: Sierra de Maracayu, Igatimi, Hassler 5511 (GH, BM); sendero
Aguara’i, Jiménez & Marin 0152 (PY); Jejui Mi, Rumbo norte, Jiménez & Marin 0174
(PY, BM. CTES, MO); Carapa, Marin & Jiménez 174 (PY); Jejui Mi, sendero Jaku-
apeti, Marin & Jiménez 299 (PY, BM, CTES, MO, RP); Jejui Mi, Rumbo Norte, Pevia-
Chocarro 248 (PY, BM); Jejui Mi, Rumbo Norte, Pefia-Chocarro 254 (BM); Jejui Mi,
sendero Aguara’1, arroyo Amambay, Peria-Chocarro 345 (BM).

Pecluma sicca (Lindm.) M. G. Price, Am. Fern J. 73. 1983.
Polypodium siccum Lindm., Ark. Bot. 1:234, t. 11, f. 4. 1903.

Epiphyte. Bosque medio, gallery forest.
Distribution: S.E. Brazil, Paraguay and N.Argentina.

Material examined: Rumbo Norte, Pefia-Chocarro 245 (BM), Carapa, camino al salto,
Penria-Chocarro 294 (BM).

Phlebodium decumanum (Willd.) J. Sm., J. Bot. (Hooker) 4:59. 1841.
Polypodium decumanum Willd., Sp. Pl. ed. 4, 5:170. 1810.

Epiphyte. Cerrado on Butia (Arecaceae).

PTERIDOPHYTES OF MBARACAYU FNR 251

Distribution: tropical America.

Material examined: Aguara Nu, camino a la comunidad Tekoha ryapu, Esquivel 24
(BM).

Pleopeltis pleopeltifolia (Raddi) Alston, Bol. Soc. Brot. ser. 2 30:21. 1956.
Polypodium pleopeltifolium Raddi, Opusc. Sci. 3:286. 1819.

Epiphyte. Bosque alto, bosque medio, bosque bajo on saturated soils.
Distribution: Brazil, Paraguay.

Material examined: Jejui Mi, sendero Aguara’i, Jiménez & Marin 0119 (PY); Jejui
Mi, sendero Aguara’i, Jiménez & Marin 0134 (PY); Jejui Mi, sendero Aguara’1,
Jiménez & Marin 0148 (PY); Jejui Mi, Rumbo Norte, Jiménez & Marin 1181 (PY);
Carapa (Yku’a), Jiménez & Marin 1624 (BM, PY, CTES, MO); Carapa, Marin &
Jiménez 181 (PY); Jejui Mi, Rumbo Norte, Pefia-Chocarro 252 (BM); Lagunita,
sendero arroyo Moroti, Pefia-Chocarro 263 (BM); Lagunita, sendero arroyo Moroti,
Pena-Chocarro 264 (BM).

Polypodium latipes Langsd. & Fisch., Pl. Voy. Russes Monde (Icones Filic.): 10
t. 10. 1810.

Polypodium loriceum sensu Humb. Bonpl. & Kunth, non L.; Polypodium harpeodes
Link, Hort. Berol. 2:97. 1833; Polypodium vacillans Link, Hort. Berol. 2:97. 1833;
Goniophlebium pectinans Fée, Crypt. Vasc. Brésil 1:109, t. 39, f. 1. 1869.

Epiphyte or terrestrial. Bosque bajo, pastures in cerrado, cerrado (on Butia
paraguayensis - Arecaceae), pastures.

Distribution: Guyana, Brazil and Paraguay.

Material examined: Jejui Mi, Jiménez & Marin 73 (BM, PY, CTES); camino a
Aguara Nu, Jiménez & Marin 1301 (PY, BM); Aguara Nu, Jiménez & Marin 1308
(PY, BM, CTES, MO, RP); Lagunita, Marin & Jiménez 268 (PY, BM, CTES, RP);
entrada a Lagunita, Pefia-Chocarro 212 (BM, PY); camino Jejui Mi a Lagunita, Km
11, Pefia-Chocarro 333 (BM).

Polypodium polypodioides (L.) Watt, Canad. Naturalist & Quart. J. Sci. ser. 2,
3:158. 1867.
Acrostichum polypodioides L., Sp. Pl. 1068. 1753.

Epiphyte. Bosque alto, gallery forest, bosque bajo on saturated soils.

Distribution: S.E. United States, Mesoamerica, Antilles and South America.

Material examined: arroyo Moroti, Marin & Jiménez 169 (PY); Jejui Mi, sendero
Jaku apeti, Marin & Jiménez 295 (PY, BM); Jejui Mi, Rumbo Norte, Pefia-Chocarro
249 (BM); camino Jejui Mi to Lagunita, sendero principal, Km 22, Pefia-Chocarro
283 (BM); Carapa, camino al salto, Pefia-Chocarro 292 (BM).

Polypodium triseriale Sw., J. Bot. (Schrad.) 1800 (2):126. 1801.

Polypodium longifolium Presl, Delic. Prag.1:167. 1822; Polypodium preslianum
Spreng., Syst. Veg. ed. 16, 5:556 (index). 1828; Gonophlebium triseriale (Sw.)
Lawalrée, Bull. Jard. Bot. Belg. 50(3-4):516. 1980.

Dad FERN GAZETTE: VOLUME 15 PART 7 (1999)

Terrestrial. Bosque alto, disturbed bosque alto, periodically flooded bosque medio.

Distribution: Mesoamerica, Antilles, Trinidad, Colombia, Venezuela, Guyana, south
to Brazil and Paraguay.

Material examined: Jejui Mi, sendero principal, Jiménez & Marin 1481la (BM); Jejui
Mi, Rumbo Norte, Marin & Jiménez 148 (PY, BM); Jejui Mi, puente Carona, Marin
& Jiménez 283 (PY, BM).

SALVINIACEAE

Salvinia minima Baker, J. Bot. 24:98. 1886.
Salvinia rotundifolia auct., non Willd.

Aquatic. Rivers, ponds.

Distribution: S.E. United States, Mesoamerica, Antilles, Bermudas, Colombia
Venezuela, south to Uruguay, Paraguay and N. Argentina.

Material examined: Jejui Mi, 1500 m N de la caseta, Marin & Jiménez 194 (PY,
BM); Jejui Mi, Puente Carona, Marin & Jiménez 294 (PY).

DISCUSSION

The diversity in the Mbaracayt region has, at least in part, been attributed to its
position on a phytogeographic ecotone and to the mixing of tropical and temperate
elements in the flowering plant flora (Keel & Herrera-MacBryde, 1997). These ideas
are based largely on angiosperm taxa and the degree to which pteridophyte
distribution and diversity patterns mirror those of angiosperms has only rarely been
discussed (Smith, 1972; Kramer, 1993). In the Neotropics four principal regions of
pteridophyte species diversity have been identified: Greater Antilles, southern
Mexico and Central America, the Andes, and southeastern Brazil (Tryon, 1972;
Tryon & Tryon, 1982). If the diversity of pteridophytes in Mbaracayu is due to the
mixing of floristic regions, we expect to see evidence of this in the taxa encountered
in the region. However, edaphic specialization may also contribute to the high
pteridophyte diversity seen in the Neotropics in general (Poulsen & Tuomisto, 1995;
Tuomisto & Poulsen, 1996), and this may also be contributing to diversity in
Mbaracayiu.

The majority of pteridophyte species present in Mbaracayu are widely distributed,
ranging from Mexico throughout tropical America to Paraguay, Brazil and northern
Argentina (62 species). One species is introduced from the Old World tropics, six
are pantropical, four widespread in the New and Old World Tropics and subtropics
and the remaining thirty six have a more restricted distribution. Of these last thirty
six, six belong to genera which are most diverse in southeastern Brazil (Doryopteris
and Anemia) and four (Phyllitis balansae (Bak.) C. Chr. and probably all three
species of Elaphoglossum) are endemic to Paraguay. In general, therefore, there is a
mixing of species from different areas of pteridophyte diversity, but the majority of
species are of relatively wide distribution. Those taxa with more restricted
distributions are generally from the southeastern Brazilian area (sensu Tryon &
Tryon, 1982), and are found in Brazil, Paraguay, Uruguay and Argentina. Some of
these also reach Peru and Bolivia, but not the majority.

PTERIDOPHYTES OF MBARACAYU FNR

253

Table 1. Habitat types in the Mbaracayu Forest Nature Reserve. Totals do not
equal the number of pteridophyte taxa in the checklist as species often occur in
more than one habitat, see text for details.

Terms
used in the
checklist
Bosque alto

Description of habitat

This forest type covers 17.000ha (30% of the total
covered area in the Reserve) and the canopy reaches
30 m in height. Soils are deep, with abundant organic
matter. The forest shows three distinct strata: the
canopy is composed of various genera, including
Aspidosperma, Tabebuia, Astronium, Peltophorum;
the middle layer is formed of species that reach c. 20
m in height and among the genera common in this
layer are Guarea, Eugenia, Ocotea, Nectandra,
Campomanesia, Chrysophyllum, etc.; in the lower
stratum Sorocea bonplandii (Baill.) W.G. Burger,
Lanjouw & Wess. Boer is dominant, other common
species are Hennecartia, Sebastiania brasiliensis
Spreng., Clavija nutans (Vell.) Stahl, Trichilia, etc.
Epiphytes occur and lianas are extremely important.

This forest type covers 30% of the total forested area )

of the reserve and reaches c. 20 m in height. Soils are
sandy and well-drained. Several types of bosque
medio can be distinguished, some dominated by
Myrcianthes pungens O. Berg., with Ocotea,
Myrciaria rivularis (Cambess.) O. Berg., Myrciaria
cuspidata O. Berg., etc., while another distinct type is
dominated by Copaifera langsdorfii Desf., with
Tabebuia, Guarea, Eugenia, etc.

Number
of
species

254 FERN GAZETTE: VOLUME 15 PART 7 (1999)

In this category we include forest types of medium to
low canopy height on sandy soils, which are on
slopes ending in water courses or along streams and
are thus subject to periodic inundation. Some of the
important canopy species in these forests are:
Copaifera langsdorfii Desf., Luehea divaricata Matt.,
Inga uruguensis Hook. & Arn., Nectandra
megapotamica (Spreng.) Mez., etc. Bambusa guadua
Bonpl. can be important on the banks of the rivers,
forming dense and often nearly pure stands. In the
middle strata are found Sebastiania sp., Myrciaria
rivularis (Cambess.) O. Berg., Esenbeckia
grandiflora Mart., Trichilia pallida Sw., etc.
Piperaceae, Rubiaceae and Melastomataceae are
abundant in the understory.
Includes those forests where the canopy is up to 10 m
in height, and usually occurs on poor soils and
depressions that are saturated, and either permanently
or periodically inundated. In the reserve, bosque bajo
covers 16.000 ha and is the third most common
habitat type. The dominant family in these forests is
Myrtaceae.
This subcommunity occurs within bosque bajo on
super saturated soils. Trees are scarce and usually
stunted. Common _ families are: Myrtaceae,
Myrsinaceae, Aquifoliaceae and Styracaceae.

Gallery
forest

Bosque
bajo

Sphagnum
swamps

Cerrado Grassland on well-drained, sandy soils. Generally
occurring on uplands, and comprises a wide variety
of mixed grass and shrub communities. Some
common families are Annonaceae, Erythroxylaceae,
Arecaceae (genera Butia and Syagrus).

Flooded
grasslands

Grassland communities on slopes which terminate in
water courses. These pastures act as water sinks, and
are on alluvial soils that are poor in organic material,
phosphorus, potassium and calcium. Dominant
familes are Poaceae and Cyperaceae. The genera
Paspalum and Setaria are among the most species-
rich.

River bank | Open habitat along river courses, usually with high
light intensity.

Pastures This community occupies a total of 1415 ha in the
Reserve (including flooded grasslands). Soils are
poor and badly drained. Pastures are dominated by
graminoids, such as Andropogon, Axonopus,
Loudetia. Common angiosperm families (in addition
to: Poaceae) are Asteracede, Turneracede,

Sterculiaceae and Fabaceae.

8

PTERIDOPHYTES OF MBARACAYU FNR a

Field observations indicate that pteridophyte species are clearly associated with
particular habitats in the Mbaracayu reserve, and that soil diversity may have a large
impact on the resulting floristic diversity in these forests, as has been shown in the
Peruvian Amazon (see Young & Leon, 1989; Poulsen & Tuomisto, 1996; Tuomisto
& Poulsen, 1996). Numbers of pteridophyte taxa found in each of our habitat
divisions are shown in Table 1 (habitats follow FMB, 1991). Associations of
pteridophyte species with habitat in Mbaracayi are shown in Table 2, where taxa
found specifically in a single habitat are listed. Taxa found in more than one habitat
are not included in Table 2. Our data are similar to those of Tuomisto & Poulsen
(1996), with species of Pteris, Bolbitis, Diplazium, Alsophila, Tectaria, Danaea and
Didymochlaena occurring on rich soils and species of Lindsaea occurring on poor
soils. In Mbaracayt the rich soils along river banks harbour many fern species; light
levels here are also high.

It is clear that at least some mixing of species from different centres of
pteridophyte diversity is occurring in Mbaracayu. However, due to generally wider
distributions of pteridophyte taxa than their angiosperm counterparts (Smith, 1972),
it is more difficult to assess whether or not this mixing accounts for the high diversity
we have observed, or if edaphic and other habitat factors are more important. It is
likely that both have contributed to the rich pteridophyte flora we document here.

Table 2. Habitat associations of Mbaracayt pteridophytes based on field
observations. See text.

Habitat Pteridophyte taxa 0h

Sphagnum | Osmunda cinnamomea L.
swamps Trichomanes cristatum Kaulf.
Doryopteris lomariacea Klotzsch
Lindsaea portoricensis Desv.
Blechnum obtusifolium (C. Presl) Ettingsh. _
Cerrado Anemia tomentosa var. anthrisifolia (Schrad.) Mickel
Anemia tomentosa (Sav.) Sw. var. tomentosa
Dicranopteris flexuosa (Schrad.) Underw.

Flooded Equisetum giganteum L.
grasslands | Lycopodiella alopecuroides (L.) Cranfill

Lycopodiella caroliniana var. meridionalis (Underw. & F.E. Lloyd)
B. Allg.

Lycopodiella cernua (L.) Pic. Serm.
Selaginella marginata (Humb. & Bonpl. ex Willd.) Spring.
Selaginella muscosa §

areas Pityrogramma trifoliata (L.) R. Tr on

Saturated Lindsaea guianensis var. lanceastrum K.U. Kramer
soils in Lindsaea lancea (L.) Bedd.

forest Lindsaea quadrangularis Raddi

Diplazium hians Kuntze
Blechnum brasiliense Desv.

256 FERN GAZETTE: VOLUME 15 PART 7 (1999)

Table 2 continued

Huperzia mandiocana (Raddi) Trevis.
Anemia phyllitidis (L.) Sw.
Adiantopsis radiata (L.) Fée
Adiantum latifolium Lam.
Doryopteris concolor (Langsd. & Fisch.) Kuhn.

Pteris deflexa Link

Thelypteris leprieurti (Hook.) R.M. Tryon

Ctenitis euriocaulis (Fée) Alston

Ctenitis falciculata (Raddi) Ching

Ctenitis submarginalis (Langsd. & Fisch.) Ching
Lastreopsis effusa (Sw.) Tindale

Tectaria incisa Cav.

Didymochlaena truncatula (Sw.) J. Sm.

Bolbitis serratifolia (Mert. Ex Kaulf.) Schott

Asplenium formosum Willd.

Asplenium serratum L.

Campyloneurum phyllitidis (L.) C. Pres]

Microgramma lindbergii (Mett.) de la Sota

Microgramma squamulosa (Schrad.) C. Pres]

Microgramma persicariifoia (Kaulf.) de la Sota
Microgramma vaccinifolia (Langsd & Fisch.) Copel.
Pecluma camptophyllaria var. lachnifera (Hieron.) Lellinger
Pecluma filicula (Kaulf.) M.G. Price

Pecluma ptilodon var. robusta (Fée) Lellinger

Pleopeltis pleopeltifolia (Raddi) Alston

Polypodium triseriale Sw.

In forests
(Bosque
alto or
alto/medio)

ACKNOWLEDGMENTS

We thank the Servicio de Parques Nacionales y Vida Silvestre for facilitating
collecting permits in Paraguay; Estela Esquivel, Alberto Madrofio, Antonio Claudio
Ferreira da Costa and the cuerpo de guardabosques of Mbaracayt for assistance in
the field; Alberto Yanosky, Claudia Mercolli and the staff of PY for logistical help in
Asuncion; curators of the herbaria cited in the text for permission to examine the
specimens in their care; Blanca Le6n and Josephine Camus for a careful review of
the manuscript; and the Darwin Initiative for the Survival of Species for funding this
work.

PTERIDOPHYTES OF MBARACAYU FNR 27

Table 2 continued

River banks | Danaea nodosa (L.) Sm.
and Gallery | Trichomanes diaphanum Humb., Bonpl. & Kunth
forest Trichomanes radicans Sw.
Trichomanes rigidum Sw.
Alsophila cuspidata (Kuntze) D.S. Conant
Adiantum petiolatum Desv.
Pteris altissima Poir.
Pteris quadriaurita Retz.
Hemionitis tomentosa (Lam.) Raddi
Thelypteris abbiatti C.F. Reed
Thelypteris pachyrhachis (Mett.) Ching
Thelypteris scabra (C. Presl) Lellinger
Lastreopsis amplissima (C. Presl) Tindale
Diplazium celtidifolium Kuntze var. puberulum Stolze
Diplazium plantaginifolium (L.) Urb.
Diplazium riedelianum (Bong. ex Kuhn) Kuhn
Lomagramma guianensis (Aubl.) Ching
Phyllitis balansae (Bak.) C. Chr.
Blechnum binervatum subsp. acutum (Desv.) R.M. Tryon & Stolze
Blechnum fraxineum Willd.
Blechnum lanceola Sw.
Blechnum occidentale L.
Campyloneurum nitidum C. Pres}

REFERENCES

BOZZANO S., B.E. & WEIK, J.H. 1992. El Avance de la Deforestacién y el
Impacto Econémico. Serie No.12: Proyecto de Planificaci6n del Manejo de
los Recursos Naturales MAG/GT - GTZ. Asuncion, Paraguay.

DAVIDSE, G., SOUSA S., M. & KNAPP, S. (eds) 1995. Flora Mesoamericana.
Volumen 1: Psilotaceae a Salvinaceae. UNAM, Mexico.

DAVIS, S.D., HEYWOOD, V.H., HERRERA-MACBRYDE, O., VILLA-
LOBOS, J., & HAMILTON, A.C. (eds) 1997. Centres of Plant Diversity.
Volume 3: The Americas. WWF and IUCN, IUCN Publishing Unit,
Cambridge.

DINERSTEIN, E., OLSON, D.M., GRAHAM, D.J., WEBSTER, A.L., PRINM,
S.A., BOOKBINDER, M.P., & LEDEC, G. 1995. Una Evaluacién del
Estado de Conservacion de las Ecoregiones Terrestres de América Latina y
el Caribe. WWF, Washington, D.C.

DIRECCION DE PARQUES NACIONALES Y VIDA SILVESTRE. 1994. Flora
Amenazada del Paraguay. Asuncion, Paraguay.

EITEN, G. 1972. The cerrado vegetation of Brazil. Bot. Rev. (Lancaster) 38: 201-
341.

258 FERN GAZETTE: VOLUME 15 PART 7 (1999)

FMB. 1991. Proyecto Trinacional de Manejo del Bosque Atldntico Interior. la
Etapa. Diagnostico de los Recursos Socio-Ambientales, Capitulo Paraguay,
Vols I y II. Fundacién Moisés Bertoni para la Conservacion de la
Naturaleza. Asuncion, Paraguay.

HASSLER, E. 1928. Pteridophytorum paraguariensium et Regionum Argentinarum
Adjacentium Conspectus criticus, por el Dr. Emilio Hassler. Inst. Bot.
Farmacologia. 45. Universidad Nacional. Buenos Aires, Argentina.

HILL, K. & HURTADO, A.M. 1996. Aché life history: the ecology and
demography of a foraging people. Aldine de Gruyter, New York.

HOLMGREN, P.K., HOLMGEN, N.H. & L. C. BARNETT. 1990. Index
Herbariorum. Part I: The Herbaria of the World. New York Botanical
Garden, New York.

HUECK, H. 1978. Ecologia, composici6n e importancia economica: Los Bosques
de Sudamérica. Sociedad Alemana de Cooperaci6n Técnica Ltda (GTZ).
Germany.

KEEL, S. & HERRERA-MACBRYDE, O. 1997. Mata Atlantica: CPD Site SA18.
Mbaracayt Reserve, Paraguay. In: Davis, S.D., Heywood, V.H., Herrera-
MacBryde, O., Villa-Lobos, J., & Hamilton, A.C. (eds) Centres of Plant
Diversity. Volume 3: The Americas: 389-392. WWF and IUCN, IUCN
Publishing Unit, Cambridge.

KRAMER, K.U. 1993. Distribution patterns of major pteridophyte taxa relative to
those of angiosperms. J. Biogeogr. 20: 287-291.

LACLAU, P. 1994. La conservacion de los recursos naturales y el hombre en la
Selva Paranaense. Bol. Técnico de la Fundacién Vida Silvestre Argentina
No. 20. Vida Silvestre Argentina/WWF, Buenos Aires, Argentina.

MORI, S.A., BOOM, B.M., CARVALHO, A.M. DE, & DOS SANTOS, T.S.
1983. Southern Bahian moist forests. Bot. Rev. (Lancaster) 49: 155-232.

MORI, S.A. 1989. Eastern, extra-Amazonian Brazil. In: Campbell, D.G. &
Hammond, H.D. (eds) Floristic Inventory of Tropical Countries: 427-454.
The New York Botanical Garden, Bronx.

PICHI SERMOLLI, R. E. G. 1996. Authors of Scientific names in Pteridophyta: a
list of authors of names of ferns and fern allies with recommended standard
forms of their names including abbreviations. Royal Botanical Gardens,
Kew.

POULSEN, A. D. & TUOMISTO, H. 1996. Small-scale to continental distribution
patterns of neotropical Pteridophytes: the role of edaphic preferences. In:
Camus, J.M., Gibby, M. & Johns, R.J. (eds) Pteridology in Perspective.
Royal Botanic Gardens, Kew.

SMITH, A.R. 1972. Comparison of fern and flowering plant distribution with some
evolutionary interpretation for ferns. Biotropica 4: 4-9.

SMITH, A.R. 1993. Phytogeographic principles and their use in understanding fern
relationships. J. Biogeogr. 20: 255-264.

SOTA, E. DE LA. 1973. La distribuci6n geografica de las pteridofitas en el cono sur
de América meridonial. Bol. Soc. Argent. Bot. 15: 23-34.

TRYON, R.M. 1972. Endemic areas and geographic speciation in tropical American
ferns. Biotropica 4: 121-131.

TRYON, R.M. & TRYON, A.F. 1982. Ferns and allied plants with special
reference to Tropical America. Springer-Verlag, New York.

PTERIDOPHYTES OF MBARACAYU FNR 22

TUOMISTO, H. & POULSEN, A.D. 1996. Influence of edaphic specialization on
pteridophyte distribution in neotropical rainforests. J. Biogeog. 23: 283-293.

YOUNG, K.R. & LEON, B. 1989. Pteridophyte species diversity in the central
Peruvian Amazon: the importance of edaphic specialization. Brittonia 41:
388-395.

BOOK REVIEW

FLORA MESOAMERICANA Volume 1 Psilotaceae a Salviniaceae Moran R. &
Riba, R. (eds); Davidse, G., Sousa, M. & Knapp, S. (gen. eds) 1995. Universidad
Nacional Aut6noma de Mexico, Missouri Botanical Garden and The Natural History
Museum, London. 470 pp. Hardback. US$85.00. ISBN 968 36 4700 6.

This is the first of seven volumes of Flora Mesoamericana, and includes 32 families,
136 genera and 1398 taxa (species, subspecies or varieties), constituting the biggest
Pteridoflora published.

Flora Mesoamericana covers a geographical area from the Istmo de Tehuantepec
(southern Mexico) to the border between Panama and Colombia. It includes five
Mexican states (Tabasco, Chiapas, Campeche, Yucatan and Quintana Roo) as well as
Belize, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica and Panama.

Families are taxonomically arranged, while the genera and species are in
alphabetical order in those cases where there is not a Satisfactory taxonomic
arrangement. Keys are included for all the families, genera, species, subspecies and
varieties. There are general descriptions for each family and genus. For each taxon a
general description, habitat, general distribution, as well as distribution within the
area are included. Types are cited only for accepted names, synonyms are cited in
case they are important for the area. The book includes as well a very helpful
glossary. This book provides an excellent guide for all pteridologists working in
Latin America, and it is worthwhile to stress that one of the great merits of the work
is due to the fact that it is written in Spanish, making it very useful for the local users.

The descriptions of the species are very complete and easy to understand.
However, the keys are not always easy to follow, and perhaps some drawings would
be helpful in order to attain identification to species level in a more easy way. (In that
respect, it can help to visit the web site http://www.mobot.org/MOBOT/fm/).

It provides a very important tool for all those people interested in the Pteridoflora,
not only because the amount of species and information included, but also because it
fills an existing gap in descriptions and keys in Spanish and it can be used for a
wider geographical area than the one covered in the book.

M.C. Pefia-Chocarro

260 FERN GAZETTE: VOLUME 15 PART 7 (1999)

BOOK REVIEW

INDEX FILICUM SUPPLEMENTUM SEPTIMUM PRO ANNIS 1991-1995.
R. J. Johns. 1997. Royal Botanic Gardens, Kew. ix + 124 pp. Price £20.00. Softback.
ISBN 1 900347 26 1.

Index Filicum Supplement 7 has been published less than one year after Supplement
6, to the delight of all Index Filicum users. Congratulations are due to all at Kew involved
in its production. Covering five years and consisting of 124 pages, it is the smallest of the
supplements apart from Supplement 2, from 1913-1916 and with 60 pages. I look forward
to seeing future supplements appearing on a regular five year basis.

In format Supplement 7 follows essentially the style of Supplement 6, unfortunately
complete with the loathed Addenda, which may have been necessary in the days of
manual type setting, but is surely an anachronism in the computer era. Users of Index
Filicum should heed the plea on page viii to send reprints of their papers to Kew and
perhaps the Addenda will no longer be needed! A convenient change from the
Supplement 6 format is the setting of invalid names in a distinctive type face so that they
are very easily recognisable. The term ‘List of species’ used in the Table of Contents in
both Supplements 6 and 7 is not accurate, nor is the actual heading to this section,
‘Enumeratio familiarum, genera, specierumque alphabetica’ because it does not take
infraspecific taxa into account.

The Catalogus litteraturae contains some references published before 1991, most of
which are also listed in the Catalogus litteraturae of Supplement 6, e. g. Burrows, J. E.
(1990), South African Ferns and Fern Allies, all of R. C. Ching’s publications, and Parris,
B. S. (1986), Kew Bull. 41(1): 69-70; the new combinations in these works are included
in Supplement 6, but not in Supplement 7. Holttum, R. E. & Grimes, J. W. (1980), Kew
Bull. 34(3): 499-516 is listed in the Catalogus litteraturae for both Supplement 6 and
Supplement 7; in the former supplement all of the new combinations and new species are
listed while in the latter supplement only the two new species have been included. One
publication, Holttum, R. E. (1972), Blumea 20(1): 105-126, is listed here and in
Supplement 5; the new taxa and new combinations appear only in the latter. Another
example of carelessness occurs in the title of a cited paper; Smith, A. R. & Palmer, D. D.
(1995), Ctenitis rubiginosa, from Hawaii, belongs in Nothosperma. Amer. Fern J. 85(2):
63 — for Nothosperma, a non-existent genus, read Nothoperanema! One new
combination, Zygophlebia longipilosa (C.Chr.) L.E.Bishop, Amer. Fern J. 79(3):109
(1989): Polypodium longipilosum C.Chr., has been omitted from both Supplements 6 and
7, although all of the other new taxa and new combinations in the article were included in
Supplement 6.

The Introduction includes a proposed change of format to provide additional data
for new taxa and new combinations for Supplement 8 (1996-2000) onwards. These will
include references to illustrations in the publication where the new taxa and new
combinations are published and information on the type specimens such as collector,
locality, altitude, date of collection and location of holotype, isotypes and paratypes.
While any extra information concerning taxa is useful, the location of authentic
illustrations being particularly desirable, my personal preference is to see other
improvements to the Index Filicum Supplements, such as removing the Addenda and
reducing the number of errors. The production of Index Filicum on CD-rom should also
be a priority, particularly if the Addenda are to become a permanent feature.

B.S. Parris

FERN GAZ. 15(7) 1999 261

NEW PHILIPPINE GONIOPHLEBIUM (POLYPODIACEAE:
PTERIDOPHYTA)

J.F. BARCELONA’ & M.G. PRICE’

‘Philippine National Herbarium, National Museum, P.O. Box 2659, Manila,
Philippines. Current address: Botany Department, Miami University,
Oxford, OH 45056, U.S.A.

*P_O. Box 468, Michigan Center, MI 49254, U.S.A.

Key Words: Batanes, fern, Goniophlebium, Philippines, Polypodiaceae.

ABSTRACT
The new species Goniophlebium coadunatum (Polypodiaceae) from the Batanes
islands, between Luzon and Taiwan, is described. An ally of G. benguetense, also
from the Philippines, it is distinguished principally by its coadunate frond apex.

INTRODUCTION

Mount Iraya of Batan Island is the highest point in the wind-swept Batanes Islands,
located nearly equidistant from northern Luzon and southern Taiwan, which are
about 350 km apart. A detailed floristic analysis of the pteridophytes of Mt Iraya
was prepared by Barcelona (1994). Previously, seven fern taxa were described from
type collections made on Mt Iraya or in the Batanes, namely Athyrium trayense
Copel. [=Diplazium doederleinii (Luerss.) Makino], Cyathea fenicis Copel.,
Cyclosorus irayensis Copel. [=Sphaerostephanos irayensis (Copel.) Holtt.],
Grammitis fenicis Copel., Lygodium mearnsii Copel. [=L. japonicum (Thunb.) Sw.],
Pteris rigidula Copel. [=P. spinescens C. Presl], and Sphaerostephanos fenixii Holtt.
[=S. irayensis (Copel.) Holtt.]|. Two of these, G. fenicis and S. irayensis, are still
recognized as endemic to Mt Iraya and Batan Island, and we are here adding a third:

Goniophlebium coadunatum Barcelona et M.G. Price, sp. nov. (Fig. 1)

Goniophlebio benguetensi Philippinarum maxime affinis autem apice frondis
coadunato potissime differt. Etiam differt soris adaxialibus prominentioribus,
paraphysibus setulosis vice clavatis in centro sororum, pinnis latioribus ambitu basis
pinnarum angustioribus, glandulis minutis marginalibus palearum deficientibus.

Holotypus: J.F. Barcelona 988, Philippines, Batanes, Mt Iraya (PNH).

Rhizome long creeping, 2-5 mm across, surface uniformly dull blackish, not chalky,
internal sclerenchyma strands over 100. Paleae dense, persistent, bright brown,
lanceate, long attenuate, to 6 mm long, 1.5 mm broad at base, with projections of the
clathrate lattice forming lateral teeth. Stipe (7.4-) 26-45 cm long, (1.4-) 2.1-3.6 mm
across at base, stramineous to brown (green above, dark below when fresh), with
numerous minute maroon palea-base remnants. Lamina lanceolate-ovate, apex
coadunate, (14.5-) 38-51 x (7.5-) 21-32 cm, bearing (12-) 16-35 pairs of pinnae, thin-
Ppapyraceous, rachis with very narrow wing joining laminar tissue of pinnae, rachis
and costae with small, toothed paleae below, glabrous above. Pinnae sessile,

262 FERN GAZETTE: VOLUME 15 PART 7 (1999)

Figure 1: Goniophlebium coadunatum Barcelona & M.G. Price.
a. Habit of adult plant; b. Portion of the lower surface of a pinna showing impressed
sori and paleae; c. rhizome palea. (Barcelona 491). [Illustrated by Nemesio Diego, Jr.

(PNH)]

PHILIPPINE GONJOPLEBIUM COADUNATUM SP. NOV. 263

articulate, opposite to subopposite, often ranging to alternate distally, patent to
slightly ascending, the largest (4.4-) 12-15 cm x (5-) 8-12 mm, basal to submedial,
base broadly cuneate, often auricled on both sides, margins serrulate, apex
acuminate, uppermost pinnae adnate, decrescent to non-articulate segments, terminal
lamina lobed at base, then pinna-like. Veins forming one series of large areoles with
a fertile free-included-veinlet on each side of costa, then with irregular small areoles,
or free. Sori medial, round to slightly oval, 1-1.5 mm across, notably impressed,
bearing conspicuous variably-branched paraphyses with shiny dark maroon setae
shorter than mature sporangia. Spores pale yellow, 42.5-52.5 x 25-30 um, exospore
smooth, perispore longitudinally winged with polar protuberances.

Type: J.F. Barcelona 988, Philippines, Batanes, Batan Island, Basco, Valugan,
Dujtak, Mt Iraya, 100-200 m, epiphytic on trunks and petrophytic, 03 December
1993 (holotype: PNH; isotypes: K, MICH, MU, US). Paratypes: J.F. Barcelona 491,
Mt Iraya, 500 m, epiphytic on trunk, April 1991 (L, PNH, T1).

This amply distinct new species is most closely related to G. benguetense (Copel.)
Copel. which also has similar rhizome sclerenchyma strands, paleae, stipes, lamina
and pinna shape, sori, paraphyses and spores. It differs most notably from G.
benguetense by the consistently coadunate apex, a character not found in any
member of the ~G. percussum-group' in which Rddl-Linder (1990) placed that
species. Whilst the absence of the coadunate apex character in all closely-related
species facilitates the recognition of G. coadunatum, its presence in this species
weakens the definition of any such group. Other differences are that G. benguetense
has a more slender rhizome, shorter paleae having a thinner and grayish-brown lattice
and bearing fewer marginal teeth but with numerous minute marginal glands, pinnae
subtruncate at base but generally narrower, sori superficial or only very slightly
impressed, and receptacular paraphyses with clavate branch-tips near the center of
sori. The two known populations of G. coadunatum occur on trunks and rocks in
shade in the remaining patches of lowland evergreen forest on the lower slopes of Mt
Iraya from about 100-500 m whereas G. benguetense is a plant of mossy forests in
Luzon at higher elevations, to 1500 m.

Also related to G. benguetense is G. demersum (Brause) R6édl-Linder of the
Moluccas and New Guinea. It differs from G. coadunatum by its conform terminal
pinna, thicker lamina, broader rhizome paleae, supramedial and only very shallowly
impressed sori, and paraphyses broadly scaly.

The East Asian G. mengtzeense (Christ) R6dl-Linder, which extends to Luzon,
also has a coadunate laminar apex (although described as conform by Rédl-Linder,
1990) but differs from G. coadunatum by its chalky rhizome, lamina paler in color,
pinnae broader, ascending, more shallowly toothed, sori superficial, and paraphyses
palea-like, among other characters.

The recent monograph of Goniophlebium by Rédl-Linder (1990) and her revision
of the Philippine species (1987) provide useful information with descriptions,
synonymy, and specimen citations, and have greatly increased our confidence in
recognising this new species. In Barcelona (1994, p. 131, f. 18) this species appeared
as an unnamed Goniophlebium.

264 FERN GAZETTE: VOLUME 15 PART 7 (1999)

The finding of a species of Goniophlebium apparently restricted to a single
Philippine volcanic mountain of medium size (only slightly over 1000 m elevation),
has the precedent of G. terrestre Copel. which has been collected only from Mt
Makiling in south-central Luzon. Like G. coadunatum, G. terrestre is a forest shade
plant surviving only as a few local populations.

ACKNOWLEDGMENTS

We are grateful to the directors and curators of MICH, MU, and PNH for their
cooperation. This paper is partly derived from JFB's project with the National
Museum of the Philippines as well as her Master's thesis at the University of Santo
Tomas, Manila. A thesis grant was provided by the National Research Council of the
Philippines. Further thanks are given to Dr Romualdo M. del Rosario and JFB's
thesis committee members, to the people of Batan Island, especially Mrs. Asuncion
Fajardo and family, for their accommodation in Basco, and to Teodoro Baldimoro,
Christy and Manuel Cordel and Leopoldo Villegas for their assistance in the field.

REFERENCES
Barcelona, J.F. 1994. Pteridophyte flora of Mt. Iraya and vicinity, Batan Island,
Northern Philippines. MS Thesis. University of Santo Tomas, Manila.
Rodl-Linder, G. 1987. Goniophlebium of the Philippines (Polypodiaceae). Philipp. J.
Sci. 116: 151-176.
Rédl-Linder, G. 1990. A monograph of the fern genus Goniophlebium
(Polypodiaceae). Blumea 34: 277-423.

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The British Pteridological <
THE FERN GAZETTE

VOLUME 15 PART 7 1999

CONTENTS

MAIN ARTICLES Page

Checklist of the pteridophytes of the Mbaracayu Forest Nature Reserve, Paraguay
M.C. Pefia-Chocorro, B. Jiménez, G. Marin & S. Knapp 221

New Philippine Goniophlebium (Polypodiaceae: Pteridophyta)
J.F. Barcelona & M.G. Price 261

BOOK REVIEWS | 259, 260

THE FERN GAZETTE Volume 15 Part 6 was published on 22 December, 1998

Published by THE BRITISH PTERIDOLOGICAL SOCIETY, c/o Department of Botany,
The Natural History Museum, London SW7 5BD

ISSN 0308-0838

_ Printed by The Electronic Document Co., 132b King Street, London W6 0QU