BLen 1993 ATaxonomicRevCampyloneurum Thesis

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A Taxonomic Revision of the Fern Genus Campyloneurum (Polypodiaceae)
Thesis · January 1993
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Blanca Leon
Afdeling for Systematisk Botanik - Biologisk Institut, Aarhus Universitet
Nordlandsvej 68, DK 8240 Aarhus, Danmark
A TAXONOMIC REVISION
OF THE FERN GENUS
CAMPYLONEURUM
(POLYPODIACEAE)
by
Blanca León
Museo de Historia Natural
Facultad de Ciencias Biológicas
Universidad Nacional Mayor de San Marcos
Lima
Peru
1993
Afhandling indleveret til det Naturvidenskabelige Fakultet ved Aarhus
Universitet til bedømmelse for Ph.D. graden
Vejleder: Lektor Benjamin Øllgaard

To my parents, Efrain León and Lucila Bocángel de León, and
to my husband, Kenneth R. Young.

León, Revision of Campyloneurum p. iii.
TABLE OF CONTENTS
List of Figures iv-v
List of Tables vi
Note vii
Danish summary - Dansk sammenfatning viii-xii
Acknowledgements xiii
I. INTRODUCTION 1
II. MATERIALS AND METHODS 1-2
III. TAXONOMIC HISTORY 3
IV. MORPHOLOGY 4-12
V. CYTOLOGY AND BIOCHEMISTRY 12-13
VI. ECOLOGY AND GEOGRAPHICAL DISTRIBUTION 13-18
VII. CONSERVATION AND USES 18-19
VIII. INFRAGENERIC CLASSIFICATION AND PHYLOGENY 19-23
IX TAXONOMY
Classification 23-93
Nomina Dubia 93-94
Excluded Taxa 94
X. CITED REFERENCES 95-99
XI. LIST OF TAXA 100-102

León, Revision of Campyloneurum p. iv.
LIST OF FIGURES
Figure 1. Stem morphology in Campyloneurum.

Figure 2. Cross section of the root in Campyloneurum.
Figure 3. Cross section of the stem in Campyloneurum.
Figure 4. Stem scale attachment in Campyloneurum.
Figure 5. Stem scale morphology in Campyloneurum.
Figure 6. Margins of stem scales in Campyloneurum.
Figure 7. Stem scale cell structure in Campyloneurum.
Figure 8. Cross section of the petiole in Campyloneurum.
Figure 9. Lamina morphology in Campyloneurum.
Figure 10. Epidermal morphology in Campyloneurum.
Figure 11. Leaf epidermis morphology in Campyloneurum.
Figure 12. Cross section of the leaf in Campyloneurum.
Figure 13. Types of hair in Campyloneurum.
Figure 14. Pattern of venation of Campyloneurum.
Figure 15. Heteroblastic development of leaves in Campyloneurum amphostenon.
Figure 16. Paraphyses in Campyloneurum.
Figure 17. Spore morphology in Campyloneurum.
Figure 21. Altitudinal range in Campyloneurum species.
Figure 22. General distribution of Campyloneurum species by countries.
Figure 23. Andean regional division and distribution of Campyloneurum species.
Figure 24. Distribution of Campyloneurum abruptum and C. tenuipes.
Figure 25. Distribution of Campyloneurum acrocarpon and C. chlorolepis.
Figure 26. Distribution of Campyloneurum aglaolepis,, C. austrobrasilianum and C.
centrobrasilianum.
Figure 27. Distribution of Campyloneurum amphostenon var. amphostenon and C. amphostenon
var. irregulare.
Figure 28. Distribution of Campyloneurum anetioides and C. aphanophlebium.
Figure 29. Distribution of Campyloneurum ensifolium and C. cochense.
León, Revision of Campyloneurum p. v.
Figure 30. Distribution of Campyloneurum angustifolium.

Figure 31. Distribution of Campyloneurum angustipaleatum.
Figure 32. Distribution of Campyloneurum asplundii.
Figure 33. Distribution of Campyloneurum brevifolium.
Figure 34. Distribution of Campyloneurum coarctatum and C. chrysopodum.
Figure 35. Distribution of Campyloneurum costatum and C. decurrens.
Figure 36. Distribution of Campyloneurum cubense, C. falcoideum and C. lorentzii.
Figure 37. External morphology of stems in Campyloneurum densifolium and C. amphostenon.
Figure 38. Distribution of Campyloneurum densifolium, and C. fallax.
Figure 39. Distribution of Campyloneurum fasciale and C. minus.
Figure 40. Distribution of Campyloneurum fuscosquamatum, C. inflatum and C. nitidum.
Figure 41. Distribution of Campyloneurum macrosorum and C. magnificum.
Figure 42. Distribution of Campyloneurum nitidissimum var. nitidissimum and var. latior.
Figure 43. Distribution of Campyloneurum ophiocaulon and C. oxypholis.
Figure 44. Distribution of Campyloneurum pascoense, C. tucumanense and C. wurdackii.
Figure 45. Distribution of Campyloneurum phyllitidis.
Figure 46. Distribution of Campyloneurum repens.
Figure 47. Distribution of Campyloneurum solutum and C. rigidum.
Figure 48. Distribution of Campyloneurum sphenodes.
Figure 49. Distribution of Campyloneurum sublucidum and C. vulpinum.
Figure 50. Distribution of Campyloneurum xalapense and C. oellgaardii.
Figure 51. Proposed evolutionary diagram for Campyloneurum species groups.
Figure 52. Consensus tree for Campyloneurum species.
Figure 53. Consensus tree for Campyloneurum groups.
León, Revision of Campyloneurum p. vi.
LIST OF TABLES
Table 1. Variablity of morphological characters in Campyloneurum.

Table. 2. Chromosome numbers reported in Campyloneurum.
Table 3. Habitat preferences in Campyloneurum species.
Table 4. Geographic distribution by country of Campyloneurum species.
Table 5. Regional distribution of Campyloneurum species.
Table 6. Regional affinities of Campyloneurum floras.
Table 7. Regional distribution of species groups.
Table 8. Uses and common names of Campyloneurum.
Table 9. Latitudinal and altitudinal distribution, habitat preference and number of collections in
Campyloneurum species.
Table 10. Characters and character states used for phylogenetic analysis in Campyloneurum species.
Table 11. Data matrix of Campyloneurum species.
Table 12. Characters and character states used for phylogenetic analysis in Campyloneurum species
groups.
Table 13. Data matrix for Campyloneurum species groups.
León, Revision of Campyloneurum p. vii.
Note: Nomenclature novelties herein are not presented for purposes of valid publication.
León, Revision of Campyloneurum p. viii.
Dansk Sammenfatning
Campyloneurum er en af de få slægter i bregnefamilien Polypodiaceae s. str.,
der kun forekommer i den Nye Verdens Troper. Slægten blev grundlagt af Prel
(1836), på basis af arternes særlige nervation; Presl medregnede både arter med hele
og pinnate blade. Definitionen af Campyloneurum forblev uændret indtil Tryon &
Tryon (1982a) udelukkede de pinnate arter og medregnede en art, der tidligere var
henført til slægten Hyalotrichopteris.
Nærværende afhandling har til formål at afklare slægtens afgrænsning, og at
gøre slægten til genstand for en taxonomisk revision, at definere arterne ved hjælp
af morfoloiske karakterer og belyse deres indbyrdes slægtskab.
Omkring 4000 eksemplarer blev unersøgt, især ved Biologisk Institut, Aarhus
Universitet (AAU), og ved det Nationale San Marcos Universitet (USM) i Lima,
Peru. Mange af disse eksemplarer blev udlånt til projektet fra de følgende herbarier:
B, BM, BR, CAY, F, GB, GH, HJBLH, LD, MICH, MO, MU, P, S, UC, US, og W, eller
blev undersøgt under besøg på folgende herbarier C, CUZ, F, GH, HUT, K, LOJA,
LPB, MO, NY, QCA og UC (herbarieakronymer ifølge Holmgren et al. 1990. Index
Herbariorum Part I. 8th ed.).

Campyloneurum er defineret af en speciel (cyrtophleboid) nervation (Kap. III
og VIII); det vigtigste traek ved denne er, at de costale areoler har en fri udløbende
nerve, og at de extra-costale areoler har (1-) 2-5 fri udløbende nerver. Slægten
omfatter 47 arter og 4 varieter (Kap. VIII), både arter med hele og med pinnate
blade, forunden en art, den har været henført til slægten Hyalotrichopteris.
Slægtens morfologi gennemgås, og visse kritiske karakterer, såsom
behåring, sporangier og parafyser, beskrives for første gang (Kap. IV).
Slægten deler mange karakterer med andre slægter i Polypodiaceae.
Stænglen (rhizomet) hos Campyloneurum er krybende, med skæl og

León, Revision of Campyloneurum p. ix.
dorsiventralt arrangerede rødder og blade. Rhizomskæl har spillet en vigtig
rolle for mange Polypodiace-slægters taxonomi, fordi de leverer mange
konstante bygningstræk. Dette gælder også Campyloneurum, hvor materiale
uden stængel ofte ikke kan bestemmes til art. Sterile og fertile blade er ens i
form og størrelse; de er i reglen anbragt i to rækker langs stænglen, og falder
af ved et distinkt løsningslag. Bladpladen er hel hos de fleste arter; kun C.
decurrens og C. magnificum har fjersnitdelte bladplader. Med undtagelse af
en enkelt art, Campyloneurum aphanophlebium, beskrives arterne som glatte.
Imidlertid er der her påvist tre typer behåring hos flere arter, samt tidligt
affaldende skæl. Fuldt udvoksede blades nervation kan inddeles i 4 typer,
der danner grundlag for en inddeling i artsgrupper (Kap. VIII).
Sporehushobene er afrundede eller elliptiske, uden indusier, og 1-2 (-3) mm
i diameter. Sori sidder på bladundersiden, på afrundede eller elliptiske
områder i niveau med bladoverfladen, oftest i to rækker mellem
primærnerverne, men undertiden tre eller flere hos C. brevifolium gruppen,
og undertiden i en enkelt række hos C. anetioides og C. falcoideum.
Forekomsten af parafyser, organer der kan findes blandt sporangierne, blev
undersøgt hos alle arterne. De findes hos 9 arter, og er gerne mindre end
sporangierne, og tyndere end sporangiestilkene; der findes to typer: simple
og dendritiske. Sporerne er ellipsoidale, de aborterede dog mere afrundede
og kollaberede, 40—100 x 30—60 µm; deres overflade er forsynet med
spredte sfæriske legemer eller granuli under under et tyndt perispor.
Exosporet er mere eller mindre vortet.
Campyloneurum er tidligere anset for at stå nærmest de andre
neotropiske slægter Microgramma og Niphidium, men anses her for at stå
særlig nær Pleopeltis (Kap. IV).

León, Revision of Campyloneurum p. x.
De fleste af arterne hører til i skov-vegetation (Kap. VI), især
stedsegrøn skov, men C. xalapense findes i delvis løvfældende skov i
Mexico. De fleste arter forekommer på mange forskellige mikrohabitater på
forstyrrede eller uforstyrrede skov-arealer. Seksogtyve arter hører til på
skyggede og fugtige mikrohabitater i sluttet skov. De fleste af disse arter er
epifyter på lavtsiddende grene eller på træstammer i de lavere etager af
skoven. Nitten arter forekommer almindeligvis i lysåbne skove eller
rydninger, på klipper, i klipperevner og i skovkanter. De er bredt tilpassede
og kan vokse som epifyter, på jorden og klipper (f.eks. C. cochense, C.
ensifolium). Det er ikke overraskende at finde de videst udbredte arter
blandt disse 19 (f.eks. C. angustifolium og C. phyllitidis). De fleste arter
vokser i højdeintervallet 1000—4500 m o.h. Kun syv er almindelige under
1000 m o.h. De fleste arter har en vid højde- og nord-syd-udbredelse.
Seksogtyve arter har en højdeudbredelse på over 1000 m; de geografisk
vidtudbredte arter er blandt disse arter (f.eks. C. angustifolium, C. brevifolium,
C. phyllitidis). Tilsyneladende har de arter, der har en højdeudbredelse på
under 1000 m, også en relativt begrænset geografisk udbredelse; de kræver
derfor særlig bevågenhed i forbindelse med naturbevarelse (Kap. VII).
De fleste af arterne forekommer i mere end 3 lande, og the artsrigeste
områder er i de tropisk sydamerikanske bjerge, mens de artsfattige områder
er i subtroperne, på små øer og i ikke-montane områder.
I fem hovedområder er slægten særlig rigt repræsenteret: Mexico,
Mellemamerika, Andes, Brasilien, og Caribien. Den Mexikanske region
omfatter 12 arter, hvoraf to er endemiske; regionens affinitet er nærmest til
Mellemamerika og den Caribiske region. Den Mellemamerikanske region
har 16 arter, hvoraf en er endemisk; denne regions affinitet er nærmest til

León, Revision of Campyloneurum p. xi.
den Mexicanske region, Andes, og den Caribiske region. I Andes
forekommer 35 arter og 14 af disse er endemiske. Det nordlige Andes har
32, det centrale Andes 23 arter og det sydlige Andes kun fire arter. I den
Brasilianske region findes ni arter, hvoraf fem er endemiske; denne regions
største affinitet er til Andes. Kun tretten arter forekommer i de
sydamrikanske subtroper; fire af disse arter forekommer også i de
nordamerikanske subtroper, og de er alle vidtudbredte. I Caribien har de
Store Antiller 10 arter, hvoraf 2 er endemiske, mens de Små Antiller kun har
fire arter, alle vidtudbredte. Denne behandling af slægten
Campyloneurum er baseret på herbarie- og feltstudier, og den morfologiske
artsopfattelse er anvendt. Kategorien varietet er anvendt i to tilfælde, for
mindre distinkte elementer i C. amphostenon and C. nitidissimum. To nye
arter er beskrevet: C. ensifolium og C. oellgaardii.
Jeg anvender en uformel inddeling af slægten i ti artsgrupper (Kap.
VIII); arterne i hver gruppe har en række fælles morfologiske karakterer og
voksestedspræferencer, og formodes at repræsentere selvstændige
udviklingslinier i slægten. I alfabetisk orden er de:
1) C. amphostenon gruppen, bestående af Campyloneurum amphostenon,
C. asplundii, C. chlorolepis, C. densifolium, C. fallax, C. lorentzii, C. rigidum og C.
solutum;
2) C. angustifolium gruppen, der består af C. aglaolepis, C. angustifolium,
C. angustipaleatum, C. angustifoliaceus, C. austrobrasilianum, C.
centrobrasilianum, og C. ensifolium.
3) C. brevifolium grupppen, der består af C. brevifolium, C. pascoense, C.
nitidissimum og C. tucumanense;
4) C. magnificum gruppen, bestående af C. decurrens og C. magnificum;

León, Revision of Campyloneurum p. xii.
5) C. aphanophlebium gruppen, bestående af C. anetioides og C.
aphanophlebium;
6) C. phyllitidis gruppen, med fire arter: C. abruptum, C. nitidum, C.
phyllitidis, C. tenuipes og C. wurdackii;
7) C. repens gruppen, bestående af C. acrocarpon, C. fasciale, C.
fuscosquamatum, C. minus, C. ophiocaulon og C. repens;
8) C. sphenodes gruppen, med C. sphenodes, C. chrysopodum, C.
coarctatum, C. falcoideum, C. inflatum, C. sublucidum, og C. oellgaardii;
9) C. vulpinum gruppen, med C. vulpinum, C. cubense, og C. oxypholis, og
10) C. xalapense gruppen, der består af C. cochense, C. costatum og C.
xalapense.
León, Revision of Campyloneurum p. xiii.
ACKNOWLEDGEMENTS
Campyloneurum became part of my life thanks to Dr. R. M. Tryon, who
suggested I study it. This thesis was begun during a DANIDA Fellowship visit
(1984-85) to the Institute of Biological Sciences, in Aarhus.
I received many suggestions and comments on the taxonomy of the genus
from, and discussed many aspects of the nomenclature and species definitions with
Drs. R. C. Moran, D. Nicolson, A.R. Smith, A. F. Tryon, and R. M. Tryon. I am
grateful to Dr. A. F. Tryon for providing me with her time and beautiful SEM
photographs. I am especially grateful to R. G. Stolze for help and encouragement.
I thank the curators for loaning or making available specimens from the
following institutions: B, BM, BR, C, CAY, CUZ, F, GB, GH, HJBLH, HUT, K, LD,
LOJA, LPB, MICH, MO, MU, NY, P, QCA, S, UC, US, W. I also thank A. Sloth and
all the personnel of the Insitute of Biological Sciences, Aarhus.

León, Revision of Campyloneurum 1
I. INTRODUCTION (1988) presented a general overview of the genus,

wherein he recognized 50 species,
Campyloneurum is one of the few genera of the including several new species, based on
Polypodiaceae (s. str.) restricted to the New characters found on the scales of the stem (cf.
World. This genus was erected by Presl (1836), Sota, 1960). Although there has been an
who included in it both entire and one-pinnate improvement on the use of more reliable
leaved species, and who defined it by its characters for species limitation, controversy in
reticulate venation . The definition of the number of species recognized in the genus,
Campyloneurum went unchanged until Tryon & and in their nomenclature has persisted. For
Tryon (1982a) proposed the exclusion of the one- these reasons, a monographic revision was badly
pinnate species and the inclusion of the genus needed in the genus.
Hyalotrichopteris based on soral position.
In this study, Campyloneurum is again defined
by its reticulate areolate venation. Important II. MATERIALS AND METHODS
characters of the venation are the presence of
costal areoles carrying one excurrent free veinlet, MATERIALS
and of non-costal areoles carrying (1-) 2-5 free About 4000 specimens were examined at the
excurrent veinlets. Herbarium of the Institute of Biological Sciences
Most species of Campyloneurum were of the University of Aarhus (AAU), Aarhus,
described during the middle of the 19th and the Denmark, and the Herbario San Marcos (USM) in
beginning of the 20th century. Circumscriptions Lima, Peru. Loans were obtained from the
of the species were based on foliar morphology following herbaria: B, BM, BR, CAY, F, GB, GH,
and/or venation (cf. Mettenius, 1864; Hooker, HJBLH, LD, MICH, MO, MU, P, S, UC, US, and
1864; Hooker & Baker, 1874; Christ, 1902; W, or were seen on visits to B, C, CUZ, F, GH,
Rosenstock, 1909; Hieronymus, 1904). During HUT, K, LOJA, LPB, MO, NY, QCA and UC
that time no monographic treatement was (abbreviations according to Holmgren et al., 1990.
attemped for the genus, although accounts for Index Herbariorum Part I. 8th ed.).
number of species in the genus were available Live material from five taxa (Campyloneurum
because of the work of Fée (1852), Moore (1861), amphostenon, C. angustifolium, C. nitissimum var.
and Smith (1875), and also because of floristic latior, C. phyllitidis, and C. repens) was examined
studies done by Mettenius (1864), Sodiro (1893), for morphological and cytological studies. These
and Christ (1902). plants were cultivated in Lima and Aarhus. The
Difficulties at the species level were indicated specimens cultivated in Lima, consisting of four
by Fée (1852, 1864) and Smith (1875). Both of the species (excluding C. angustifolium), came
authors mentioned problems in defining the from my collection in Peru, while the specimens
species based on leaf morphology (Fée, 1852, cultivated in Aarhus (C. angustifolium and C.
1864) and/or pattern of venation (Smith, 1875). phyllitidis) were collected by Danish botanists in
During the last 30 years, several pteridologists Ecuador.
have contributed to a better understanding of
species circumscription in Campyloneurum. Sota METHODS
(1960) treated the meridional South American Measurements of macroscopic characters
species. He was one of the first pteridologists to (stem diameter, distance between phyllopodia,
recognize the value of stem scale characteristics petiole length and width, lamina width and
in the genus. He recognized 36 species. Later, length, angle of divergence of the primary veins,
Meyer (1964) made a first attempt to review the and the distance between primary veins) were
genus; based mostly on foliar morphology, she made on five to thirty complete representative
recognized 29 species. Lellinger (1977, 1984) herbarium specimens of each species.
called attention to several Andean and Central Microscopic characters (width and length of
American species when he made new the stem scales, width of the cell walls of selected
nomenclatural combinations. Recently Lellinger representative stem scales, width and length of
the cellular lumen of stem scales, sporangia following the acetolysis technique (Erdtman,
length, number of cells of the sporangial annulus, 1943). Pretreated spores were mounted in
width and length of the spore) were measured on ethylacetate for observation under a compound
ten representative specimens from each species microscope. Others were treated for observation
using a compound microscope. under the scanning microscope (JEOL JSM-840 at
Material for anatomical observations was the University of Aarhus) using gold coating;
obtained from both live and herbarium photographs were made using AGFA PAN 100
specimens. Sections were made from the roots, film. Additional spore material was studied with
stems, and leaves of seven species: Dr. Alice F. Tryon at the Gray Herbarium of the
Campyloneurum amphostenon, C. angustifolium, C. University of Harvard. Photographs of spores at
aphanophlebium, C. densifolium, C. fuscosquamatum, Harvard were processed with Polaroid film.
C. sphenodes, and C. vulpinum. Dried material Microphotographs were taken of most of the
was rehydrated in an aqueous solution of ethanol morphological characters and were procesed at
before it was embedded in paraffin. Sometimes the laboratory of the Biological Institute in
before embedding, the samples required Aarhus, Denmark.
extraction of air bubbles using a vacuum Phylogenetic analyses were performed using
chamber. Sections were made using an automatic PAUP version 3.0 (Swofford, 1990) in a
microtome. For fresh material, small samples Macintosh II computer.
were first cut using a freezing microtome; then Two cladistic analyses were performed using
the sections were dyed with Chlorazol-black and the heuristic search method, and without ordered
mounted in Euparal (cf. Radford et al. 1964). states or weighted characters (Wiley et al., 1991).
Stem indument was obtained from herbarium In this way "a priori" inferences were avoided.
specimens. Scales were mounted directly in The first analysis was done for the species based
Hoyer's solution or Glycerine jelly. Some on a matrix of 23 morphological characters. The
mounted samples in Hoyer's solution were left in second analysis was done for the species groups,
the oven at 60°C for two days allowing liberation based on a matrix of 12 characters. In both cases
of air bubbles from the sample. an ancestor was defined.
Foliar epidermal samples were obtained from Descriptive morphological terms are used
herbarium specimens for indument and stomata according to the Systematics Association
examination. Small sections were taken from the Committee (Taxon 9: 245-257. 1960), except for
middle of the lamina, and then treated with the use of linear-lanceolate instead of narrow
elliptic and lanceolate instead of elliptic, when
fuchsin-KOH at 60oC for two to four days,
both ends are gradually narrowed.
depending on the thickness of the lamina. The
In this study, the species are presented in
sections were cleaned in absolute ethylic alcohol,
alphabetical order. Lists of selected revised
then small drops of hydrocloric acid were added
specimens are included after the morphological
before they were cleaned again in alcohol and
descriptions and distribution information.
mounted in Euparal.
Abbreviations of bibliographic references
The pattern of lamina venation was examined
follow those of Stafleu & Cowan (1976) for books,
in adult samples from herbarium specimens.
and those of the Botanico Periodicum Huntianum
Cleared leaves were obtained following the
(Lawrence et al.,1968) for periodicals. Author's
fuchsin-KOH technique used for epidermal
name abbreviations follow those of the Authors
studies, as mentioned above. Heteroblastic
of Plant Names (Brummit & Powell, 1992).
development was studied in Campyloneurum
amphostenon, based on specimens collected in
Peru.
Measurements of sporangia, spores, and
paraphyses were made on material mounted in
Hoyer's solution. This material was collected
from herbaria specimens. Spores were treated
III. TAXONOMIC HISTORY Hoshizaki (1982). It was considered as a genus

by Ching (1940), Copeland (1947), Sota (1960,
The genus Campyloneurum ("kampylos" arched 1973), Duek (1971), Long & Lakela (1971), Crabbe
and "neuron" nerve) was described by Presl et al. (1975), Pichi-Sermolli (1977), Lellinger
(1836) in his work Tentamen Pteridographiae, (1977, 1984, 1985, 1988), Smith (1981), Tryon &
together with other genera considered before as Tryon (1982a, 1982b), Proctor (1985), Mickel &
Polypodium. Presl characterized his new genus by Beitel (1988), and León (1993).
areolate venation with two or more free excurrent The interpretation of Campyloneurum as a
veinlets per areole and by mainly entire leaves. genus including species with one-pinnate and
Brown (Bennet & Brown, 1838) proposed in entire leaves has been used by most
Polypodium the section Cyrtophlebium, placing in pteridologists. However, Tryon & Tryon (1982a)
it the American plants with the characters of the proposed a new generic concept based on the
genus Campyloneurum proposed by Presl, i.e. soral position, which allowed them to exclude
leaves mostly entire, reticulate venation, areoles species with 1-pinnate leaves and to include the
formed by arching veins, and always two series monotypic genus Hyalotrichopteris W. Wagner.
of sori except next to the costa. Smith (1841) The exclusion by Tryon & Tryon (1982a) of the
raised Brown's Cyrtophlebium to genus level, one-pinnate species (Campyloneurum decurrens
citing Campyloneurum as a synonym. Thus, the and C. magnificum) was based on the sori, which
name proposed by Smith was nomenclaturally are born at the tip of the excurrent veinlets. This
illegitimate. Mettenius (1856) proposed character was considered by Tryon & Tryon
Cyrtophlebium as a subgenus, but he included (1982 a) to ally those species with Polypodium.
within it species from tropical America and Asia However, this exclusion has not been widely
that belong today to such different genera as accepted (e.g. Lellinger, 1988; Hennipman et al.,
Pyrrosia, Polypodium, and Campyloneurum. Later, 1990; León, in press) because other characters
however, Mettenius (1857) used the subgenus such as the areolate venation and stem scales ally
Cyrtophlebium as the section proposed by Brown, the one-pinnate species with Campyloneurum .
i.e. for tropical American plants. The inclusion of Hyalotrichopteris anetioides by
Among the first authors to accept Tryon & Tryon (1982a, b) was due to the medial
Campyloneurum as a genus were Link (1841), position of the sori on the excurrent veinlets,
Hooker & Bauer (1842), Fée (1852, 1857, 1869), together with the presence of reticulate venation
Smith (1854, 1875), Hooker (1859), and Moore and leaf indument formed by branched hairs.
(1861), among others. Fée (1852) added some Lellinger (1988) did not accept this inclusion,
species previously in Marginaria, under the although he did accept a close link between both
ortographic variant Campyloneuron. genera, calling Hyalotrichopteris a "satellite genus"
Presl (1836) had published Campyloneurum of Campyloneurum. However, Hennipman et al.
without designating a type. In 1875, Smith made (1990) and León (in press) have included
the typification, selecting Polypodium repens (C. Hyalotrichopteris in the synonymy of
repens). Campyloneurum.
During the 19th century, other authors placed In this treatment, Campyloneurum is
the species of Campyloneurum in different considered a genus defined by its reticulate
subgeneric categories, thus as a section of venation, with primary parallel veins from which
Polypodium by Klotzsch (1847) and Diels (1899); curved veins arise and anastomose to form
and as a subgenus of Polypodium by Kunze (1850, primary areoles, and by costal areoles bearing
1853), Eaton (1860), Hooker (1864), Baker (1870), one included excurrent veinlet, and non-costal
Hooker & Baker (1874), Sodiro (1893), and Christ areoles, entire or divided, with most included
(1897). free veinlets excurrent. The genus consists of
In this century, there were still diverse criteria species with entire and one-pinnate leaves.
for the treatment of the group. It was considered
a subgenus of Polypodium by Christensen (1906),
Kramer (1954), Proctor (1977), Stolze (1981), and
IV. MORPHOLOGY persistent in most herbarium specimens.

Pruinosity is present in some species; a persistent
ROOTS whitish wax covers the surface of the stem of C.
The roots are located opposite the leaves on amphostenon, C. angustifolium, C. densifolium, and
the ventral side of the stem (Fig. 1 a-c). The roots C. solutum, while in C. cochense and C. lorentzii it
usually form two rows, although three can be is less common.
found in some individuals of Campyloneurum The distance between neighboring
amphostenon. The diameter and branching of the phyllopodia is variable (Table 1), for example in
roots are variable, as is the distance between Campyloneurum angustifolium the distance is 1-4
roots. The abundance of roots appears indirectly mm, and in C. coarctatum it is 10-15 mm. As in
related to the degree of creeping; also habitat other genera of the Polypodiaceae (e.g. in
conditions may influence it, similar to the case of Pyrrosia, Hovenkamp, 1986), this distance allows
Pyrrosia as described by Hovenkamp (1986). the recognition of two states of stem morphology:
Short-creeping stems in close contact with the long-creeping and short-creeping, based on the
substrate tend to develop abundant, spongy- relation between phyllopodium distance and
looking roots (e.g. C. brevifolium, C. pascoense, C. stem diameter. Long-creeping stems (Fig. 1 b)
phyllitidis ), while long-creeping stems in loose are here defined by the smallest phyllopodium
contact with the substrate (e.g. C. coarctatum, distance being larger than the smallest stem
C.fasciale , C. repens ) tend to have fewer roots. diameter (e.g. C. falcoideum and C. sphenodes). In
The roots are endogenous (cf. Ogura, 1972) short-creeping stems (Fig. 1 a), the largest
and the epidermis has numerous root hairs. The phyllopodium distance is equal to or smaller than
vascular strand is diarch (Figs. 2 a, b) and is the largest stem diameter (e.g. C. angustifolium
surrounded by a cortex or stereom (sensu Ogura, and C. costatum). A few species (C. acrocarpon, C.
1972), (Fig. 2 b), which is differentiated into an amphostenon, C. asplundii, C. densifolium, C.
external parenchymatous cortex and an internal lorentzii, C. minus, and C. nitidum) may exhibit
schlerenchymatous cortex. The anatomy of the both states, which can be interpreted as an
roots in Campyloneurum is similar to that of other adaptation to different habitat conditions, such as
genera in the Polypodiaceae, such as rock crevices or tree branches.
Microgramma, Niphidium and Pleopeltis, as was The stem frequently branches laterally along
shown by Zlotnik (1991). its axis. Branching was observed in
Campyloneurum amphostenon, C. asplundii, C.
STEM coarctatum, C. cochense, C. densifolium, C.
The stem in Campyloneurum is characterized falcoideum, C. lorentzii, C. ophiocaulon, C. pascoense,
by a creeping habit, by its dorsiventrality, and by C. solutum, C. sphenodes , and C. vulpinum. Also,
having an indument of scales (Fig. 1 a-c). The Sota (1960) observed branching of the stem in C.
stem is terete and usually has two rows of aglaolepis and C. tucumanense, and Wagner &
alternate phyllopodia on the dorsal side, while Farrar (1976) in C. anetioides.
the ventral side supports the roots. Three The stem in all species of Campyloneurum has
discontinuous rows of phyllopodia are unusual the cortex and pith consisting mostly of
in the genus. This phenomenon was observed parenchyma, a group or nest of sclereids in the
only in C. angustifolium ; similar observations in cortical parenchyma, and a dictyostele (Fig. 3 a-
this species were reported by Hovenkamp (1990). d). The stele in the genus was defined as a
The diameter of the stem ranges from 1 to 20 perforated dictyostele by Ogura (1972).
mm (Table 1). For example, in Campyloneurum The epidermis is formed by a single layer of
vulpinum the stem is 1-2 mm wide and in C. regular, oblong cells with a thin cuticle (Fig. 3 a).
pascoense it is 15-20 mm. The scales are appendages of the epidermis.
The color of the stem in most species is green, The parenchyma in Campyloneurum is found
and usually turns brown or atropurpureus when in the cortex and pith, and is characterized by
dry. However in some species, such as cells with thin walls (Fig. 3 b, d), bearing granules
Campyloneurum sphenodes, the green color is of starch and tanines (Fig. 3 a-d). These features
were also observed by Zlotnik (1990). Sometimes Similarly, the stem scales are very valuable in
a collenchyma occurs only in the cortex, beneath Campyloneurum, as was recently corraborated by
the epidermis; it has thicker walls, as observed in Lellinger (1988).
C. fuscosquamatum and C. sphenodes (Fig. 3 a-c). In Campyloneurum, the persistence of the scales
The presence of nests of sclereids is one of the stem varies among species. Those species
striking character in the cortex and pith. The with long-creeping stems have mostly caducous
color of these sclereid cells is brown as a result of scales, although they persist or are more
a pigment called phlobaphene, which may abundant on young parts, such as the stem apex
prevent the decay of the tissue (Ogura, 1972). and lateral branches (e.g. C. repens and related
The presence of nests of sclereids has been species). The stem scales partially overlap each
related to the age of the stem by Wagner & Farrar other in three or four layers, but the apices are
(1976) in Campyloneurum anetioides, with young usually spread or, in a few species, adpressed.
plants having fewer nests. In this study it was The scales of the stem have a wide range of
observed, however, that the size and abundance variation in shape, from linear, as in
of nests are variable among examined species Campyloneurum angustipaleatum and C. coarctatum
(Fig. 3 a-d). It appears that the patterns observed (Fig. 5 i), to broadly ovate, as in C. fallax and C.
may be related instead to the thickness of the nitidum. The most common shapes in the genus
endodermis. Sometimes nest of sclereids are few, are narrowly ovate, as for example in C. costatum
small, formed by 2-4 cells, and scattered among and C. nitidissimum, or ovate as in C. aglaolepis, C.
the parenchyma cells in those species with amphostenon, and C. solutum (Fig. 5 h). The shape
vascular bundles surrounded by thick of the scale of the stem is usually constant within
endodermis (C. angustifolium, C. densifolium, and a species. Rarely two shapes can be found in the
C. sphenodes, Fig. 3 a). Nests of sclereids are more stem of an individual, as was observed in some
numerous and formed by more than 4 cells in individuals of C. xalapense with both linear and
species with vascular bundles without a thick narrowly ovate scales. Rare in the genus is the
endodermis (C. fuscosquamatum and C. vulpinum, presence of bullate scales, as in C. minus, or
Fig. 3 b-d). It remains to be clarified if the slightly bullate, as in C. acrocarpon.
arrangements observed here have some The area of attachment of the scale to the
taxonomic value, as was shown by Hovenkamp epidermis is differentiated in cell structure and
(1986) in Pyrrosia. coloration from the rest of the body of the scale
In cross section, the perforated dictyostele (Fig. 4 a, b). The basal projection of the scale
shows (3-) 5-11 meristeles arranged in an lamina and the position of the attachment allows
elliptical ring (Figs. 1 c; 3 a, b). The vascular the recognition of three types of scales:
bundle is composed of xylem surrounded by "pseudopeltate", "peltate", and "basifix", as was
phloem. The pericycle consists of three layers of proposed by Hovenkamp (1986) for Pyrrosia.
cells (Fig. 3 b). The endodermis usually appears Pseudopeltate scales are the most common type
as a protecting sheath with strongly in the genus (Fig. 4 a, 5 e), as are found in
differentiated thick cell walls surrounding the Niphidium, Pleopeltis, and some species of
vascular bundle (Fig. 3 a, b). However, in Polypodium. These scales have two separate
Campyloneurum vulpinum (Fig. 3 d) the auricles, usually overlapping. The two auricles
endodermis appears slightly differentiated from may be the same size, or one of them may be
the pericycle. longer than the other. Peltate scales are those
with only one auricle; they are found in
INDUMENT OF THE STEM Campyloneurum falcoideum, and sometimes in C.
Scales of the stem have played an important sphenodes and C. vulpinum (Fig. 4 b, 5 e). Finally,
role in the taxonomy of many genera of the the basifix scales, i.e. those without basal auricles,
Polypodiaceae because they are the main source rarely occur in the genus. This type of scale was
of characters with taxonomic value (e.g. Sota, observed in young parts of the stem in several
1960, 1973; Lellinger, 1972; Bir & Trikha, 1979; specimens of C. amphostenon, probably indicating
Hennipman & Roos, 1983; Hovenkamp, 1986). an incomplete development of the scale.
The length of stem scales varies from 2 mm, as by Lellinger (1972).

in Campyloneurum chrysopodum, to 15 mm, as in C. In mass the scales of the stem are almost
magnificum. The base of the scale can be abruptly concolorous, and for most species the color of the
wide, as in C. aphanophlebium, C. centrobrasilianum scales is brown, except in Campyloneurum
and C. costatum. The apex varies from obtuse as chlorolepis, which has stramineous scales due to
in C. fallax, C. minus, C. nitidum, C. ophiocaulon, the hyaline cell walls. The color in the walls of
and C. wurdackii (Fig. 5 a, b), to acute, as in C. the cells is usually homogeneously distributed,
densifolium, and acuminate, as in most of the although along the scale margins, the cells have
remaining species (Fig. 5 e-j). very thin and colorless outer walls. The lumen of
The margin of the scale in Campyloneurum can the cells is for most species translucent and
be entire to slightly dentate (Fig. 6 a-c). The teeth colorless, but in some species all cells or some of
are composed of one to several protruding cells them may have dark yellow brown or yellowish
(Fig. 6 a-c). Some of the teeth look like the lumen, as in C. asplundii, C. inflatum, C.
indument found in the leaves (Fig. 6 b); this type macrosorum, C. nitidissimum, C. solutum, and C.
of tooth is formed of 1-3 cells, with apical cells as vulpinum, or completly occluded, as in C.
dark as glandular hair. Teeth along the margin fuscosquamatum.
of the stem scale are not restricted to any group Three main types of scales can be recognized
of species or species in particular. However, they based on the thickness and coloration of the cell
are more frequently found in C. cochense, C. wall: clathrate, slightly clathrate and non-
nitidissimum, and C. xalapense. clathrate. In the clathrate scale type
The scales are one layer of cell, except at the (heterotoechous of Pichi-Sermolli, 1972), the cell
point of attachment. The shape of the cells of the wall is usually brown and has a clearly defined
scales varies from narrowly oblong (Fig. 7 a, b), lumen. This clathrate type scale is the most
such as in Campyloneurum aglaolepis and C. common in the genus, and occurs in
coarctatum, to roundish (Fig. 7 c), as in C. differentiated and nondifferentiated stem scales
ensifolium and C. fallax. Narrowly oblong cells are (Fig. 7 a). In marginal differentiated scales the
the most common type. The cell disposition and well-defined cells are located in the middle of the
cell shape differentiation between margin and scale. Slightly clathrate scales occur in
center of the scale can be used to recognize two Campyloneurum macrosorum and C. vulpinum (Fig.
types of scales: marginally nondifferentiated 7 b), where the brownish cell walls are not clearly
scales, i.e, those without major differences in defined from a colored lumen. Non-clathrate
shape and disposition between marginal and scales (isotoechous, Pichi-Sermolli, 1972) occur in
central cells (Fig. 7 a); and marginally C. asplundii, C. chlorolepis, and C. fuscosquamatum;
differentiated scales, which have two or more of these species, C. chlorolepis has stramineous
rows of cells distinctive from the central cells in scales with hyaline cell walls.
shape, cell wall thickness, and disposition (Fig. 7 As mentioned by Moran (1987), there is not a
c). Nondifferentiated scales are widely satisfactory explanation for the adaptive
distributed in the genus (Table 1). advantages of stem scales. Their most common
Most of the nondifferentiated scales have the attribute has been claimed to be protection. Thus
cells arranged along the main axes, but for some Campyloneurum stem scales are mostly restricted
species the narrow rectangular cells appear to the young parts. However, Muller et al. (1981,
irregularly arranged, either in the main body of in Bosman, 1991) has attributed to them a
the scales, as in Campyloneurum amphostenon var. physiological role influencing evaporation, water
irregulare (Fig. 5 f), or at the base of the scale, as in absorption, and temperature regulation. In the
C. centrobrasilianum. Differentiated scales occur case of pruinose stems with caducous scales,
in all species groups except in C. angustifolium Bosman (1991) has suggested the same role with
and related species. The differentiated scale type regard to the whitish wax.
shows the presence of heterocellularity in
Campyloneurum. This is similar to Microgramma
as discussed by Sota et al. (1982) and Niphidium LEAVES
In Campyloneurum, sterile and fertile leaves are (1990), based on a study of three Mexican species.
similar in shape and size. Leaves are articulate to
the stem, and they are usually arranged in two
rows on the adaxial side. Leaves are frequently Lamina
curved in most species; sometimes they are erect, In Campyloneurum, the lamina of most species
as in C. abruptum, overhanging as in C. anetioides is simple and entire (Fig. 9 a-i, k-l); only in C.
or pendent as in C. sublucidum. decurrens and C. magnificum is the lamina one-
pinnate (Fig. 9 j). Pinnate leaves are here
Petiole interpreted as a primitive character in the genus.
The petiole is always present. It ranges in As with other genera in the Polypodiaceae
length from 0.3 mm, as in Campyloneurum (Hovenkamp, 1986; Bosman, 1991), aberrant
anetioides, to 95 cm in C. magnificum. The petiole is lamina shapes are occasionally found, as in
usually terete and wider at its base, becoming Campyloneurum amphostenon (Fig. 9 e). The
slightly ranurate and narrow toward the distal aberrant laminae occur in simple-leaved species,
portion. and are characterized by irregular lobes either
In Campyloneurum, distal portions of the laterally or distally. Aberrants were described by
petiole have narrow lateral expansions that result Mettenius (1864) as Polypodium angustifolium var.
from the decurrency of the lamina. These lateral monstruosum (=C. cochense) and by Poiret (1804)
lamina expansions may consist of parenchyma, as P. conjugatum (=C. phyllitidis).
usually at the most distal parts, or sclerenchyma, The size of the lamina varies within a species;
in the proximal ones, similar to other fern genera thus it has reduced taxonomic value. The
(Lin & De Vol, 1977). In most species the lamina smallest leaves in the genus are found in
expansions are not strikingly conspicuous, except Campyloneurum anetioides, and in C. chrysopodum
in C. abruptum. and C. falcoideum with leaves sometimes less than
In herbarium specimens the color of the 10 cm long. The largest leaves are found both in
petiole varies from brown to stramineous; in one-pinnate species, as in C. magnificum (2.5-3 m
living plants it is usually a deeper green on the long), and in simple-leaved species, as in C.
adaxial side. pascoense (2 m) and C. oellgaardii (1.5 m).
The indument of the petiole is composed of Shape varies little within a species, and is
deciduous and scattered scales and hairs. Scales therefore more valuable taxonomically (Fig. 9).
are similar in cell structure and shape to those of The most common shapes are linear-lanceolate to
the stem. Hairs are found during juvenile stages lanceolate (Fig. a-d, f-l), as in Campyloneurum
in Campyloneurum amphostenon, C. anetioides, and amphostenon, C. angustifolium, and related species.
C. aphanophlebium. In C. amphostenon hairs are Other shapes include oblong leaves, as in C.
deciduous, but in C. anetioides and C. coarctatum and C. inflatum; narrowly obovate
aphanophlebium they are persistent. Hairs may leaves, as in C. aphanophlebium, and spathulate
also occur in juvenile leaves of other species with leaves, as in C. anetioides. Spathulate is the basic
hairs in adult stages, such as for example in C. shape found in young leaves, as illustrated in
cochense and C. repens (Table 1). Figure 15 (a-c) and by Mitsuta (1983). Spathulate
In general, the petiole has anatomical features leaves of adult C. anetioides may represent a
similar to those of the stem (Fig. 8 a, b). The neotenic condition.
epidermis is formed by a single layer of cells, The base of the lamina is most often narrowly
with a very thick cuticle. The parenchyma, cuneate or attenuate, and is usually unequal. In
however, does not present nests of sclereids, and Campyloneurum abruptum, the base is cuneate,
it is spongy with many intercellular spaces. before abruptly becoming long decurrent on the
Under the epidermis lies a 6-15 cell-thick layer of petiole (Fig. 9 l).
sclerenchyma. There are 3-7 meristeles; two of The margin of the lamina is well defined by a
them are usually larger than the rest, and occupy cartilaginous tissue. It is usually entire and
a dorsal position. The anatomical structure of the slightly sinuate. Most of the species have a plane
petiole has recently been described by Zlotnik margin; in Campyloneurm angustifolium and
closely related species however, the margin can contiguous stomata had not been previously
become revolute, probably as a response to observed in the family (Sen & Hennipman, 1981),
environmental factors, such as low humidity and however, during this study contiguous stomata
wind exposure. were found in C. cubense (Fig. 10 b).
The apex for most species is acuminate to Hydathodes occur in all species of the genus.
subcaudate (Figs. 9 f-l), forming a "drip-tip". They appear at the tip of the free veins, on the
Only in Campyloneurum anetioides and juvenile adaxial surface of the lamina (Fig. 11 b). The
specimens is it obtuse. enlarged tip consists of numerous tracheids and
The texture of the lamina is usually cells of the epidermis arranged concentrically.
herbaceous-chartaceous. Leaves are thickly Active hydathodes are recognizable by the
chartaceous or sub-coriaceous, as in whitish surface covering. This probably results
Campyloneurum nitidissimum and C. rigidum. from the secretion of water and salts from root
Rarely leaves are thinly papyraceous, as in C. pressure, as was observed in Blechnum by Sperry
tucumanense . (1983). Although functional hydathodes are
In most species the abaxial surface of the present during juvenile stages of leaves, only
lamina is usually a dull green, while the adaxial rarely do they continue functioning in adult
side is bright and usually shiny. In leaves, as sometimes occurs in Campyloneurum
Campyloneurum rigidum and C. sublucidum brevifolium and C. xalapense. As Bosman (1991)
however, both sides have the same degree of observed for Microsorum, non-functioning
shininess. hydathodes are usually sunken into the
The indument of the lamina is composed of epidermis.
hairs and scales. The lamina scales are scattered Foliar nectaries were observed in recently
and deciduous along the costa. The hairs, developed adult leaves of Campyloneurum
however, tend to persist in most species either on phyllitidis and C. xalapense, and have also been
the lamina surface or along the costa (see below reported in C. repens by Mickel and Beitel (1988).
for a description of hairs). These structures are located abaxially, on the
acroscopic side at the joint of the costa with
Leaf anatomy primary veins. In herbarium specimens these
The leaves have a single layer of epidermis structures appear as dark spots. Koptur et al.
cells with a thick cuticle; the outline shape of the (1982), working with other polypodiaceous
cells is sinuose (Fig. 10 a-d), similar to other species, found that the secretion is composed of
polypodiaceous genera (e.g. Barrera, 1981; aminoacids and sugars. Although no chemical
Bosman, 1991). tests were done, the exudate of C. phyllitidis had a
The leaves in Campyloneurum are sweet taste. It was observed both in the field and
hypostomatic, i.e. the stomata are located on the in cultivation that during the production of
abaxial surface. Following Van Cotthem's (1970, exudates C. phyllitidis was visited by ants, but no
1973) definitions, three types of stomata were damage was noticed on the leaves. The role of
found in the genus: polocytic, copolocytic (Figs. nectaries in ferns is still unresolved; as discussed
10 a-c, 11 a-b) and anomocytic (Fig. 10 d). in Koptur et al. (1982), foliar nectaries may be
Polocytic stomata is the most common type associated with protection from predators or due
(Table 1), and it occurs in single strands (e.g. in C. to some other adaptation to the environment.
abruptum) or mixed with copolocytic stomata (e.g. The mesophyll in Campyloneurum is composed
in C. angustifolium, C. phyllitidis, and C. tenuipes). of 4-7 cell layers of parenchyma. On the adaxial
Copolocytic stomata predominate only in C. side of the leaf the parenchyma has cells closely
decurrens (Fig. 11 a). Anomocytic stomata are arranged, with small intercellular spaces; these
rare; they have been observed in C. cubense (Fig. cells contain numerous chloroplasts. On the
10 d) and C. nitidum (Sen & Hennipman, 1981). abaxial side is a spongy parenchyma,
Abortive stomata have been observed in characterized by more loosely cells arranged,
Campyloneurum cubense (Fig. 10 c), which may with conspicuous intercellular space. Some
indicate a hybrid origin for this species. Polar individuals of C. phyllitidis growing in a
greenhouse, however, showed cells of the amphostenon, numerous hairs cover the tip of the
parenchyma from both adaxial and abaxial sides lamina on the abaxial side, and are sometimes
that resembled one another in shape and scattered along the costa on the adaxial side.
disposition. Yet another type of parenchyma, The types of hairs found in Campyloneurum are
with large polygonal spaces was reported in C. not exclusive to the genus. The same type of
anetioides by Wagner & Farrar (1976). hairs are found in Grammitis (Parris, pers. comm.,
Schlerenchyma is found at the costa and margins 1985), Microgramma and some species of
of the lamina. Polypodium (Sota 1960), Pecluma (Baayen &
The vascular tissue is represented by 1-2 large Hennipman, 1987), Platycerium (Hennipman &
meristeles located in the costa (Fig. 12 b). In the Roos, 1982) and Pyrrosia (Hovenkamp, 1986).
lamina (Fig. 12 a), there are small meristeles that
correspond to the subsidiary veins. Venation
The pattern of venation has been a very
Indument of the lamina important feature in the taxonomy of the
Leaves in Campyloneurum species, except C. Polypodiaceae (e.g. Presl, 1836; Pichi-Sermolli,
aphanophlebium (as C. occultum) are commonly 1977). This character is a basis for establishing
described as glabrous (e.g. Mickel & Beitel, 1988; generic limits in Campyloneurum (Chapter III).
Lellinger, 1988). During this study, however, In Campyloneurum the pattern of venation of
indument consisting of scattered hairs and scales adult leaves has been briefly described by
was observed in several species. various authors, while the venation found in
Scales are only found along the costa, and juvenile plants has been described for a very few
their features are similar to those of the stem, species by Wagner & Farrar (1976) and Mitsuta
although size in costal scales is very reduced. (1981, 1983).
Because most of these scales are deciduous, their Campyloneurum is characterized by a reticulate
taxonomic value is limited. They are, however, venation, with areoles between parallel primary
usually persistent in Campyloneurum lorentzii. veins, and by excurrent free veinlets included in
They can be basifix, as was also observed by most areoles.
Baayen & Hennipman (1987) in C. angustifolium. The costa is prominent in most species (Fig.
During this study, representatives of three 14), and can be rounded, angular abaxially, and
types of hairs were found in the genus. They are slightly sulcate adaxially. In Campyloneurum
described according to their complexity. The first anetioides, it is only conspicuous at the base of the
type (Fig. 13 a) consists of two cells with the lamina and then becomes immersed in the lamina
apical cell slightly enlarged, as in Campyloneurum tissue.
decurrens and C. nitidum. The second type (Fig. The primary veins (Fig. 14) run parallel,
13 b) is branched, composed of three cells, one slightly sinuate or straight from costa towards the
forms the base, another is small, lateral and margin. The distance between two neighboring
glandular, while the third is an enlarged cell with primary veins varies from 4-10 (-15) mm. The
a gradually obtuse apex. This was observed for angle of divergence between the costa and
example in C. aphanophlebium, C. repens , and C. primary veins at the middle of an adult leaf
sphenodes. The third type (Fig. 13 c) is branched, ranges from 40 to 75o. There is no aparent
and also has a basal glandular cell, plus one or relationship between lamina width and this
two pluricellular branches, as in C. anetioides. angle, as was also observed in Microsorum
In most species, inconspicuous and scattered (Bosman, 1991).
hairs are found on the abaxial surface of the In most species, primary veins become
lamina (Table 1). Campyloneurum anetioides and immersed in the tissue toward the margin, except
C. aphanophlebium have hairs scattered on both in Campyloneurum nitidissimum var. nitidissimum
surfaces. In a few species, such as C. and C. pascoense where they are prominulous
amphostenon, C. angustifolium and closely related from the costa to the margin. Primary veins can
species, hairs are usually absent on the lamina be totally immersed, becoming inconspicuous,
tissue of adult leaves. But in juvenile leaves of C. except in dry herbarium specimens, as in C.
angustifolium and related species. They can also anetioides (Fig. 14 i), C. angustifolium (Fig. 14 c), C.
be slightly prominulous, or prominulous, with brevifolium (Fig. 14 j), and C. decurrens (Fig. 14 h).
dark green or stramineous color on one side of The development of the venation in juvenile
the lamina. These primary veins may have plants was studied in Campyloneurum
different degrees of prominence on each side of angustifolium, C. caudatum (here underC.
the lamina, as in C. amphostenon and related costatum),C. phyllitidis, and Polypodium vexatum
species, or similar degrees of prominence on both (here under C. cubense) by Mitsuta (1981). Figure
sides, as in C. repens and C. phyllitidis. 15 shows a similar pattern of the development in
Primary veins always branch into secondary C. amphostenon. Collectively, these studies show
veins. Secondary veins run almost parallel to the that the costal vein is formed by forked veinlets,
costa, and they are mostly slightly arched. The where one of the branches run along the main
acroscopic secondary vein unites with the axis of the lamina (Figs. 15 a-d); that the lateral
basiscopic vein of the neighboring primary vein; veins continue branching and anastomose to
the fusion of these secondary veins creates form areoles (Fig. 15 e-f); that free excurrent
primary areoles, which vary in number from two included veinlets are present in the areoles (Fig.
to more than 15 between the costa and margin. 15 g, h); and that the more complex anastomosing
The primary areoles are uniform in size and occurs later in more mature leaves.
shape, although those close to the costa (costal In adult leaves, the pattern of venation is an
areoles) are usually different from those found important taxonomic character. The pattern of
elsewhere (non-costal areoles). These costal venation together with other important feautures
areoles can be larger and more polygonal than has allowed the recognition of ten species groups
the non-costal areoles, as in Campyloneurum during this study (see Chapter VIII), based on
angustifolium and related species, or they can be four types of venation in adult leaves. These
shorter and wider as in C. magnificum, C. repens, types are used for descriptive purposes only.
and related species. In most species, secondary However, some evolutionary tendencies were
veins are immersed in the tissue, except in C. inferred during their recognition, and they will
nitidissimum var. nitidissimum and C. pascoense, be discussed in the chapter on phylogeny.
where they are prominulous. Type 1 is characterized by immersed primary
Secondary veins usually form tertiary veins. veins, less than two rows of non-costal areoles on
Tertiary veins are veinlets inside the primary each side of the lamina, and one free excurrent
areoles. Tertiary veins or veinlets may be simple veinlet in each areole (Fig. 14 a-c). The areoles
or furcate. They are mostly excurrent and are usually longer than wide. Type 1 is mostly
parallel to the primary veins, especially at the found in narrow leaves, as in Campyloneurum
marginal areoles, however, they may also be angustifolium, C. vulpinum, and allied species.
recurrent (Fig. 14 k). Type 2 is characterized by slightly
Tertiary veins may be free as in prominulous or prominulous primary veins,
Campyloneurum magnificum, C. repens, C. more than two rows of non-costal areoles,
sphenodes, and allied species, or may connect with primary areoles divided symmetrically into
the arched secondary veins forming secondary secondary areoles with one or two free excurrent
areoles. veinlets (Fig. 14 d-f). Type 2 is found in
Some secondary marginal areoles do not have Campyloneurum amphostenon,C. phyllitidis, C.
included veinlets (Fig. 14 a, b). Costal areoles xalapense, and allied species.
without veinlets are rare in the genus. Alston Type 3 resembles type 2 in having more than
(1957) showed this type of areoles in two rows of non-costal areoles. However, it
Campyloneurum chlorolepis, Sota (1960) observed differs by the immersed or prominulous primary
them at the base of the lamina of C. tucumanense veins, by the predominance of undivided
and Mitsuta (1983) in C. angustifolium. primary areoles, usually with 2 (-5) excurrent
Marginal free veinlets are excurrent and veinlets, and by areoles usually wider than long
usually undivided. They may occur in several (Fig. 14 g-i). Type 3 occurs in Campyloneurum
unrelated species, such as Campyloneurum aphanophlebium, C. magnificum, C. repens, C.
sphenodes, and allied species, and it may represent position is considered of limited value for generic
a less advanced type of venation in the genus. definition.
Three species (C. anetioides, C. chrysopodum and C. Paraphyses
falcoideum) that also belong to this venation type, The presence of paraphyses (sensu Wagner,
have non-costal areoles usually with only one 1964) in Campyloneurum was noticed earlier by
free excurrent veinlet, and they appear to Mettenius (1864) in Polypodium lindigii (=C.
represent a reduced pattern of this type of macrosorum). Wagner & Farrar (1976) observed
venation. them in C. anetioides; Tryon & Tryon (1982a) in
Type 4 is characterized by prominent primary Campyloneurum occultum (=C. aphanophlebium),
veins, more than two rows of non-costal areoles, and Lellinger (1988) in C. cooperi (=C.
and asymmetrical secondary and tertiary areoles amphostenon). Baayen & Hennipman (1987),
with more than two free mostly excurrent however, did not find these structures in C.
veinlets in each primary areole (Fig. 14 j-k). Type angustifolium and C. phyllitidis.
4 corresponds to the venation found in During this study, each species was examined
Campyloneurum brevifolium and allied species. for paraphyses. They were observed in nine
Some general features of the pattern of species (Table 1). Paraphyses are most often
venation in Campyloneurum, such as costal areoles smaller than the sporangia, and more slender
with one excurrent veinlet, or the presence of than the stalk. Two types of paraphyses are
excurrent and sometimes recurrent veinlets in recognized here: simple and dendritic. Simple
non-costal areoles, are shared with three other paraphyses are composed of one unbranched
neotropical polypodiaceous genera, Pleopeltis, row of 2-3 cells (Fig. 16 a), as occurs in
Microgramma and Niphidium, and these may Campyloneurum acrocarpon, C. pascoense, and some
reflect the closest affinities of Campyloneurum. individuals of C. amphostenon and C.
angustifolium. Dendritic paraphyses are
SORI composed of a row of 5-7 cells with 3-5
In Campyloneurum, sori are round to elliptical, unicellular branches (Fig. 16 b), as seen in C.
1-2 (-3) mm across, and on the surface of the aglaolepis, C. aphanophlebium, C. nitidum, and C.
lamina. They usually occur in two rows between vulpinum. Dendritic paraphyses in
primary veins, but three or more are common in Campyloneurum resemble those found in
C. brevifolium and allied species, while one row Microgramma ciliata, as described by Baayen &
of sori may occur in C. anetioides and C. Hennipman (1987). In C. macrosorum, simple
falcoideum. Each sorus is innervated by a solitary paraphyses are common in addition to dendritic
excurrent free veinlet, however, rarely two or ones.
three sori occur together in C. brevifolium. Sporangiasters or abortive sporangia (Wagner,
The position of sori on the veinlet is subapical 1964) are frequently found on the receptacle.
or medial for most species. Terminal sori occur
in the one-pinnate species, rarely in some Sporangia
individuals of Campyloneurum aphanophlebium Sporangia are characterized by a spherical
and C. falcoideum, and it was shown by Mitsuta capsule supported by a slender stalk of 2 (-3)
(1983) in C. angustifolium. The terminal position rows of cells. Sporangia are numerous in the
of sori is considered to be a primitive character sori. They appear to mature in a centripetal
state in the genus. Sori at the junction of two order. They are mixed with abortive sporangia
veinlets may occur in C. brevifolium and related and sometimes with paraphyses. Abortive
species, correlated with the more complex sporangia are easily recognized by the small size
venation (Fig. 14 k), although it was also of the tannin-filled capsule.
observed by Mitsuta (1983) in C. angustifolium. The length of the capsule ranges from 330-460
The range of variation of sorus position (medial, µm. The annulus is composed by 12-18 bow cells.
subapical or terminal) in the genus is similar to The faces of the capsule are similar to other
that in other genera in the family (e.g. Pyrrosia, genera in the family s.str., as described by Wilson
Hovenkamp, 1986), and in this study the soral (1959). The stalk is thin and well developed; it is
composed of two rows of cells, except below the spore morphology are shared, such as shape and
capsule, where there are three rows. surface.
Spores GAMETOPHYTES
Earlier observations of spores of As Atkinson (1973) mentioned, features found
Campyloneurum date back to Fée (1869). Since in gametophytes may help to demonstrate the
then, there have been several studies of the spore cohesiveness of a genus and to understand
morphology in Campyloneurum, especially during relationships. In Campyloneurum, little is known,
the last 30 years, such as those of Nayar (1962), however, of the development and characteristics
Tschudy & Tschudy (1965), Pal & Pal (1970), of the gametophytes.
Kremp & Kawasaki (1972), Murillo & Bless Spore germination has only been observed in
(1978), Lloyd (1981), and Tryon & Lugardon Campyloneurum anetioides by Wagner & Farrar
(1991), among others. Campyloneurum spore (1976).
morphology was examined here for each species The development and characteristics of the
when available (Figs. 17, 18, 19). Spores are gametophyte has been observed in
reniform and ellipsoidal, although abortive Campyloneurum anetioides (Wagner & Farrar,
spores may be rounded and collapsed. Size 1976) and C. angustifolium (Nayar, 1962). In both
varies between 40-100 µm long and 30-60 µm species the shape of the gametophyte was found
wide (Table 1). For most species, spore sizes to be thalloid, with a cordate apex and with a
varies little intraspecifically; large differences in continuous or discontinuous midrib. The
size, however, were observed in C. angustifolium, indument of the gametophyte was composed of
C. repens, and C. vulpinum. This variation may bicellular glandular branched hairs, similar to
suggest the occurrence of different ploidy levels those found in the sporophyte of most species of
(cf. Tryon & Lugardon, 1991). Campyloneurum.
The laesura characterizes the monolete spore
in the genus. It always protrudes from the surface
(Fig 17, 18). Laesura sizes are usually constant V. CYTOLOGY AND BIOCHEMISTRY
among species. The laesura occupies between
1/4 of the spore length, as in Campyloneurum CYTOLOGY
costatum (Fig. 17 c) and 3/4 of the length of the Chromosome numbers in Campyloneurum
spore, as in C. amphostenon (Fig. 17 a). Rarely this have been reported in at least ten taxa (Table 2),
range of size may occur within a single species, by several authors (see Fabbri, 1963, 1965;
as in C. phyllitidis (Fig. 18 c, d). Walker, 1985). For most cases the basic
The surface of the spore is covered by chromosome number is x=37.
scattered spherical bodies or granules, which are Three species (Campyloneurum costatum, C.
located under a thin perispore (Fig. 19 b, 20 a, b). tenuipes, and C. xalapense) are diploids with n=37
The exospore can be slightly verrucate or and 2n=74; five species (C. acrocarpon, C.
verrucate (Fig. 17, 18, 19). In a few cases, as in anetioides, C. minus, C. nitidum, and C. repens) are
Campyloneurum anetioides, C. aphanophlebium, and tetraploids with n=74 and 2n=148; and in two
C. falcoideum, the surface is rather smooth (Figs. species (C. angustifolium and C. phyllitidis) both
17 c, 18 e), supporting a probable affinity of these tetraploids and diploids are found.
species. The presence of both diploid and polyploid
Tryon & Tryon (1982a) and Tryon & populations within a species is not uncommon in
Lugardon (1991), based on spore characteristics, ferns (e.g. Walker, 1985) and may occur in
have suggested generic relationships between Campyloneurum. In C. angustifolium there is
Campyloneurum and Niphidium, and also with evidence of a wide variation in spore sizes, that
some species of Polypodium. In this study these may support the findings of two ploidy levels. A
relationships are considered, and it is suggested diploid was once reported by Evans (1963) from
that the genus Pleopeltis is closely related to material collected in Peru, while tetraploids were
Campyloneurum, because many characteristics of recorded from specimens collected in Costa Rica
and Florida (Evans, 1963; Wagner, 1963). there has been an increase in the study of fern
However, the evidence of different cytotypes biochemistry (Soeder, 1985).
within this species is not conclusive, due to the Several works in different genera of the
poor taxonomic understanding of the species at Polypodiaceae s.str., as mentioned by Soeder
that time and the fact that vouchers specimens (1985), have revealed the presence of alkaloids,
from those cytological studies were not examined flavonoids, terpenoids, sterols, and other
for this treatment. compounds. According to Swain & Cooper-
Campyloneurum brevifolium is a diploid species, Driver (1973), flavonoids and terpenoids are
as was reported by Evans (1963) for southern widespread in ferns.
U.S.A. and by Smith & Foster (1984) and Walker Very little is known about the biochemistry in
(1985) for Trinidad. This is the only species thus Campyloneurum. Lynch et al. (1970) reported the
far that is known to present two chromosomal presence of saponins (terpenoid) and the absence
satellites (Walker, 1985). of anthocyanin (flavonoid) and triterpenoids in
Campyloneurum phyllitidis is tetraploid, as Polypodium latum (=C. brevifolium).
reported by Evans (1963) and Wagner (1963) for
southern U.S.A., by Jarrett et al. (1968) for the
Galápagos in Ecuador, and by Walker (1973, VI. ECOLOGY AND DISTRIBUTION
1985) for Jamaica and Trinidad. Similar to the
case of C. angustifolium, C. phyllitidis appears to ECOLOGY
have two cytotypes, since Sorsa (in Fabbri,1965) Most Campyloneurum species are found in
registered a diploid from Peru, but no forests, in the tropics and subtropics of the New
conclusions can be certain since no voucher World. These forests are often spatially
specimens were cited. Allopolyploidy has been discontinuous and have been classified in
suggested by Walker (1985) to occur in C. numerous life zones and with many types of
phyllitidis from Jamaica based on the bimodal vegetation (cf. Gómez-Pompa, 1965, 1973; Walter,
distribution of chromosomal length; however, no 1971; Oldeman 1990). Most Campyloneurum
further information is available in relation to the species inhabit evergreen forest, but C. xalapense
probable parental species. is also found in drier semideciduous forests in
Campyloneurum nitidum is tetraploid, as Mexico.
reported by Smith & Foster (1984) from Most species in Campyloneurum may occur in
Paraguay. disturbed sites, both natural, such as landslides,
In Campyloneurum, aneuploidy has been treefalls and streamsides, and those made by
reported once (Pal, 1961 as cited by Fabbri, 1963) humans, such as roadcuts. Some species,
for a cultivated plant of C. phyllitidis. This however, such as C. anetioides, C. aphanophlebium,
phenomenon is not uncommon among ferns C. falcoideum, C. inflatum, C. macrosorum, C.
(Walker, 1985), however, further studies are oellgaardii, and C. sublucidum occur only in forest
neccesary to clarify its occurrence in the genus. environments not modified by humans.
Based on the present cytological information, Twenty-six species occur in shady and moist
some Campyloneurum species present different microhabitats inside forests (Table 3). Most of
cytotypes that may correlate with the these species are low branch epiphytes or trunk
morphological variation found in their leaves and climbers, and they occupy the lower zones of the
spores. If allopolyploidy proves to be more forest strata. Epiphytes or climbers usually have
widely distributed among species of erect leaves, rarely pendent leaves, as in
Campyloneurum, it may be useful character for Campyloneurum sublucidum, are found. Among
testing affinities among species. terrestrial species are C. abruptum, C. decurrens,
and C. magnificum, which are usually distributed
BIOCHEMISTRY in lowland forests below 1000 m elevation, while
The biochemistry of ferns has been recognized C. pascoense and C. tucumanense are frequently
as less variable than that of angiosperms (Swain found as terrestrials, but in montane forests.
& Cooper-Driver, 1973). During the last 20 years, Other species may occur as terrestrials, but only
on forest floors covered by thick layers of debris latitudinal range is more restricted than for those
and mosses. species having a large elevational range.
Nineteen species generally inhabit either Fourteen species (C. abruptum, C. acrocarpon, C.
exposed areas or open forests (Table 3). These austrobrasilianum, C. chrysopodum, C. coarctatum,
species occur on cliffs, in rock crevices, and in the C. costatum, C. decurrens, C. fallax, C. macrosorum,
ecotone between forests and grasslands. In one C. nitidum, C. ophiocaulon, C. pascoense, C. rigidum,
locality in Costa Rica, Campyloneurum and C. sublucidum) have an altitudinal range
angustifolium has been recorded inhabiting tree between 600 and 1000 m, and most of them have
canopies (Grayum & Churchill, 1989). These 19 a latitudinal range of 15o to 40o. On the other
species have wide environmental adaptations, hand, ten species in the genus have a narrow
and they can grow as epiphytes, epipetrics or altitudinal range (Fig. 21), thus C. anetioides, C.
terrestrials (e.g. C. cochense, C. densifolium). It is centrobrasilianum, C. cubense, C. inflatum, C.
not surprising to find among these species the oelgaardii, C. oxypholis, C. tenuipes, C. tucumanense,
most widely distributed species (e.g. C. C. vulpinum, and C. wurdackii occur with less than
angustifolium and C. phyllitidis). a 500 m elevational amplitude, and all are
Two species, Campylonerum cubense and C.
latitudinally restricted species, with less than 15o
oxypholis (Table 3) occur only outside forests, i.e.
of range.
cliffs in partially protected microsites.
Adaptations to strongly mesic conditions are
DISTRIBUTION
found in Campyloneurum anetioides, the only
Campyloneurum ranges from southern Florida,
species in the genus with spongy mesophyll
the Caribbean Islands and Mexico, throughout
(Wagner & Farrar, 1976). Most species occurring
Central America, the Andes to northern
in forested areas have a "drip-tip" and/or curvate
Argentina, eastern Brazil and Uruguay (Fig. 22).
or hanging leaves; these adaptations probably
This genus is predominantly tropical, with
allow draining of the leaf surface (Jungner, 1891;
occurrences in adjacent subtropics.
Richards, 1952; Dean & Smith, 1978). In species
Most species are widespread (Table 4). The
occurring in exposed areas, leaves are usually
countries with more than five species are located
thickly chartaceous, with well-developed
in mountainous tropical areas; while the
cartilaginous margins. In some species
countries with five or fewer species are located in
inhabiting different kinds of exposed areas,
non-mountainous areas, mostly in the subtropics
margins of the leaf are partially revolute (e.g. C.
and/or on small islands (Fig. 22). Only ten
angustifolium, C. densifolium). Some species
species (Campyloneurum acrocarpon, C.
growing above 3000 m elevation (C. amphostenon,
austrobrasilianum, C. centrobrasilianum, C.
C. cochense, C. densifolium, and C. lorentzii) have
chrysopodum, C. cubense, C. fallax, C. oellgaardii, C.
stems covered by a white wax, which, as
oxypholis, C. rigidum, and C. wurdackii) are
suggested by Bosman (1991) for Microsorum, may
geographically restricted (i.e. found within a
protect the stem from dessication.
single political boundary).
Most Campyloneurum species inhabit montane
Disjunct distributions occur in C. inflatum and
areas between 1000 and 4500 m elevation (Fig.
C. sublucidum. These disjunctions may be
21). Only eight species (C. abruptum, C.
explained by a lack of collections, or by
acrocarpon, C. austrobrasilianum, C. coarctatum, C.
specialized biological and habitat requirements.
cubense, C. decurrens, C. rigidum, and C. wurdackii)
For example, C. sublucidum is found on
frequently occur below 1000 m.
calcareous rocks.
Elevational range differs among species (Fig.
21). Twenty-three species have a greater than
Patterns of distribution of species
1000 m altitudinal range (Fig. 21; Table 9); among
The distribution of fern species has been used
these species are the widely distributed species
by Tryon (1972, 1979, 1985, 1986) to discuss the
(e.g. Campyloneurum angustifolium, C. brevifolium,
relation of geographic features to speciation. For
C. phyllitidis). It appears that for species growing
tropical American ferns, those features were
with a less than 1000 m elevational range, their
discussed in terms of the concentration of species Central American region has a Campyloneurum
and of endemism in regions or regional centers. flora of 18 species (Table 5), with one endemic (C.
Here the geographic information of anetioides). Eleven of these species (C.
Campyloneurum species is presented according to amphostenon, C. angustifolium, C. aphanophlebium,
biogeographical regions (modified from Tryon, C. brevifolium, C. coarctatum, C. costatum, C.
1972), relating their floristic affinities with the densifolium, C. fasciale, C. magnificum, C. phyllitidis,
history of the area, and thus allowing speculation and C. repens) occur in another two or more
on geographic speciation. regions. This region has greater affinities with
The six biogeographical regions are: Mexico, the Mexican and Andean, than with the
Central America, Caribbean Islands, Andes, Caribbean and Guayanan regions (Table 6). In
Guayana and Brazil. the Central American region, two main
The Mexican region lies north of the Isthmus subdivisions were recognized by Savage (1966,
of Tehuantepec. Present conditions of climate 1982), based on the herpetofauna: a Nuclear or
and vegetation in the area are diverse (see Upper Central America (from southern Mexico to
Rzedowski, 1978). Campyloneurum species occur northern Nicaragua) and the Isthmian link or
mainly in conifer, Quercus-Liquidambar, and Lower Central America (from southern
tropical evergreen forests. This region contains Nicaragua to Panama), with a different geological
12 species and no endemics (Table 5). Nine of history (cf. Coney, 1982). These two subdivisions
these species (C. amphostenon, C. angustifolium, C. appear to be also important in the analysis of
aphanophlebium, C. brevifolium, C. costatum, C. Campyloneurum regional distribution and
densifolium, C. fasciale, C. phyllitidis, and C. repens) speciation. Two species (C. ensifolium and C.
are widely distributed, occurring in another two tenuipes) are only found in the northern
or three regions more. On the other hand, three subdivision. These two species occur also in the
species (C. ensifolium, C. tenuipes, and C. xalapense) Mexican region, reflecting perhaps the early
occur in one other region, the Central American. Eocene connection between this region and the
This region's affinities are closer with the Central northern Central American subdivision (cf.
American and Caribbean than with the Andean Taylor, 1991). The northern subdivision may also
and Guayanan regions (Table 6). These affinities have been an important source area for the flora
and distribution patterns may be interpreted as a of the southern subdivision. Campyloneurum
result of geological events in the past. Data on anetioides, is endemic to the whole region, and
the geological history of the area suggest that probably arose in the oldest northern subdivision
from the Permian to the early Cretaceous the area not long before the connection occurred between
was separated from South America, which the northern subdivision and southern Central
caused a separation of the floras. During the American subdivisions during the Pliocene. On
early Eocene, the Mexican region was closer to the other hand, three species (C. falcoideum, C.
the Greater Antilles, and probaby, due to sphenodes, and C. sublucidum) are restricted to the
orogenesis and volcanism, cycles of contact may Isthmian link subdivision. These species are
have occurred with northern Central America (cf. Andean in origin and they reflect the Pliocene
Coney, 1982; Savage, 1982). These Cenozoic contact between South and Central America.
events and changes in climate conditions, as The Caribbean region comprises the islands of
summarized by Savage (1982), may account for the Greater Antilles, excluding the Lesser
the closer affinities with the northern Central Antilles. There are four main islands (Cuba,
American region. Hispaniola, Jamaica and Puerto Rico) which have
The Central American region runs between a wide variety of environments due to changes in
the south of the Isthmus of Tehuantepec and the altitude, soil types, temperature ranges and
Isthmus of Panama. The present climate and vegetation as mentioned by Howard (1973). In
vegetation in the area are also as varied as those this region Campyloneurum is represented by 10
found in the previous region, and they can be species of which two are endemics (C. cubense, C.
related to the presence of two important oxypholis). Seven species (C. amphostenon, C.
Cordilleras: Guatemalan and Costa Rican. The angustifolium, C. brevifolium, C. costatum, C.
densifolium, C. phyllitidis, C. repens, and C. Tertiary, 3) the cordilleras that form the Andes
vulpinum) are distributed in another two or more today had extensive uplift in the Oligocene, and
regions. The floristic affinities of this region are 4) the last phase of the uplift of the Andes began
closer to the Mexican and Central American in the Upper Pliocene. In the northern
regions. These affinities may be a result of subdivision, Campyloneurum is represented by 31
connections among these regions during the late species, five of them (C. chrysopodum, C. cochense,
Cretaceous (Coney, 1982). The richness of this C. inflatum, C. macrosorum, and C. oellgaardii) only
zone appears to be related to its diversified occur in this subdivision (Fig. 23). The Middle
habitats (Tryon, 1979) and its geological history subdivision has 24 species and the Southern
of isolation after the late Eocene from the Lesser subdivision 4, both subdivisions without
Antilles-South America and the north Central restricted species (Fig. 23). These results differ
America. from the general patterns provided by Berry
The Andean region, as defined by Tryon (1982) for Fuchsia sect. Fuchsia, and by Molau
(1972), includes the area along the Andes from (1988) for Calceolaria. The richness of the
10oN to 25oS. The implications of the geological Northern Division in the Andes may be a result
history of the Andean region related to of the presence of younger zones in the area
biogeography and evolution have been discussed compared to rest of the Andes (e.g. Taylor, 1991).
for different organisms (e.g. angiosperms, Besides, the Northern Division is characterized
Simpson, 1975, 1979; Berry, 1982; Gentry, 1982; by three cordilleras with humid montane
Molau, 1988; reptiles and amphibians, Duellman, vegetation that could have offered more suitable
1979). Campyloneurum has wind-dispersed spores environments for diversification of
and belongs to an older group of plants, and thus Campyloneurum. The affinities of the whole
it differs from many other organisms studied in region are closer to the Mexican and Central
the Andes. These and other intrinsic American, although these affinities are low (less
characteristics of the genus (e.g. the possibility of than 25%), which may reflect the long isolation of
polyploidy), together with dynamic changes in South America.
the past, may have offered a wide range of The Brazilian region is here interpreted to
possibilities for speciation. In the Andes, include the central Brazilian shield and the south-
Campyloneurum is represented by 33 species of eastern mountains ranging to 30oS. This region
which 15 are endemics (C. aglaolepis, C. asplundii, is characterized by the presence of nine species,
C. chrysopodum, C. cochense, C. fuscosquamatum, C. five of which are endemics (Fig. 22). The highest
inflatum, C. lorentzii, C. macrosorum, C. affinity of this region is with the Guayana,
nitidissimum, C. oellgaardii, C. ophiocaulon, C. although it is low (Table 6). The high endemism
pascoense, C. solutum, and C. tucumanense ). Both in the region has also been observed in other
total number of species and number of endemics groups of plants (angiosperms, Smith, 1962;
are higher than in any other region (Table 5). In pteridophytes in general, Brade, 1942; Tryon,
order to understand the richness of this area 1972; and in the fern genus Polybotrya Moran,
Campyloneurum species distribution is analyzed 1987), and may also be related to the high
within three subdivisions in the Andes: Northern ecological diversity in the area as shown by
(10°N-6°S), Middle (6-18°S) and Southern (south Brade (1942). Another important factor that may
allow the explanation of the richness of
of 18°S). Each of the these subdivisions have a
endemism in the Brazilian region is its geological
complex geology, mountain building history and
history, which dates back from the Cretaceous
paleofloristic history, as discussed in detail by
(cf. Novaes Pinto, 1990) and also its isolation
Taylor (1991). Major events presented by Taylor
from other mountain systems in the continent.
(1991) that may have influenced speciation events
The Guayanan region, as defined by Tryon
in this genus are 1) in the middle and southern
(1972), is located on the Roraima shield. It
subdivisions mountains probably existed from
contains six species (Table 5) of which one is
the early Cretaceous, 2) the northern subdivision
endemic (Campyloneurum wurdackii). Its closest
had mountains probably later, during the early
affinities are with the Central American and
Mexican regions. The importance of the region observed, no endemics are found in these islands,
for species diversification has been shown for which are located less than 1500 km from the
flowering plants (Maguire, 1970; Huber, 1988) mainland. These islands are also relatively
and for certain fern genera (Lellinger, 1967; young in geologic age, which may have not
Tryon, 1972). For Campyloneurum speciation allowed time for speciation.
events appear to have occurred in the more
isolated areas. Distribution of species groups and a general
The Amazon area, not included in the above overview of speciation
regions, has a flora of widely distributed In this study, ten informal species groups are
Campyloneurum species, which mostly inhabit the recognized in Campyloneurum (Chapter VIII) that
foothills of the Andes and the Cordilleras of are interpreted as evolutionary lineages.
Central America. Most of the discussion of the Each of the biogeographical regions differ in
geological history of the basin is centered around the number of species groups. The Mexican
the last five million years, associated with region has seven species groups, the Central
changes in climate and vegetation in the American has nine, the Caribbean has seven, the
Pleistocene (see Prance, 1982). Assuming that Andean has all ten, the Brazilian has five and the
time is a important factor for speciation in the Guayanan has four (Table 7).
genus, then the evidence from the Pleistocene The common species groups for the regions
may account for the low endemism reported by are the Campyloneurum phyllitidis and the C.
Tryon & Conant (1975) for the Amazonian fern repens groups, which represent two succesful
flora in general. In the case also of groups regarding dispersal, with numerous
Campyloneurum, the Amazon area does not have species. The C. phyllitidis group seems to have
endemics. originated in continental America; four species in
Only thirteen species occur in the subtropics this group are restricted to one of the oldest
of South America. Four of these species also biogeographical regions. The Brazilian hasC.
inhabit the subtropics of North America and all nitidum, the Andean has C. abruptum, the Mexican
of them are widely distributed. In the subtropics has C. tenuipes and the Guayanan has C. wurdackii
of South America, nine species (Campyloneurum . The group is presently associated with low to
acrocarpon, C. aglaolepis, C. austrobrasilianum, C. middle elevations and forested areas. It seems
fallax, C. minus, C. nitidum, C. lorentzii, C. rigidum, that the environment suitable for this group was
and C. tucumanense) have a restricted geographic partially available in most parts of America since
distribution. the late Cretaceous, as can be inferred from the
Other non-continental areas in the Neotropics work of Gentry (1982) for angiosperms; although
are the Lesser Antilles and other small islands of in some parts the origin of the present vegetation
the West Indies. The Lesser Antilles have only where the C. phyllitidis group is found may be
four species (Campyloneurum angustifolium, C. recent in origin (e.g. Toledo, 1982).
brevifolium , C. fasciale, and C. phyllitidis), all of The Campyloneurum repens group probably
them widely distributed elsewhere. The had a continental origin and successfully reached
Campyloneurum flora of the West Indies islands is the Antillean region like the C. phyllitidis group.
depauperate, with only three species (C. The former group seems to have radiated in two
angustifolium, C. brevifolium, and C. phyllitidis). main mountain systems, the Andes and the
These species are the ones that can grow at low eastern Brazilian Cordillera, with an ancient
altitudes in their range, as was also mentioned by geological history. The Andean region has more
Correll (1976). species (C. fuscosquamatum, C. macrosorum, C.
In the Neotropics there are few islands ophiocaulon) than the Brazilian (C. acrocarpon and
isolated from the continent. The Campyloneurum C. minus). Since the former region has suffered
flora of the Cocos has one species, C. phyllitidis more continuous dramatic geological changes
(Gómez, 1975) and the Galápagos islands have since the early Cretaceous, it seems that
two widely distributed species (C. amphostenon speciation in this group can be due to geographic
and C. phyllitidis). As Tryon (1970; 1972) isolation. However, speciation by peripheral
divergence (Tryon, 1972), may have also ocurred rate for speciation and high capacity for
within each region. dispersal. These features, combined with
The available information indicates that the regional isolation and geological changes has
elements of the Campyloneurum brevifolium group resulted in the presence of vicarious species
occur widely in all regions, except the Brazilian. between the oldest mountainous regions in South
This group appears to have been succesful in America: the Brazilian (C. fallax) and the Andean
radiating in different parts of tropical continental (C. lorentzii) regions. These features, together
and insular America. However, its absence from with the geologic dynamism of the areas, may
the Brazilian region may be interpreted as the account for the presence of extreme variation
result of a recent origin outside of this region, within some taxa, as for example in C.
since it is assumed here that all biogeographic amphostenon and C. densifolium.
regions were available for ancestral populations The C. angustifolium group seems to have
of the genus before they became isolated. The C. similar intrinsic characteristics as discussed for
brevifolium group appears also to have a high rate the C. amphostenon group. Vicarious species are
for speciation, especially in the dynamic Andean found between the Mexican-north Central
region, where most of its endemic species are American (C. ensifolium ) and the central
found. Brazilian Plateau (C. centrobrasilianum). High
Two species groups, the Campyloneurum speciation rate appears to occur in the Andes,
aphanophlebium and the C. xalapense, are presently but is not necessarily related to geographic
distributed in the Mexican, the Central American isolation, as seen in the case of C. aglaolepis and
and the Andean regions. The C. aphanophlebium closely related species (C. angustipaleatum, C.
group has a clear continental origin; its present austrobrasilianum).
distribution is limited to the Central American The elements of the Campyloneurum sphenodes
and Andean regions. The center of endemism in group occur in three regions, Central American,
this group seems to be the north Central Andean and Guayanan, all of them in continental
American-Mexican region, an area emergent America. This group has a great number of
since the early Eocene. One feature of this group species in the Andean region, where they
is the paucity of species, that can be interpreted presently occur from middle to low elevations. It
as the result of a recent origin and the short time seems that this group migrated to the Guayanan
available for speciation. and Central American regions recently since
The Campyloneurum xalapense group is a there are no endemics there.
heterogeneous one. The pattern of regional The Campyloneurum vulpinum group has an
distribution appears similar to that found in the unclear origin. It may be the only lineage that
C. phyllitidis group. The C. xalapense group may have arisen in the Caribbean region.
occurs in all regions except in the Brazilian and Campyloneurum vulpinum may later have
Guayanan. Its elements presently grow at high- migrated to South America. On the other hand, if
middle (C. cochense and C. xalapense) or low this lineage was of South American origin, it may
altitudes (C. costatum). It appears that this group had enough time since the late Eocene for
diverged in the Mexican-Caribbean (C. costatum dispersal and speciation by isolation. This group
and C. xalapense) and in the Andes (C. cochense). apparently had a low rate of speciation.
The elements of the Campyloneurum The Campyloneurum magnificum group has
amphostenon and the C. angustifolium groups several primitive characteristics in the genus,
show a similar pattern of distribution in five of such as division of the lamina, undivided
the regions (Brazilian, Andean, Caribbean, primary areoles and terminal position of the sori,
Central American and Mexican). Both are more as was discussed in Chapter IV. The elements of
speciose in the Andean and Brazilian region. this group occur today in the Andean and
Both groups seem to have speciated and radiated Central American region, mostly in mesic
in middle to high altitude environments. environments. The present distribution of the
TheCampyloneurum amphostenon group seems to two species may support the idea that this group
combine two important intrinsic features, high migrated recently from South America to the
Lower Central America and Lesser Antilles, areas forests. However, other habitats in the region,
geological younger (Figs. 35, 41). It appears that including disturbed sites, should be incorporated
this group has not been succesful in colonizing into efforts to protect and preserve biotic
different habitats, and that his capacity of resources. One reason for this approach is the
dispersal is rather low. fact that species such as those of the genus
In conclusion, the distribution of the species Campyloneurum occur in a great variety of
groups in the regions suggests that the genus mountainous environments from rock crevices in
may have arose before the late Jurassic. Post grasslands to evergreen or semi-deciduous
Jurassic geological and environmental changes forests, as was discussed in Chapter VI.
may have allowed the speciation by isolation in Based on taxonomic and biogeographical
the regions. The genus appears to have a middle information, conservation issues for
elevation origin, and may have had a high rate of Campyloneurum are here tentatively addressed,
speciation, at least in some of the lineages. The recognizing endangered species, species with an
Pliocene changes allowing the connection of unknown conservation status, and species with
South with Central America have allowed the probably adequate and unthreatened
exchange of floras between regions, but the lapse populations. In the future, it will be necessary to
of time has probably not been sufficient for reevaluate these categories with empirical data
numerous speciation events to have occurred in on the size and status of populations.
the newly connected areas. Those species with small latitudinal and
altitudinal ranges, and growing in habitats that
are being destroyed are considered to be
VII. USES AND CONSERVATION probably endangered. Twelve species belong to
this category: Campyloneurum acrocarpon, C.
USES anetioides, C. centrobrasilianum, C. chrysopodum, C.
Campyloneurum species are occasionaly used fallax, C. inflatum, C. macrosorum, C. oellgaardii, C.
for decorative, therapeutical or religious oxypholis, C. sublucidum, C. tucumanense, and C.
purposes. wurdackii. All these species appear to be
The use of Campyloneurum as a decorative restricted to forested areas, except C. acrocarpon,
plant has not reached the degree of acceptance which occurs in exposed slopes (Table 9). Some
found with polypodiaceous Asian genera. Only species, such as C. oellgaardii, C. sublucidum, and
two species of the 47 in the genus are cultivated C. wurdackii are currently known from fewer than
as garden or interior plants; Hoshizaki (1982) and three localities. Among these endangered species
Jones (1987) listed C. amphostenon and C. are also those that inhabit the forest in
phyllitidis in cultivation in temperate zones. southeastern Brazil, an area with high
Only four Campyloneurum species have deforestation rates (Mori et al., 1988; Prance &
popular names or are probably used as medicine Campbell, 1988).
for a disparate assortment of illnesses. The Six species (Campyloneurum decurrens, C.
common names and uses of some species of falcoideum, C. lorentzii, C. magnificum, C. minus,
Campyloneurum were recorded from herbaria and C. rigidum) are here considered species with
labels (Table 8). All these are widespread, populations of unknown status. They have
lowland species, common in the areas that they larger altitudinal and latitudinal ranges than
are used. The common name "calaguala" is those of the previous category, but are known
applied in the Andes to different genera of from a rather small number of localities. They
Polypodiaceae with entire leaves. In Paraguay inhabit either forests or open forested areas (as
this name is used for C. nitidum. defined in Chapter VI).
The majority of species do not appear to be
CONSERVATION endangered since they have large altitudinal and
In the Neotropics, major efforts in latitudinal distributions , and occur in a wide
conservation and protection of the environment variety of habitats.
have been focused on the tropical lowland rain
state among Polypodiaceae (e.g. Hovenkamp,

1986), and this view is probably true for
VIII. INFRAGENERIC GROUPS AND Campyloneurum, too.
PHYLOGENY
2. The Campyloneurum repens group.
SPECIES CONCEPT Campyloneurum acrocarpon, C. fasciale, C.
The available data for Campyloneurum comes fuscosquamatum, C. macrosorum, C. minus, C.
from the study of field and herbarium specimens, ophiocaulon and C. repens have long-creeping
and from distributional and ecological stems, usually less than 5 mm wide, distant
information. Based on these sources, the leaves (except in C. acrocarpon and C. minus),
morphological species definition (Haufler, 1989) short petiolate lamina with decurrent lamina
is employed in this treatment. bases, and undivided non-costal primary areoles.
The infraspecific category of variety has been This group occurs today from low to middle
used here in two cases, for Campyloneurum elevations, and most species inhabit disturbed
amphostenon and C. nitidissimum. The use of areas. The undivided primary areoles are
variety has been applied when some considered to be a less advanced character in the
morphological characters differences are obvious genus. However, in the C. repens group other
but still overlap. characters such as the medial position of the sori,
entire leaves and climbing habit are considered to
INFRAGENERIC GROUPS be more advanced. Here, this group is
Campyloneurum has never been formally considered closer to the C. sphenodes and the C.
subdivided, and that is not the intention here aphanophlebium groups than to the C. magnificum
either. Herein are presented ten informal species group.
groups, based on the sharing of at least three
morphological characters. 3. The Campyloneurum sphenodes group.
The groups are intended to represent a This group includes Campyloneurum chrysopodum,
hypothesis of evolutionary relationships within C. coarctatum, C. falcoideum, C. inflatum, C.
the genus (Fig. 51). In this study, it is considered oelgaardii, C. sphenodes, and C. sublucidum , which
that all descend from a common ancestor are characterized by a stem diameter less than 5
characterized by pinnatifid leaves, and mm wide, more than 3 areoles between costa and
undivided primary areoles with 2-4 excurrent margin, areoles rarely undivided and usually
free veinlets. This ancestor may have been a carrying two excurrent veinlets, and broadly
terrestrial plant, growing in humid low to middle oblong-lanceolate leaves with cuneate bases.
elevations. Each group then independently This group shows some morphological
followed speciation at different rates and with similarities with the C. repens species group, but
individualistic patterns. differs by the long petioles, broadly oblong-
lanceolate lamina, and cuneate bases. All species
1. The Campyloneurum magnificum group. are restricted to continental tropical America,
This group is composed of Campyloneurum where they occur in mountainous areas of
decurrens and C. magnificum. These species are Central and South America as climbing
the only ones with pinnatifid leaves. Their stem epiphytes. The position of this group appears to
diameter is more than 10 mm wide, short- be close to the C. repens group, but more
creeping, and the stem scales are broadly ovate, advanced.
clathrate with differentiated margins. The
venation in this group is that described as type 4, 4. The Campyloneurum aphanophlebium group.
which is interpreted as the least advanced in the This includes Campyloneurum anetioides and C.
genus. This species group is restricted to low aphanophlebium. They both have undivided
elevation forest habitats, and might represent an primary non-costal areoles, closely spaced leaves,
old evolutionary line in the genus. Pinnate leaves free veinlets along the margin, and medial sori.
are commonly considered a primitive character This group is also characterized by a specialized
indument of branched pluricellular hairs that continental and insular tropical America, where it
occupy both sides of the leaf. It appears from the occurs from middle to high elevations. Because
venation pattern and indument features that this of the dynamic geological history of mountain
group has affinities with the C. repens and C. environments in the Neotropics, populations of
sphenodes groups. this group may have repeatedly had contacts and
isolations, which is now reflected in the high
5. The Campyloneurum phyllitidis group. number of species. The Campyloneurum
Campyloneurum abruptum, C. nitidum, C. amphostenon group is considered to be one of the
phyllitidis, C. tenuipes, and C. wurdackii are advanced groups in the genus.
characterized by a short-creeping stem, stem
diameter more than 4 mm wide, prominent 8. The Campyloneurum xalapense group.
primary veins, more than 5 rows of areoles This group includes Campyloneurum costatum, C.
between costa and margin, and usually cochense, and C. xalapense. This is a
symmetrically divided primary areoles, or rarely heterogeneous group; most of the characters
undivided. All species occur typically at low appear intermediate between C. amphostenon and
elevations as terrestrials or low trunk climbers. C. phyllitidis, and for this reason is located among
This group is interpreted as a line independent them in the evolutionary diagram (Fig. 51). The
from the C. amphostenon group because of the C. xalapense group is characterized by having the
more complex venation, and separate from the C. stem usually more than 5 mm wide, leaves with
brevifolium group because of the equal division of more than 5 areoles between the costa and
areoles. margin, and primary veins inconspicuous or
partially prominulous on one side of the lamina.
6. The Campyloneurum brevifolium group.
This includes Campyloneurum brevifolium, C. 9. The Campyloneurum angustifolium group.
nitidissimum, C. pascoense, and C. tucumanense. This group includes Campyloneurum aglaolepis, C.
These species are terrestrial or creeping trunk angustifolium, C. angustipaleatum, C.
epiphytes, characterized by a stem diameter more austrobrasilianum, C. centrobrasilianum and C.
than 6 mm wide; large leaves, non-costal areoles ensifolium. This is a very homogeneous group that
asymmetrically divided, with prominent primary shares stem diameters less than 4 mm wide,
and secondary veins. Campyloneurum lamina with 1-3 rows of areoles between costa
nitidissimum is the only species with non-clathrate and margin, and inconspicuous primary veins.
scales. This group resembles the C. phyllitidis They are epiphytes that occur mostly at middle
group in stem diameter and stem scale elevations.
characters. The complexi pattern of venation of
the C. brevifolium group is interpreted as highly 10. The Campyloneurum vulpinum group.
specialized. This is composed of Campyloneurum cubense, C.
oxypholis, and C. vulpinum, and is related to the C.
7. The Campyloneurum amphostenon group. angustifolium and the C. repens groups. The three
Campyloneurum amphostenon, C. asplundii, C. species are characterized by primary veins 50-65o
chlorolepis, C. densifolium, C. fallax, C. lorentzii, C. divergent from the costa, with undivided
rigidum, and C. solutum have a stem diameter less primary non-costal areoles, and narrowly
than 6 mm wide; lamina with 2-5 rows of areoles lanceolate, thin chartaceous leaves.
between costa and margin, divergent angle of
primary veins less than 50o, and primary non- PHYLOGENY
costal areoles symmetrically divided with one During the last thirty years many genealogical
veinlet per areole. In C. amphostenon, C. rigidum relationships in different kinds of organisms have
and C. solutum stem scales are clathrate or been discussed on the basis of hypotheses
slightly clathrate, while in C. asplundii and C. generated by cladistic analyses under the
chlorolepis they are non-clathrate. This group is assumption of parsimony. The results of this
widespread in mountainous environments of kind of analysis have been compared to the
current classification and/or to other hypotheses. near to C. anetioides.

Here a cladistic analysis is attempted for the first Other groups are different. For instance
time for Campyloneurum. Campyloneurum aglaolepis, C. ensifolium, C.
angustipaleatum, C. austrobrasilianum, and C.
Species level centrobrasilianum are separated from C.
Twenty-three characters were used in the angustifolium, which on the other hand is placed
analysis (Table 10). A hypothetical closer to C. amphostenon, perhaps indicating a
polypodiaceous ancestor was included in the close relationship .
analysis as an outgroup, because earlier attempts In the cladogram Campyloneurum phyllitidis
using Microgramma, Niphidium, Pleopeltis, and seems closer to the clade of C. brevifolium. Two
Pyrrosia as outgroups produced more than 1000 of the species (C. nitidissimum, C. tenuipes) that
unresolved polytomic trees. were assumed to be allies of C. phyllitidis are
The characters used in this analysis were located closer, while others are scattered in the
mostly qualitative. Most characters had binary tree: C. abruptum, C. nitidum, and C. wurdackii.
states and only six characters had multistate Campyloneurum xalapense is located close to C.
conditions (Table 10). Not all characters found in tenuipes as was anticipated (Fig. 51). Allied
the genus were used because of a lack of species of C. xalapense, however, are separated: C.
information for the majority of species. Examples cochense comes out closer to C. densifolium and C.
include, for example, chromosome number, spore nitidum , and C. costatum between C. amphostenon
morphology, and anatomical details of the stem and allied species, and C. cubense, C. leuconeuron
and leaf. The data matrix used for this analysis and C. rigidum.
(Table 11) has very few missing data, and these In the cladogram (Fig. 52), the species with
correspond to characters where the states were undivided primary areoles (Campyloneurum
defined with a binary condition rather than a acrocarpon, C. chrysopodum, C. coarctatum, C.
multistate. falcoideum, C. fasciale, C. fuscosquamatum, C.
A consensus tree for the species (Fig. 52) was inflatum, C. minus, C. oellgaardii, C. ophiocaulon, C.
obtained from the first 600 trees; the length of the repens, C. sphenodes and C. sublucidum) are located
shortest tree was 86, and the consistency index closely together, as was previously anticipated;
was 0.326. The low value of the consistency however, they come out far away from the
index may suggest high homoplasy in the ancestor. Species with undivided areoles come
characters used. Homoplasy is one of the out together with species considered to be closely
problems that was encountered by Hovenkamp related to other species: C. abruptum, C. cochense,
(1986) and Bosman (1991) while performing C. densifolium, C. lorentzii, C. oxypholis, and C.
cladistic analyses in other polypodiaceous fern wurdackii. Some of these species were difficult to
genera. assess in regards to their affinities, such as C.
The consensus tree for the species shows a cochense and C. oxypholis.
major polytomy at the base of the tree, with one Although the cladistic analysis was not
branch going to the hypothetical ancestor, one to completely satisfactory due to homoplasy, in
Campyloneurum magnificum, another to C. general the results support the classification
decurrens and another to the entire-leaf species presented in this revision.
(Fig. 52). Other polytomies are found among the
species with narrow leaves, those with divided Species-group level
areoles and others with undivided areoles. Twelve characters were used for this analysis
When the ideas of phenetic relationships, as (Table 12). These characters were chosen
presented before (Fig. 51) are compared to the because most of them only allowed two
results of the cladistic analysis, consistency is conditions in six groups in the data matrix (Table
found to some extent. For example, according to 13). Three of these characters had a multistate
both approaches Campyloneurum brevifolium condition (Table 12). Similar to the species
comes out with C. pascoense and C. tucumanense, analysis, a hypothetical ancestor was defined and
and Campyloneurum aphanophlebium comes out the character states were given based on
comparisons to other genera of the of Campyloneurum has been discussed among

Polypodiaceae. pteridologists (cf. Tryon & Tryon, 1982 b;
A consensus tree (Fig. 53) was obtained from a Lellinger, 1988, León, in press). The scope of this
total of 149 trees; the length of the shortest tree discussion has implications in the phylogenetic
found was 28, and the consistency index was interpretation of the genus because it deals with
0.536. Polytomies were found at the base of the its monophyletic status. The two cladistic
tree and in two other places. analyses made here showed that the pinnate
The information from the tree shows that the species are not totally incorporated in the clade of
Campyloneurum magnificum group is closer to the the entire-leaf species. This can be used to
ancestral type of the genus than any other group support the idea of the exclusion of those species
(Fig. 53). This agrees with the analysis with pinnate leaves. However, both analyses are
performed for the species (Fig. 52), and also with problematic because of homoplasy and
the phenetic hypothesis (Fig. 51). The positions polytomies. This classification, as presented in
of the C. angustifolium and C. amphostenon groups Chapter IX and as was discussed earlier, accepts
are interesting. The former appears closer to the the entire genus as monophyletic, based on
ancestral type than the C. amphostenon group, morphological similarities, without considering
which previously had four of its components that only apomorphies have to define the clade.
closer to the species of the former group (Fig. 52).
The position of the C. angustifolium group may be
interpreted as representing ancestral features for
all simple leaf species; this idea was intuitively IX. TAXONOMIC TREATMENT
proposed by Lellinger (1988).
The Campyloneurum xalapense group was Campyloneurum C. Presl, Tent. Pterid. 189. 1836.
placed twice in the data matrix (Table 13) because Lectotype: (chosen by J. Smith, 1875)
character #6, stem diameter, was variable within Campyloneurum repens (Aublet) C. Presl.
the group. It is difficult to interpret the position Polypodium subg. Cyrtophlebium R. Brown, in
of this group in the tree: the group was Bennet & Brown, Pl. Jav. Rar. 4. 1838.
heterogeneous, with features found in both C. Cyrtophlebium (R. Br.) J. Sm., J. Bot. 4: 58. 1841.
amphostenon and C. phyllitidis groups, but it did Hyalotricha Copel., Amer. Fern J. 43: 12. 1953.
not occur among them (Fig. 53). Type: Hyalotricha anetioides (Christ) Copel.
The closeness of the groups with long- (Polypodium anetioides Christ), not Dennis,
creeping stems and mostly undivided areoles 1949.
(Campyloneurum aphanophlebium, C. repens, C. Hyalotrichopteris W. Wagner, Taxon 27: 548.
sphenodes, and C. vulpinum groups) shows that 1978. Nom. nov. for Hyalotricha Copel.
they in fact represent close lineages, as Stem short or long creeping, sometimes
previously anticipated. The position of the C. branched, with clathrate or non-clathrate scales,
aphanophlebium group closer to the C. sphenodes phyllopodia present. Leaves simple or 1-pinnate,
group than to the C. repens group is very similar homophyllous, 10-300 cm long, glabrate or
to the taxonomic interpretation at the species pubescent, or scales on the costa; petiole usually
level (Chapter IX). present and articulate. Costa usually prominent
The Campyloneurum phyllitidis and the C. on both sides of the lamina, veins areolate,
brevifolium groups appear together and far away primary veins usually parallel, divergent from
from the ancestor of the groups (Fig. 53). These the costa, secondary veins transverse, forming
positions differ from that of the species-level primary areoles, these with 1-6 included veinlets,
analysis (Fig. 52), where most of the species tertiary veins or veinlets free or anastomosing,
belonging to both groups appeared near the simple or furcate, excurrent, transverse, rarely
hypothetical ancestor. recurrent; excurrent veinlets connected
sometimes to secondary veins forming secondary
Phylogenetic interpretation areoles, in a few cases veinlets in secondary
During the last ten years the generic concept areoles forming tertiary areoles, apex of free
veinlets with hydatodes. Sori exindusiate, side of the lamina, or prominulous near the
usually medial to apical on the free veinlet, or at costa on both sides of the lamina. 5
the junction of two veinlets, in 2-4 (-6) series 4. Primary veins conspicuous, prominulous or
between secondary veins; paraphyses simple, prominent to the same degree on each side of
filamentous or dendritic, or absent. Spores the lamina. 9
monolete, slightly verrucate or verrucate. Basic 5. Lamina lanceolate or narrowly obovate; bases
chromosome number: 2n= 74, 148. narrow cuneate (southern U.S.A. to Panama,
Greater Antilles, Trinidad, Venezuela and
The genus comprises epiphytic, creeping or Ecuador). 17. C. costatum
terrestrial plants. It grows from southern U.S.A., 5. Lamina linear-lanceolate or narrowly
Mexico, Central America, Antilles, Colombia, lanceolate, bases attenuate. 6
Venezuela to Bolivia and Brazil. In this 6. Primary veins conspicuous, prominulous
treatment, forty-seven species are recognized, from the costa to the margin on one side of the
most of them from the American tropics. lamina; 4-7 primary areoles between the
margin and costa. 7
The characters of the key apply to mature 6. Primary veins inconspicuous; 2-5 primary
leaves and individuals. Complete specimens are areoles between the margin and costa. 8
required, especially among narrow leaved 7. Stem scales subadpressed, with well
species, where stem scales have many valuable differentiated margins, 2-3 (-4) mm long; stem
characters. usually pruinose (Colombia to Ecuador).
The number of areoles between the costa and 16. C. cochense
the leaf margin includes both costal and non- 7. Stem scales spreading, without differentiated
costal areoles. Prominence of the veins and angle margins, 3-6 mm long; stem not pruinose
of divergence of the primary veins from the costa (Mexico to Costa Rica).
should be examined at the central part of the leaf, 47. C. xalapense
and does not usually require the use of a 8. Stem scales with acuminate apices, 4.5-7 mm
microscope. However, a stereomicroscope will long, cells oblong or broadly oblong (Mexico
be required to examine the indument and stem to Bolivia). 20. C. densifolium
scales. 8. Stem scales with acute or obtuse apices, 3-4
Some species are mentioned two or more mm long, cells broadly ovate (Bolivia to
times in the key when characters may overlap. Argentina). 27. C. lorentzii
9. Stem scales narrowly ovate, three times or
more longer than broader. 10
Key to species 9. Stem scales ovate or broadly ovate, less than
three times longer than broader. 12
1. Leaves 1-pinnate 2 10. Lamina bases cuneate; lamina long petiolate,
1. Leaves simple. 3 petiole length 1/4-1/2 of the lamina (Mexico
2. Areoles of the pinna with 2-3 excurrent to Honduras).
veinlets; pinna width less than 4 cm 43. C. tenuipes
(Colombia, Venezuela, Martinica, southern 10. Lamina bases abruptly cuneate then
Brazil). 19. C. decurrens decurrent or attenuate; lamina short petiolate,
2. Areoles of the pinna with 3-5 excurrent petiole length less than 1/5 of the lamina. 11
veinlets; pinna width more than 5 cm (Panama 11. Stem scales distinctly clathrate, cell lumen
to Bolivia, Trinidad, southern Brazil). translucent; lamina herbaceous-chartaceous;
29. C. magnificum primary non-costal areoles usually undivided,
3. Stem usually more than 6 mm wide. 4 rarely symmetrically divided. (Colombia to
3. Stem usually equal or less than 5 mm wide. Bolivia and western Brazil).
17 1. C. abruptum
4. Primary veins inconspicuous, or conspicuous 11. Stem scales non-clathrate, cell lumen dark
and prominulous in different degrees on each yellow brown; lamina subcoriaceous or
coriaceous; primary non-costal areoles between the costa and margin (Costa Rica and
asymmetrically divided (Colombia to Bolivia).32. C. Ecuador).
nitidissimum 42. C. sublucidum
12. Primary non-costal areoles undivided or 19. Lamina lanceolate or narrowly lanceolate;
symmetrically divided only to secondary dull on one side of the lamina; 2-5 primary
areoles. 13 areoles between the costa and margin. 20
12. Primary non-costal areoles asymmetrically 20. Primary non-costal areoles mostly
divided to secondary and tertiary areoles. 15 undivided; stem 1-2 mm wide. 21
13. Lamina bases cuneate, long petiolate; petiole 20. Primary non-costal areoles divided; stem
length more than 1/3 of the lamina; more than more than 2 mm wide. 22
16 areoles between the costa and margin o
21. Primary veins 50-70 divergent from the
(northwestern Ecuador). costa; stem scales clathrate, peltate, falcate
33. C. oellgaardii (Costa Rica, Panama, southwestern Colombia).22. C. falcoideum
13. Lamina bases attenuate, short petiolate;
21. Primary veins 30-40o divergent from the
petiole length usually less than 1/3 of the
costa; stem scales non-clathrate, mostly
lamina; equal or less than 16 areoles between
pseudopeltate, narrowly ovate (Haiti,
the costa and margin. 14
Dominican Republic, and from Ecuador to
14. Primary areoles undivided; lamina apices
Bolivia, and central Brazil).
and bases attenuate; stem scales slightly
45. C. vulpinum
bullate, 1-2 (-3) mm long (southeastern Brazil).
22. Stem scales spreading, narrowly ovate. 23
2. C. acrocarpon
22. Stem scales subadpressed or adpressed,
14. Primary areoles usually divided; lamina
ovate or ovate lanceolate. 24
apices acuminate and bases attenuate; stem
23. Stem scales distinctly clathrate, cell lumen
scales flattish, 4-8 mm long (southern U.S.A.,
usually transparent; scales with oblong cells
Mexico, Central America, Antilles to Bolivia
(Mexico, Central America, Antilles, Venezuela
and central Brazil).
to northern Argentina).
37. C. phyllitidis
4. C. amphostenon
15. Secondary veins inconspicuous or slightly
23. Stem scales slightly clathrate, cell lumen
prominulous, same color as the leaf tissue.
usually yellowish; scales with narrowly
(Mexico, Central America, Antilles, Venezuela
oblong cells (Colombia, Venezuela to Peru).
to Bolivia). 11. C. brevifolium
40. C. solutum
15. Secondary veins conspicuous, prominulous,
24. Apices of stem scales acute or obtuse; stem
usually stramineous. 16
scales 3-4 mm long (Bolivia and northern
16. Stem scales 8-10 mm long, more than 2.5 mm
Argentina). 27. C. lorentzii
wide; laminae chartaceous or subcoriaceous
24. Apices of stem scales acuminate; stem scales
(Ecuador to Bolivia). 36. C. pascoense
4-7 mm long. 25
16. Stem scales 5-6 mm long, 1-2.5 mm wide;
25. Lamina linear-lanceolate; cells of stem scales
laminae herbaceous (northern Argentina).
oblong or broadly oblong; stem scales light
44. C. tucumanense
brown in mass (Mexico to Panama, Cuba, and
17. Phyllopodia distance more than 7 mm apart.
from Colombia to Bolivia).
18
20. C. densifolium
17. Phyllopodia distance 6 mm apart or less. 36
25. Lamina lanceolate; cells of stem scales
18. Primary veins 40-55o (-60o) divergent from roundish or broadly oblong; stem scales
the costa, inconspicuous or sometimes slightly pinkish brown in mass (southeastern Brazil).
prominulous abaxially. 19 23. C. fallax
18. Primary veins 60-80o divergent from the 26. Stem scales with obtuse apices. 27
costa, prominulous or prominent to the same 26. Stem scales with acuminate apices. 28
degree on both sides of the lamina. 26 27. Stem scales slightly bullate, with non-
19. Lamina broadly lanceolate, shining on both differentiated margins; stem scales broadly
sides of the lamina; 5-6 primary areoles ovate (southeastern Brazil).
2. C. acrocarpon yellow or obsolete (Colombia to Bolivia).

27. Stem scales flat, with differentiated margins; 25. C. fuscosquamatum
stem scales ovate (Colombia and Venezuela to 36. Primary areoles 6 or more between the costa
Bolivia). 34. C. ophiocaulon and margin. 37
28. Margins of stem scales differentiated in color 36. Primary areoles 5 or less between the costa
and cell disposition. 29 and margin. 46
28. Margins of stem scales not differentiated. 32 37. Leaves puberulous; hairs spreading on both
29. Petiole length more than 1/3 of the lamina; sides of the lamina (Belize to Bolivia and
lamina base cuneate, if narrowly cuneate then western Brazil). 8. C. aphanophlebium
stem scales narrowly ovate. 30 37. Leaves glabrous or glabrescent; hairs if
29. Petiole length less than 1/3 of the lamina; present spreading abaxially. 38
lamina base decurrent or narrowly cuneate. 38. Primary veins inconspicuous or slightly
31 prominulous near the costa, and with same
30. Leaves more than 50 cm long; stem scales 2 color as the leaf tissue. 39
mm or more wide (northwestern Ecuador). 38. Primary veins prominulous or prominent;
33. C. oellgaardii stramineous or darker than the leaf tissue. 42
30. Leaves less than 50 cm long; stem scales less 39. Lamina oblanceolate or narrowly obovate
than 1 mm wide (Costa Rica to Peru). (southern U.S.A. to Panama, Greater Antilles,
41. C. sphenodes Trinidad, Venezuela and Ecuador).
31. Stem scales clathrate, subadpressed, 17. C. costatum
caducous; sori without paraphyses (Mexico, 39. Lamina linear-lanceolate or narrowly
Central America, Greater Antilles to Bolivia lanceolate. 40
and central Brazil). 38. C. repens 40. Scales of the stem with acuminate apices;
31. Stem scales slightly clathrate, spreading, cells of stem scales narrowly oblong (Mexico
persistent; sori with long filamentous to Costa Rica). 47. C. xalapense
paraphyses (Colombia and Venezuela). 40. Scales of the stem with obtuse or short
28. C. macrosorum acuminate apices; cells of stem scales oblong
32. Lamina ovate, long petiolate, petiole usually or roundish. 41
more than 10 cm long. 33 41. Scales of the stem slightly bullate, with
32. Lamina lanceolate, short petiolate, petiole obtuse apices, 2-2.5 mm long; primary non-
usually less than 5 cm long. 34 costal areoles undivided (southeastern Brazil,
33. Cell lumen of stem scales translucent Argentina and Paraguay).
colorless; cells narrowly oblong; stem scales 30. C. minus
dark brown in mass. Costa Rica to Bolivia 41. Scales of the stem flattish, with short
dark brown in mass (Costa Rica to Bolivia and acuminate apices, ca.4 mm long; primary non-
western Brazil). 15. C. coarctatum costal areoles usually divided (Cuba, Jamaica
33. Cell lumen of stem scales yellowish; cells and Haiti. 18. C. cubense
roundish; stem scales brown in mass 42. Apices of the scales of the stem obtuse or
(Colombia and Peru). 26. C. inflatum acute. 43
34. Stem scales peltate, base with one long 42. Apices of the scales of the stem long
auricle; stem 1-2 mm wide; lamina base acuminate. 44
narrowly cuneate (Costa Rica, Panama, 43. Stem scales ovate; primary veins 65-70 o
southwestern Colombia). 22. C. falcoideum divergent from the costa (southeastern Brazil.
34. Stem scales pseudopeltate, base with two 2. C. acrocarpon
symmetrical auricles; stem 2-3 mm wide; 43. Stem scales broadly ovate; primary veins 50-
lamina base decurrent. 35
60° divergent from the costa (southeastern
35. Stem scales distinctly clathrate, cell lumen
Brazil, Argentina, Uruguay, and Paraguay).
translucent (Mexico to Bolivia).
31. C. nitidum
24. C. fasciale
44. Petiole length 1/4-1/2 of the lamina; lamina
35. Stem scales non-clathrate; cell lumen dark
base cuneate (Mexico, Guatemala and 4-7 mm apart (Venezuela to Bolivia). 9. C.

Honduras). 43. C. tenuipes asplundii
44. Petiole length less than 1/8 of the lamina; 53. Stem scales whitish or light brown in mass;
lamina base attenuate. 45 phyllopodia 1-2 mm apart (Colombia,
45. Primary veins prominulous or prominent to Venezuela to Bolivia, and central-southern
the same degree on both sides of the lamina, Brazil). 13. C. chlorolepis
approximately straight (southern U.S.A., 54. Stem scales broadly ovate, with obtuse
Central America, Antilles, Colombia to Bolivia apices. 55
and central Brazil). 54. Stem scales ovate or narrowly ovate, with
37. C. phyllitidis acute or acuminate apices 56
45. Primary veins prominulous usually 55. Lamina dull, dark green adaxially; usually 5
adaxially, slightly prominulous abaxially, or more areoles between costa and margin
conspicuously sinuous (Mexico to Costa Rica). (southern Brazil to Argentina).
47. C. xalapense 31. C. nitidum
46. Leaves puberulous; hairs scattered on both 55. Lamina bright light green on both sides;
sides of the lamina. 47 usually 3 or less areoles between costa and
46. Leaves glabrescent; hairs if present scattered margin (southeastern Brazil). 39. C. rigidum
abaxially. 48 56. Primary veins 60-75o divergent from the
47. Lamina spathulate, apex obtuse; plants costa; primary areoles 4-7 between the costa
usually less than 10 cm long (Nicaragua, Costa and margin. Mexico to Costa Rica.
Rica and Panama). 5. C. anetioides 47. C. xalapense
47. Lamina narrowly obovate, apex acuminate or
56. Primary veins 40-55o (-60o) divergent from
subcaudate; plants usually more than 10 cm
the costa; primary areoles usually 1-5 between
long (Belize to Bolivia and western Brazil).
costa and margin. 57
8. C. aphanophlenium
57. Stem scales adpressed, broadly lanceolate;
48. Primary non-costal areoles undivided; stem
2.5-3 mm wide (Mexico, Central America to
1-2 mm wide. 49
Bolivia). 20. C. densifolium
48. Primary non-costal areoles mostly divided; if
57. Stem scales spreading or subadpressed, ovate
primary areoles undivided then stem more
or narrowly ovate. 58
than 3 mm wide. 50
58. Stem usually long-creeping; stem scales dark
49. Stem scales ovate, slightly bullate (Brazil,
brown, bright, cell lumen yellowish; lamina
Argentina and Paraguay). 30. C. minus
base usually narrowly cuneate (Venezuela to
49. Stem scales narrowly ovate, plane (western
Bolivia). 40. C. solutum
Venezuela). 14. C. chrysopodum
58. Stem usually short-creeping; stem scales
50. Lamina ovate with a cuneate then decurrent
brown, dull; cell lumen usually translucent;
base; some primary areoles asymmetrical
lamina base decurrent. 59
divided (southern Venezuela).
59. Stem scales with differentiated margins
46. C. wurdackii
(Cuba, Jamaica and Haiti). 18. C. cubense
50. Lamina linear or linear-lanceolate, base
59. Stem scales without differentiated margins.
attenuate; primary areoles symmetrically
60
divided. 51
60. Lamina linear or linear lanceolate; primary
51. Lamina linear-lanceolate, more than 1.5 cm
areoles 1-3 between costa and margin
wide. 52
(southern U.S.A. to Bolivia, Antilles, central
51. Lamina linear or narrowly obovate, less than
Brazil). 6. C. angustifolium
1.5 cm wide. 62
60. Lamina lanceolate or narrow lanceolate;
52. Stem scales non-clathrate; cell lumen same
primary areoles 2-5 between costa and margin.
color as the cell wall. 53
61
52. Stem scales slightly or distinctly clathrate;
61. Stem scales 1.5-2 mm wide; stem usually
cell lumen translucent or yellowish. 54
pruinose, 3-5 mm wide (Mexico, Central
53. Stem scales dark brown in mass; phyllopodia
America, Antilles, Venezuela to Argentina). oblong; cell walls ferrugineus (southeastern

4. C. amphostenon Brazil). 10. C. austrobrasilianum
61. Stem scales 1-1.5 mm wide; stem never 71. Stem scale cells roundish; stem scale apex
pruinose, 2-2.5 mm wide (Haiti). shortly acuminate (Mexico to Guatemala,
35. C. oxypholis Nicaragua). 21. C. ensifolium
62. Lamina yellow bright green to the same 71. Stem scale cells oblong; stem scale apex long
degree on both sides (southeastern Brazil). acuminate. 72
39. C. rigidum 72. Stem scales 6-10 mm long, 1.5-2 mm wide,
62. Lamina dark bright green or dull to different persistent, cells oblong usually with iridescent
degrees on either side. 63 lumen (central Ecuador, southern Peru to
63. Lamina lanceolate or linear-lanceolate, more Argentina). 3. C. aglaolepis
than 1 cm wide. 64 72. Stem scales 3-6 mm long, 1-1.5 mm wide,
63. Lamina linear, less than 1 cm wide. 68 caducous, cells narrowly oblong, lumen
64. Lamina narrowly obovate; stem scales with usually not iridescent (southern U.S.A.,
differentiated margins; stem never pruinose Antilles to Bolivia, and central Brazil).
(Cuba, Jamaica and Haiti). 18. C. cubense 6. C. angustifolium
64. Lamina linear-lanceolate; stem scales with or
without differentiated margins; stem usually
pruinose. 65 1. Campyloneurum abruptum (Lindman) B. León,
65. Stem scales whitish (Colombia, Venezuela to Fieldiana Bot. n.s. 1993: in press.
Bolivia and central-southern Brazil). Polypodium repens Aublet var. abruptum
13. C. chlorolepis Lindman, Ark. Bot. 1: 245. 1903. Lectotype
65. Stem scales brown. 66 (chosen here). Brazil: Matto Grosso, Serra do
66. Stem scales adpressed, broadly ovate or Itapirapuam, ad arbores, 28 Apr. 1894,
ovate (Mexico, Central America to Bolivia). Lindman (Regnell Exped. I) 3345 (type S!,
20. C. densifolium isotype K!).
66. Stem scales spreading, narrowly ovate. 67 Campyloneurum nitidissimum var. abruptum
67. Stem scales 3-5 mm long, 1-1.5 mm wide, cell (Lindman) B. León, Ann. Missouri Bot. Gard.
lumen occluded; lamina base usually 77: 212. 1990. p.p.
decurrent (Venezuela, Ecuador to Bolivia). Stem creeping, not pruinose, 5-10 mm wide.
9. C. asplundii Stem scales dark brown in mass, linear-
67. Stem scales 4-7 mm long, 1.5-2.5 mm wide, lanceolate or narrowly-ovate, 5-7 mm long, 1-1.5
cell lumen yellowish or transparent; lamina mm wide; scale bases short auriculate, apices
base usually narrowly cuneate (Colombia, acuminate, scales distinctly clathrate, cells
Venezuela to Peru). 40. C. solutum narrowly oblong, cell walls 6-13 µm wide, some
68. Base of scales of stem 1 mm or less wide. times with spreading hair-like teeth on the
69 margins. Phyllopodia 1-2 mm long, 4-5 mm
68. Base of scales of stem more than 1 mm wide. wide, 2-7 mm apart. Leaves erect, 60-110 cm
71 long; petiole 2-7 cm long, stramineous, slightly
69. Base of stem scales with outline of cells ranurate adaxially; lamina lanceolate or ovate,
contorted (central Brazil). (4.5-) 6.5-13.5 cm wide, chartaceous or
12. C. centrobrasilianum herbaceous-chartaceous, lamina base abruptly
69. Bases of stem scales without contorted cells cuneate or attenuate, then long-decurrent, apex
(Mexico to Panama, Greater Antilles, and from usually acuminate, margins of the lamina
Colombia to Bolivia, southeastern Brazil). 70 slightly sinuate, cartilaginous, lamina
70. Stem scales subulate from a roundish base; puberulous, indument of inconspicuous,
scale cells narrowly oblong; cell walls usually bicellular hairs, 70-75 µm long, scattered
brown (Costa Rica, Ecuador to Bolivia). abaxially, stomata polocytic; costa prominent,
7. C. angustipaleatum slightly sulcate adaxially, indument of caducous
70. Stem scales narrowly ovate; scale cells scales at the base; primary veins prominent,
diverging (60o-) 65-70o from the costa, straight, MADRE DE DIOS: Río Manu, Cocha Cashu Biological
station, Jul. 1978, Foster & Terborgh 6559 (F);
lighter in color than the adjacent tissue, (3-) 5-7 (-
Tambopata, SE Puerto Maldonado, Albergue "Cuzco
10) mm apart, with 9-15 areoles between costa Amazónico", Apr. 1986, León 884 (F, USM).
and margin, with 2-4 excurrent veinlets in each BOLIVIA. LA PAZ: Provincia Larecaja, Casanavi, 27.8
non-costal areole, veinlets entire or furcate, km Guanay, 28 Nov. 1980, Beck 3784 (F, LPB);
usually free, rarely forming regular secondary Caranavi and Guanay, 28 Nov. 1980, Croat 51658 (MO,
areoles. Sori subterminal on the excurrent UC); Nor Yungas, 4.5 km below Yolosa, 19-20 Oct.
veinlet, paraphyses inconspicuous or lacking; 1982, Solomon 8549 (MO).
only abortive spores seen. Figs. 9 a; 19 a. BRAZIL. RORAIMA: Posto Mucajaí, Rio Mucajaí, vic.
Mucajaí airstrip, 13 Mar. 1971, Prance et al. 10923 (K,
MO, NY, S, US). AMAZONAS: Matto do Curupira, 18
Feb. 1894, Lindman 3075 (S, B); Rio Curuquete,
South American species, distributed from Cachoeira República, 25 Jul. 1971, Prance et al. 14584
Colombia and Venezuela, south to Bolivia and (NY, US). PARA: W bank of Rio Maicurú, ca. 23 km
Brazil, where is found between 100 m and 650 m from Lageira, 29 Jul. 1981, Strudwick et al. 3702 (F, NY).
elevation. It grows mostly as terrestrial in shady RONDONIA: basin of Rio Madeira, 2 km below
and humid places, on abundant organic debris, confluence of Río Abunã, 12 Nov. 1968, Prance et al.
and sometimes on rocks. 8341 (K, MO, S, US). MATTO GROSSO: between S.
Cruz and Jairip, 1891-92, Moore 370 (BM); gorge of Véu
de Noiva, Chapada dos Guimarães, 17 Oct. 1973,
Prance et al. 19105 (K, US). GOIAS: Mun. Jataí,
REPRESENTATIVE SPECIMENS: COLOMBIA.
Bálsamo, 9 Feb. 1895, Macedo 2148 (S, US). MINAS
MAGDALENA: La Jagua, 11 Sep. 1924, Allen 655
GERAIS: Distrito Rio Branco, 13 Nov. 1930, Mexia 5298
(MO); on Quebrada Santa Cruz, 5 km N of La Jagua, 3
(BM, NY, S). Without locality: Glaziou 15755 (B).
Aug. 1943, Haught 3582 (GH, US). BOLIVAR: Boca
Verde, on Río Sinu, 13-14 feb. 1918, Pennell 4216 (NY).
SUR DE SANTANDER: vicinity of Barranca Bermeja, Campyloneurum abruptum can be
Magdalena valley, between Sogamoso and Colorado distinguished by the abruptly cuneate or
rivers, 19 Feb. 1935, Haught 1567 (US). BOYACA: attenuate leaf bases that are then long-decurrent,
Casanare, Tauramena, 13 Apr. 1963, Uribe 4281 (US). and by the stem scales dark brown in mass,
META: Río Guatiquia, near Villavicencio, 18 Mar. distinctly clathrate, with a clear cell lumen.
1939, Alston 7597 (US). WHITOUT EXACT Campyloneurum abruptum belongs to the C.
LOCALITY: Llanos de San Martín, Stübel 598 (B). phyllitidis group.
VENEZUELA. MERIDA: s.d. Karsteinley s.n. (W); ESE
of Santa Bárbara, 9 Mar. 1980, Liesner & Gonzales 9236
(MO). TACHIRA: ESE of la Fundación, 0-3 km below
Represa Dorada, 29 Apr. 1981, Liesner & Guariglia 2. Campyloneurum acrocarpon Fée, Cr. Vasc. Br. 1.
11539 (MO, UC); W of Pinal, W of bridge over Río 35., t. 115. 1869. Type. Brazil: Serra dos
Frío, 27-30 Aug. 1966, Steyermark & Rabe 96710 (GH); Orgãos, Glaziou 2801 (probably P).
dist. Uribante, from La Siberia to entrance to Represa Campyloneurum wacketii Lellinger, Amer. Fern J.
Las Cuevas, 10 Jul. 1983, Werff & Gonzalez 5241 (MO). 78: 27. 1988. Type. Brazil: São Paulo, Río
BARINAS: dist. Pedraza, above El Algarrobo, 3 Aug. Grande, Wacket (Ros. Fil. Austrob. Exsc. 213)
1983, Werff & Ortiz 5810 (MO, UC).
(holotype US!, isotypes B!, S!, UC!).
ECUADOR. NAPO: 1.1 km E of Río Conejo on road to
Lago Agrio, 31 Mar. 1972, Dwyer & Mac Bryde 9769
Stem long-creeping, not pruinose, black or
(MO); Napo-Pastaza, Cantón Napo, Zatzayacu, 22 stramineous, 3-6 mm wide. Stem scales partly
Mar. 1935, Mexia 7065a (UC). persistent, light brown in mass, 1-2 (-3) mm long,
PERU. SAN MARTIN: Chazuta, Río Huallaga, Apr. 0.5-1 (-1.5) mm wide, ovate, slightly bullate, base
1935, Klug 4080 (F, GH, MO, S, UC, US); 15 km E of auriculate, apex obtuse, clathrate, the cells
Shapaja on road to Chazuta, along Quebrada Chumia oblong, irregularly arranged, cell walls 8-10 µm
4 Aug. 1986, Knapp 7867 (F, MO, USM); Mariscal wide, scales sometimes with teeth on the
Cáceres, dist. Campanilla, Mashuyacu, 12 Aug. 1970,
margins. Phyllopodia 2-4 mm long, 4 mm wide,
Schunke V. 4226 (F, US); Moyobamba, Huallaga,
3-8 mm apart. Leaves 45-92 cm long; petiole 1-3
Potrero to Tabalosos, Stübel 1101 (B). HUANUCO:
Pachitea, Panguana, 1983, Seidenschwarz 100/34 (USM). cm long, stramineous, slightly ranurate
adaxially; lamina narrowly lanceolate to 3. Campyloneurum aglaolepis (Alston) Sota, Opera

oblanceolate, 4-7 cm wide, herbaceous- Lilloana 5: 96. 1960.
chartaceous to chartaceous, lamina bases and Polypodium aglaolepis Alston, J. Bot. 77: 346. 1939.
apices attenuate, lamina margins sinuate, Type. Argentina: Tucumán, Siambón, Sierra
cartilaginous, indument of inconspicuous simple de Tucumán, Lorentz & Hieronymus 948
glandular hairs, scattered abaxially, stomata (holotype BM!; isotype B! CORD).
polocytic; costa prominent; primary veins Stem short-creeping, sometimes slightly
prominulous on both sides of the lamina in the pruinose, 1-3 mm wide. Stem scales brown or
same degree, slightly stramineous, 65-70o dark brown, iridescent in mass, (4.5-) 6-10 mm
divergent from the costa, slightly sinuate, (4-) 5-8 long, (1-) 1.5-2 (-2.5) mm wide, narrowly ovate,
mm apart, (5-) 7-12 areoles between costa and scale base auriculate, apex long acuminate,
margin, 2-3 (-4) excurrent veinlets in each clathrate, the cells oblong, cell walls 10-15 µm
noncostal areole, veinlets entire or furcate, thick. Phyllopodia 1-2 mm long, 1-1.5 mm wide,
usually free; sori medial or subterminal, 2-4 (-6) mm apart. Leaves (10-) 40-60 cm long;
paraphysis scarce, simple, 11 µm long; spores 45- petiole green stramineous, (0.2-) 0.5-3 (-8) cm
50 µm long, 25-30 µm wide. Fig. 25. long, slightly ranurate adaxially, convex
abaxially, glabrate; lamina linear, bases and
Species restricted to southeastern Brazil, apices attenuate, 0.3-0.8 (-1.2) cm wide,
where it is usually found from sea level to 800 m. herbaceous-chartaceous, lamina margins
It grows mostly terrestrial in disturbed forests. cartilaginous, slightly revolute, indument absent,
stomata polocytic; costa prominent on both
REPRESENTATIVES SPECIMENS: BRAZIL. SÃO surfaces; venation areolate, primary veins
PAULO: Arariba, 1926, Brade 8443 (UC); Tietê, 16 Oct. inconspicuous, 2 areoles between the costa and
1904, Gerdes 39a (UC); Estação Sagrado do Coração, margin, undivided, marginal areoles smaller
Linha Sorocobana, 3 Oct. 1979, Mizoguchi 1043 (MO); than costal areoles, excurrent veinlets one in each
Santos, 1 Apr. 1875, Mosen 3744 (B). PARANA:
areole, sometimes marginal free veinlets. Sori
Yacarehy, 20 Mar. 1914, Dusen 97a (MO); 18 Jul. 1914,
subterminal or medial, paraphyses scarce,
Dusen 15311 (B); Yacarehy, 7 Aug. 1914, Dusen 15345
(BM, MO, S); Mun. Guaraqueçaba, Serrinha, dendritic, smaller than the sporangia; spores (60-
Hatschbach 16314 (F); Mun. Paranaguá, Ilha do Mel, 28 ) 65-70 µm long, 40-42 µm wide. Figs. 6 c; 7 a; 14
Nov. 1970, Hatschbach & Guimarães 25680 (UC); Mun. b; 26.
Morretes, Ilha do Turco, 2 Dec. 1975, Hatschbach 37834
(MU); Mun. Morretes, Ilha do Malha, 12 Nov. 1975, South American species known from central
Kummrow 988 (MU). MINAS GERAIS: dist. Rio Ecuador, southern Peru to Argentina and
Branco, 13 Nov. 1930, Mexia 5298 (MO). SANTA southern Brazil, where is found between 1000 m
CATARINA: Itajubá, 29 Jan. 1988, Krapovickas &
and 3600 m elevation. It grows as an epiphyte
Cristóbal 42140 (F); Santa Catarina, 10 Jul. 1901, Schmalz
13 (F); Antonio Carlos, Biguassú, Feb. 1943, Reitz 274 or in crevices among rocks.
(US); Brusque, Mata Hoffman, 10 Oct. 1949, Reitz 3091
(S, US); Indayal, 1904, Vierdel 15 (S). REPRESENTATIVE SPECIMENS: ECUADOR:
PICHINCHA: Hacienda Margarita, 35 km S of Santo
Domingo at Puerto Ila, 3 Jun. 1980, Balslev & Quintana
Campyloneurum acrocarpon is characterized by
24163 (AAU).
its long-creeping stem, with stem scales ovate, PERU. CUSCO: Piñasniocj, Panticalla (Pantiacolla)
slightly bullate, and by the herbaceous leaves, Pass, 14 Jul. 1915, Cook & Gilbert 1826 (US);
which are lanceolate with a long decurrent base. Quispicanchis, Marcapata, Cadena, 24 Jul. 1957, Vargas
Because of the characters in the pattern of 11665 (F). PUNO: Carabaya, Ollachea, Vargas 6885
venation and those of the stem, this species is (CUZ, MO).
placed in the C. repens group. Although the type BOLIVIA. LA PAZ: Sud Yungas, La Paz-Calacoto, 31
of C. acrocarpon was not located, its description Dec. 1980, Beck 3909 (F, LPB); Quime, 5 May 1949,
Brooke 5441 (BM, F); Bautista Saavedra, Charazani,
and illustration undoubtedly apply to this taxon.
Charazani to Khata, 19 Apr. 1982, Feurer 11258a (F);
Charazani, Río Curva, 17 May 1985, Feurer 22332a (F);
Bautista Saavedra, Curva, 30 Oct. 1979, Krach & Feuerer (W); Meyer et al. 21988 (W); Cochuna, 4 Mar. 1941,
6563a (F) Larecaja, Sorata, 7 Dec. 1981, Sperling & King Ousset 135 (F); Ousset 136 (F); San Javier, Cumbres
5386 (MO); Sorata, 1901-1902, Williams 2574 (MO). Calchaquies-Amaicha to Siambón, 25 Jan. 1933, Parodi
COCHABAMBA: Ayopaya, ca. 10 km NW 10659 (S); Quebrada de Las Sosas (Los Nogales); 12
Independencia, 9 May 1988, Beck & Seidel 14476 (LPB); Nov. 1952, Petersen & Hjerting 608 (BM, C); Sierra de
Choro, ca. 100 mi NW of Cochabamba, 18 Jan. 1950, Aconquija, 14 Nov. 1948, Skottsberg s.n. (GB);
Brooke 6005 (BM, F, S); Sailapata, Dec. 1934, Cárdenas Quebrada de los Sosa, km 25, 2 Sep. 1957, Sota 1668
3084 (US); Carrasco, 19 km W from Epizana,11 Feb. (S).
1987, Solomon & Nee 16048 (MO, USM); Socaba, 11 Oct.
1921, Steinbach 5847 (F, MO); Steinbach 5848 (F, MO). Campyloneurum aglaolepis is characterized by
SANTA CRUZ: Caballero, above Tunal, 30 km NE of its brown stem scales with oblong cells and
El Tambo School, 11 Jun. 1987, Killen 2556 (F, MO);
iridiscent cell lumen. The stem scales are
Vallegrande, Vallegrande to Piraimiri, 31 Jan. 1987,
Nee & Coimbra 33913 (MO). CHUQUISACA: Luis
persistent and numerous, slightly spreading.
Calco, 5 km N of Muyumpampa (Vaca Guzman), Sep. This species has dendritic paraphyses among the
1986, Willian 275 (F). TARIJA: Pinos near Tarija, 6 Jan. sporangia. It belongs to the Campyloneurum
1904, Fiebrig 2474 (BM, F, MO, S); O'Connor, Rancho angustifolium group.
Huayco, W of Entre Ríos, 4 Sep. 1987, Killeen 2705 (F,
MO); O'Connor, 73.1 km E of Tarija-Padcaya road, ca.
1 km below Narvaez, 1-2 May 1983, Solomon 10350 4. Campyloneurum amphostenon (Kunze ex
(LPB, MO, UC). Klotzsch) Fée, Gen. Fil. 258. 1852.
BRAZIL. PARANA: Mun. Bocaiuva do Sul,
Polypodium amphostenon Kunze ex Klotzsch,
Bacaetava, 30 Dec. 1980, Kummrow 1423 (MO). SÃO
PAULO: Campos do Jordão, Apr. 1947, Leite 3467 (GH, Linnaea 20: 399. 1847. Type. Venezuela:
MO, UC). MINAS GERAIS: Mun. Ouro Preto, São Mérida, Moritz 120b (holotype B!; isotypes
Sebastião, Macedo 3046 (MO). GOIÁS: Mun. Jataí, BM!).
Fazenda Queizadu, 13 Dec. 1948, Macedo 1480 (MO). Stem long-creeping, usually pruinose, (2-) 3-5
RIO DE JANEIRO: Itatiaya, 23 Oct. 1927, Zerny s.n. mm wide. Stem scales dark brown in mass,
(W). ovate, adpressed at their bases, with spreading
ARGENTINA. JUJUY: Selva Río Grande, Troncoso, 17 apices, (3-) 5-6 (-8) mm long, (1-) 1.5-2 (-2.5) mm
Feb. 1940, Burkart & Troncoso 11255 (MO); Jujuy, 6 Mar.
wide, the cell walls 8-10 µm thick. Phyllopodia
1937, Castellanos 19972 (BM); Laguna de Yala, 25 km
NW of Jujuy, 26 Sep. 1938, Eyerdam & Beetle 22230 (F,
1-2 mm long, 2-3 mm wide, 2-5 (-10) mm
MO, UC); dist. Santa Bárbara, 11 Feb. 1964, Fabris 5102 apart. Leaves 30-70 cm long; petiole stramineous
(US); Jujuy de Yala to Laguna de Yala, 8 Apr. 1945, or brown stramineous, (2-) 5-10 (-30) cm
O'Donnell 2899 (MO); road Lozano de Tiraxi, 3 Nov. long; lamina linear-lanceolate or lanceolate,
1974, Schinini et al. 10211 (UC); Tiraxi, 17 Mar. 1982, bases attenuate, apices acuminate, (1-) 2-3 (-5) cm
Schinini & Vanni 22459 (F). ROSARIO DE LA wide, chartaceous or subcoriaceous, margins
FRONTERA: Los Baños, 13 Jul. 1929, Venturi 9198 slightly revolute or plane, cartilaginous,
(MO). SALTA: Salta,16 Sep. 1985, Bonavia 74 (MO);
indument of simple trichomes scarce abaxially,
Salta, Mar. 1907, Hauman 339 (BM); San Lorenzo, Feb.
stomata polocytic; costa prominent, with
1936, Schulz 906 (GH). CATAMARCA: Andalgalá, 2
May 1915, Jorgensen 337 (BM). TUCUMAN: Quebrada indument of caducous scales, primary veins
Lules, Jul. 1928, Burkart 3283 (GH); Valle Tafí, Dec. obscure or prominulous adaxially, slightly
1911, Castillón s.n. (GH); 1-5 km S of Anta Muerta, on prominulous abaxially, often concolorous with
road to Villa Nougués, 7 Feb. 1974, Conrad & Dietrich the adjacent tissue, straight or slightly flexuosus,
2599 (MO, UC); La Banderita, 14 Feb. 1971, Ellenberg
5-7 mm apart, 40-50o divergent from the costa,
4480 (LPB); Tafí, road Apachiri-Alto del Clavillo, 20
transverse secondary veins forming 2-4 (-5)
Sep. 1946, Hunziker 6811 (US); valley of Río Cochuna,
W vicinity of Concepción, 16 Oct. 1989, Kramer et al. areoles between the costa and margin, 1-2 free
10681 (F); Quebrada Monteros, 5 Apr. 1872, Lorentz 791 excurrent veinlets in each noncostal areole; sori
(F); Quebrada de la Angostura, May 1945, Lourteig medial or subterminal on the excurrent veinlets,
1045 (BM, MO); Tafí, Tafí del Valle to Los Nogales, 6 paraphyses not seen, spores 62-70 µm long, 40-45
May 1947, Meyer 12111 (BM); Chicligasta, Las Pavas, µm wide.
16 Jun. 1949, Meyer 15137 (BM); Chicligasta, Las Pavas,
15 Mar. 1961, Meyer et al. 21977 (W); Meyer et al. 21984
Campyloneurum amphostenon is characterized Polypodium pittieri Christ in Pittier, Prim. Fl.

by linear lanceolate leaves with four or fewer Costa Rica 3: 16. 1901. Type. Costa Rica: El
rows of areoles between the costa and margin, by Páramo, E Cerro Buena Vista, Jan. 1897,
stems being frequently pruinose, and by its Pittier 10479 (holotype P ; isotypes US!, photo
lanceolate stem scales with spreading apices. of P, BM!).
This species is closely related to Campyloneurum pittieri (Christ) Ching,
Campyloneurum densifolium and C. fallax. They Sunyatsenia 5: 263. 1940.
are different from C. amphostenon by the Polypodium leuconeuron var. latifolia (ibid.)
adpressed, broadly ovate stem scales. Rosenst., Fedde Rep. 11: 58. 1912. Type.
At middle elevational levels some individuals Bolivia: La Paz, Yungas septentrionalis,
of C. amphostenon with narrow leaves and Unduavi, Nov. 1910, Buchtien 2759 (holotype,
narrow stem scales are difficult to distinguish not seen; isotypes S!, US; photo of US at F!,
from C. angustifolium. They can be USM!)
distinguished, however, because C. amphostenon Polypodium poloense Rosenst., Repert. Sp. Nov.
presents more than three costal areoles between Fedde 12: 473. 1913. Type. Bolivia: La Paz,
the margin and costa regardless of the width of near Coroico, Polo Polo, Oct.-Nov. 1912,
the leaf. Buchtien 3525 (holotype, S!).
Campyloneurum amphostenon is recognized Campyloneurum cooperi Lellinger, Amer. Fern J.
here as a complex of populations that may have 78: 19. 1988. Type. Costa Rica, Cartago,
been subject to recurrent isolation and Cartago, Cooper 6053 (holotype, US!).
reconnection through time. This complexity is Stem scales lanceolate (3-) 5-7 mm long, (1-)
shown in the wide range of variation in stem 1.5-2 mm wide; the cells oblong, arranged along
morphology (short or long creeping stems) and the main axes, cell lumen transparent. Figs. 9 e;
structure of the stem scale (cell wall width and 10 d; 15; 27; 35.
cell arrangement), especially in Central America
and the north-central Andes. In this treatment Widely distributed in tropical America, from
two varieties are recognized. Mexico and the Antilles to Argentina. It usually
grows above 1500 m elevation, in remnant
–– Most of stem scales with elongate cells forested areas or in shady rock crevices.
parallel to the main axis.
var. amphostenon REPRESENTATIVE SPECIMENS. MEXICO.
–– Most of stem scales with cells irregularly HIDALGO: Agua Blanca-Iturbide, 22 Jul. 1973, Gimate
ordered. var. irregulare 1065 (F). VERACRUZ: El Volcancillo, Las Vigas, 30
Oct. 1975, Dorantes et al. 5120 (BM, F, NY); road
Huayacocotla-Viborillas, 1 km from Huayacocotla, 14
4A. Campyloneurum amphostenon var. Jul. 1977, Fay & Calzada 898 (F, NY); 42 km W of
amphostenon. Escola, on road to Jalcal, 12 Jan. 1981, Nee & Schatz
Polypodium angustifolium var. amphostenon 19773 (F); crater of volcano Rafael Ramirez, 21 Jan.
(Kunze ex Klotzsch) Baker in C. Martius, Fl. 1976, Ortega et al. 132 (F); Mun. Chiconquiaco, Planta
Bras. 1(2): 530. 1870. el Pie, 23 Mar. 1972, Ventura 5123 (F, NY). CHIAPAS:
Campyloneurum angustifolium var. amphostenon Mun. San Cristóbal de las Cajas, 9 mi SE of San
(Kunze) Farwell, Amer. Midl. Naturalist 12: Cristobal, 20 Aug. 1966, Breedlove 15109 (F); Mun.
296. 1931. Tenejapa, Banabil, 7 Nov. 1971, Breedlove & Smith
22021 (F, MO); Mun. San Cristóbal de las Casas, S of
Polypodium leucorhizon Kunze ex Klotzsch,
Zonthuitz, 8 Nov. 1971, Breedlove & Smith 22066 (F);
Linnaea 20: 400. 1847. Cyted Syntypes. Mun. San Cristóbal de las Casas, 5 Dec. 1971, Breedlove
Venezuela: Moritz 83 (B!); Moritz 135 (n.v.); 22998 (F, MO); Cerro Huitepec, 20 Aug. 1972, Breedlove
Moritz 136b (B!, BM!). British Guiana: 27219 (F); Cerro Huitepec, 4 Dec. 1983, Cabrera &
Schomburgk 1145 (B!). Lectotype (chosen Cabrera 6000 (MO); above San Juan Chamula, 9 Jul.
here). Peru: Ruiz 10 (B!). 1977, Croat 40672 (AAU, MO); from Motozintla de
Campyloneurum leucorhizon (Kunze ex Klotzsch) Mendoza to Siltepec, 11 Feb. 1979, Croat 47280 (MO);
Fée, Gen. Fil. 258. 1852. Croat 47320 (MO); between Chiapa de Corzo and
Pichualco, 17 Feb. 1987, Croat & Hannon 65142 (MO); 5 Jun. 1942, Steyermark 46945 (F); 5-10 km W of Los
NE side of Cerro Huiytepec, 19 Apr. 1945, Little & Encuentros, Cerro María Tecum, 24 Dec. 1972, L.O.
Sharp 9885 (US); creek at Rincón Chamula, 12 km NW Williams et al. 41730 (AAU, F). CHIMALTENANGO:
of Pueblo Nuevo Solistahuacan, 23 Jan. 1965, Raven & Santa Elena, 10 Aug. 1932, Johnson s.n. (F); Santa Elena,
Breedlove 19785 (F); Mun. Tenejapa, Paraje Balum 12 May 1936, Johnston 420 (F); Santa Elena, cerro
k'anal, 13 Apr. 1966, Ton 814 (F, US). OAXACA: Tecpám, 4 Dec. 1938, Standley 58780 (F, MO); 26 Dec.
Cumbre de los Frailes, 11 Dec. 1907, Conzatti 2130 (F); 1938, Standley 61081 (F); slopes of Volcán de
Santa Ana, Cuicatlán, 23 Jun. 1909, Conzatti 2364 (F); Acatenango, above Las Calderas, 3 Jan. 1939, Standley
along Oaxaca-Tuxtepec road, beyond Cerro Pelón, 3 61951 (F); cerro Chichoy, 26-27 Jan. 1949, L.O. Williams
Jan. 1974, Carlson 4151 (F); between Oaxaca and & Molina 15403 (F). SACATEPEQUEZ: San Rafael,
Pochutla, 19 Jan. 1979, Croat 46052 (MO); vic. of Cerro Feb. 1892, Smith 2741 (GH, NY). GUATEMALA:
Zempoaltepetl, E slopes at Patio de Arena, 8 Aug. slopes of Volcán de Pacaya, 20 Dec. 1940, Standley
1950, Hallberg 845 (US); dist. Ixtlan, Sierra de Juárez, 80764 (F, UC). Volcán de Agua, 22 Mar. 1905, Maxon &
Llano Verde, ca. 15 km NE de Calpulalpán, 9 Apr. Hay 3742 (US).
1981, Lorence et al. 3267 (MO); dist. Central, N slope of HONDURAS. LEMPIRA: Montaña Celaque, 18-22
Cerro San Felipe, 13 Oct. 1969, Mickel & Hellwig 4038 Nov. 1974, Hazlett 2316 (F). San Pedro de Sula, 35 km
(UC); dist. Teotitlán, 26-29 km NE of Teotitlán del NE Nuevo Ocotepeque, 12 Jun. 1985, Martinez & Tellez
Camino, 16 Oct. 1969, Mickel & Hellwig 4127 (UC); 12954 (MO).
between Teotitlán del Camino and Huautla de Juárez, EL SALVADOR. SANTA ANA: Forest Montecristo,
8 Jul. 1972, Webster et al. 17274 (UC); ca. 24 km N of 15 Sep. 1977, Seiler 89 (F); forest Montecristo, 19 Sep.
San Gabriel Mixtepec, 18 Jul. 1985, Yatskievych et al. 85- 1977, Seiler 113 (F); forest Montecristo-Los Planes del
205 (MO). Miramundo, 6 Nov. 1977, Seiler 187 (F, NY); Cerro El
GUATEMALA. HUEHUETENANGO: between Pital, 10 Jun. 1978, Seiler 392 (F), Seiler 393 (F), Seiler
Paquix and Todos Santos, 25 Sep. 1944, Melhus & 395 (F); forest Montecristo, 10 Oct. 1978, Seiler 656 (F,
Goodman 3569 (F). QUICHE: 1942, Aguilar 1131; NY); forest Montecristo, 11 Oct. 1978, Seiler 669 (F, NY,
Aguilar 1165 (F). ALTA VERAPAZ: San Juan UC); Cerro El Pital, 16 Nov. 1978, Seiler 755 (F); forest
Chamelco, 12 Dic. 1973, Dary-Rivera 1713 (F). BAJA Montecristo, 31 Jan. 1979, Seiler 848 (MO); forest
VERAPAZ: San Rafael Chilascó, Salama, 10 Sep. 1973, Montecristo, 7 May 1979, Seiler 1124 (NY, UC).
Dary-Rivera 541 (F); 10 Oct. 1973, Dary-Rivera 970 (F); CHALATENANGO: E slope of Los Esesmiles, 27 Mar.
10 Nov. 1973, Dary-Rivera 1208 (F), Dary-Rivera 1228 1942, Tucker 1151 (UC).
(F). SAN MARCOS: 10 mi S of San Marcos, along road COSTA RICA. ALAJUELA: Varablanco de Sarapiquí,
from San Rafael, 13 Jul. 1977, Croat 41013 (MO), Croat N slope of Cordillera Central, Feb. 1938, Skutch 3539
41303 (MO); Volcán Tacaná, 6 Feb. 1987, Martinez et al. (NY, S). HEREDIA: Sacramento, Volcán Barba, 18
19557 (F); Martinez et al. 19576 (F), Martinez & Ramirez Dec. 1983, Givens 3372 (F). LIMON: Cordillera
19588 (F); Barranco Eminencia, San Marcos-San Rafael Talamanca, at cerro Bekom, 21-27 Mar. 1984, Davidse et
Pie de la Cuesta, 6 Feb. 1941, Standley 86545 (F); San al. 25728 (MO); Cordillera Talamanca, Atlantic slope,
Sebastián at km 21 and km 8, 15 Feb. 1940, Steyermark Valle de Silencio, along the Río Terbi, 9 Sep. 1984,
35658 (F); slopes of Cerro Tumbador, 15 Dec. 1962, Davidse et al. 28746 (MO); Cordillera between Río
L.O. Williams et al. 23046 (F); near Aldea Fraternidad, Terbi and Río Siní, 13 Sep. 1984, Davidse 29003 (MO);
between San Rafael Pie de la Cuesta and Palo Gordo, Kámuk massif, NE of the main Kámuk peak, 17-18
10-18 Dec. 1963, L.O. Williams et al. 25745 (F); Sierra Sep. 1984, Davidse & Herrera 29328 (MO).
Madre Mountains, 13 Dec. 1963, L.O. Williams 25875 PUNTARENAS: Cordillera Talamanca, upper slopes
(F); between San Rafael Pie de la Cuesta and Palo of Cerro Echandi, 23 Aug. 1983, Davidse et al. 23965
Gordo, 10-18 Dec. 1963, L.O. Williams et al. 26276 (F); (MO); slopes between cerro Echandi and cerro Burú,
on outer slope of Tujumulco volcano, ca. 10 km W of 24 Aug. 1983, Davidse et al. 24023 (UC). SAN JOSE:
San Marcos, 3 Jan. 1965, L.O. Williams et al. 27166 (F, trail from Canaán to Chirripó, N of Río Talari, 19-22
NY), L.O. Williams et al. 27206 (F). Jan. 1970, Burger & Liesner 7440 (F, GH, NY); from
QUEZALTENANGO: Quebrada El Pocito, S of San Canaán to Chirripó, Burger 8323 (F); SW slopes from
Martín Chile Verde, 27 Jan. 1941, Standley 85078 (F, Canaan to summit, 23 Aug. 1967, Lellinger et al. 92
UC), Standley 85108 (F); vic, of Fuentes Georginas, (MO); upper slopes of Cerro Daser (Azulillo), 5 km S
slopes of Volcán Zunil, 3 Feb. 1941, Standley 86032 (F, of Aserrí, 19 Mar. 1973, Stolze 1405 (F); along
UC); above los Bahos, Cerro Quemado, 5 Feb. 1941, Panamerican highway, Villa Mills, 15 Sep. 1961, Weber
Standley 86087 (F); Volcán Zunil, 22 Jan. 1940, 6239 (MO, US). CARTAGO: Cerro de la Muerte, near
Steyermark 34711 (F). TOTONICAPAN: Sierra Madre km 97 marker, 8-10 Aug. 1967, Evans & Bowers 3169
mountains, S of Totonicapán, 13 Dec. 1962, L.O. (MO); side of Turrialba volcano, 26 Jul. 1965, Lent 684
Williams et al. 22922 (F). SOLOLA: Volcán Santa Clara, (F, NY); Cerro de la Muerte, 24.5 km NW of La
Asunción, 11 Aug. 1967, Mickel 3330 (NY, US); 1 km N GH, US). CUNDINAMARCA: Cordillera Oriental,
of Carretera Interamericana, 10 km SE of Empalme, 17 Macizo de Bogotá, Quebrada de San Cristóbal, 4 Feb.
Mar. 1973, Stolze 1388 (F), Stolze 1390 (F); N of Irazú, 27 1940, Cuatrecasas 8019 (F); about 5 km SW of Bogotá,
Mar. 1928, Stork 1293 (UC); E of Irazú, 17 May 1928, on road to Usme, 4 Aug. 1950, G. Smith & Idrobo 1319
Stork 2027 (UC). SAN JOSE-CARTAGO: near El (MO). DEL VALLE: Cordillera Occidental, Hoya del
Trinidad and km 72, 25 May-19 Jun. 1968, Burger & Río Sanquininí, 10-20 Dec. 1943, Cuatrecasas 15428 (F);
Stolze 5244 (F); near La Asunción, 21 Nov. 1969, Burger Los Farallones, Cerro Alto del Buey, 11-12 Oct. 1944,
& Liesner 6302 (F), Burger & Liesner 6337 (F); SE of El Cuatrecasas 17969 (F, S, US); Río Palo, Quebrada de
Empalme, 27 Nov. 1969, Burger & Stolze 6501 (F); Santo Domingo, 13 Dec. 1944, Cuatrecasas 19266 (F).
upper slopes of Talamanca, W ridge of cerros Cuericí, VENEZUELA. ANZOATEGUI: dist. Libertad, NE of
15 Sep. 1983, Davidse 24661 (MO); Davidse 24721 (AAU, Bergantín, NE of Buenos Aires, Serranía de
UC); Parque Nacional Chirripó, 16 Sep. 1983, Davidse Turimiquire, 28 Nov. 1981, Davidse & González 19622
24813 (AAU, MO, UC); road Cartago-San Isidro del (MO). TERRITORIO FEDERAL AMAZONAS: Río
General, ca. 1 km NW of Asunción, 29 Jan. 1986, Smith Negro, Cerro de la Neblina, 5.1 km NE Pico Phelps, 2
& Beliz 2022 (MO). PUNTARENAS-LIMON: between Dec. 1984, Bell 396 (UC); Departamento Atabapo,
cerro Kasir and cerro Nai, 22 Mar. 1984, Davidse et al. Cerro Marahuaca, 26-27 Feb. 1985, Steyermark & Holst
25837 (MO). 130749 (MO), Steyermark & Holst 130817 (MO).
PANAMA. CHIRIQUI: Monte Azul, 1.4 mi N of Entre MERIDA: Páramo de La Negra, near Pregonero road
Ríos on E slopes of Cerro Punta, 22 Nov. 1979, Antonio junction, 21 Mar. 1947, Box 3750 (BM), Box 3751 (BM);
2738 (MO); Volcán Chiriquí, 7.3 mi from Boquete, 24 Páramo de Los Leones, La Lagunita, W de Mucurubá,
Jul. 1970, Armond 533 (F); Guadalupe, Cerro Punta, 6 31 May 1930, Gehriger 141 (F); dist. Campo Elías, S de
Mar. 1982, Caballero 55 (MO); dist. Boquete, W end of Pueblo Nuevo, entre Mucusus y Cerro La Becerrera, 18
Paso de Respingo, Cochrane et al. 6313 (MO); Las Sep. 1984, Ortega & Diaz 2142 (UC); dist. Campo Elías,
Cumbres, N of Quebrada Iglesia, near town Cerro La Carbonera, El Palmarito, 24 Nov. 1985, Ortega &
Punta, 22 Jul. 1971, Croat & Porter 16170 (AAU, MO); Werff 2939 (MO); dist. Libertador, Laguna Coromoto,
12 mi above Boquete on road to Volcán Baru, 18 May 22 Feb. 1971, Ruiz-Terán & López 1534 (UC); dist.
1976, Croat 34879 (MO); 7 km NW of Cerro Punta, Las Rangel, Quebrada Las Escaleras, ca. 10 km SE from
Nubes region, 11 Feb. 1978, Hammel 1395 (AAU, MO); San Rafael de Mucuchíes, 16-17 May 1972, Ruiz-Terán
lower N slope of Baru, E bajo Choro region, 7 May 7239 (UC). ZULIA: dist. Mara, Puesto Guardia
1978, Hammel 2995 (MO); W slopes of Cerro Respingo, Nacional, 10-15 Nov. 1982, Bunting et al. 12135 (UC).
Ne of Cerro Punta, ca. 15 Jun. 1971, Webster & Breckon TACHIRA: Quebrada La Lejía, S of Quebrada Agua
16578 (UC). BOCAS DEL TORO: Cordillera Azul, 25 Jul. 1979, Steyermark & Liesner 118580 (MO).
Talamanca, 7-8 Mar. 1984, Davidse et al. 25333 (MO); LARA: dist. Moran, trail from Humocaro to Buenos
between Itamut and Bine, Fabrega massiff, 5-9 Mar. Aires, 25 Jun. 1979, Liesner et al. 7974 (MO); dist.
1984, Gómez et al. 22547 (MO); 1-2 km SWW of Itamut Moran, Páramo Las Rosas, 5 Dec. 1984, Rivero 755
camp, 6-7 Mar. 1984, Gómez et al. 22615 (MO). Isla (UC). MONAGAS: Cerro Negro, above La Sabana de
Potrero, Changuinola valley, 6 Aug. 1923, Dunlap 71 las Piedras, NW of Caripe, 15 Apr. 1945, Steyermark
(F). 62070 (F, NY). SUCRE: Cerro Turumuquire, W of
CUBA. GUANTANAMO: Sierra de Imías, Cabezadas Boquerón, 8 May 1945, Steyermark 62671 (F, MO, US).
del Río Jojo, 20 Aug. 1975, Bisse & Meyer 27692 (HAJB); MIRANDA: Pico de Naiguatá, above Los Chorros, 16-
San Antonio del Sur, Puriales de Caujeri, Sierra del 17 Jun. 1945, Steyermark 62975 (F, GH). BOLIVAR:
Purial, 30 May 1982, Bisse et al. 47263 (HAJB); Cerro Roraima, S border Guyana, Brasil
Palenque, Sierra de Frijol, loma Bernardo, 21 May andVenezuela, Río Arabapó, 26 Aug.-2 Sep. 1976,
1983, Bisse et al. 50051 (HAJB). Steyermark et al. 112602 (NY, UC); dist. Piar, Macizo del
COLOMBIA. MAGDALENA: Sierra Nevada de Santa Chimantá, NE of Chimantá-tepui, 26-29 Jan. 1983,
Marta, 1 km NW Quebrada de Laguna Río Trío, Forero Steyermark et al. 128057 (F, MO). TRUJILLO: Páramo
& Kirkbride 628 (F, MO); Sierra Nevada de Santa de Guaracamal-Vega de Guaramacal, 23-24 Jul. 1984,
Marta, 3 Aug. 1972, Forero & Kirkbride 655 (MO), Forero Ortega et al. 2022 (MO); Páramo de Guaracamal, 22
& Kirbride 656 (MO); Serranía de Perijá, Cerro El Nov. 1984, Ortega & Werff 2274 (MO); dist. Boconó, ca.
Avión, 3 Mar. 1959, Romero-Castañeda 7364 (MO); 10 mi SW Batatal, 3 Nov. 1982, Smith et al. 921 (MO);
Santa Marta, 1898-1901, H.H. Smith 945 (BM, F, S, US). Páramo de Guaramacal, 3 Feb. 1987, Werff et al. 8805
NORTE DE SANTANDER: between Pamplona and (UC). Without locality: Vogl 186 (F) Colonia Tovar,
Chorro Colorado, 4 May 1983, Croat 56416 (MO); 1854-1855, Fendler 226 (MO).
Quebrada Samaria, drainage of Río Chitagá, 19 Nov. ECUADOR. CARCHI: El Angel-Tulcán, 14 May 1973,
1942, Fosberg 19200 (US); Pamplona, S of García, 18 Holm-Nielsen et al. 5235 (AAU, F, GB, MO); Valle de
Mar. 1945, Garganta 975 (F). Cipacoa, Lindig 241 (BM, Maldonado, km 71 on road Tulcán-Maldonado, 20
May 1973, Holm-Nielsen et al. 6092 (AAU, USM); Holm- Huamachuco-Cajabamba road, Sausacocha-
Nielsen et al. 6100 (AAU, USM); Páramo El Angel, 12 Cajabamba, 15 Feb. 1983, D.N. Smith & Vásquez 3378
Jan. 1980, Holm-Nielsen 20986 (AAU, USM). (F, UC); Pataz, 1985, Young 2933 (USM). SAN
IMBABURA: SW slopes of volcano Cotacachi, 7 Nov. MARTIN: Mariscal Cáceres, Parque Nacional Río
1983, Boysen-Larsen et al. 45583 (QCA); Cantón Ibarra, Abiseo, Puerta del Monte, patch of forest P2, 1985,
SW of Ibarra, 30 Jun. 1935, Mexia 7402 (F). Young 1685 (USM), Young 1686 (USM); Parque
PICHINCHA: Los Alpes, 19 Jan. 1944, Acosta-Solis Nacional Río Abiseo, forest patch P3, 1985, Young 1722
7094 (F); road Olmedo-Laguna San Marcos, 10 Jul. (USM); forest patch P11, Young 1994 (USM); forest
1980, Øllgaard et al. 34363 (AAU, USM), Øllgaard et al. patch C6, 25 Nov. 1985, Young 2153 (USM); forest
34438 (AAU, USM). NAPO: junction of Río Borja and patch P5, Young 2134 (USM); Parque Nacional Río
Río Quijos, 19 Sep. 1980, Holm-Nielsen et al. 26230 Abiseo, forest patch C10, 24 Nov. 1985, Young 2534 (F,
(QCA); 6.6 km W of Papallacta, 26 Mar. 1972, Mac USM); Chochos valley, Young 3691 (USM); NW corner
Bryde & Dwyer 1242 (QCA). CAÑAR: north rim of the of Río Abiseo Nat. Park, Chochos, 14 Jul. 1987, Young
valley of Río Cañar, 25 Apr. 1945, Camp 2883 (F). & León 4562 (USM); Parque Nacional Río Abiseo,
AZUAY: Gualaceo-Sigsig, 31 Ago. 1984, Jaramillo 7156 Laguna de Chochos, Jul. 1987, Young & León 4848
(QCA); Hacienda Tarqui, 20 Ago. 1987, Jaramillo 9998 (USM, HUT). ANCASH: Cordillera Blanca, Quebrada
(QCA); road Gima-Gualaquiza km 20.1, 28 Dec. 1990, Honda, small vally between Toqllarju and Pallkaraju,
Øllgaard et al. 98628 (QCA). MORONA-SANTIAGO: 8 Jul. 1979, Gibby & Barrett 173 (BM); Yungay,
trail Alao-Huamboya, 7 May 1982, Øllgaard et al. 38277 Llanganuco, 9 Ago. 1986, Mostacero et al. 1385 (F, MO,
(AAU, QCA). LOJA: muletrack Amaluza-Palanda, NY); Santa, Jalca Ultu Cruz (Jumbe), 3 May 1987,
western slope Cerro Amarillo, 22 Sep. 1976, Øllgaard & Mostacero et al. 1869 (F, NY); Corongo, Nueva Victoria,
Balslev 9610 (AAU, QCA, UC, USM); Podocarpus 7 May 1987, Mostacero et al. 2001 (F, UC); Huari, slopes
National Park, Nudo de Cajanuma, around Centro de valley of Laguna Ichicpotrero, 8 May 1986, D.N. Smith
Información, 25-26 May 1988, Øllgaard et al. 74487 et al. 12412 (F, MO); Huari, P. N. Huascarán, Quebrada
(QCA). GALAPAGOS ISLANDS: Santa Cruz Pachachaca, lateral valley of Quebrada Rurichinchay,
(Indefatigable), Cerro Colorado, 7 Feb. 1974, Adsersen 12 Jun. 1986, D.N. Smith et al. 12539 (MO).
& Adsersen 205 (QCA) HUANUCO: Mito, 8-18 Apr. 1923, Bryan 204 (F);
PERU. PIURA: Huancabamba, Cuello El Indio, 13 Cushi, trail to Tambo de Vaca, 19-23 Jun. 1923, Bryan
Nov. 1981, López et al. 8894 (GH, MO, UC). 620 (F); Tambo de Vaca, Bryan 646 (F); Mito, 23 Jul. 14
CAJAMARCA: Cajamarca-Celendín, 18 Oct. 1986, Aug. 1922, Macbride & Featherstone 1919 (US);
Díaz 2159 (MO, NY); Santa Cruz, upper Río Zaña Huacachi, near Muña, 20 May-1 Jun. 1923, Macbride
valley, 2-4 May 1987, Dillon et al. 4896 (F, UC); jalca 4127 (F); 32 km de Huánuco, Huánuco-La Unión, 25
Kumulca, road to Celendín, 17 Jun. 1975, Sagástegui et Jul. 1982, D.N. Smith 2191 (F, USM). PASCO:
al. 8117 (NY); San Miguel, Llapa-Uchuquinua, 14 May Oxapampa, Río San Alberto, 28 Jun. 1985, Foster et al.
1977, Sagástegui et al. 8907 (F, MO, UC); Contumazá, 10302 (F); Huariaca-Cerro de Pasco, 25 Jun. 1953,
Guzmango, Cerro Chungarrán, 24 May 1978, Ferreyra 9502 (GH); 95 km S from Huánuco, 15 Jul.
Sagástegui & Mostacero 9180 (F, MO, NY); Contumazá, 1982, Gentry et al. 37489 (F); trail to summit of
Lledén, 3 Nov. 1979, Sagástegui et al. 9404 (F, MO, NY); Cordillera Yanachaga, via Río San Daniel, 17 Jul. 1984,
Contumazá, around Pozo Kuan, 13 Jun. 1981, D.N. Smith et al. 7856a (USM). JUNIN: Tarma,
Sagástegui et al. 10068 (GH, UC); Quebrada Honda, Incatacuna, Tarmatambo-Acolla, 29 Jun. 1954,
Santiago-Yumal, 13 Jun. 1983, Sagástegui & López 10616 Constance & Tovar 2348 (UC); vicinity of La Oroya, 14
(MO); Celendín, Sendamal, 17 Aug. 1984, Sagástegui et Jul. 1914, Rose & Rose 18691 (US); Pampa Hermosa,
al. 12093 (MO, NY); Contumazá, 6 Apr. 1985, Satipo, 17 Apr. 1965, Saunders 1062 (GH), Saunders
Sagástegui 12639 (NY, UC); Cajamarca, Cerro Cumbe 1068 (GH); Tarma, Incatacuna, 29 Jun. 1954, Tovar 2351
Mayo, 9 Aug. 1969, Sanchez Vega & Ruiz 7351 (GH). (USM). LIMA: Huarochirí, Infiernillo, 17 Jan. 1949,
Chota, Chota-Tacabamba road, 20 Feb. 1983, D.N. Ferreyra 5294 (GH); between San Mateo and Casapalca,
Smith & R. Vásquez 3618 (MO). AMAZONAS: 8 Aug. 1949, Ferreyra 6236 (BM, USM); Huarochirí,
Chachapoyas, jalca de Calla Calla, 23 Oct. 1965, Viso, 22 Apr. 1939, Goodspeed et al. 11547 (GH, UC,
Sagástegui 6070 (GH, MO); Mendoza, 30 Jul. 1963, US); Lima-La Oroya road, 76 km W of La Oroya, 29
Woytkowski 8062 (MO); Chachapoyas, Puma-urcu, ESE Jan. 1983, Gentry et al. 39747a (MO); Río Blanco, 15-17
of Chachapoyas, 1 Jun. 1962, Wurdack 707 (F, GH, NY, Apr. 1929, Killip & Smith 21603 (NY, US); San Mateo,
US, USM); Caño Santa Lucía, E of Chachapoyas, 1/2 km before Río Blanco, 1973, Saunders 1161 (GH).
Wurdack 736 (US, USM); Bongará, hill WNW of HUANCAVELICA: Tayacaja, Huaribamba, 1km
Pomacocha, 19 Jun. 1962, Wurdack 948 (US); summit of before Huari, 28 Jul. 1968, Saunders 1147 (GH);
Cerro Campanario, 3 Aug. 1962, Wurdack 1574 (GH, Conaica, above Alauma, 22 Mar. 1952, Tovar 796 (GH,
NY, US). LA LIBERTAD: Sánchez Carrión, USM); Pachaspampa, below Huando, 3 Apr. 1953,
Tovar 1230 (USM); Tayacaja, above Tocas, 20 Apr.

1954, Tovar 2006 (GH); Tayacaja, Chuspi, near Tocas, Campyloneurum amphostenon var. amphostenon
22 Apr. 1954, Tovar 2039 (GH, USM); Tayacaja, Yacu- is characterized by its pruinose stem with
huanay, 16 Apr. 1962, Tovar 3680 (GH). AYACUCHO:
caducous brown, clathrate scales, and the cells
La Mar, E massif of the Cordillera Central, between
Tambo San Miguel, Ayna and the Hacienda Luisiana,
oblong arranged along the main axes of the scale.
24 Aug. 1968, Dudley 12029 (GH, US); La Mar, road Among the synonyms Polypodium leucorhizon
from Tambo to Ayna, above Jano, 3 Jan. 1975, Plowman is included. The protologue of this basionym is
& Davis 4676 (GH). CUSCO: Anta, El Chaccan, 3 Jan. based on five syntypes, four of them were seen at
1973, Brunel 236 (F, MO); Urubamba, trail from the herbarium in Berlin. Three of those
Chincheros to Antakillqa, 13 Jan. 1982, Davis et al. 1454 specimens (Ruiz 10, Moritz 83, Moritz 136b)
(F); Urubamba, Quebrada above Pojpoj waterfall, 14 clearly represent Campyloneurum amphostenon,
Jan. 1982, Davis et al. 1477 (F, NY, USM); Abra de and the Ruiz's specimen appears to be the one on
Málaga, 15 km Quillabamba, 9 Mar. 1971, Ellenberg
which the description was based. Only one
4777 (LPB); Cusco, Herrera 2591 (F); Tres Cruces, upper
edge of Parque Nacional de Manu, 29 Jun. 1978, Gentry specimen (Schomburgk 1145) appears to belong to
et al. 23450 (F); Urubamba, Chincheros, Titiqaqa the taxon here called Campyloneurum asplundii.
Ch'impa, 3 Feb. 1982, King et al. 128 (F); prov.
Paucartambo, km 130 road to Kosñipata, 30 Oct. 1987, 4B. Campyloneurum amphostenon var. irregulare
Nuñez 8500 (F, NY, UC); Machu Picchu, above (Lellinger) B. León, stat. nov. Fieldiana Bot.
Pauqarcancha, 3 May 1982, Peyton & Peyton 142 (MO); n.s. 1993: in press.
Machu Picchu, in Urcoscancha, above the village of Campyloneurum irregulare Lellinger, Amer. Fern J.
Palcay, 4 Jul. 1982, Peyton & Peyton 761 (GH, MO);
78: 24. 1988. Type. Ecuador: Pichincha,
Machu Picchu, Llactapampa, below Palcay, Acobamba
river, 7 Jul. 1982, Peyton & Peyton 810 (GH, MO);
Holdridge 1580 (holotype US!).
Machu Picchu, in the Pacasmayo river, 22 Aug. 1982, Polypodium crassifolium f. angustissimum Rosenst.
Peyton & Peyton 1084 (GH); Quillabamba, Santa Teresa, Mém. Soc. Sci. Nat. Neuchâtel 5: 45. 1912.
Mandonilloc, 5 Sep. 1982, Peyton & Peyton 1153 (GH, Type. Colombia: Cundinamarca, Sabana de
MO); Cusco, Soukup 159 (F, GH, US); Hacienda Urco, Bogotá, Mayor 40 (holotype not found;
18 Sep. 1939, Schmidt s.n. (F). isotypes S!, US!).
BOLIVIA: Camacho, Ambana, 19 Dec. 1980, Beck 4171 Stem scales ovate, 4-6 mm long, 2-2.5 mm
(LPB); Larecaja, Sorata, 16 Dec. 1981, Beck 4985 (LPB);
wide; cells oblong irregularly arranged. Figs. 6
Nor Yungas, Chuspipata, 5 km vía Unduavi, 2 Apr.
b; 17 a; 27.
1982, Beck 7634 (LPB); Nor Yungas, between Unduavi
and Chulumani, 25 Nov. 1980, Croat 51465 (MO);
Larecaja, 7 Mar. 1982, Fernández-Casas 6537 (MO); It is known from Mexico, Guatemala, Costa
Bautista Saavedra, Chullina, 30 Jan. 1979, Krach & Rica, Panama, and from Ecuador to Bolivia,
Feuerer 6573 (F, US); Inquisivi, between Pongo Chico where it grows above 2500 m elevation. It is
and Laguna Naranjani, 27 Jun. 1988, Lewis 88960 (F); usually found as a terrestrial in open habitats.
Sud Yungas, Yanacachi, 3 Jan. 1981, Liberman 241 (F);
Yungas, 1885, Rusby 350 (F, US); Nor Yungas, 22 km REPRESENTATIVE SPECIMENS: MEXICO.
NE Unduavi, on road to Yolosa, 29 Feb. 1980, Solomon VERACRUZ: Mun. Atzalan, vicinity Puente de Rieles,
5181 (UC); Murillo, 27.4 km N of dam at Lago Zongo, 4 km NE of Altotonga, 28 Jun. 1980, Nee & Hansen
27-28 Nov. 1982, Solomon 8970 (LPB, MO, NY, UC); 18701 (F).
Bautista Saavedra, Chajaya, near Charazani, 30 Mar. GUATEMALA. HUEHUETENANGO: between San
1985, Solomon 13345 (LPB); Sud Yungas, 1.4 km W of Mateo Ixtatán, at Cruz de Limón, 31 Jul. 1942,
Unduavi, between Chuspipata and La Paz, 2 Jul. 1986, Steyermark 49793 (F).
Solomon 15403 (MO); Murillo, Valle of Río Zongo, 18 COSTA RICA. SAN JOSE-CARTAGO: NW of La
Mar. 1987, Solomon 16384 (MO); Larecaja, Sorata, 7 Asunción, 5 Feb. 1982, Burger & Barringer 11506 (AAU,
Dec. 1981, Sperling & King 5387 (MO). F); ca. 15-20 km SE from El Empalme, 15 Jul. 1970,
COCHABAMBA: Ayopaya, above Independencia, 20 Lellinger & White 1177 (F).
Oct. 1986, Linke 12 (LPB); Yungas de San Mateo, Pojos, PANAMA. CHIRIQUI: Volcán de Chiriquí, 7.3 mi
25 Oct. 1928, Steinbach 8547 (MO). BENI: Ballivián, from Boquete, 24 Jul. 1970, Armond 533 (F).
Puente Río Quiquibey, 14 Jul. 1979, Beck 3899 (LPB). ECUADOR. CARCHI: Cunquer-Cuesaca, 17 Jul. 1945,
ARGENTINA. JUJUY: Santa Bárbara, Cerro Centinela, Acosta-Solís 10456 (F). PICHINCHA: around Quito, 28
11 Feb. 1964, Fabris et al. 5135 (UC, US). Mar. 1942, Paredes s.n. (F). COTOPAXI: Latacunga,
Humbles 6289 (MO); road Pilaló-Zumbagua, 10 km Jan. 1982, Davis et al. 1384 (F); Saxaihuaman, Dec. 1928,
above Pilaló, 28 Jul. 1980, Holm-Nielsen & Quintana Herrera 204 (F, GH, US); Saxaihuaman, Mar. 1929,
24650 (AAU, QCA). TUNGURAHUA: Pasa-San Herrera 2371 (F); Cusco, Herrera 2585 (F); Urubamba,
Fernando, 27 Oct. 1944, Acosta-Solís 8709 (F). Chincheros, 26 Jan. 1982, King et al. 112 (F); Huayoccari
CHIMBORAZO: Riobamba, Schimpff 831 (F). to Yanacocha, 14 Feb. 1987, Nuñez et al. 7035 (MO);
CAÑAR: Partidero El Corte-San Miguel de Porotos- Cusco, near Sacsaihuaman, 23 Sep. 1956, R. Tryon & A.
Parcialidad Jatumpamba, 26 May 1979, Jaramillo 936 Tryon 5345 (BM, F, GH, US, USM); Lauca, Espinar, 10
(AAU, QCA); between Cañar and Biblian, 29 Aug. Jan. 1957, Vargas 11506 (F). PUNO: Baja Isla in Lago
1984, Laegaard 52747 (QCA). LOJA: Parque Nacional Titicaca, 26 Nov. 1935, Mexia 7787 (BM, F, GB, GH,
Podocarpus, above Nudo de Cajanuma, 14-15 May MO, S, UC); Huancané, Moho, 20 Dec. 1919, Shepard 54
1988, Øllgaard et al. 74138 (QCA); Øllgaard et al. 74145 (GH); Granja Salcedo, near Puno, Oct. 1935, Soukup 10
(QCA). (F); Soukup 82 (F).
PERU. PIURA: Huancabamba, above Canchaque, 10 BOLIVIA. La Paz: Sud Yungas, Calacoto, 69 km E,
Oct. 1957, Hutchison 1646 (UC). CAJAMARCA: pasando nevado Illimani, 31 Dec. 1980, Beck 3899 (F);
Huancabamba, trail from Las Huaringas to Franz Tamayo, Pelechuco, 5 Mar. 1980, Krach & Feuerer
Huancabamba, 23 Feb. 1981, Davis & Turner 717 (GH); 9143a (F). COCHABAMBA: Chapare, Llantas-Aduana,
Celendín, canyon of the Río Marañón, above Balsas, 23 9 Mar. 1929, Steinbach 9557 (F, S, UC).
May 1964, Hutchison & Wright 5282 (F, GH, UC, USM);
Contumazá, Las Tres Cruces, Guzmango-Contumazá, Campyloneurum amphostenon var. irregulare is
7 May 1965, Sagástegui & Fukushima 5122 (GH); characterized by the irregular arrangement of the
Guzmango, 31 Jul. 1961, Sagástegui 3381 (GH);
cells in most of the stem scales. Some studied
Trinidad, Juque, 19 Jun. 1962, Sagástegui 3787 (GH);
Cajamarca, above La Encañada, Celendín, 18 May
material show stem scales with both regular and
1976, Sagástegui et al. 8394 (MO, NY, US); Contumazá, irregular arrangement of cells (e. g. Armond 533,
Las Quinuas-El Mojón, 14 Jun. 1981, Sagástegui et al. Herrera 204, Smith & Vasquez 3519 and Smith et al.
10102 (MO, UC); Cerro Cumbe Mayo, 13 Jun. 1966, 12243), for this reason recognition of this taxon
Sánchez Vega 36 (GH, US); Lluscapampa, 12 Sep. 1965, as a variety instead of a species is preferred.
Sánchez Vega 109 (GH); Hualgayoc, pass above
Hualgayoc, 17 Feb. 1983, D.N. Smith & R. Vásquez 3519
(F, MO, UC); Hualgayoc, 28 Jun. 1968, Soukup & 5. Campyloneurum anetioides (Christ) R. Tryon &
Carmona 5010 (US). LA LIBERTAD: Otuzco, Chilte,
A. F. Tryon, Rhodora 84: 125. 1982.
Hacienda Lalguen, 2 Jun. 1951, Lopez-Miranda 661 (BM,
US); near km 214 from Trujillo, between Huamachuco Polypodium anetioides Christ, Bull. Soc. Bot.
and Cajabamba, 27 Mar. 1960, Correll & Smith 916 Genève 2: 219. 1909. Type. Costa Rica:
(GH); about 3 km W of Huamachuco, Correl & Smith Alajuela, Candelaria, 4 Aug. 1908, A. C. Brade
937 (GH); Santiago de Chuco, Huacás, Cachicadán, 15 177 (holotype S!; isotypes BM!, K!, S!, US!).
Jun. 1984, Sagástegui et al. 11917 (MO, NY); Santiago de Hyalotricha anetioides (Christ) Copeland, Amer.
Chuco, Cahicadán, 25 Nov. 1973, Stork & Horton 9969 Fern J. 42: 12. 1953.
(F, UC, US). ANCASH: Carhuaz, Parque Nacional Hyalotrichopteris anetioides (Christ) W. Wagner,
Huascarán, Quebrada Ishinca, 13 Feb. 1985, D.N. Smith
Taxon 27: 548. 1978.
et al. 9516 (F); Yungay, Parque Nacional Huascarán,
Quebrada Ranincuray, 18 Apr. 1985, D.N. Smith et al.
Stem creeping, not pruinose, 1-2 mm wide.
10407 (F, MO); Huari, P. N. Huascarán, 6 May 1986, Stem scales brownish in mass, 3-6 mm long, 1.5-2
D.N. Smith et al. 12243 (F). LIMA: Río Blanco, 8-19 mm wide, narrowly ovate, clathrate, the cells
Mar. 1922, Macbride & Featherstone 716 (F, S, US). oblong, cell walls 6-8 µm thick. Phyllopodia
JUNIN: near turn off to Huasahuasi, 17 Mar. 1960, rarely present, 1-5 mm apart. Leaves 3-10 cm
Correll & Smith 779 (GH, US); Huancayo, Mar. 1947, long; petiole 0.5-1.5 cm long, usually not
Soukup 3143 (BM, F, GH, US); Quebrada Ocopilla, Feb. articulate, greenish-stramineous; lamina simple,
1948, Soukup 3646 (BM, F, US); Acopalca, 16 Jan. 1969,
entire, spathulate, bases decurrent, apices obtuse,
Soukup 6099 (US); Palián, 2 May 1961, Tovar 3351 (GH).
0.6-2 cm wide, herbaceous, repand margins,
APURIMAC: Quebrada of Juccuchic-chupan, on trail
Andahuaylas-Chincheros, 3 Nov. 1935, West 3723 indument of multicellular furcate hairs, 1.5 mm
(UC). HUANCAVELICA: North of Mejorada, km 362 long, the basal branch usually unicellular, short,
on the Carretera Central, 28 Oct. 1957, Hutchison 1673 sometimes apical and basal branches
(UC). CUSCO: Cusco, Saxaihuaman, 28 Feb. 1954, multicellular; stomata polocytic or copolocytic;
Coronado 156 (GH, UC); Urubamba, Maranqaqa, 12 costa inconspicuous on the adaxial side, slightly
prominulous abaxially, slightly flexuous, angustifolium and C. phyllitidis (Mitsuda 1981,

primary and secondary veins inconspicuous, 3 1983).
areoles between the costa and margin, tertiary The closest affinities of Campyloneurum
excurrent veinlet one per areole. Sori anetioides are with C. aphanophlebium, and they
subterminal or terminal, paraphyses present, both in turn might be related to the open areolate
filamentosous, spores slightly verrucate, 80-100 species.
µm de long, 50-60 µm wide. 2n= 148. Figs. 13 c;
14 i; 28.
6. Campyloneurum angustifolium (Sw.) Fée, Gen.
Central American species known from Fil. 257. 1852.
Nicaragua, Costa Rica and Panama, between Polypodium angustifolium Sw., Prodr. Veg. Ind.
1000 m and 1450 m elevation. It grows with Occ. 130. 1788. Type Jamaica, Swartz s.n.
hanging leaves among mosses on rocks, in shady (holotype not found, isotypes BM!, LD!, S!).
and humid sites in tropical forest. Marginaria angustifolia (Sw.) C. Presl, Tent.
Pterid. 188. 1836.
EXAMINED SPECIMENS: NICARAGUA. Grammitis angustifolia (Sw.) Heward, Mag. Nat.
MATAGALPA: 6-10 km NE of Matagalpa, road to El Hist. 458. 1838.
Tuma, 14-16 Jan. 1963, Williams et al. 23835 (F). Cyrtophlebium angustifolium (Sw.) J. Sm. Bot.
COSTA RICA. CARTAGO: Tapantí, Río Grande de
Mag. 72: 12. 1846.
Orosí, 19 Jan. 1964, Jimenez 1601 (F); Orosí, Dec. 1923,
Lankester 702 (BM). SAN JOSE: vicinity of El General,
Goniophlebium angustifolium (Sw.) Brack. in
Jul. 1936, Skutch 2753 (GH, K, MO, S). Without Wilkes, U.S. Explor. Exped. 16: 33. 1854.
locality: Apr. 1987, Stevens 15350 (MO). Polypodium taeniosum Willd. Sp. 5. 155. 1810.
PANAMA. CHIRIQUI: valley of Río Caldera, from El Type. Venezuela: Caripe, Humboldt (Herb.
Boquete to the Cordillera, 5-19 Feb. 1918, Killip 5013 Willdenow 19631) B!; photos BM!, USM!.
(BM); Chiriquí Viejo, vicinity of Monte Lirio, Seibert Campyloneurum taeniosum (Willd.) Fée, Gen. Fil.
310 (K). 257. 1852.
Cyrtophlebium difforme Loddiges, Cat. Pl. 1849.
Campyloneurum anetioides is characterized by Type. Cultivated. Nom. nudum
its spathulate leaves with inconspicuous Polypodium difforme (Loddiges) Kunze, Linnaea
venation and multicellular hairs. 23: 69. 1850.
This species was considered by Lellinger Campyloneurum difforme T. Moore, Index Fil. 224.
(1988) within the "satellite genus" 1861. Nom. nov. for Polypodium difforme
Hyalotrichopteris based on size, habit, and (Loddiges) Kunze. Non Polypodium difforme
indument features, which he considered rare in Blume 1828.
Campyloneurum. However, leaf size is not an Polypodium angustifolium var. gramineum Sodiro,
important taxonomic character for these species. Crypt. Vasc. Quit. 366. 1893. Type. Ecuador,
Small leaves, as in C. anetioides, also occur in C. Napo, bosque de Los Colorados, Río Zuma,
falcoideum and C. chrysopodum. The hairs of C. 1892, Sodiro s.n. (holotype not found, photo of
anetioides are branched and multicellular. They P, BM!).
are thought to be derived from the hair type Stem creeping, usually pruinose, (2-) 3-4 (-5)
found in C. aphanophlebium and C. repens (see mm wide. Stem scales dark brown or brown in
Chapter IV). The venation of C. anetioides is the mass, spreading, 3-6 mm long, 0.8-1 (-1.5) mm
typical Campyloneurum venation. It is similar to wide, narrowly ovate, scale bases auriculate,
that found in C. aphanophlebium and C. falcoideum, apices long acuminate, clathrate, the cells
where non-costal areoles bears one excurrent free narrowly oblong, regularly arranged, cell walls
veinlet. In addition, the presence of excurrent 10 µm thick, cell lumen transparent. Phyllopodia
free veinlets along the margin in C. anetioides 1-2 mm long, 1.5-2 mm wide, 1-4 mm apart.
occurs in mature plants of other species, such as Leaves 30-100 cm long; petiole stramineous or
C. brevifolium and C. falcoideum, and it is also dark stramineous, 0.5-5 (-7) cm long, slightly
present in juvenile leaves of such species as C. ranurate adaxially, convex abaxially, glabrate;
laminae linear or narrowly lanceolate, bases and hwy. 140, 26.5 km NW of Jalapa, 1.7 km of La Jolla, 19
apices attenuate, (0.5-) 1-2 (-3) cm wide, Jul. 1978, Caughlan et al. 320 (F, NY); near Fortín, 27
herbaceous-chartaceous, lamina margins Jun. 1977, Croat 39459 (MO); 3 km WNW of Cuichapa
on road to Coetzala, 3 Jul. 1982, Diggs t al. 2720 (F);
cartilaginous, plane or slightly revolute, slightly
road Plan de Arroyos-Alvaro Obregón, Hidalgotitlán,
sinuate, indument of inconspicuous hairs, 13 Apr. 1974, Dorantes et al. 2766 (F); road Cedillo-Río
scattered abaxially, stomata polocytic; costa Alegre, 18 Jan. 1975, Dorantes et al. 3916 (F); Agaltepec
prominent, sometimes with scales similar to Island, lake Catemaco, 5 Aug. 1976, Faden et al. 134 (F);
those on the stem, primary veins inconspicuos, 1- Cuahutlalpan, Ixtaczoquitlán, Jul. 1976, Faden et al. 168
2 (-3) areoles between the costa and margin, (AAU, F, US); road Huayacocotla-Viborillas, 14 Jul.
marginal areoles usually smaller than costal 1977, Fay & Calzada 898 (F); Orizaba, 9 Aug. 1924,
ones, excurrent veinlet one per areole. Sori Fischer s.n. (F, MO); Mun. Huatusco, Puente Adolfo
medial, paraphyses inconspicuous, scarce, Ruiz Cortines, 7 km NE of Huatusco, 28 Mar. 1979,
García & Delgado 950 (F, MO, NY); near Catemaco, 22
simple, shorter than the sporangia; spores (40-)
Jan. 1972, Hernandez 1386 (F); Montepio, Estación
50-70 Biológica Los Tuxtlas, 29 Jul. 1976, Kennedey & Horvitz
(-80) µm long, 40-45 µm wide. Chromosome 3676 (F); Cordoba, 5 mi E of Cordoba, 26 Oct. 1957,
number 2n=74. Figs. 9 b; 10 a; 12 a, b; 14 c; 16; Knobloch 710 (F); San Andrés Tuxtla, between
30. Montepio and Sontecomapan, 12 Jun. 1981, Lorence
3477 (MO); 2 km NE of La Joya, Acajete, 25 Nov. 1975,
Marquez et al. 447 (F); La Joya, 28 Sep. 1940, Moore 59
This species grows from southern U.S.A. to (MO); 10 km N of Altotonga, on road to Tlapacoyan,
28 Jun. 1980, Nee & Hansen 18668 (F); 7 km NW of
Bolivia, central Brazil and the Antilles; it is found
Coscomatepec, on road to Escola, 12 Jan. 1981, Nee &
between sea level and 2400 m elevation, mostly Schatz 19828 (F); Mun. Pajapan, 3 Nov. 1981, Nee &
as an epiphyte. Calzada 22768 (F); Mun. Chocaman, 8.5 km W of
Chocomán, on road to Xocotla, 18 Nov. 1981, Nee
REPRESENTATIVE SPECIMENS: USA. FLORIDA: 23226 (F); Sanborn, 10 Mar. 1910, Orcutt 2994 (BM,
near Brown's, 10 Dec. 1903, Eaton s.n. (F). MO); 1 mi W of Fortín, 4 Aug. 1947, Paxson et al.
MEXICO. SAN LUIS POTOSI: km 338 on route 85, 25 17M680 (F); Córdoba, May 1927, Reko 5129 (F); Region
Mar. 1961, Hewitson 67 (MU); barranca Las Canoas, 8 de los Tuxtlas, Salto de Eyipantla, 16 Mar. 1974, Riba &
Aug. 1891, Pringle 3821 (MU); Jamazunchale, 2 Aug. Perez 836 (MO); Mun. Xalapa, Salto del Gato, 29 Feb.
1937, Taylor 923 (F). COLIMA: rancho El Jabali, NNW 1976, Ronzon 6 (F); road from Catemaco to
of Colima, in the SW foothills of the volcán de Colima, Sontecomapan, around lake Catemaco, 14 Jan. 1981,
28 Oct. 1990, Lott et al. 2953 (F). QUERETARO: 74 mi Schatz & Nee 227 (F); Jalapa, 21 Dec. 1894, C. Smith 2001
NE of Zimapan, 22 Aug. 1957, Waterfall & Wallis 14271 (F, MO, UC); Veracruz-Coattacoalcos highway, Cerro
(F). HIDALGO: Mun. Xochicoatlán, 3 km S de Cimtepec, 13 Jul. 1974, Sohmer 9465 (F); 3 km N of
Jalamelco, 21 Nov. 1982, Acosta & Barrios 306 (MO); Catmaco, 15 Jun. 1982, Solheim & Powers 848 (F); Mun.
Jacala, 15 Nov. 1937, Kenoyer 650 (F); Mun. Tenango de Huayacocotla, 17 km NNE of Huayacocotla, NE of
Doria, 3 km SW de Tenango de Doria, 17 Mar. 1979, Agua de Calabaza, 27 Jan. 1984, Taylor & Nee 256 (F);
Koch 798 (F, NY); below Trinidad Iron Works, 1904, Mun. Altotonga, Nahualaco, 24 Nov. 1969, Ventura 110
Pringle 8972 (F, MO); 12 mi SW of Chapulhuacán, 4 (F, MO); Mun. Acatlán, El Balneario, 14 Nov. 1981,
May 1983, Yatskievych & Wollenweber 83-125 (F). Ventura 19134 (MO); Mun. Jesús Carranza, 14 Jul. 1984,
MICHOACAN: Montes de Oca, Pasión, 7 Oct. 1937, Wendt & Villalobos 4454 (MO). OAXACA: Teotitlán del
Hinton 10775 (F, NY). PUEBLA: Huauchinango, 9 Feb. Camino-Chilchotla, 23 Feb. 1979, Croat 48406 (MO);
1932, Fröderström & Hulten 704 (S); deroute to near Arroyo Sangre, ca. 2 km E of Santa María,
Atotocoyan, Teteles-Mazatepec, 24 Mar. 1976, Marquez Hernández 369 (MO); dist. Ixtlán, 3 km E of Ixtlán, 24
et al. 718 (F). VERACRUZ: Veracruz, 2 Aug. 1979, Jul. 1971, Mickel 5542 (UC); Mun. Valle Nacional, 21
Avendaño & Calzada 429 (F); Mun. Teocelo, La Cuetería, km N of Vista Hermosa, SW of Vista Hermosa, 15
2 Apr. 1980, Avendaño 703 (F); Jalapa, 13 Sep. 1906, Dec. 1985, Nee 32158 (UC); dist. Tuxtepec, Mun. Valle
Barnes et al. 93 (F), Barnes et al. 94 (F); 19 Sep. 1906, Nacional, 4 km SW of Valle Nacional, 15 Dec. 1985,
Barnes et al 95 (F); along the Mexican railway, above Nee 32158 (UC). CHIAPAS: boundary between
Fortín, 15 Nov. 1908, Barnes & Land 644 (F); Pedregal Zinacantán and Chamula, 20 Jan. 1965, Breedlove &
Las Vigas, 22 km NW of Jalapa, 3 Jan. 1982, Bohs et al. Raven 8129 (F); Mun. La Trinitaria, along the Comitán
1770 (F); hill de la Cima, La Sombra-Tierra Blanca, 10 river, Lagos de Montebello, NE of La Trinitaria, 13
Apr. 1981, Castillo & Vazquez 1534 (F); along Mexican Nov. 1971, Breedlove & Smith 22360 (F, NY); Mun.
Ocozocoautla de Espinosa, 26-28 km N of 69774 (F); below Patal, 4 Apr. 1941, Standley 91156 (F,
Ocozocoautla, 15 Nov. 1971, Breedlove & Smith 22445 UC), Standley 91244 (F); Baja Verapaz, 7 Feb. 1969, L.O.
(F); Mun. Cintalapa, 4 km W of La Cienaga, 10 May Williams et al. 40678 (F). IZABAL: along Río Frío and
1972, Breedlove 25134 (F); Mun. Tenejapa, Paraje of tributaries, 18 Dec. 1941, Steyermark 39992 (F). SAN
Mahosik, 5 Jun. 1972, Breedlove 25526 (F, NY); Mun. MARCOS: Volcán Tajumulco, NW of San Marcos, 15
Ocosingo, 10 km SW of Ocosingo , road to San Feb. 1940, Steyermark 35678 (F); between San Rafael Pie
Cristóbal, 23 Sep. 1972, Breedlove 27855 (F); Mun. San de la Cuesta and Palo Gordo, 10-18 Dec. 1963, L.O.
Andres Larrainzar, summit of Chuchil Ton, NE of Williams et al. 25745 (F); Sierra Madre Mountains, N of
Bochil, 17 Oct. 1972, Breedlove 29245 (F); Mun. San San Marcos, 13 Dec. 1963, L.O. Williams et al. 25867 (F,
Cristobal de las Casas, 17.5 km SE of San Cristóbal, 14 NY). QUEZALTENANGO: 17 Feb. 1939, Standley
Jan. 1973, Breedlove & Smith 31511 (F); Ocozocoautla, 65432 (F); Quezaltenango, 11 Mar. 1939, Standley 68419
above 2 mi N Roblada, 23 Mar. 1949, Carlson 1558 (F); (F); above Los Vahos, Cerro Quemado, 5 Feb. 1941,
between Tapachula and Unión Juárez, 1-3 mi N of Standley 86087 (UC); along old road between Finca
Trinitaria, 10 Feb. 1979, Croat 47205 (MO); Honduras, Pirineos and Patzulín, 9 Feb. 1941, Standley 86975 (F,
ca. Siltepec, 9 Jul. 1941, Matuda 4371 (MO); Mun. UC), Standley 87028 (F). TOTONICAPAN:
Ocozocoautla, Reserva Ecológica El Ocote, 17 Feb. Totonicapán, 8 Dec. 1962, L.O. Williams et al. 22582 (F);
1986, Palacios-Ríos 2884 (UC). Without locality: Sierra Madre Mountains, S of Totonicapán, 13 Dec.
Temascaltepec, Cucha, 19 Nov. 1934, Hinton 6833 1962, L.O. Williams 22922 (F). SOLOLA: Sierra Madre
(MO); Orizaba, 1905, Lemmon 325 (UC); Ruina mountains, near Nahuala, 17 Dec. 1962, L.O. Williams
Palenque, 10-12 Jul. 1939, Matuda 3705 (F); 10 Mar. et al. 23173 (F). SACATEPEQUEZ: 2.3 mi SW
1910, Orcutt 2994 (BM, MO); Oxtacihuat, Feb. 1905, Alotenango, from Antigua to Escuintla, 26 Jul. 1977,
Purpus 1841 (F); Orizaba, 17 Feb. 1982, J. G. Smith 78 Croat 41956 (MO); above Barranco Hondo, 11 Mar.
(MO); Orizaba, 29 Mar. 1890, Stone 104 (MO); Stone 107 1941, Standley 88933 (F). EL PROGRESO: hills SE of
(MO). Finca Piamonte, 4 Feb. 1942, Steyermark 43402 (F).
GUATEMALA. PETEN: La Cumbre, km 139-139 JALAPA: Jalapa, 1 Dec. 1939, Steyermark 32355 (F).
Cadenas road, 27 Jul. 1969, Contreras 8861 (F, MO). ZACAPA: Sierra Las Minas, between Río Hondo and
HUEHUETENANGO: Huehuetenango, 25 Sep. 1944, waterfall, 10 Oct. 1939, Steyermark 29428 (F), Steyermark
Melhus & Goodman 3569 (F); vic. of Maxbal, ca. 17 mi N 29429 (F). RETALHULEU: Retalhuleu, along Río
of Barillas, Sierra de los Cuchumatanes, 15-16 Jul. Coyote, W of Retalhuleu, 24 Feb. 1941, Standley 88398
1942, Steyermark 48763 (F); Huehuetenango, 22 Jul. (F, UC). ESCUINTLA: between Río Jute and Río
1942, Steyermark 49153 (F); above La Libertad, on Cerro Pantaleón, on road between Escuintla and Santa Lucía,
Pueblo Viejo, 20 Aug. 1942, Steyermark 51004 (F). 24 Jan. 1939, Standley 13479 (F). SANTA ROSA: Santa
QUICHE: W of Chichicastenango, 12-23 Jan. 1966, Rosa, near Cuilapilla, 23 Nov. 1940, Standley 78056 (F,
Molina et al. 16309 (F). ALTA VERAPAZ: between San UC).
Pedro Carcha and Campur, 20 Mar. 1970, Harmon & EL SALVADOR: Volcán San Vicente, Dec. 1930,
Fuentes 2166 (MO); Alta Verapaz, 10 May 1963, Molina Iglesias 9 (F). CHAL: Cerro El Pital, 10 Jun. 1978, Seiler
& Molina 12019 (F); Chamal, 13 May 1963, Molina & 394 (F). AHUACHAPAN: vic. of Ahuachapán, 9-27
Molina 12152 (F); Alta Verapaz, 26 Mar.-15 Apr. 1939, Jan. 1922, Standley 19932 (F, NY, US).
Standley 69268 (F), Standley 69431 (F); region of Chicoj, HONDURAS. Olancho, Los Zapotes, 8 Oct. 1979,
NE of Carchá, 2 Apr. 1939, Standley 70059 (F); Alta Andino 86 (MO). FRANCISCO MORAZAN: Valle de
Verapaz, 5 Apr. 1939, Standley 70744 (F), Standley 70940 Angeles, 15 km NE of Tegucigalpa, 17 Sep. 1983,
(F); Alta Verapaz, 26-27 Mar. 1941, Standley 89854 (F); Clotter 71 (UC); Quebrada de Zambrano, Zambrano-La
along Río Carchá, between Cobán and San Pedro Pirámide, 26 Jun. 1964, Molina 14249 (F); road Cerro
Carchá, 26-27 Mar. 1941, Standley 89906 (F); near Uyuca, 8 km of El Zamorano, 10 Sep. 1982, Montoya 64
Tactic, above bridge Río Frío, 30 Mar. 1941, Standley (MO); 14 km NE Tegucigalpa, 6 Aug. 1983, Morales 34
90488 (F); Alta Verapaz, 10 Apr. 1941, Standley 92030a (MO); Cerro Uyuca, 15 km SE of Tegucigalpa, 30 Oct.
(F); Alta Verapaz, 27 Jan. 1969, L.O. Williams et al. 1987; Moreno 185 (F); Cerro El Picacho, 13 Nov. 1982,
40193 (F); Alta Verapaz, 4 Jan. 1973, L.O. Williams et al. Ochoa 64 (MO); 20 km NE of Tegucigalpa, 14 Nov.
42022 (F); N of Cobán, 4 Jan. 1973, L.O. Williams et al. 1982, Padilla 76 (MO); along and near Río Agua
42111 (F); 6 km NE of Cobán, 3 Jan. 1974, L.O. Williams Amarilla, above El Zamorano, Oct.-Nov. 1948, Standley
et al. 42225 (F, GH); near San Cristóbal Verapaz, 6 Jan. 13956 (F); between Peña Blanca and Ponce, 5 Feb. 1950,
1973, L.O. Willliams et al. 42207 (AAU, F); 21 Jan. 1974, L.O. Williams & Molina 17143 (F). COMAYAGUA: Río
L.O. Williams et al. 43627 (F); along Río Cobán, 4 km E San Marcos, Intibucá, 24 May 1970, Barkley &
of Cobán, 21 Jan. 1974, L.O. Williams et al. 43627 (F); Hernández 40420 (MO); Montaña Comayagua, 29 Jan.
cerro Sielab, 23 Feb. 1939, Wilson 244 (F). BAJA 1975, Hazlett 2448 (F); Intibucá, between Siguatepeque
VERAPAZ: N of Santa Rosa, 30 Mar. 1939, Standley and Jesús de Otoro, 2 Nov. 1974, Horwath 135 (F);
Coyocute, 23 May 1956, Molina 7151 (US); Intibucá, La 13 Aug. 1977, Stevens 3227 (MO).
Esperanza, 12 Sep. 1981, Segovia 118 (MO); COSTA RICA. GUANACASTE: Parque Nacional
Siguatepeque, 5 Apr. 1947, Standley & Chacón 6706 (F); Rincón de Vieja, SE slopes Volcán Santa María, 27-28
vic. Siguatepeque, 14-27 Feb. 1928, Standley 56335 (F); Jan. 1983, Davidse et al. 23326 (MO); Parque Rincón de
vic. Siguatepeque, Jun.-Aug. 1936, Yuncker et al. 5607 la Vieja, Herrera 662 (F), Herrera 728 (MO); 4.5 km W
(MO), Yuncker et al. 5650 (F, MO); vic. Siguatepeque, 8 of Tilarán, 26 Jul. 1967, Mickel 2905 (NY, UC); Rincón
Jul. 1936, Yuncker et al. 5728 (F, S) . EL PARAISO: de la Vieja National Park, slopes of Volcán Santa
Güinope, El Paraíso, Dec. 1943, Valerio 1868 (F); El María, 26 Jan. 1986, A.R. Smith et al. 1963 (UC).
Paraíso, 28 Dec. 1962, Molina & Williams 11199 (F). ALAJUELA: Fila volcán Muerte, above headwaters
SANTA BARBARA: Trinidad, 25 Jun. 1982, Salguero 33 Río San Lorenzo, 15-17 Apr. 1982, Barringer & Gomez-
(MO). CORTES: Puerto Cortés, 23 Aug. 1984, Gómez Laurito 2545 (F); La Palma de San Ramón, 2 Nov. 1924,
49 (F); Montaña de la Nieve, caserío El Tapiquilar, 20 Brenes 4133 (F); San Pedro-San Ramón, 22 Jun. 1926,
km S San Antonio Cortes, 23 Feb. 1982, Nelson et al. Brenes 4880 (F); Santiago de San Ramón, El Piñón, 5
7897 (MO). Without locality: 1852, Hjalmarson s.n. (S). Dec. 1928, Brenes 6459 (F); La Balsa, 4 Mar. 1983,
NICARAGUA. MATAGALPA: between Jinotega and Carvajal 476 (MO); 14 km NE Ciudad Quesada, San
Matgalapa, 1 Apr. 1964, Bunting & Licht 1004 (F); Finca Carlos, 22 Jul. 1963, Jiménez 950 (F); La Marina-Aguas
Tepeuac, 18 km N de Matagalpa, 25 Feb. 1982, Castro Zarcas, 24 Jun. 1966, Jiménez 4084 (F); S of San Ramón,
2488 (UC); Santa María de Ostuma, N of Matagalpa, 25 ca. 3 km above San Rafael, 26 Jul. 1970, Lellinger &
Aug. 1976, Hall & Bockus 7897 (BM, NY); Santa María White 1350 (F); above Angel falls, Río La Paz Grande,
de Ostuma, Cordillera Central de Nicaragua, 1960- 23 Mar. 1969, Lent 1501 (F); above Río Toro, 3 Sep.
1961, Heller 6 (F); Matagalpa, 1963, Heller s.n. (F); 1972, Lent 2813 (F); lower slopes of Volcán Arenal, 17
Fuente Pura, 26 Aug. 1982, Moreno 17001 (MO); Cerro Sep. 1972, Lent 2945 (F); Cordillera Central, about 2 km
Matapalo, 9 km of Matagalpa, 23 Feb. 1983, Moreno & E of Zarcero, 15 Feb. 1966, Molina et al. 17098 (F, NY);
Robleto 20498 (MO); Matagalpa, along route 5 toward Zarcero, Jun. 1983, A. Smith H25 (F); region of Zarcero,
TUMA, 28 Feb. 1971, Seymour 4037 (MO); Matagalpa, 16 Jan. 1938, A. Smith H134 (F); region of Zarcero, 9
8-15 Jan. 1963, L.O. Williams et al. 23313 (F), L.O. Feb. 1938, A. Smith 301 (F); region of Zarcero, 13 Mar.
Williams et al. 23516 (F), L.O. Williams et al. 23596 (F); 1938, A. Smith 305 (F); Alajuela, 10 Jan. 1939, A. Smith
road to El Tuma, NE of Matagalpa, 14-16 Jan. 1963, 1429 (F); slopes of Volcán Poas, 1 May 1949, L.O.
L.O. Williams et al. 24053 (F, NY, US); Matagalpa, 19-21 Williams 16591 (F); N of Zarcero, ca. Tapesco river, 6
Feb. 1963, L.O. Williams et al. 24790 (F), L.O. Williams et Feb. 1965, L.O. Williams et al. 28926 (F). HEREDIA:
al. 24794 (F); near Santa María de Osuma, 20, 24 Feb. Sarapiquí Cantón, between Cariblanco and San
1963, L.O. Williams et al.25055 (F); Matagalpa, 15 Jan. Miguel, 12 Jul. 1983, Barringer et al. 3738 (F); El Gallito,
1965, L.O. Williams et al. 27659 (F); Matagalpa, 16 Jan. 19 Dec. 1933, Brenes 21711 (F); between Bajo La
1965, L.O. Williams et al. 27715 (F); Matagalpa, 18 Jan. Hondura and Alto La Palma, 19 Jul. 1983, Barringer et
1965, L.O. Williams et al. 27911 (F); Hacienda Santa al. 4002 (F); Río Puerto Viejo, near 2 km confluence Río
María de Ostuma, 11 Feb. 1965, L.O. Williams et al. Sarapiquí, 14-17 Jun. 1968, Burger & Stolze 5810 (F);
29151 (F); Cordillera Central de Nicaragua, near near Porrosati on the southern slopes of volcán Barba,
Xelaju, 12 Feb. 1965, L.O. Williams et al. 29243 (F); 22 Jun. 1968, Burger & Stolze 6009 (F), Burger & Stolze
Finca Santa María de Ostuma, 30 Nov-4 Dec. 1973, 6027 (F); Volcán Barba, 13 Jul. 1967, Evans & Bowers
L.O. Williams & Molina 42615 (F). JINOTEGA: Laguna 2698 (MO); forest of Río Vueltas, 23 May 1969, Gómez
Miraflores, ca. 26.1 km NE of hwy 1 at Estelí, 10-11 2287 (F); Monte de la Cruz, 19 Mar. 1963, Jimenez 474
Jun. 1981, Henrich & Stevens 339 (MO); Jinotega, 23 Jun. (F); between San Miguel and Cariblanca, 11 Jul. 1983,
1947, Standley 9905 (F); Jinotega, Finca Aventina, E of Moran 3155 (F). HEREDIA-SAN JOSE: slopes along
Jinotega, 23 Jun. 1947, Standley 9956 (F); region of La Río Para Blanca, Cerros de Zurqui, 13 Sep. 1978,
Montañita, W of Jinotega, 29 Jun. 1947, Standley 10360 Burger & Antonio 11034 (F). SAN JOSE: Tablazo, above
(F); vic. of Finca San Roque, E of Jinotega, 5 Jul. 1947, San Lorenzo de Tres Ríos, 25 Jun. 1985, Barringer &
Standley 10843 (F); between La Danta and La Luna, E Christenson 3313 (F); Parque Nacional Braulio Castillo,
of Esquipulas, 25 Jan. 1979, Stevens 11863 (MO); 19 Feb. 1983, Chacón 384 (MO); Las Tablas, Río
Ocotillo near Santa Lastenia, Cordillera Central de Cotoncito, 10 Dec. 1983, Chacón et al. 1783 (MO); San
Nicaragua, 17 Jan. 1965, L.O. Williamas et al. 27801 (F), Isidro del General, 29 Jul. 1940, Chrysler 5309 (MO);
L.O. Williams et al. 27864 (F, NY). ESTELI: Cerro San José, Cedrus plantation, 15 Aug. 1982, Moran 2393
Quiabú, 2 Jul. 1982, Moreno 16775 (MO); 0.6 km from (MO); ca. 8 km San José at La Carpintera, 24 Jun. 1983,
hwy. 3 on road to Aranjuez, 9 Apr. 1978, Stevens 7543 Moran 3030 (F), Moran 3045 (F, MO); Parque Nacional
(MO); Peñas Blancas, above Río El Gusaneras, 13-18 Braulio Carrillo, 19 Jul. 1983, Moran 3294 (F); vic. of El
Jan. 1979, Stevens 11692 (MO). Nueva Segovia, ca. 5.2 General, Jan. 1936, Skutch 2351 (MO, NY, S); about 0.7
km N of San Fernando, NE to Portillo los Coyotes, 10- km N of Tarbaca on road to Aserrí, 26 Aug. 1979,
Stevens 13676 (F, MO); S slopes of Cerro Zurquí, 5 km (F). DARIEN: Punta Guayabo Grande, 20 Apr. 1980,
N of San Luis Norte, 28 Mar.-4 Apr. 1973, Stolze 1532 Antonio & Hahn 4219 (MO); Parque Nacional del
(AAU, F); El Copey, 6 mi E of Santa María de Dota, 19 Darien, ridge between Río Topalisa and Río Pucuro,
Apr. 1928, Stork 1599 (UC). CARTAGO: Río Grande ca. 17 km E of Pucuro,, 15 Oct. 1987, Hammel et al.
de Orosí, between Orosí and Tapantí, 14 Apr. 1973, 16222 (F); Cruce del Mono Field Station, ca. 20 km
Burger & Gentry 9216 (F); Agua Caliente, 6 Aug. 1940, SSW of Boca de Cupe, 29 Apr. 1990, Moran 4176 (F).
Chrysler 5396 (MO); Cartago, Apr. 1888, Cooper 6053 PUERTO RICO. Alto de la Bandera, near Adjuntas, 14
(F); between Motavia and Quebrada Platanillo, 30 Jun. Mar. 1913, Britton & Schafer 2081 (F); Reserva Forestal
1976, Croat 36614 (MO); lower slopes of La Carpintera, Maricao, 22 mi SE of Maricao, 9 Nov. 1983, Sauleda et
14 Aug. 1925, Dodge 3446 (S); vic. Quebrada Casa al. 8551 (F). Without Locality: 28 Apr. 1886, Sintenis
Blanca, 30 Sep. 1984, Grayum 3958 (MO); crossing Río 4276 (F).
Tuis-Río Perlas, near Humo village, 25 Nov. 1986, CUBA. LAS VILLAS: Cienfuegos, Cumanayagua, Las
Hennipman et al. 7172 (MO); Valley of Río Reventazón, Vegas, 29 Oct. 1985, Berazaín et al. 57973 (HAJB);
in back of Turrialba hospital, 18 Jul. 1949, Holm & Iltis Sancti Spiritus, Fomento, road from Pedreras to
435 (BM, F, NY); Tapantí, Río Grande de Orosí, 19 Jan. Gavilán, 10 Nov. 1982, Bisse & Diaz 48477 (HAJB);
1964, Jiménez 1603 (F); Cartago, 7 Oct. 1964, Jiménez Trinidad mountains, San Blas-Buenos Aires, trail to
2437a (F); NW of Turrialba, near Pastora, ca. 0.3 km Naranjal, Aug. 1941, Howard 6419 (MO); hill behind
from the highway on the road to Finca Central, 5 Aug. Gavinas, Aug. 1941, Howard 6504 (F, MO); San Blas,
1970, Lellinger & White 1455 (F, MO, US); 20 mi S of Trinidad mountains, 29 Jul. 1936, L.B. Smith et al. 3255
Cartago, 10 May 1928, Stork 1906 (UC); Orosí, 23 Jun. (F, MO); Mina Carlota, in Trinidad hills, SE of
1928, Stork 2744 (UC); between Puente Negro (Río Cumanayagua, 24 Jul. 1947, Wood & Atchison 7467
Agua Caliente) and Río Sombrero, 25 Aug. 1975, Utley (GH). SANTIAGO: El Yunque, 30 Jan. 1902, Pollard &
& Utley 2979 (F); near Tuis, 3 May 1956, L.O. Williams Palmer 178 (F, MO, NY). EL ORIENTE: La Prenda, Jul
19547 (F). PUNTARENAS: road to Talamanca, Río 1919, Hioram 2452 (MO); Sierra del Nipe, Río Piedra,
Cotón, 2 Sep. 1983, Davidse 24498 (MO); foothills of ad Loma de Estrella, 3 Jul. 1914, Ekman 1754 (S); Sierra
Cordillera de Talamanca, Sitio Cotón, 3-4 Sep. 1983, Maestra, N spur of Pico Turquino, 16 Apr. 1915, Ekman
Davidse 24552 (AAU, MO); upper Río Burú, 19 Aug. 5412 (S); Baconas, María Pilar, 5 Nov. 1916, Ekman
1983, Gómez et al. 21431 (MO); Monteverde, 13 Nov. 8222 (LD, S); Sierra Maestra, between Río Yara and Río
1979, Koptur 223 (UC). ALAJUELA-PUNTARENAS- Palmamocha, 18 Jul. 1922, Ekman 14425 (S); N spur of
GUANACASTE: Cordillera de Tilarán, cerca de Sierra Maestra, W of Río Yao, 24-30 Oct. 1941, Morton
Reserva, 22 Jun. 1976, Dryer 351 (F). LIMON: along & Acuña 3390 (F, GH, NY); on top of El Yunque, 19-20
road between Limón and Shiroles, along Río Sixaola, 9 Dec. 1910, Shafer 7985 (F); camp La Gloria, S of Sierra
mi SW of Bambu, 12 Aug. 1977, Croat 43304a (MO); Moa, N of Río Jaguani, 24 Dec. 1910, Shafer 8049 (F);
Cordillera de Talamanca, Atlantic slope, Valle del Monte Verde, 13 Feb. 1911, Shafer 8691 (F); Firmeza to
Silencio, N of Cerro Hoffman, 8 Sep. 1984, Davidse et Gran Piedra, 3 Mar. 1911, Shafer 8983 (F); Monte
al. 28649 (MO). Verde, Jun.-Jul. 1859, Wright 797 (BM, NY, S), Wright
PANAMA. CHIRIQUI: vic. of "New Switzerland", 800 (BM, S, UC, US). Without locality: Aug. 1923,
central valley of Río Chiriquí Viejo, 6-14 Jan. 1939, Clement 956 (F); Eggers 4820 (F).
Allen 1357 (F); Allen 1358 (F, NY); La Fortuna dam site, JAMAICA. ST. ANN: Cedar valley, 13 Feb. 1900, Clute
20.9 mi from bridge over Río Estí, 27 Nov. 1979, 165 (F); 1.1. mi E of Clarksonville, St. Ann, 3 Aug.
Antonio 2821 (MO); vic. of El Boquete, 24 Jul. 1966, 1977, Goodfriend s.n. (F); 8.5 mi W by road of Albion, 10
Blum & Dwyer 2572 (MO); vic. of El Boquete, 9 Feb. Aug. 1965, Hespenheide et al. 1032 (GH); Mt. Diabolo, 6
1918, Cornman 902 (US); 2 mi N of El Hato del Volcán, Aug. 1957, Walker & Walker 1263 (BM). Between
30 May 1970, Croat 10649 (MO); El Boquete, pastures Claremont and Moneague, 7 Nov. 1902, Fawcett 8404
on Cerro Azul, near Quebrada Jaramillo, 11 Aug. 1974, (BM, F). MANCHESTER: hill at Banana Ground, 14
Croat 26868 (MO); slopes of Cerro Horqueta, 13 Aug. Jul. 1963, Crosby et al. 704 (F, NY). CLARENDON: S of
1974, Croat 26943 (MO); ca. 9 km WNW of El Boquete, Aenon town, 16 Jul. 1963, Crosby et al. 739 (F, GH, NY,
21 Nov. 1975, Davidse & D'Arcy 10319 (AAU); Nueva UC). PORTLAND: vic. of Mill Bank, 16-17 Feb. 1920,
California, 31 May 1986, Lara & Chavarria 5 (F). Maxon & Killip 206 (F); near Green River, on trail from
COLON: hills just N of the Río Guanche, 16 Nov. 1975, Cinchona, 22 Mar. 1920, Maxon & Killip 1354 (F); gorge
Davidse & D'Arcy 10063 (MO). COCLE: road from La of the Stony river above junction of the Macungo river,
Pintada to Coclesito, 7 Feb. 1983, Hamilton & Davidse 25 Jul. 1967, Proctor 28337 (F). Blue Mountains, 12 Dec.
2851 (MO); El Valle de Antón, along Río Indio trail, 30 1890, Rothrock 411 (F); Trelawny, Cockpit county, ca. 3
Jan. 1935, Hunter & Allen 315 (F, MO, S); El Valle de mi W of Troy, 20 Jun. 1959, Webster et al. 8390 (S).
Antón, 2-3 Dec. 1967, Lewis et al. 2660 (MO); La Mesa HAITI. Massif de la Selle, Morne Trauchant, 6 Sep.
del Valle de Antón, 12 Jun. 1977, Moreno & Vasquez 9 1914, Ekman 1791 (S); Petionville, Tête de l'Eau, 27
Nov. 1927, Ekman 9369 (S, US). Massif du Nord, Aug. 1979, Holm-Nielsen 19337 (AAU, QCA); Napo
Marmelade, Morme Belle-Terre, 22 May 1927, Ekman creek, 3.5 km NW of Borja, 20 Sep. 1980, Holm-Nielsen
8200 (F, S); vic. Marmelade, 18 Dec. 1925, Leonard 8088 et al. 26334 (QCA); Añangu, 30 Jun. 1983, Lawesson et
(F). Massif de la Holle, Jérémie, between Mafrand and al. 39660 (QCA); Zatzayacu, 22-28 Mar. 1935, Mexia
Maron, 11 Jul. 1928, Ekman 10292 (S). Mt. Vincent, 14 7063 (F, GH, US); Añangu, in the Parque Nacional
Dec. 1944, Holdridge 2058 (F, NY). L'Artibonite, vic. of Yasuní, 30 May-21 Jun. 1982, Øllgaard et al. 39209
Ennery, 20 Jan. 1926, Leonard 9047 (F, GH). Vicinity of (AAU, QCA), Øllgaard et al. 38853 (AAU, QCA).
Furcy, 26 May-15 Jun. 1920, Leonard 4493 (F, UC). CAÑAR: between Suscal and Chontamarca, 25 Apr.
DOMINICAN REPUBLIC. AZUA: Santo Domingo, 1945, Camp E-2883 (F). AZUAY: between Paute-
Cordillera Central, top of La Pelona, 3 Oct. 1929, Guarumales road, sector Amaluisa, 9 Aug. 1983,
Ekman 13646 (C, S). LA VEGA: Santo Domingo, Pico Jaramillo & Winnerskjold 5633 (QCA). LOJA: Alamor-
del Valle Nuevo, 15 Oct. 1929, Ekman 13774 (F, S). Celica, 2-3 km S of Río Alamor, 5 Apr. 1980, Harling &
PACIFICADOR: vic. of San Francisco de Macorís, La Andersson 17939 (QCA); Celica-Zapotilla, ca. 4 km
Bracita, 5-17 Apr. 1922, Abbott 2086 (F). Peravia, below Pozul, 22 Feb. 1985, Harling & Andersson 22428
Dieciseis, 6.1 km SW of San Juan Aldian on Piedra (QCA). ZAMORA-CHINCHIPE: new road Loja-
Blanca to Rancho Arriba road, 10 Jun. 1980, Mejía & Zamora, km 12.5 E of the pass, 14 Feb. 1991, Øllgaard et
Zanoni 6838 (MO). SANTIAGO: Cordillera Central, al. 98795 (QCA). Without locality: 12 Aug. 1893,
SW spur of Monte Gallos, 19 Jun. 1929, Ekman 12915 Eggers 14899 (F).
(S); San José de las Matas, Arroyo Jicomé, 21 Dec. 1929, PERU. CAJAMARCA: Río Aacra, 30 Jun. 1975,
Valeur 302 (F). Espinoza 305 (USM); Jaen, Chontalí, 28 Jun. 1979, Llanos
GUADELOUPE. Matorba, 1868, Husnot 297 (F). & Chimoy s.n. (USM). AMAZONAS: Bagua, ca. 1 km
COLOMBIA. ANTIOQUIA: San Luis, 12.4 km from NE of Quebrada Chinganza, 11 Jun. 1986, Knapp &
San Luis, 15 Sep. 1988, Betancur et al. 611 (MO); Mun. Alcorn 7732 (F, NY); between Aramango and
Tarazá, 201 km NE de Medellín, road to Barroblanco, Montenegro, 26 May 1963, Lopez et al. 4220 (GH);
19 Aug. 1986, Callejas et al. 2449 (F); banks of the Río mountains E of Balsas, 22 May 1912, Osgood &
Cauca, at Puerto Valdivia, 1942, Metcalf & Cuatrecasas Anderson 85b (F). SAN MARTIN: Mariscal Cáceres, 6
30052 (F). SANTANDER: vic. Puerto Berrio, between km NE of Tingo María, Cerro Azul, 4 Sep. 1956, Tryon
Carare and Magdalena rivres, 2 May 1935, Haught 1691 & Tryon 5273 (GH, US). LORETO: between
(F). BOYACA: road to Casanare, Pajarito, Uribe 6354 Yurimaguas and Balsapuerto, 26-31 Aug. 1929, Killip &
(US). CUNDINAMARCA: Suba, Sep. 1941, Uribe 211 Smith 28349 (NY, US); Santa Rosa, lower río Huallaga,
(F). below Yurimaguas, 1-5 Sep. 1929, Killip & Smith 28862
VENEZUELA. FALCON: Sierra de San Luis, arriba de (GH, NY, US); Maynas, Indiana, Yanayacu, 30 Dec.
La Chapa, 24 Mar. 1979, Werff 3405 (UC). MONAGAS: 1982, McDaniel & Rimachi 26564 (MO, NY); above
Caripe, N of El Guácharo cave, 5 Jan. 1987, Hahn & Pongo de Manseriche, left bank of Río Santiago, 3 Jan.
Grifo 3387 (F, MO). YARACUY: Sierra de Aroa, Cerro 1932, Mexia 6369a (BM, GH, NY, UC, US); Maynas,
Tigre, ridge W of Río Carabobo, 3 Apr. 1980, Liesner & Lupuna cocha, 10 Aug. 1956, Tryon & Tryon 5187 (BM,
Gonzalez 9961 (UC). F, GH, US, USM); Maynas, Río Manití, NE of Iquitos,
ECUADOR. CARCHI: Maldonado, 1 Jun. 1978, 22 Dec. 1980, Vásquez & Jaramillo 1119 (F, MO).
Madison et al. 4844 (F). IMBABURA: between El Pajón LAMBAYEQUE: Ferreñafe, Tute, 25 Jun. 1989, Llatas-
and Cachaco, 2 Jun. 1949, Acosta-Solís 12706 (F) Cantón Quiroz 2514 (F). HUANUCO: Muña, 23 May-4 Jun.
Ibarra, SW of Ibarra, 30 Jun. 1935, Mexia 7402 (F). 1923, Bryan 547 (F); Pachitea, Honoria, Bosque
PICHINCHA: 35 km S of Santo Domingo, at Puerto Nacional Iparia, Miel de Abeja, 18 Jan. 1967, Schunke
Isla farmland, 3 Jun. 1980, Balslev & Quintana 24163 V. 1526 (F); Río Pachitea, near Miel de Abeja camp, 11
(AAU, QCA); Santo Domingo de los Colorados, Apr. 1967, Schunke V. 1847 (F); Isla del Pacanasi, 5 km
Fagerlind & Wibom 1652 (S); road Nanegalito-Pacto, above Iparía camp, 14 Nov. 1967, Schunke V. 2322 (F,
archaelogical site at Tulipe, 21 Jul. 1980, Holm-Nielsen GH, NY); Panguana, 11 Jun. 1983, Seidenschwarz 447
et al. 24498 (AAU, USM). MANABI: Jul. 1893, Eggers (US); Tingo María, at confluence of Huallaga and
14899 (F, LD). COTOPAXI: 3 km E of El Empalme, on Monzón rivers, 25 Oct. 1938, Stork & Horton 9493 (UC,
road Quevedo-Latacunga, 5 Apr. 1980, Dodson & US). PASCO: Oxapampa, Ulcumanu, SW of
Gentry 10200 (MO); trail from El Corazón to Facundo Oxapampa, 31 Dec. 1983, Foster et al. 7695 (F);
Vela, 1-3 km from El Corazón, 17 May 1980, Harling & Oxapampa, forest S of city , 31 Jan. 1983, León 494
Andersson 19194 (F); Quevedo-Latacunga road, 6 Apr. (USM); Oxapampa, 1944, Soukup 2343 (F, GH). JUNIN:
1973, Holm-Nielsen et al. 3099 (AAU, F). BOLIVAR: Chanchamayo valley, above La Merced, at cumbre
Charquiyacu, 3 Oct. 1943, Acosta-Solís 6090 (F). NAPO: Yacunay, 15 Aug. 1957, Hutchison 1889 (F, GH, NY,
Reserva Biológica Jatun Sacha, Cerón 1461 (QCA) UC); Río Pinedo, N of La Merced, 30 May 1929, Killip
Misahuallí, at junction Río Misahuallí-Rio Napo, 13-14 & Smith 23647 (F, GH, NY, US); colonia Perené, 14-22
Jun. 1929, Killip & Smith 25089 (F, NY, US); brown in mass, 3-5 (-6) mm long, 0.4-0.8 mm
Chanchamayo, Jun. 1908, Schunke s.n. (BM, US); La wide, subulate or linear from a round base, scale
Merced-Chanchamayo, Feb. 1939, Soukup 1013 (F), bases shortly auriculate, apex long acuminate,
Soukup 1016 (F). AYACUCHO: between Huanta and
clathrate, without differentiated margins, the
Río Apurimac, 7,17 May 1929, Killip & Smith 22588 (F,
NY). CUSCO: ntre Mistiana y Keros, valle de
cells with walls 10-12 µm thick, sometimes with
Cosñipata, 23-31 Jul. 1948, Scolnik 882 (US). marginal trichomes. Phyllopodia 1-2 mm wide,
BOLIVIA. LA PAZ: Milluguaya, Dec. 1917, Buchtien 2-3 mm long, less than 5 mm apart. Leaves 12-77
4229 (BM, F, US); along road between Unduavi and cm long; petiole 1.5-2 cm long, stramineous;
Caranavi, 27 Nov. 1980, Croat 51553 (LPB, UC); lamina linear, bases decurrent, apices acuminate,
Canamina, 19 Jul. 1921, White 525 (F); Río Tipuani, 0.3-0.8 (-1.5) cm wide, herbaceous-chartaceous,
Okara, Apr. 1926, Tate 925 (LPB). BENI: Ballivian, margins cartilaginous, sometimes slightly
Serranía Pilón Lajas, 13-15 km de Yucumo, 20 May revolute, indument of simple, inconspicuous
1989, D.N. Smith et al. 13290 (F). SANTA CRUZ: Cerro
hairs, scattered abaxially, stomata polocytic;
Tres Cruces, 9 Oct. 1928, Steinbach 8205 (F).
BRAZIL. ACRE: vic. of Tarauacá, 18 Sep. 1968, Prance costa prominent, primary veins inconspicuous,
et al. 7384 (F, NY, S). PARA: Lageira, airstrip on Río 60-65o divergent from the costa, 1-2 areoles
Maicuru, 18 Jul. 1981, Strudwick et al. 3133 (F, NY); between the costa and margin. Sori medial or
Lageira, airstrip on Río Maicuru, 19 Jul. 1981, subterminal, paraphyses not seen; spores 50-55 (-
Strudwick et al. 3222 (F); Macau airstrip on Río 70) µm long, 40-45 µm wide. Figs. 4; 14 a; 31.
Maicuru, 25 Jul. 1981, Strudwick et al. 3555 (F); west
bank of Río Maicuru, ca. 23 km upstream from Lageira
airstrip, 29 Jul. 1981, Strudwick et al. 3744 (F, NY). This species is found from Costa Rica,
Colombia to Bolivia, where it grows between 800
Campyloneurum angustifolium is distinctive in m and 3000 m elevation, mostly as an epiphyte.
having narrowly lanceolate leaves and closely
REPRESENTATIVE SPECIMENS:
spaced phyllopodia. The stem scales are mostly
COSTA RICA. CARTAGO: San Herrano, 11 Aug.
caducous, persisting only at the growing points 1982. Moran 2358 (MO).
of the short creeping stem; the scales are COLOMBIA. EL VALLE: along hwy. Cali-
characterized by having narrowly oblong cells Buenaventura, 20 km W of village Borrero Ayerbe, 28
and transparent cell lumina. It can be Aug. 1976, Croat 38620 (MO).
differentiated from C. aglaolepis, C. VENEZUELA. ZULIA: dist. Mara, 10-15 Nov. 1982,
angustipaleatum, C. austrobrasilianum, C. Bunting et al. 12135 (MO).
centrobrasilianum, and C. ensifolium by the ECUADOR. CARCHI: ascent of Río Gualpi Chico,
characters of the stem scales used in the key. Hoover et al. 3564 (MO). NAPO: Río Napo, Huiririma,
Harling et al. 7516 (GB, MO). ZAMORA-CHINCHIPE:
Some populations of Campyloneurum.
road Loja-Zamora, km 24-25, Holm-Nielsen et al. 3526
angustifolium, growing mainly at middle (AAU, F, MO, UC).
elevations, have leaves that resemble those of C. PERU. CAJAMARCA: Colasay, Woytkowski 6941 (GH,
amphostenon. However, they can be MO, US). AMAZONAS: Bagua, 12 km E of La Peca,
distinguished by the number of areoles between Barbour 2565 (F, MO, NY, UC); Chachapoyas, Ingenio-
the costa and margin, which in the former Pomacocha, 28 May 1963, López et al. 4310 (GH); NW
species are 1-2 (-3), while in the latter they are 2-4 of Jumbilla, laguna Pomacocha, 23 Jan. 1965, Soukup
(-5); further more the stem scales in the latter 5258 (GH); Bongará, hills WNW of Pomacocha, 19 Jun.
1962, Wurdack 946 (US, USM); Chachapoyas, along Río
species are wider, as mentioned in the key.
Ventilla, 1-2 km W of Molinopampa, Wurdack 1503 (F,
7. Campyloneurum angustipaleatum (Alston) Meyer GH, NY, UC, US, USM); Mendoza, Woytkowski 8167
ex Lellinger, Amer. Fern J. 74: 56. 1984. (GH, MO, US). SAN MARTIN: Rioja, Pedro Ruiz,
Polypodium angustipaleatum Alston, J. Bot. 77: 346. Moyobamba road, km 390 Venceremos, D.N. Smith
1939. Type. Bolivia: near Cochabamba, 1891, 4358 (MO), D.N. Smith & Vásquez 4630 (MO, NY).
Bang 1288 (holotype BM!; isotypes B!, F!, MO!, HUANUCO: Muña, 23 May-4 Jun. 1923, Bryan 516 (F);
US) Monzón, "Cuevas de las Lechuzas", 11 Jul. 1958,
Stem short creeping, sometimes branched, 1-2 Ferreyra 13217 (USM); 32 km from Huánuco, road
Huánuco-La Unión, 25 Jul. 1982, D.N. Smith 2191
(-4) mm wide, pruinose. Stem scales dark or red
(USM). PASCO: Ulcumanu, SW of Oxapampa, road to
María Teresa and Llaupi, 31 Dec. 1983, Foster et al. (Berlin) 288. 1845. Type. Venezuela: Caracas,
7695 (MO); aprox. 3 km del puente sobre el Río Moritz 17 (B!).
Paucartambo, camino a Oxapampa, León 473 (USM); Campyloneurum occultum (Christ) L. D. Gómez,
Oxapampa, Fundo La Esperanza, 11 Aug. 1985, León
Brenesia 8: 46. 1976.
624 (F, USM); 8 km N of Huancabamba, on the
Oxapampa-Pozuzo road, 26 May 1978, Skog et al. 5075
Polypodium occultum Christ, Bull. Herb. Boissier
(US); 5 km SE of Oxapampa, D.N. Smith & W. Brack 2: 7. 1905. Type. Costa Rica: Cartago, Río de
2796 (F, MO); D.N. Smith 3653 (F, MO, NY, USM). las Vueltas, Tucurrique, Nov. 1898, Tonduz
JUNIN: Huacapistana, 5-8 Jun. 1929, Killip & Smith 12752 (holotype, P!; isotypes B!, BM!). Christ
24222 (NY, US); Chanchamayo valley, C. Schunke 29 in the protologue of the species mentioned
(F); Yaupe, Woytkowski 6468 (MO), Woytkowski 6480 Tonduz 12756, as the type collection, but this
(GH, MO, US), Woytkowski 6481 (MO); Yunguy, may represent a typographic error.
Woytkowski 6603 (MO); Tarma, Utcuyacu, Woytkowski Polypodium trichiatum Rosenst., Repert. Spec.
37006 (MO, S, UC). CUSCO: Convención, Tupitari,
Nov. Regni Veg. 7: 148. 1909. Type. Ecuador:
Bües 5447 (MO, US); San Miguel, Urubamba valley, 10
Jul. 1915, Cook & Gilbert 17599 GH, NY, US), Cook & Riobamba, Cordillera Occidental, Rimbach 87
Gilbert 1760 (US); Urubamba, Machu Picchu, (holotype, not found; isotypes, S!, US!, photo
Doppelbaur & Doppelbaur s.n. (MO); Machu Picchu, 25 of S, BM!).
Jan. 1983, León 454 (USM); Río Marcapata, 60 km Campyloneurum trichiatum (Rosenst.) Ching,
above Quincemil, 17 Jan. 1973, Madison 999 (GH); La Sunyatsenia 5: 263. 1940.
Convención, Maranura, Chaullay, Nuñez 8150 (MO, Stem creeping, black, dark stramineous, 1-3 (-
NY); valle de Santa Ana, above Quillabamba, Plowman 4) mm wide. Stem scales dark brown in mass, 2-
& Davis 4800 (F, GH); Potrero, 8 km W of
4 (-5) mm long, 0.75-1 (-1.2) mm wide, narrowly
Quillabamba, Tryon & Tryon 5385 (GH, US, USM);
Quispicanchis, Inambari, 2 Sep. 1965, Vargas 16427
ovate, bases auriculate, apices acuminate,
(GH); La Convención, Itma, 28 Jun. 1967, Vargas 19831 clathrate, slightly differentiated margins, cells
(GH); Apr. 1976, Vargas 22746 (GH). PUNO: oblong, except the rondish cells at the base of the
Carabaya, Ollachea-San Gabán road, 25 Aug. 1980, scale, central cell walls 16-20 µm wide, marginal
Boeke & Boeke 3208 (MO, NY). cell walls 8-12 µm wide. Phyllopodia 1-1.5 mm
BOLIVIA. LA PAZ: Nordyungas, Polo Polo bei long, 2-2.5 mm wide, 2-4 (-6) mm apart. Leaves
Coroico, Buchtien 3529 (F, S, US); Buchtien 3530 (F, S). (10-) 20-40 (-55) cm long; petiole stramineous or
COCHABAMBA: Chapare, Paractí, 83 km from
dark stramineous, (0.3-) 0.7-3 (-0.6) cm long,
Cochabamba, on road to villa Tunan, M. Foster 79-174
slightly ranurate adaxially, puberulent, rarely
(MO); Sacaba, Incachaca, 11 Oct. 1921, Steinbach 5848
(F, GH); Chapare, Steinbach 9073 (GH); Llanta-Aduana, with scattered scales similar to those on the stem;
Steinbach 9599 (F, MO, S). SANTA CRUZ: Ichilo, laminae oblanceolate, rarely narrow lanceolate,
Parque Nacional Amboro, along Río Saguayo, 20 Dec. bases attenuate, apices acuminate or caudate,
1988, Nee & Saldias 37289 (MO). Without locality: (1.2-) 2-4 (-4.7) cm wide, herbaceous-chartaceous,
Rusby 2461 (F). margins cartilaginous, plane or slightly revolute,
sinuate, indument of pluricellular hairs, with one
Campyloneurum angustipaleatum is closely small basal branch, glandular and the other long;
related to C. angustifolium. These species , stomata polocytic; costa prominent on both
surfaces of the lamina, primary veins slightly
have similar leaf morphologies and patterns of prominulous or inconspicuous, same color as the
venation. They differ in the characteristics of the leaf tissue or different adaxially, slightly
stem scales, which in the former are less than 1 flexuosous, (-50o) 60-65o divergent from the
mm wide and are linear from a wide base, while costa, (3-) 4-7 (-9) mm apart, secondary veins
in the latter they are lanceolate, with the base inconspicuous, transverse veins sometimes
more than 1 mm wide. darker than the leaf tissue, forming 3-6 primary
areoles between the costa and margin, primary
areoles usually undivided, 2 (-3) excurrent
8. Campyloneurum aphanophlebium (Kunze) T. veinlets in each primary non-costal areole; sori
Moore, Index Fil. 223. 1861. medial or subterminal, paraphyses dendritics,
Polypodium aphanophlebium Kunze, Bot. Zeitung spores 50-60 (-65) µm long, 30-40 µm wide. Figs.
9; 13 b; 18 a; 28. EL VALLE: Río Digüa valley, La Margarita, 4-5 Apr.

1939, Killip 34889 (GH, US). SUR DE SANTANDER:
This species is distributed from Belize to vic. of Puerto Berrío, between Carare and Magdalena
river, 14 Jun. 1935, Haught 1777 (US). CAUCA: valle
Brazil and Bolivia. It grows in forests, from sea
del Cauca, Duque-Jaramillo 1985 (F).
level to 1500 m, usually as an epiphyte. VENEZUELA. Colonia Tovar, 1856-1857, Fendler 409
(K). Encanto, 1929, Vogl s.n. (S).
REPRESENTATIVE SPECIMENS: BELIZE. Edwards FRENCH GUIANA: Without locality, Jan. 1836,
road, beyond Columbia, 13 May 1951, Gentle 7327 Leprieur s.n. (B).
(US). ECUADOR. NAPO: San Pablo de los Secoyas, Río
NICARAGUA. ZELAYA: between cerro La Pimienta Wai-si-aya, northern tributary to Río Aguarico, 7 Aug.
and El Hormiguero, 15 Mar. 1980, Pipoly 5971 (MO, 1980, Brandbyge et al. 32635 (AAU, QCA); 6 km upriver
UC); Caño El Hormiguero, 17 Mar. 1980, Pipoly 6106 from San Pablo, 10 Aug. 1980, Brandbyge & Asanza
(MO). CHONTALES: Chontales, Jun. 1868, Tate s.n. 32734 (AAU, QCA); W of confluence with Río Napo, at
(BM). Without locality: Jun. 1868, Tate 57 (K). Bimbino, 21 Oct. 1960, Whitmore 787 (BM).
COSTA RICA. GUANACASTE-ALAJUELA: slopes of PICHINCHA: San José de Toachi, 100 km W of Quito,
Miravalles, above Bijagua, Nov. 1982, Gómez et al. 3 Apr. 1951, Bell 217 (S); forest of the Cooperativa
19050 (AAU, UC). ALAJUELA: NW of Volcán Arenal, Santa Marta No. 2, at km 3 of bypass, around Santo
ca. 2 km NE of Tabacón, 16 Aug. 1970, Lellinger & Domingo, 22 Jul. 1979, Dodson et al. 8505 (US); road
White 1650 (F, US). CARTAGO: behind main building Aloag-Santo Domingo, Alluriquin, 14 Mar. 1967,
of CATIE, Turrialba, 30 Jul. 1985, Grayum & Hammel Sparre 14819 (S); Quito, Spruce 5738 (BM, K).
5748 (MO, USM); near Turrialba, slope of Río PASTAZA: Ceilán, Pica, from Ceilán to Río Cononaco,
Reventazón, behind the Instituto Interamericano de N side of the Río Curaray, 6 Jun. 1980, Brandbyge &
Ciencias Agrícolas, 20 Aug. 1967, Mickel 3362 (NY, Asanza 31659 (AAU, QCA); oil exploration camp
US); bords du Río Colorado, Golfo Dulce, Mar. 1896, Chirirota, on the Río Bobonaza, 26 Jul. 1980, Ølllgaard
Pittier 9993 (US); Turrialba, near Interamerican et al. 35298 (AAU, QCA). GUAYAS: Teresita, 3 km W
Institute, 30-31 Mar. 1953, Scamman 7244 (GH). of Bucay, 5-7 Jul. 1923, Hitchcock 20410 (GH, US).
HEREDIA: La Selva, along Río Viejo, 1 Feb. 1988, MORONA-SANTIAGO: Tukupi, near the military
Hennipman et al. 7156 (MO); Puerto Viejo, Finca La camp, 25 Jun. 1980, Brandbyge & Asanza 32276 (AAU,
Selva, 18 Jun. 1967, Sota 5120 (US). PUNTARENAS: QCA); 35 km NE of Montalvo, 2-12 Jul. 1989, Zak &
about 5 km W of Rincón de Osa, Osa Península, 24-30 Espinoza 4646 (MO). SANTIAGO-ZAMORA: valley of
Mar. 1973, Burger & Gentry 8875 (F). Río Negro and río Chupianza, 2-3 km W of Méndez,
PANAMA. COLON: E Santa Rita ridge, 23 Feb. 1968, 13 Dec. 1944, Camp E-1495 (US). Without locality, Sep.
Correa & Dressler 754 (MO). COCLE: El Valle de 1884, Sodiro s.n. (UC).
Antón, 4 Jun. 1939, Alston 8712 (BM). CANAL ZONE: PERU. AMAZONAS: Bagua, ca. 1 km NE of
Barro Colorado Island, Shannon trail 100, 17 Jun. 1970, Quebrada Chinganza, 10 km NE of Mayo, 11 Jun.
Croat 10906 (F, NY); Armour trail 1040, 29 Jul. 1970, 1986, Knapp & Alcorn 7733 (F, NY); Bagua,
Croat 11634 (F, NY); Isthmo de Panama, 1859-61, Hayes Montenegro-Chiriaco, 16 Oct. 1965, Sagástegui 5924
54 (B); trail along Río Petitpie, from road to Ft. (HUT). SAN MARTIN: Mariscal Cáceres, dist.
Sherman from Gatun Locks, 22 Oct. 1974, Mori & Campanilla, Cajón Pericote, 21 Aug. 1970, Schunke V.
Kalluncki 2676 (US); 6 mi N of Gamboa, on ridge S of 4288 (F, UC, US, USM); Río Marañón, Apr. 1855,
Río Frijol, 6 Oct. 1965, Tyson 1514 (GH). DARIEN: Río Spruce 3912 (K); Tarapoto, 1855-1856, Spruce 3912 (K).
Morti, drill site 7, 15 Sep. 1967, Duke 14138 (F); vic. of LORETO: Balsapuerto, lower Río Huallaga basin, 28-
Caná, 24 Jun. 1959, Stern et al. 687 (US). GOLFO DE 30 Aug. 1929, Killip & Smith 28642 (NY, US); San José
PANAMA: San José Island, Perlas Archipielago, about de Parinari (Río Marañón) , 10 Aug. 1981, Vásquez et al.
55 mi SSE of Balboa, 25 Oct. 1944, Johsnton 285 (BM, 2274 (NY) San Martín de Tipishca (Río Samiria), 13
GH, K). May 1985, Vásquez & Jaramillo 6511 (F, MO); Maynas,
COLOMBIA. ANTIOQUIA: around Villa Arteaga, 10 Indiana, Explorama reserve, 10 Nov. 1989, Vásquez &
Oct. 1947, Gutierrez & Barcley 17C101 (BM); Villa Jaramillo 13167 (MO). HUANUCO: Tingo María, 30
Arteaga, 4-8 Aug. 1947, Hodge 6988 (GH); Mun. Oct. 1949-19 Feb. 1950, Allard 21520 (US), Allard 21982
Zaragoza, Corregimiento de Providencia, vicinity of (US); Fundo Chela, Sinchono, 3 Aug. 1948, Aguilar 946
Tirana, 12 Feb. 1971, Soejarto & Villa 2817 (GH). (USM); above Río Huallaga, at Tingo María, 4 Oct.
CHOCO: W of Puerto Mutis, Bahía Solano, 25 Jan. 1972, Croat 21034 (USM); Pachitea, dist. Puerto Inca,
1971, Lellinger & Sota 12 (US); Chocó, Schott s.n. (MO). Agua Dulce, Bosque Nacional Iparía, 6 Dec. 1968,
INTENDENCIA DEL META: Sierra de la Macarena, Schunke V. 2818 (F, GH, US); Tingo María, by Río
Caño Estrada, 3 Jan. 1950, Philipson & Idrobo 2017 (BM). Huallaga, 16 Sep. 1956, Tryon & Tryon 5335 (GH, US,
USM); Cachicoto, 5 Apr. 1963, Woytkowski 7868 (UC). diverging 50-60o from the costa, with 2-4 areoles
PASCO: Puerto Bermudez, 14-17 Jul. 1929, Killip &
between costa and margin, one, rarely two,
Smith 26647 (US). JUNIN: La Merced, 29 May-4 Jun.
1929, Killip & Smith 25268 (S); Killip & Smith 23781 veinlet in each areole. Sori subterminal,
(GH, NY, US). MADRE DE DIOS: Parque Nacional paraphyses not seen, spores 60-65 µm long, 40-42
Manu, Cocha Cashu, 8 Nov. 1984, M. Foster 84-64 (F); µm wide. Fig. 32.
Manu, Cocha Cashu Station, 15 Aug. 1978, Foster &
Terborgh 6630 (F). This species is distributed from Venezuela,
BOLIVIA. Yumupasa, 10 Jun. 1902, Williams 1064 (US). Ecuador, Peru to Bolivia, where it grows
BRAZIL. ACRE: Cruzeiro do Sul, Rio Juruá and Rio between 1000 m and 3500 m elevation, among
Moa, vic. of Porangaba, Rio Juruá-Mirim, 21 May 1971,
rocks and sometimes as an epiphyte.
Maas et al. P131381 (S, US); Mun. Tarauacá, vic. of
Tarauacá, 14 Sep. 1968, Prance et al. 7267 (GH, K, S,
REPRESENTATIVE SPECIMENS: VENEZUELA.
US). PARA: W bank of Rio Maicuru, ca. 23 km
MERIDA: Andrés Bello, Zerpa, La Carbonara, bosque
upstream from Lageira airstrip,29 Jul. 1981, Strudwick
San Eusebio, 14 Nov. 1981, Marin et al. 115 (F).
et al. 3709 (F).
ECUADOR. PICHINCHA: Chaparro de Sebritana, 9
Jul. 1944, Acosta-Solis 8364 (F). CHIMBORAZO: Cerro
Campyloneurum aphanophlebium is Chiguazo, ca. 15 km from Penipe, 24 Sep. 1968, Lugo
characterized by its dense puberulent leaves and 474 (F).
inconspicuous secondary veins. Campyloneurum PERU. SAN MARTIN: Mariscal Cáceres, Parque
aphanophlebium and C. anetioides belong to the Nacional Río Abiseo, trail between La Playa and
same group, sharing morphological features, Papayas, 25 Jul. 1987, Young & León 4995 (F, USM);
such as stem scales characters, closely spaced Huallaga, valley of Río Apisoncho, 30 km above
leaves on the stem, and the presence of foliar Jucusbamba, 9 Sep. 1965, Hamilton & Holligan 903 (US),
Hamilton 905 (US). HUANUCO: Carpish, 12 Dec.
indument on both sides of the lamina.
1953, Coronado 62 (GH, UC, US), Coronado 67 (US); 6
mi S of Mito, 1-5 Aug. 1922, Macbride & Featherstone
1835 (US); Yanano, 13-16 May 1923, Macbride 3818 (F,
9. Campyloneurum asplundii (C. Chr.) Ching, GH, US); Muña, 23 May-4 Jun. 1923, Macbride 3911 (F,
Sunyatsenia 5: 263. 1940. US); Muña, 10 Mar. 1959, Woytkowski 5181 (GH, US).
Polypodium asplundii C. Chr., Ark. Bot. 20A (7): PASCO: Oxapampa, 31 Jan. 1983, León 491 (F, USM); 1
24. 1926. TYPE: Bolivia, Prov. Sur Yungas, La Feb. 1983, León 506 (F, USM); canyon of Huancabamba,
Sirena, ca. 2300 m, 1920, Asplund 282 Fundo La Esperanza, 11 Aug. 1985, León 617 (F, GH,
USM); border of Parque Nacional Yanachaga, 21 Jun.
(holotype S!, isotype BM!).
1986, León et al. 946 (F, USM); Río El Tunqui, 2 Jan.
Stem creeping, (2-)3-4 (-4.5) mm wide,
1984, D.N. Smith & Alban 5526 (F, MO, NY); Santa
sometimes pruinose, black. Stem scales narrowly Bárbara, 3 Aug. 1984, D.N. Smith 8160 (F); border Prov.
ovate, non-clathrate, shining dark brown in Oxapampa and Pasco, below San Gotardo, 7 Mar.
mass, 3-5 mm long, 1-1.5 mm wide; with cell 1986, Werff et al. 8521 (MO, NY, UC). JUNIN:
lumen yellow brownish, scale base short Carpapata, above Huacapistana, 7 Jun. 1929, Killip &
biauriculate. Phyllopodia 1.5-3 mm long, 1.5-2 Smith 24429 (NY, US); Chanchamayo, Jun. 1908, C.
mm wide, 4-7 mm apart. Leaves 35-60 cm long; Schunke s.n. (BM, F); Chanchamayo, Río Rondayacu, 45
petiole stramineous, (1-) 4-12 cm long, indument km from San Ramón, 2 Jun. 1982, D.N. Smith 2117
(USM); La Merced, Aug. 1947, Soukup 3400 (F); Yaupe,
of hairs; lamina linear to narrowly lanceolate,
9 Jul. 1961, Woytkowski 6497 (US). LIMA: Chancay,
bases and apices attenuate, (0.6-) 1.5-3.5 (-5.8) cm Huaccaña, E of Supe, 4 Jan. 1952, Cerrate 1062 (USM).
wide, chartaceous, lamina margins cartilaginous, AYACUCHO: Huanta-La Mar, Tambo, 37 km from
slightly revolute, indument of bicellular hairs, Ayna, 23 Mar. 1977, Ellenberg 7021 (LPB). CUSCO:
scattered abaxially; stomata polocytic; costa Cerro Huacontoy, Hacienda Muy Muy, valle de Lares,
prominent on both surfaces, with indument of Mar. 1932, Bues 1868 (US); La Convención,
often persistent scales, primary veins obscure or Vilcabamba, trail from Yupanqui to Apurimac, 4 Jul.
slightly prominulous to different degrees on 1981, Davis et al. 1231 (GH); Machu Picchu, 18 Nov.
1947, Ferreyra 2707 (BM, USM); along Inca trail, above
either side of the lamina, sometimes to the same
Machu Picchu, 13 Apr. 1977, Gentry et al. 19422 (F),
degree, slightly flexuous, 4-5 mm apart, near Cusco, Apr. 1923, Herrera 5 (US); Río Urubamba,
22 May 1958, Humbert 30662 (GH); Urubamba, Winay inconspicuous, 1-2 areoles between costa and
Wayna, 26 Oct. 1986, Nuñez et al. 8422 (F, MO, NY, margin, one veinlet in each areole; sori medial,
UC); Kosñipata, between Yanamayo and Santa Isabel, paraphyses not seen, spores 60-65 µm long, 40-42
23-31 Jul. 1948, Scolnik 847 (US); Urubamba, Machu
µm wide. Fig. 26.
Picchu, Tancarpata, 20 Jul. 1961, Vargas 13591 (GH); La
Convención, Potrero, Sapansayoc, 23 Sep. 1961, Vargas
13645 (GH). PUNO: Sandia, bajando a Valle Grande, 7 This species is restricted to central east and
Aug. 1957, Vargas 11864 (GH); Sandia, Limbani a southeast Brazil, where it occurs below 1500 m
Chacani, 13 Oct. 1963, Vargas 14936 (GH). elevation, mostly as an epiphyte in forests.
BOLIVIA. LA PAZ: Murillo, Zongo valley, above
Jarca, Río Chuchulluni, 31 May 1980, Beck 3590 (F); REPRESENTATIVE SPECIMENS: BRAZIL. MINAS
Zongo valley, Cambaya, 14 Dec. 1982, Liberman 490 (F). GERAIS: Mun. Jabaticatubas, Serra do Cipó,
Hatschbach 30048 (UC); Serra do Espinhaço, lower
Campyloneurum asplundii is characterized by slopes of Serra do Caraça, ca. 12 km W of Barão de
having linear-lanceolate, non-clathrate, shining Cocais, 27 Jan. 1971, Irwin et al. 29288 (F, NY); Mun.
dark brown stem scales, with an almost occluded Ouro Preto, Macedo 3046 (S); Caldas, Mosen 2218 (S);
Caldas, Pedra Branca, Regnell 317 (S). MATO
lumina.
GROSSO: Palmeiras, Lindman A2527 (S). GOIAS: ca.
One syntype of Polypodium leucorhizon Kunze 30 km N of Alto Paraíso, Chapada dos Veadeiros, 23
ex Klotzsch, Schomburgk 1145 from British Mar. 1971, Irwin et al. 33063 (F, NY). PARANA: Mun.
Guiana, has scales of the stem similar to Lapa. Fundo São Sebastião, Kummrow 1396 (MU, UC);
Campyloneurum asplundii. However, it seems Mun. Bocaiuva do Sul, Bacaetava, Kummrow 1423
uncertain that this specimen from lowland areas (UC); Mun. Quatro Barras, Borda do Campo,
belong in this taxa. Kummrow 1959 (UC). RIO GRANDE DO SUL: Sep.
This species belongs to the C. amphostenon 1940, Eugenio & Leopoldo 1773 (F); Paso da Mangueira,
Santa Cruz, Jürgens s.n. (Rosenstock 96) (BM, F, S, UC);
group. Campyloneurum asplundii is closely
Cachoeira, Lindman 1169 (S); Silveira Martins, Lindman
related to C. solutum; these species are similar in 1169b (LD, S); Porto Alegre, Cañoas, Lindman 1169c (S);
shape and color of the scales. The former, Esmeralda, Est. Ecol. Aracauri, 8 Oct. 1980, Waechter
however, has non-clathrate scales with yellow 1726 (F). SANTA CATARINA: Lages, Spannagre 7
brown cell lumina, almost occluded. (UC); Joinville, Schmalz 175 (F, MO); Mun. Cacador,
Pinheiral, Smith & Reitz 8965 (US). SÃO PAULO:
Campos de Jordão, 20 Feb. 1937, Campos Porto 3211 (F,
10. Campyloneurum austrobrasilianum (Alston) US). RIO DE JANEIRO: Itatiaya, Maromba-Macieiras,
Sota, Op. Lilloana 5: 99. 1960. Zerny s.n. (W). Without exact locality: Sehnem 6321
(US).
Polypodium austrobrasilianum Alston, J. Bot. 77:
347. 1939. Type. Brazil: Paraná, Curitiba,
Campyloneurum austrobrasilianum is
Hoehne 24365 (holotype, BM!, isotype GH!).
characterized by brown, linear stem scales, with
Stem short-creeping, not pruinose, 1-3 mm
cell walls clearly defined. This species belongs to
wide. Stem scales red brown or brown in mass,
the C. angustifolium group.
2.5-4.5 mm long, 0.3-0.8 (-1) mm wide, narrowly
ovate, bases auriculate with open or superposed
auricles, margins sometimes with hairs 12 µm
11. Campyloneurum brevifolium (Lodd. ex Link)
long, scales clathrate, with nondifferentiated
Link, Fil. Spec. 124. 1841.
margins, the cells oblong, cell walls 12-16 µm
Polypodium brevifolium Lodd. ex Link, Hort.
wide. Phyllopodia 0.5 mm long, 1 mm wide, 2-4
Berol. 90. 1833. Type. Habitatio ignota.
mm apart. Leaves 15-50 cm long, petiole
Cultivated. (holotype B!; photo BM!).
greenish or stramineous, 0.3-0.8 (-3.5) cm long;
Campyloneurum latum T. Moore, Index Fil. 225.
laminae linear, bases and apices attenuate, 0.3-
1861. Type. Windward Islands. St. Vincent:
0.6 cm wide, herbaceous-chartaceous, margins
s.d. Guilding s.n. (Lectotype chosen by Proctor
cartilaginous, sometimes revolute, indument not
in R. Howard 2: 342, 1977, K!).
seen, stomata polocytic; costa prominent on both
Polypodium phyllitidis Linn. var. lata (T. Moore)
surfaces, without indument, primary veins
Hook. Sp. Fil. 5: 38. 1864.
Polypodium latum (T. Moore) T. Moore ex Sodiro, grows on abundant organic matter on the
Crypt. Vasc. Quit. 371. 1893. ground or as a hemiepiphyte in tropical forest.
Campyloneurum phyllitidis var.latum (T. Moore)
Farwell, Amer. Midl. Naturalist 12: 297. 1931. REPRESENTATIVE SPECIMENS: U.S.A. FLORIDA:
Polypodium phyllitidis f. latum (T. Moore) Proctor, Dade County, 13 Dec. 1903, Eaton 562 (GH, US).
Bull. Inst. Jamaica Sci. Ser. 5: 49. 1953. MEXICO. JALISCO: Los Chorritos, La Rinconada,
Purificación, 450 m, 5 Feb. 1979, Alcocer 11111 (UC).
Stem creeping, not pruinose, (4-) 6-17 (-24)
PUEBLA: between Pahuatlan and Honey, 16 Dec.
mm wide. Stem scales light brown in mass, 4-6 1942, Martinez 101 (F). OAXACA: Tuxtepec, hills 9 km
mm long, 2-3 mm wide, broadly ovate, S of Tuxtepec and 4 km W of route 175, 30 Jul. 1971,
pseudopeltate, auriculate, apices acuminate, Mickel 5782 (UC); 4-9 km S of Valle Nacional, 31 Jul.
clathrate, with differentiated margins, the cells 1971, Mickel 5867 (UC); 1-3 km S of Usila, 27 Sep. 1973,
oblong, central cells along the main axis of the Mickel 7375 (UC).
scale, marginal cells transverse, these usually BELIZE. TOLEDO: 1.5 mi S of Mayan village of San
smaller than the central ones, central cell walls José, 12 Jun. 1973, Croat 24351 (F, US); near Jacinto
16-20 µm wide, marginal cell walls 6-8 µm wide. Creek, 17 Nov. 1944, Gentle 4982 (F, S); on hilltop
beyond Carmelita camp, Edwards road beyond
Phyllopodia 2-5 mm long, 3-6 mm wide, 2-4 (-10)
Columbia, 27 Jun. 1951, Gentle 7385 (F, S, US);
mm apart. Leaves (35-) 40-120 cm long; petiole Salamanca camp, 26 May 1979, Whitefoord 1887 (BM).
stramineous or brownish, (2-) 5-28 cm long, EL CAYO: Valentin, Jun.-Jul. 1936, Lundell 6231 (US).
ranurate adaxially, convex abaxially, glabrate, Middlesex, 16 Sep. 1929, Schipp 8-46 (F).
indument of broadly ovate scattered scales GUATEMALA. PETEN: Vaxactum, 20 Mar. 1931,
similar to those in the stem; lamina Bartlett 12148 (F, UC, US). IZABAL: S shore of Lake
elliptic-lanceolate, bases attenuate or subcuneate Isabel, between Izabal and Mariscos, 28 May 1966,
then short or long decurrent, apices acuminate or Jones & Facey 3498 (F). Quirigua, lawn at United Fruit
Co. Hotel, 8 Feb. 1945, Weatherwax 228 (UC).
caudate, 5-13.5 cm wide, chartaceous or
HONDURAS. CORTES: N of Lago de Yojoa, SW of
coriaceous, margins cartilaginous, sinuate or Santa Cruz de Yojoa, 4 Aug 1977, Croat 42737 (UC);
undulate, indument of inconspicous simple Isla de La Guamita, N shore of Lake Yojoa, 29 May
hairs, scattered abaxially, stomata polocytic; 1974, Horwath 44 (F). COMAYAGUA: 10 km N of
costa prominent on both surfaces, slightly Talube, 28 May 1974, Horwath 18 (F). ATLANTIDA:
ranurate abaxially, convex or slightly angulate Lancetilla, Tela, 17 Jun. 1964, Lent 7 (F, US); near Tela,
adaxially, sometimes with oval-lanceolate Lancetilla valley, 6 Dec 1927-20 Mar. 1928, Standley
scattered scales, primary veins prominent on 53540 (US); Tela river, above Lancetilla, 1 Aug. 1951,
Steeves & Ray 413 (GH, UC, US); above Lancetilla, 12
both surfaces, stramineous, straight or slightly
Jul. 1934, Yuncker 4548 (F). GRACIAS A DIOS: La
flexuosous at the margin, 70-75o divergent from Mosquita, alrededores del Río Plátano, 17-23 May
the costa, (4-) 6-9 (-13) mm apart, secondary 1973, Clewell & Cruz 4138 (US). OLANCHO: vicinity
veins inconspicuous or slightly prominulous of Juticalpa, 5-16 Mar. 1949, Standley 18043 (F).
abaxially, rarely adaxially, transverse veinlets EL SALVADOR. SAN SALVADOR: Botanical Garden
forming 8-18 primary areoles between the costa La Laguna, 2 Mar. 1979, Seiler 971 (F).
NICARAGUA. CHONTALES: between Santo Tomas
and margin, excurrent veinlets 2-5, entires or
and Villa Somozo, 7 Apr. 1961, Bunting & Licht 1098
furcate, free or forming 2-5 secondary areoles;
(F, GH); vic. La Libertad, 29 May-1 Jun. 1947, Standley
decurrent veinlets 1-2 in each primary areole; 9002 (F). MATAGALPA: summit of Matagalpa-Tuma
sori medial, subterminal or at the junction of the road, 24 May 1981, Stevens & Henrich 20316 (UC).
secondary veins, 2-4 series between the ZELAYA: Sector Mina Nueva America, 22 Sep. 1984,
secondary veins; paraphyses not seen, spores Ortiz 2129 (MO). BLUEFIELDS: SE slopes of Cerro
50-60 (-70) µm long, 35-40 µm wide, verrucate San Isidro, 3.6 km SE Cerro San Isidro, 25 Mar. 1966,
surface. 2n= 74. Figs. 14 j; 33. Proctor et al. 27254 (F, NY). MANAGUA: vic. Casa
Colorada, near El Crucero, summit of Sierra de
Managua, 14-25 May 1947, Standley 8425 (F). Without
Campyloneurum brevifolium is distributed from
exact locality: Cabo Gracias a Dios, 1 May 1923,
Mexico, Central America, the Antilles Venezuela Schramm s.n. (US); Castillo, Mar. 1893, Shimek s.n. (F);
to Bolivia, rarely in southern U.S.A. (Florida). It s.l. 1853-1856, Wright s.n. (US).
occurs from sea level to 2000 m elevation. It
COSTA RICA. GUANACASTE: Rincón de la Vieja (MO). DARIEN: Serranía del Darién, trail from Cerro
National park, ridge SE of Quebrada Zopilote, lower Mali to Río Pocuro, 20 Jul. 1976, Gentry, A. et al. 16827
SE slope of Volcán Santa María, 24 Jan. 1986, Smith et (US); trail NW of Caná, 28 Jul. 1976, Sullivan 687 (MO).
al. 1925 (UC). ALAJUELA: ca. 7 km E of Ciudad ISLA COLON: 16 May 1940, Wedel 131 (MO).
Quesada, 17-18 May 1968, Burger & Stolze 4934 (F); CUBA. SANTA CLARA: SE of Cuamanayagua, Sierra
wooded area above Río Aguas Zarcas, S of Aguas de San Juan, 21-23 Mar. 1938, Senn 189 (GH); Senn 367
Zarcas, 20 May 1968, Burger & Stolze 5111 (F); near (GH). CIENFUEGOS: around Topes de Collantes, 16
Artezalea, ca. 8 km NE of Villa Quesada, 16 Feb. 1966, Jul. 1974, Areces & Berazain s.n. (HAJB 25095).
Molina et al. 17220 (F, US); Florencia, 14 Jul. 1963, ORIENTE: Santiago de Cuba, loma del Gato, Sierra
Rickson 208 (GH). HEREDIA: forest near Río Puerto Maestra, Jul. 1923, Clement 917 (BM); Sierra Maestra,
Viejo, 14-17 Jun. 1968, Burger & Stolze 5819 (GH); Florida above Daiquiri, 28-29 Jun. 1914, Ekman 1593
Burger & Stolze 5914 (F, NY); Finca of Dr. Holdridge, (S); Bayate, 5 Jun. 1915, Ekman 5911 (S).
on the Río Puerto Viejo, 18-28 Feb. 1955, Scamman 7511 GUANTANAMO: s.l., Hioram 6491 (F). Without
(GH); Cerros Sardinal, ca. 2-2.5 km N of Chilamate de locality: Apr. 1881, Herb. Small s.n. (F); Eggers 4953 (F).
Sarapiquí, 21 Jan. 1986, Smith et al. 1800 (UC); Finca La JAMAICA. Dollwood, 6 Aug. 1898, Harris 7273 (BM,
Selva, Sarapiquí region, 30 Aug. 1961, Weber 6121 (GH, F); near Mocho, above Catadupa, 3 Apr. 1920, Maxon
US). PUNTARENAS: about 5 km W of the Rincón de & Killip 1558a (US); along the trail from Bath to Cuna
Osa, Osa Peninsula, 24-30 Mar. 1973, Burger & Gentry Cuna Pass, 1 May 1903, Maxon 1713 (US); near Bath, 2
8889 (F); foothills of the Cordillera de Talamanca, vic. Jun. 1904, Maxon 2438 (US). Gorge of the Stony River
of Helechales, 29 Mar. 1984, Davidse & Herrera 26275 below junction of the Macungo River, 24 Jul. 1967,
(MO); Rincón de Osa, ridge between Quebrada Proctor 28322 (F).
Aparicio and Quebrada Aguabuena, 7 Oct. 1984, REPUBLICA DOMINICANA. SAMANA: Samana
Grayum et al. 3989 (UC); Monteverde, Turrialba, 26 Peninsula, vic. Sanchez, 29 Nov.-12 Dec. 1920, Abbott
Apr. 1980, Koptur 322 (UC); Osa Peninsula, ca. 20 km S 141 (US); Península de Samana, slope of Pan de
of Rincón de Osa, 18 Jul. 1967, Mickel 2784 (NY, UC); Azúcar, 4 May 1930, Ekman 14861 (S). LA VEGA:
Turrialba, near Interamerican Inst. Scamman 6166.5 Cotuy, 28 Jan.-7 Feb. 1921, Abbott 762 (US). Sánchez
(GH). SAN JOSE: Bajo La Hondura, 4 Jul. 1972, Mc Ramirez, 1 km from Hernando Alonso on road to
Alpin et al. 1186 (F); valley La Palma, above the La Palmarito, on side of Loma El Diviso, 28 Jan. 1981,
Hondura, ca. 9 km NE of San Jerónimo, 23 Mar. 1973, Mejía et al. 10460 (MO).
Stolze 1434 (F); Stolze 1446 (F). CARTAGO: ravine of PUERTO RICO. N of Bayamón, 26 Apr. 1944, Wagner
Río Reventazón, Interamerican Institute Turrialba, 1 s.n. (US); slopes of El Yunque, 28 May 1944, Wagner
Aug. 1961, Brown 211 (US); Cartago, May 1954, s.n. (US).
Carpenter 639 (F, US). LIMON: banana and cacao VIRGIN ISLANDS. St. THOMAS: Charlotte Amalia,
plantations between Siquerres and the Río Pacuaré, 20- 17-18 Jan. 1899, Millspaugh 546 (F).
22 Dec. 1969, Burger & Liesner 6942 (F). ISLA DEL LESSER ANTILLES. LEEWARD ISLANDS: Antigua,
COCO: Chatham Bay, 13 Apr. 1965, Jimenez 3165 (F, 4-16 Feb. 1913, Rose et al. 3336 (F, GH, US).
NY). Without exact locality: Carrillo, 400 m, 18 Jun. Guadeloupe, s.f, Blanchard s.n. (F); 1893, Duss 4104 (F).
1909, Brade & Brade 715 (S). Bois de Tremedal, San WINDWARD ISLANDS: Montserrat, 18 Feb. 1907,
Ramón, Apr. 1913, Tonduz 17575 (F, GH, S, US). Shaffer 752 (F, US). Saint Lucia, 21 Jun. 1945, Beard
PANAMA. BOCAS DEL TORO: Laguna Chiriquí, 1109 (F, US).
Bocas del Toro, Nov-Dec. 1885, Hart 49 (US); along COLOMBIA. MAGDALENA: Sierra Nevada de Santa
road to Chiriquí Grande, 24 Jun. 1986, McPherson & Marta, 28 Aug. 1972, Kirbride 1957 (UC). GUAJIRA:
Allen 9620 (UC). CHIRIQUI: vic. of El Boquete, 5 Feb. Serrania de Macuira, Arroyo Chichimahu, 2 Aug. 1975,
1918, Cornman 813 (US); district Boquete, Bajo Chorro, Sugden 13 (UC). NORTE DE SANTANDER: camp 84
11 Jan. 1938, Davidson 106 (F, S). PANAMA: Cerro on pipeline, 16 Sep. 1946, Foster & Foster 1714 (GH);
Campana, on road to Sun Lin, 3 Jan. 1973, Kennedy et road from Pamplona de Toledo, crossing divide
al. 2039 (MO); 12-16 km above Pan-Am hwy. from El between Río La Teja and Río Mesme, 27-28 Feb. 1927,
Llano to Carti-Tupile, 5 May 1973, Kennedy et al. 3114 Killip & Smith 19989 (US). CHOCO: 0.5-2.5 km N of
(MO); on trail to Cerro Campana, 23 Aug. 1967, the Inderena camp, 3 Mar. 1971, Lellinger & Sota 547
Kirkbride & Hayden 284 (MO). CANAL ZONE: W of (US). ANTIOQUIA: Mun. Caramanta, 9.8 km from
Río Chagres, opposite Bohio, 12 Feb. 1911, Maxon 4780 Caramanta to Supia, Cerro Viringa, 15 Oct. 1988,
(S, US); Barro Colorado Island, Pearson trail, 20 Nov. Betancur et al. 1051 (MO); road tol Villa Arteaga, 4-8
1931, Shattuck 541 (F). COLON: bridge over Río Apr. 1947, Hodge 7050 (F, GH); Anori, Providence
Guanche, 0-15 m, 27 May 1980, Antonio 4814 (MO); hydroelectrical station, 6 Jun. 1971, Soejarto 2901 (F).
Santa Rita ridge, 26 May 1987, McPherson 10989 (MO, SANTANDER: upper Río Lebrija valley, NW of
USM); Santa Rita ridge, 25 Sep. 1980, Sytsma 1315 Bucaramanga, 29 Dec. 1926, Killip & Smith 16284 (GH,
US). CALDAS: Mun. Salamina, Río San Lorenzo, 12 Gualaquiza, 26-27 Apr. 1973, Holm-Nielsen et al. 4620
Aug. 1975, Acosta-Arteaga 981 (AAU). (AAU, USM). AZUAY: between Cruzpamba and
CUNDINAMARCA: Cordillera Oriental, Sebastopol, 3 Loma de Canela, in region of Río Sadacray, s.f.,
Sep. 1944, Little & Little 8603 (F, GH, US); Cordillera Steyermark 52959 (F, US). SANTIAGO-ZAMORA:
Oriental, at railroad station Tablanca, 31 Dec. 1944, Región Oriental, between Río Sordo and La Esperanza,
Little & Little 9151 (F, US); Santandercito, near Río 13 Feb. 1944, Acosta-Solis 7318 (F). GALAPAGOS
Bogotá, Sep. 1941, Uribe 202 (F). EL VALLE: Cisneros, ISLANDS: Indefatigable Island, 6 mi N of Academy
5 May 1939, Killip 35570 (US). CAUCA: Cordillera Bay, 6 Apr. 1930, Svenson 114 (UC). Without locality:
Occidental, Cerro Munchique, Hoya del Río Tambito, slope of western Cordillera, Rimbach 79 (F); Río Cristal,
16 Jul. 1939, Pérez Arbelaez & Cuatrecasas 6244 (F, US). 26 Oct. 1933, Schimpff 301 (F, MO); 1857-1859, Spruce
Without exact locality: 9 Aug. 1910, Mayor 119 (US); 5249 (BM).
Santa Marta, 1898-1901, H. Smith 1013 (F, S). PERU. SAN MARTIN: Mariscal Cáceres, Campanilla,
VENEZUELA. ZULIA: Dist. Mara, NW side of Cerro Mashuyacu, 12 Aug. 1970, Schunke V. 4225 (GH, US);
Negro, 28 May 1980, Steyermark et al. 122672 (UC). Rioja, Río Negro, 20 Jan. 1965, Soukup 5146 (GH).
FALCON: Cerro Santa Ana, S de Santa Ana, 24 Jan. LORETO: 12 km SW of Iquitos, 18 Jul. 1972, Croat
1966, Steyermark & Braun 94264 (US). LARA: Dist. 18257 (F, UC); Río Ampiyacu, Brillo Nuevo-Río
Moran, Quebrada El Guairon, 2 km Guarico, 1 Apr. Yaguasyacu, 13 Mar. 1981, Davis et al. 900 (F);
1983, Rivero & Ortega 283 (UC). YARACUY: Sierra de Yurimaguas, lower Río Huallaga, 23 Aug.-7 Sep. 1929,
Aroa, 15 km NW of Cocorote and 1 km SW of Los Killip & Smith 27668 (F, S, US); Mishuyacu, near
Cruceros, 4 Apr. 1980, Liesner & Gonzalez 10052 (UC). Iquitos, Oct.-Nov. 1929, Klug 238 (F, NY, US).
SUCRE: Dist. Sucre, El Guayabito, along Río Guayabo, HUANUCO: Sinchono, Fundo Chela, 3 Aug. 1948,
20-22 Nov. 1981, Davidse & Gonzalez 19111 (MO, UC). Aguilar 943 (USM); Pozuzo, 20-22 Jun. 1923, Macbride
MIRANDA: Colonia Tovar, 1854-1857, Fendler 230 4581 (F); Huánuco, near confluence of Río Cayumba
(BM, F); between Panaquire and El Peñón, 20 Sep. with Río Huallaga, 10 Oct. 1936, Mexia 8272 (BM, F, S,
1977, Fernández 3281 (F); Distrito Paez, fila La Tigra, 18 UC). PASCO: Oxapampa, 5 km SE of Oxapampa, 9-11
km SW of Cupira, 2-7 Sep. 1977, Ortega & Gonzalez 413 Dec. 1982, D.N. Smith 2917 (MO); Palcazú valley, near
(MO); Ortega & Gonzalez 424 (UC). BOLIVAR: Río the confluence of Río Palcazú and Río Iscozacin, 23
Caura from foot of gorge below Salto Para, 250 m, 14 Apr. 1983, D.N. Smith 3874 (MO, UC). JUNIN: Puente
Aug. 1985, Horner et al. 237 (MO). Perené, 31 Oct. 1954, Coronado 253 (GH, UC, US); E of
TRINIDAD. Lalaja, Blanchisseus road, 28 Sep. 1969, Quimiri bridge, 1-3 Jun. 1929, Killip & Smith 23896 (F,
Fay 364 (BM). Mararas bay, 1903, Othmer 423 (S); GH); Colonia Perené, 14-22 Jun. 1929, Killip & Smith
Aripo, 2 Oct. 1953, Pickering s.n. (BM). Without 24917 (F); Pichis trail, between Meriatiriani and
locality: 1877-1880, Fendler 117 (BM, UC, US). Yessup, 28 Jun.-8 Jul. 1929, Killip & Smith 26221 (US);
TOBAGO. Near Memma forests, 9 Nov. 1932, La Merced, 28 Jun. 1982, León 242a (USM);
Broadway 9062 (BM). Chanchamayo, Feb. 1939, Soukup 1100 (F); La Merced,
GUYANA: Kamakusa, 19 Dec. 1922, Lang & Persaud Aug. 1947, Soukup 3403 (BM). CUSCO: Prov.
398 (F). Convención, Río Apurimac, 20 minutes below Puerto
ECUADOR. ESMERALDAS: Río Onzolé, 3 Sep. 1980, Capiro, 9 Jul. 1981, Davis et al. 1302 (F, GH); Potrero, 8
Holm-Nielsen et al. 25741 (AAU); Río Santiago, at km W of Quillabamba, 7 Oct. 1956, Tryon & Tryon
Concepción, 4 Sep. 1980, Holm-Nielsen et al. 25980 5393b (BM, F, GH, US). PUNO: Carabaya, Hacienda
(AAU). IMBABURA: Lita, 4 Oct. 1980, Maas & Cobb Palmera, Inambari, 4 Mar. 1965, Vargas 16149 (GH).
4681 (QCA). PICHINCHA: road Aloag-Santo MADRE DE DIOS: Tambopata, SSW of Puerto
Domingo, Chitoa, 28 Oct. 1980, Holm-Nielsen et al. Maldonado at effluence of Río La Torre, 25 Apr. 1980,
28001 (QCA). NAPO: Cantón Aguarico, Parque Barbour 4967 (F, MO); Prov. Manu, Río Palotoa,
Nacional Yasuní, laguna Garza Cocha, 22 Sep. 1988, tributary of Alto Madre de Dios, NW of Shintuya, 26-
Cerón & Gallo 4934 (MO); 1.1 km of Río Conejo, on 28 Aug. 1978, Foster & Terborgh 6759 (F); Cocha Cashu
road to Lago Agrio, 31 Mar. 1972, Dwyer & Mac Bryde Camp, Río Manu, Parque Nacional del Manu, 16 Oct.
9769 (QCA); creek 3 km NW of Borja, 20 Sep. 1980, 1979, Gentry et al. 26784 (F).
Holm-Nielsen 26280 (QCA). COTOPAXI: Quevedo- BOLIVIA. PANDO: SW of Cobija on the Río
Latacunga hwy., 17 Dec. 1976, Boeke 510 (QCA, UC). Naraueda, 1 Aug. 1982, Sperling & King 6456 (F, UC).
PASTAZA: from Ceilan to Río Cononaco, 6 Jun. 1980, LA PAZ: Prov. Sur Yungas, Chungamayo, La Sirena,
Brandbyge & Asanza 31633 (AAU); Puyo-Arajuno road, 31 Aug. 1920, Asplund 283 (S); Uchimachi, 22 Aug.
1-5 km SW Diez de Agosto, 4 Mar. 1980, Harling & 1894, Bang 2395 (F); Murillo, valle de Zongo, Cahua, 7
Andersson 16896 (GB); Hacienda San Antonio, 2 km N Apr. 1979, Beck 1219 (F); Polo Polo bei Coroico, 1912,
de Mera, 5-19 Mar. 1985, Baker et al. 5613 (MO). Buchtien 3532 (F, S, UC); Hacienda Casana, sobre el
MORONA-SANTIAGO: Pachicutza, km 140 rd. Loja- camino a Tipuani, 6 Oct. 1922, Buchtien 7078 (NY, S,
UC); Sur Yungas, basin of Río Bopi, near Calisaya, 1-22 broadly auriculate, apices acuminate, clathrate
Jul. 1939, Krukoff 10203 (F, GH, MO); Bopi river valley, with differentiated margins mainly at the base of
6 Aug. 1921, Rusby 721 (F, US); Larecaja, 6 km N of the scale, cells narrowly oblong along the main
Consata, 15 Dec. 1981, Solomon et al. 6579 (UC).
axes of the scale, cells at the base of the scale
BRAZIL. RONDONIA: Rodovia 399, a 13 km de
Vilhena, 4 Nov. 1979, Vieira et al. 897 (F).
irregularly arranged, cell lumen translucent,
AMAZONAS: near Livramento, 12 Oct.-6 Nov. 1934, central cell walls 12-18 µm thick, marginal cell
Krukoff 6756 (MO, S). walls 9-12 µm thick. Phyllopodia 1 mm long, 1-
1.5 mm wide, tightly closed less than 2 mm
Campyloneurum brevifolium is characterized by apart. Leaves 15-50 cm long; petiole
its pattern of venation of irregularly divided stramineous, 0.5-3 cm long; lamina linear, rarely
areoles, and by the presence of costal scales. narrow lanceolate, bases and apices attenuate,
Campyloneurum brevifolium is the correct 0.3-1 cm wide, herbaceous-chartaceous, margins
basyonym for the taxon here defined. It replaces slightly revolute, sinuate, cartilaginous,
C. latum, a name which has been used commonly indument of inconspicuous, simple and
in the literature. bicellular hairs, spreading abaxially, along the
Some problems arose in distinguishing this costa; stomata polocytic; costa prominent,
species from Campyloneurum phyllytidis, mainly usually castaneous abaxially, primary veins
because of the presence of similar venation, and inconspicuous, with 1-2 primary areoles between
similar shape and size of leaves. Lellinger (1988) the costa and margin, with one excurrent veinlet;
suggested that the pattern of venation of C. sori subterminal, paraphyses not seen, spores 60-
phyllitidis is an intermediate state of the "loosely 65 µm long, 35-40 µm wide. Fig. 26.
organized" pattern of venation found in C.
brevifolium. Here, the pattern of venation in C. This species is found in central eastern Brazil,
brevifolium is interpreted as specialized, because where it grows between 800 and 1500 m
it is more complex than that of C. phyllitidis. elevation. The species occurs in campo rupestre
These species probably belong to different vegetation of the Brazilian Planalto.
lineages in the genus. Pattern of venation
appears is a good character to define EXAMINED SPECIMENS: BRAZIL. GOIAS: SW of
Campyloneurum brevifolium as a distinct species. Brasília D.F. 20 Feb. 1966, Irwin et al. 13057 (F); Serra
Although, still in some cases symmetrical and dos Pirineus, 50 km N of Corumbá de Goiás on road to
asymmetrical divided primary areoles occur Niquelândia, valley of Rio Maranhão, 25 Jan. 1968,
within an individual. Irwin et al. 19184 (F); ca. 30 km N of Alto do Paraíso,
Chapada dos Veadeiros, 23 Mar. 1971, Irwin et al.
In the Caribbean region, leaf shape can be
33063 (F, NY); Serra dos Pirineus, ca. 20 km E de
used to differentiate C. brevifolium from C.
Pirenópolis, 16 Jan. 1972, Irwin et al. 34302 (F, NY).
phyllitidis, as was shown by Proctor (1977, 1985). MINAS GERAIS: Serra do Espinhaço, rocky hillside
Campyloneurum brevifolium has broader leaves ca. 7 km N of São João da Chapada, road to Inhaí, 29
than C. phyllitidis, although this character seems Mar. 1970, Irwin et al. 28607 (F); Serra do Espinhaço ca.
restricted to specimens from the Antilles. 12 km W of Barão de Cocais, 27 Jan. 1971, Irwin et al.
Campyloneurum brevifolium is closely related 29288 (F, NY).
to C. nitidissimum, C. pascoense, and C.
tucumanense. Campyloneurum centrobrasilianum is
characterized by having stem scales with a broad
base and the cells irregularly arranged at the
12. Campylonerum centrobrasilianum Lellinger, base of the scale, but regularly arranged along
Amer. Fern J. 78: 16-17. 1988. Type. Brazil, the main axis. This species belongs to the
Minas Gerais, Viçosa, Kuhlmann 1898 Campyloneurum angustifolium group.
(holotype, US!).
Stem creeping, not pruinose, 2-3 (-4) mm
wide. Stem scales dark brown in mass, 3-4 mm 13. Campyloneurum chlorolepis Alston, Bull. Jard.
long, 0.5-1 mm wide, narrowly ovate, bases Bot. Etat 27: 56. 1957. Type. Colombia:
Caldas, Chinchina, Pereira-Manizales, 1400 CUNDINAMARCA: Mun. San Cayetano, road to

m, s. f., Køie 5208 (holotype C, photo BM!). Hondura, 10 May 1977, Acosta 1198 (B); Salto de
Polypodium angustifolium Sw. var. heterolepis Tequendama, 1-3 Oct. 1938, Cuatrecasas 73 (F);
Santardecito, Sep. 1941, Uribe 201 (F); Bogotá, at
Rosenst., Mem.Soc. Sci. Nat. Neuchâtel 5: 54.
railroad station Tablanca, 21 Dec. 1944, Little & Little
1914. Type. Colombia:Antioquia, near 9143 (F). VALLE: Hoya del Río Sanquininí, 10-20 Dec.
Angelopolis, ca. 1800 m, Mayor 140 (holotype, 1943, Cuatrecasas 15428 (F); Hoya el Río Cali, Pichinde,
P!; isotypes S!, UC!, US!, photo of P, BM!). Morro Pelado, 17 Oct. 1944, Cuatrecasas 18147 (F, S,
Campyloneurum heterolepis (Rosenst.) Lellinger, US); Mun. Sevilla, vía Sevilla-Barragán, 26 Sep. 1981,
Amer. Fern J. 67: 58. 1977. Silverstone 697 (MO). WIithout locality: Lindig 358 (B).
Stem creeping, rarely pruinose, 3-4 (-5) mm VENEZUELA. BARINAS: road Barinitos-La Soledad,
wide. Stem scales whitish, 5-7 mm long, 0.75-1 Stergios 1631 (UC). MIRANDA: Tumerito a Ocumare
mm wide, narrowly ovate, bases auriculate, del Tuy, 15 Oct. 1939, Williams 12410 (F, UC).
PORTUGUESA: Guanare, San José de la Montaña, 27
apices acuminate, non-clathrate, the cells
Sep. 1981, Ortega & Stergios 1290 (F, UC); Ortega &
narrowly oblong along the main axes of the Stergios 1320 (UC); Sucre-La Divisoria de la
scale, cell walls usually well defined, but without Concepción, 23-26 Oct. 1985, Ortega et al. 2798 (MO,
pigmentation, 8 µm wide, lumen translucent. UC); Sucre, Palo Alzao-Guayabital, 3 Nov. 1981,
Phyllopodia 1 mm long, 2-3 mm wide, tightly Ortega & Aymard 1437 (UC); San José de la Montaña, 8
closed less than 2 mm apart. Leaves 40-85 cm Nov. 1982, A.R. Smith et al. 1075 (MO, UC); dist.
long; petiole stramineous or brownish, 4-8 (-10) Ospino, 20 km W of La Estación, 10 Nov. 1982, A.R.
cm long; laminae narrowly lanceolate or linear- Smith et al. 1159 (MO, UC); 17.8 km de la Estación, N
de Ospino, 1 Nov. 1982, Steyermark et al. 126976 (AAU,
lanceolate, bases narrowly cuneate or attenuate,
MO, UC). LARA: dist. Iribarren, vía Guamasire, 16
apices acuminate, 1-3 (-5) cm wide, coriaceous, Jul. 1983, Rivero & Ortega 346 (UC); dist. Jiménez,
margins slightly revolute, cartilaginous, apices Parque Nacional Yacambú, Quebrada El Blanco, 24
acute to acuminate, indument of inconspicuous, Oct. 1982, Davidse & González 21076 (MO). Colonia
simple and multicellular hairs, scattered Tovar, 1854-1855, Fendler 225 (MO).
abaxially; stomata polocytic; costa prominent, ECUADOR. CARCHI: Chical, Colombian side of Río
usually castaneous abaxially, primary veins San Juan, along trail leading to Altaquer, 26 Feb. 1983,
inconspicuous, however when dry the color of Barfod & Blicher-Mathisen 41574 (QCA). PICHINCHA:
road Nanegalito-Pacto, Tulipe, 21 Jul. 1980, Holm-
the veins are darker than the leaf tissue, 5-8 mm
Nielsen et al. 24498 (AAU, QCA, USM). NAPO: Ceilán,
apart, 50-60o divergent from the costa, path from Ceilán to Río Cononaco, 6 Jun. 1980,
transverse secondary veins forming 2-4 primary Brandbyge & Asanza 31695 (AAU, USM); Río Wai si
areoles between the costa and margin, excurrent ayá, a northern tributary of Río Aguarico, about 6 km
veinlets 3-4 free or anastomosed to the transverse upriver from San Pablo, 10 Aug. 1980, Brandbyge &
veins forming secondary areoles, sometimes Asanza 32757 (AAU, USM); Añangu, Río Napo, 30
Jun. 1983, Lawesson et al. 39660 (AAU); Lawesson et al.
asymmetrically; sori subterminal, paraphyses not
39662 (AAU); Cantón Napo, Zatzayacu, 22 Mar. 1935,
seen, spores 60 µm long, 30-35 µm wide. Fig. 25.
Mexia 7059 (F, US). LOS RIOS: Río Palenque
Biological Station, km 56 road Quevedo-Santo
This species is found from Colombia, Domingo, 30 Mar. 1980, Dodson & Gentry 10045 (MO).
Venezuela to Bolivia and central Brazil, where it PASTAZA: Cantón Pastaza, 50 km SSE de Curaray, 1-
grows between 260 m and 3000 m elevation. It is 19 Oct. 1990, Rubio & Coba 878 (MO); Río Bobonaza,
mainly distributed along the eastern side of the between Cachitama and the outleet of Río Bufeo, 19
Andes. Jul. 1980, Øllgaard et al. 34678 (AAU, USM);
Destacamento Chiriboga and Apachi Entza, 24 Jul.
REPRESENTATIVE SPECIMENS: COLOMBIA. 1980, Øllgaard et al. 35177 (AAU, USM); Río Bobonaza,
ANTIOQUIA: Mun. Amalfi, 1-4 km from Amalfi to Destacamento Cabo Pozo, 20 Jul.1980, Øllgaard et al.
Rumezón, 27 Sep. 1988, Betancur et al. 738 (MO); Río 34878 (AAU, USM), Øllgaard et al. 34883 (AAU, USM).
Verde, 12 Jul. 1880, Kalbreyer 1770 (B, S). EL ORO: 11 km W of Pinas on new road to Santa Rosa,
SANTANDER: vic. of Las Vegas, 21-23 Dec. 1926, 8 Oct. 1979, Dodson et al. 9148 (MO). MORONA-
Killip & Smith 15994 (BM, F). CALDAS: Cannan, S of SANTIAGO: Bomboiza, 17 km SE Gualaquiza, 25 Jul.
Salento, 31 Jul. 1922, Pennel 9079 (US). 1985, Palacios 562 (MO); 35 km NE of Montalvo, 2-12
Jul. 1989, Zak & Espinoza 4474 (MO).
PERU. AMAZONAS: Huampaní, 18 Jul. 1974, Kayap not pruinose. Stem scales dark brown in mass,
1206 (MO). SAN MARTIN: Tingo María, along (1-) 1.2-2 (-3) mm long, 0.5-0.7 mm wide,
Huallaga about 20 km from Tingo María on road to narrowly ovate, bases peltate or slightly
Huánuco, 30 Oct-19 Feb. 1950, Allard 21974 (US); km
auriculate, apices acuminate, clathrate, with
21-22 of Tarapoto-Yurimaguas road, 7 Aug. 1986,
Knapp et al. 7883 (MO, NY, USM); San Martín, km 28 of
differentiated margins, cells oblong, marginal
Tarapoto-Yurimaguas road, 17 Aug. 1986, Knapp 8037 cells with transparent lumen, central cells with
(MO); Prov. Mariscal Cáceres, dist. Uchiza, NW of brownish lumen, cell walls 10-13 µm thick.
caserío Nuevo Progreso, 25 Jun. 1969, Schunke V. 3241 Phyllopodia 1-2 mm long, 1.5-2 mm wide, 2.5-3
(F, GH, NY, US, USM); dist. Tocache Nuevo, cm apart. Leaves 17-25 cm long; petiole
Quebrada de Almendras, 2 Sep. 1970, Schunke V. 4453 stramineous, 1.5-4 cm long, slightly ranurate
(F, GH, US); Lamas, Alonso de Alvarado, fundo Las adaxially, convex abaxially, glabrous; lamina
Flores, E of San Juan de Pacayzapa, 11 May 1973, narrowly lanceolate, bases and apices attenuate,
Schunke V. 6243 (NY). LORETO: Río Corrientes at the
1-3 cm wide, herbaceous-chartaceous, margins
Ecuador border, between Teniente López and Puesto
Avanzado, 4 Apr. 1977, Gentry et al. 19057 (F, MO, cartilaginous, slightly sinuate, indument of
USM). HUANUCO: Tingo María, W side of Río inconspicuous simple hairs; stomata polocytic;
Huallaga, 1 Jul. 1977, Solomon 3389 (MO). PASCO: costa prominent on both sides of the lamina,
Colonia Perené, 14-22 Jun. 1929, Killip & Smith 25089 indument of scarce scales, similar to those on the
(F). JUNIN: Tarma, Ruiz 11 (B); Chanchamayo valley, stem, primary veins slightly prominulous on
Schunke 124 (US); s.l., Soukup 1102 (F). AYACUCHO:
both surfaces, 65-70o divergent from the costa,
Río Apurímac valley, near Kimpitiriki, 10-11 May
1929, Killip & Smith 22890 (NY, US). CUSCO: flexuous, secondary transverse veins forming 3-4
Pilcopata, Atalaya, Paucartambo, 15 Jan. 1987, Núñez primary areoles between the costa and margin, 1-
6866 (MO). MADRE DE DIOS: Manu, Atalaya, 3 excurrent veinlets, primary areoles sometimes
Hacienda Amazonía, Foster & Wachter 7417 (USM). symmetrically divided; sori subterminal or
BOLIVIA. LA PAZ: Yumpasa, 10 Jan. 1902, Williams terminal, paraphyses not seen, spores 45-50 µm
1067 (GH, US). BENI: Ballivian, Serranía del Pilón long, 30-35 µm wide. Fig. 34.
Lajas, 13-15 km de Yucumo, 20 May 1989, D.N. Smith
et al. 13294 (F).
This species is found only in Venezuela,
BRAZIL. MATO GROSSO: Santa Anna da Chapada,
14 Oct. 1902, Malme s.n. (S).
between 1200-1800 m elevation, where it grows
as an epiphyte.
Campyloneurum chlorolepis is a very distinctive
REPRESENTATIVE SPECIMENS: VENEZUELA.
species characterized by the whitish non-
TACHIRA: Córdoba, fila de Paramito, N of Mesa de
clathrate stem scales. The width of leaves in C. Tigre, 16 Nov. 1982, Davidse & Gonzalez 22407 (MO,
chlorolepis varies markedly within and between UC). PORTUGUESA: Sucre, La Divisoria de la
individuals. Concepción, 23-26 Oct. 1985, Ortega et al. 2749 (MO,
This species belongs to the Campyloneurum UC). FALCON: arriba en La Chapa, Sierra de San Luis,
amphostenon group. 18 Jan. 1979, Werff van der et al. 189 (UC). TRUJILLO:
en las cercanías de Vitú, Cerro El Zamuro, Quebrada
El Limón, 23 Nov. 1984, Ortega & Werff van der 2292
(UC).
14. Campyloneurum chrysopodum (Klotzsch) Fée,
Gen. Filic. 258. 1852.
Campyloneurum chrysopodum is characterized
Polypodium chrysopodum Klotzsch, Linnaea 20:
by long creeping stems less than 3 mm wide, by
401-402. 1847. Type. Venezuela: Monagas,
stem scales less than 3 mm long, and by well-
Cumanacoa, Moritz 134 (holotype B!; isotypes
spaced leaves. It belongs to the C. repens group.
BM!, K!, photo BM!).
Campyloneurum fendleri T. Moore, Index Fil. 224.
1861. Type.. Venezuela: Colonia Tovar, 1854-
15. Campyloneurum coarctatum (Kunze) Fée, Gen.
1855, Fendler 228 (holotype, not found;
Filic. 258. 1852.
isotypes B!, BM!, K!, MO!).
Polypodium coarctatum Kunze, Linnaea 9: 39.
Stem 1-2 mm wide, stramineous or greenish,
1834. Type.. Peru: Huánuco, Cucheros, Jul.
1829, Poeppig s. n. (holotype, LZ, probably (MO, UC); valley of Río Piarnasta, about 5 mi E of El
destroyed; isotypes P!, W!; photo of W, BM!). Boquete, 9-22 Feb. 1918, Killip 5142 (GH, MO, S).
Stem long-creeping, greenish, black or dark DARIEN: N Punta Guayabo, 21 Apr. 1980, Antonio &
Hahn 4319 (MO); Serranía del Darien, trail from Cerro
stramineous, 2-3 mm wide. Stem scales dark
Mali to Río Pucuro, 20 Jul. 1976, Gentry et al. 16827
brown or brown in mass, 1.5-4 mm long, 0.3-0.7 (MO); Rancho Frío, 9 Aug. 1986, Mc Donagh et al. 594
(-1) mm wide, linear or narrowly ovate, (MO).
pseudopeltate, clathrate, cells oblong-elongate, COLOMBIA. CHOCO: Mun. San José del Palmar,
cell walls (8-) 10-15 µm wide. Phyllopodia 2 mm basin of Río Torito, 7 Mar. 1980, Forero et al. 6861 (MO);
long, 2-3 mm wide, 1-1.5 cm apart. Leaves 45-85 W slope S ridge of Cerro Mecana, 7 Jan. 1984, Juncosa
cm long; petiole stramineous or dark 1762 (MO, UC). Llanos de San Martín, Villavicencio,
stramineous, (10-) 13-28 cm long, ranurate Stübel 639 (B).
abaxially, indument of caducous scales similar to GUYANE. Haut Tapoc: Crique Alice, 4 Apr. 1977,
Cremers 4625 (CAY); Monts Atachi Bacca, Granville 745
those on the stem; lamina broadly elliptic to
(CAY); Tumuc Humac, SE of Toukouchipann', 21 Aug.
ovate elliptic, bases narrowly cuneate or 1972, Granville 1323 (CAY); sources de la Mana, Monts
acuminate, sometimes shortly decurrent, apices Galbao, 11 May 1973, Granville 1614 (CAY); Grand
acuminate or subcaudate, (6-) 8-13 cm wide, Tamouri river, affluent of Camopi, 15 Mar. 1974,
herbaceous-chartaceous, margins slightly Granville 2124 (CAY); Petit Tamouri river, 19 Mar.
sinuate, cartilaginous, indument of 1974, Granville 2174 (CAY); Haut Oyapock, crique
inconspicuous hairs abaxially, hairs 50-70 µm Takululi, 17 Jul. 1975, Granville 2470 (CAY).
long; hydatodes sometimes present adaxially; ECUADOR. PICHINCHA: Santo Domingo de los
Colorados, Dodson & Duke 7713 (MO). PASTAZA:
stomata polocytic; costa prominent on both
Montalvo, on the Río Bobonaza, 28 Jul. 1980, Øllgaard
surfaces, primary veins prominulous to et al. 35411 (AAU, QCA, UC). MORONA-
prominent at same degree on both surfaces, SANTIAGO: Pachicutza, km 140 on road Loja-
straight, (60o-) 65-70o divergent from the costa, Gualaquiza, 26-27 Apr. 1973, Holm-Nielsen et al. 4620
5-6 mm apart, transversal veins forming 9-19 (AAU, F, MO). NAPO: Nuevo Rocafuerte, al SW de la
primary areoles between the costa and margin, población, 2 Mar. 1981, Jaramillo & Coello 4629 (AAU);
Añangu, Río Napo, 6 Jul. 1983, Lawesson et al. 39775
primary areoles undivided, (1-) 2 excurrent
(AAU, QCA); Añangu, Parque Nacional Yasuní, 30
veinlets in each areole; sori subterminal, May-21 Jun. 1982, Øllgaard et al. 38827 (AAU, UC).
paraphyses not seen, spores not seen. Fig. 34. ZAMORA-CHINCHIPE: road La Saquea Yacuambi, 9
Apr. 1985, Harling & Andersson 23859 (QCA); 4 km W
This species is found from Costa Rica to of Panguintza, 14 Apr. 1985, Harling & Andersson 24156
Bolivia, and Amazonian Brazil, where it is (QCA). Without locality: Andes Quitenses, Spruce
usually found between 100 m and 2000 m 5644 (BM, W).
elevation, in forested areas. PERU. CAJAMARCA: Santa Cruz, Catache, upper Río
Zaña valley, ca. 5 km above Monteseco on path to
Chorro Blanco, 16-18 Mar. 1986, Dillon et al. 4355 (F,
REPRESENTATIVE SPECIMENS: COSTA RICA.
GH); Santa Cruz, ENE of Monteseco, 7 May 1987,
PUNTARENAS: San Vito de Coto Brus to Ciudad
Santisteban & Guevara 31 (F). SAN MARTIN: Tarapoto,
Neily, 11 Jul. 1985, Hammel 14163 (UC). CARTAGO:
1855-1856, Spruce 4646 (BM, NY, W). LORETO: Río
Navarro, 6-8 mi SW of Cartago, 24 Jul. 1924, Stork 4263
Marañón, above Saramuro, 22 Jan. 1979, Diaz & Ruiz
(UC). W de lago Dabagri, 4 Nov. 1984, Gómez et al.
875 (MO); Yurimaguas, 23 Aug.-7 Sep. 1929, Killip &
23178 (UC). Tucurrique, Torres de las Vueltas, Dec.
Smith 27668 (F, MO, NY, S, US); Balsa Puerto, 28-30
1898, Tonduz 12914 (B).
Aug. 1929, Killip & Smith 28427 (F, US), Killip & Smith
PANAMA. PANAMA: 16 km above Pan-am hwy.
28472 (NY, US); Santa Rosa, lower Río Huallaga,
from El Llano to Carti Tupile, Kennedy & Dressler 2921
below Yurimaguas, 1-5 Sep. 1929, Killip & Smith 28854
(MO); 6-7 mi from Pan-am. hwy. on El Llano-Carti
(F); above Pongo de Manseriche, left bank of Río
road, 26 Feb. 1982, Knapp & Mallet 3859 (MO, UC); El
Santiago, 23 Nov. 1931, Mexia 6141a (F, MO, UC); from
Llano-Carti road, 21 Mar. 1975, Mori & Kalluncki 5131
Marañon valley to Iquitos, crossingis Santiago-
(MO). CHIRIQUI: shoulder of El Barú, above Bajo
Morona, at Pongo de Manseriche, 7 Dec. 1924,
Grande, Folsom et al. 2138 (MO); along Río Colorado,
Tessmann 4885 (B). HUANUCO: Sinchono, 3 Aug.
11 Jul. 1983, Hamilton & Krager 3818 (MO); along Río
1948, Aguilar 943 (F, USM); Tingo María, at Las cuevas
Colorado, 17 Mar. 1983, Hamilton & Stockwell 3534
de los Pavos, 30 Oct. 1949-19 Feb. 1950, Allard 20527
(US); Tingo María-Pucallpa, 1971, Ellenberg 3886 (GH); margins cartilaginous, sometimes slightly
Huánuco, gorge of Río Chinchao, 11 Sep. 1956, Tryon revolute; costa prominent, indument of scales
& Tryon 5302 (F, GH, NY, UC, US). JUNIN: E of similar to those on the stem, primary veins
Quimiri bridge, near La Merced, Killip & Smith 23896
(F, GH); La Merced, 28 Jun. 1982, León 242a (USM); prominulous, usually stramineous, (50o-) 55-60o
Chanchamayo, Aug.-Oct. 1923, C. Schunke 167 (US); divergent from the costa, 5-7 mm apart,
Chanchamayo, 24-27 Sep., C. Schunke 512 (F); secondary veins sometimes slightly
Chanchamayo, Jul. 1929, C. Schunke 978 (F); La prominulous, same color as the leaf tissue,
Merced, Chanchamayo, Feb. 1939, Soukup 1100 (F); forming 5-6 primary areoles between costa and
Tarma, Agua Dulce, 16 Apr. 1948, Woytkowski 37025 margin, regularly or irregularly divided,
(MO, UC). UCAYALI: Coronel Portillo, Bosque von
excurrent veinlets 1-2 in each secondary areole,
Humboldt, Young & Salazar 1015 (F, MO). MADRE DE
DIOS: Parque Nacional Manu, Cocha Cashu, 7 Sep.
sometimes free veinlets along the margin; sori
1984, Foster 84-16 (F). subterminal or medial, paraphyses not seen,
BOLIVIA. COCHABAMBA: Antahuacana, Jun. 1909, spores 45-50 µm long, 28-30 µm wide. Figs. 1 a;
Buchtien 2153 (UC). 14 e; 29.
BRAZIL. ACRE: Mun. Canamari Amazonas, Rio
Jurúa, N of Cruzeiro do Sul, lake Cigana, S of Porto This Andean species is known from Colombia
Alvaro Mestrinho, 22 Aug. 1986, Croat 62525 (MO); and Ecuador, where it grows above 2400 m
Mun. Canamari Amazonas, vic. of Floresta, 23 Aug. altitude. It is found in the paramo and in
1986, Croat 62550 (MO). AMAZONAS: Juruá Mury,
montane forest vegetation types, in crevices of
Jun. 1901, Ule 5607 (B).
rocks or rarely as an epiphyte.
Campyloneurum coarctatum is recognized by its
REPRESENTATIVE SPECIMENS: COLOMBIA.
linear and clathrate stem scales. The leaf is TOLIMA: Boquerón de Quindiu, 27 Mar. 1939, Alston
elliptical, with undivided primary areoles. It 7747 (MO). CALDAS: carretera entre Manizales y
belongs to the C. sphenodes group. hotel "Termales del Ruiz", 8 Jun. 1966, Forero et al. 552
(F). CUNDINAMARCA: Páramo de Guasca, 15 Dec.
1938, Balls 5726 (UC, US). CAUCA: southern slope
16. Campyloneurum cochense (Hieron.) Ching, above Carpintería, 24 Apr. 1939, Alston 8250 (MO);
Sunyatsenia 5: 263. 1940. Río Palo, Quebrada de Santo Domingo, 13 Dec. 1944,
Polypodium cochense Hieron., Hedwigia 48: 269. Cuatrecasas 19266 (F). NARIÑO: vicinity of Cordoba,
28 Sep. 1944, Ewan 16236 (BM, S, UC). PUTUMAYO:
1909. Type. Colombia: Pasto, lake Cocha,
Sibundoy, 4 May 1939, Alston 8395 (MO); S of la
Stübel 250 (holotype B!, photo BM!). Cocha lake, 8 Jan. 1941, Cuatrecasas 11805 (F, US);
Polypodium angustifolium Sw. var. monstruosum Laguna de la Cocha, 29 Oct. 1944, Ewan 16373 (BM,
Mett., Ann. Sci. Nat. (Paris) V, 2: 258. 1864. GH, S, UC).
Type. Colombia: Cundinamarca, Cipacón, ECUADOR. CARCHI: road Julio Andrade-El
Lindig 241 (B!, photo F!). Carmelo, km 7-10, 16 May 1982, Balslev et al. 2558
Stem creeping, dark stramineous, black, (AAU, B, NY, QCA). Balslev et al. 2628 (AAU, B, NY,
usually pruinose, (5-) 6-8 mm wide. Stem scales QCA); Paramo El Angel, on road El Angel-Tulcán, 14
May 1973, Holm-Nielsen et al. 5336 (AAU, MO, UC);
grey brown in mass, adpressed, ovate, 2-3 (-4)
Tulcan-El Angel, 24 Feb. 1984, Juncosa 2395 (MO).
mm long, (1-) 1.5-2 mm wide, bases auriculate, IMBABURA: Shanshipamba, Macnoloma, 14 Nov.
apices acuminate, scales clathrate, the cells 1949, Acosta-Solís 14295 (F); Acosta-Solís 14317 (F);
oblong, along the main axes of the scale, central Lake Cuicocha, Islote Chica, 23 Jun. 1939, Asplund 7128
cell walls 9-12 µm thick, margins differentiated, (S, US); Otavalo-Hacienda Perugachi, NW of Peñas
marginal cell walls 6-9 µm thick. Phyllopodia 4- Blancas, 5 Jan. 1980, Jaramillo et al. 1891 (AAU, QCA).
6 mm long, (4-) 5-7 mm wide, (6-) 10-15 (-30) mm PICHINCHA: Quito-Santo Domingom de los
apart. Leaves (35-) 70-92 (117) cm long; petiole Colorados road, 11 Jun. 1977, Ayala 1 (QCA); carretera
Chillogallo-Chiriboga, km 36, 18 Oct. 1981, Balslev 2107
dark stramineous, 4-12 (-24) cm long, ranurate
(QCA); Concepción, 28 Mar. 1951, Bell 18 (S);
adaxially, concave abaxially; lamina narrowly
Parroquia Calacali, Reserva Geobotánica Pululahua, 16
lanceolate, bases and apices attenuate, (1.5-) 2.5- Nov. 1987, Cerón & Cerón 2756 (QCA); Cantón
4.5 (-7) cm wide, chartaceous-subcoriaceous, Ruminiahui, Parroquia Amaguaña, Pasochoa, Cerón &
Alarcón 3544 (MO); Lloa valley, 18 May 1980, Holm- Polypodium costale Jenm., Bull. Bot. Dep. 4: 140.
Nielsen 23536 (AAU); Chillogallo-San Juan, 23 Jun. 1897. (err. script. for P. costatum).
1980, Jaramillo & Lascano 2552 (AAU, QCA); road Campyloneurum phyllitidis var. costatum (Kunze)
between Calacali and San José de Niebli, Zogg &
Farwell, Amer. Midl. Nat. 12: 297. 1931.
Gassner 13045 (QCA). NAPO: upper slopes of Guagra
Urcu, 26 Sep. 1980, Holm-Nielsen et al. 27124 (AAU);
Stem short-creeping, not pruinose, 4-6 mm
road Tena-Baeza, 12 Jan. 1981, Jaramillo 4085 (AAU, wide. Stem scales brown in mass, 3-5 mm long,
QCA); SE slopes of Cordillera Huacamayos, 12 Jan. 0.8-1 (-1.5) mm wide, narrowly ovate, bases
1981, Proctor 38725 (QCA); Quebrada Violetas, about 4 auriculate, apices acuminate, clathrate, the cells
km W of Aloag, on road Aloag-Santo Domingo, 25 oblong, along the main axes of the scale, cell
Mar. 1967, Sparre 14967 (S); road betwen Quito and walls 10-15 µm thick, sometimes hairs at the
Baeza, Río Chalpi, 18 Sep. 1989, Zogg & Gassner 13007 margin. Phyllopodia 1 mm long, 2 mm wide, 1-4
(QCA). BOLIVAR. Urcu corral, Chillanes, 3 Nov. mm apart. Leaves 30-55 (-75) cm long; petiole
1943, Acosta-Solís 6609 (F) TUNGURAHUA: Chaupi, 4
stramineous or brownish, (4-) 7-13 cm long,
Jan. 1962, Dodson & Thien 1847 (MO, US), Dodson &
Thien 2054 (MO); road Patate-El Triunfo, 4 Mar. 1989, ranurate adaxially, convex abaxially; lamina
Buitrion 475 (QCA). CHIMBORAZO: sector Las lanceolate or narrowly obovate-lanceolate, bases
Chorreras, Hacienda La Carmela, Cord. Occidental assymetrically narrowly cuneate or attenuate,
Sibambe, 20 Aug. 1943, Acosta-Solís 5466 (F). AZUAY: apices subcaudate, rarely attenuate, 3-7 cm wide,
1-8 km N of Sevilla de Oro, 27 Jul.-12 Aug. 1945, Camp herbaceous-chartaceous, chartaceous, margins
E-4571 (MO, S, US); road Sigsig-Ludo, 16 Nov. 1983, cartilaginous, slightly sinuate, indument of
Eriksen & Boysen-Larsen 45676 (QCA). MORONA- inconspicuous simple hairs, scattered abaxially;
SANTIAGO: trail Alao-Huamboya, 8 May 1982,
stomata polocytic; costa prominent, primary
Øllgaard et al. 38449 (AAU, QCA). LOJA: Parque
Nacional Podocarpus, Cerro Toledo, E of Yangana, 4
veins inconspicuous or slightly prominulous
May 1987, Werff & Palacios 9323 (MO); along road adaxially, 60-65o divergent from the costa, 4-6
between Loja and Zamora, 2 Aug. 1978, Zarucchi & mm apart, secondary transversal veins forming
Andrade 2301 (MO, S, US). ZAMORA-CHINCHIPE: 8-10 primary areoles between the costa and
Loja-Zamora road, at the pass, 12 Feb. 1985, Harling & margin, primary areoles usually symmetrically
Andersson 21990 (QCA); road Loja-Zamora, 16 Apr.
divided, excurrent veinlets 1-2 in each secondary
1973, Holm-Nielsen et al. 3628 (AAU, F, GB, UC).
areole; sori medial, paraphyses and spores not
seen. Figs. 17 b; 35.
Campyloneurum cochense is characterized by its
linear-lanceolate leaves, which resembles those
This species is known from southern United
from C. amphostenon and C. densifolium. It also
States (Florida) to Panama, Greater Antilles,
has more than 5 areoles on each side of the costa.
Trinidad, Venezuela and Ecuador, where it is
In addition, C. cochense has adpressed stem
found below 1000 m elevation. It grows as
scales, usually with obtuse apices, and margins
terrestrial or as low trunk epiphyte.
clearly differentiated from the center of the scale.
It belongs to the Campyloneurum xalapense group.
REPRESENTATIVE SPECIMENS: UNITED STATES.
FLORIDA: Collier County, Fahkahatchie Cypress,
about 7 mi NW of Copeland, Nov. 1958, Darling, s.n.
17. Campyloneurum costatum (Kunze) C. Presl, (US); near Everglade, 11 Jul. 1904, Eaton 1135 (GH).
Tent. Pterid. 190. 1836. MEXICO. VERACRUZ: near the summit of
Polypodium costatum Kunze, Linnaea 9: 38. 1834. Ejecatepetl, Zongolica, 6 Jun. 1944, Vera-Santos 3010
Type. Cuba: Limonar, Poeppig s.n. (holotype (US). CHIAPAS: Mun. Ocozocoautla, Reserva
LZ, probably destroyed; isotypes B, BM!, C!, Ecológica El Ocote, 14 Feb. 1986, Palacios-Ríos 2802
(UC).
K!, P!, photo BM! from B).
BELIZE. TOLEDO: Punta Gorda and Joe Taylor Creek,
Cyrtophlebium costatum (Kunze) J. Sm., London J. 2 Jul. 1949, Gentle 6792 (US).
Bot. 1: 196. 1842. GUATEMALA. Chamá, 26 Feb. 1920, Johnson 358
Campyloneurum immersum J. Sm., Bot. Voy. (US).
Herald 231. 1854. Type. Panama: Darién, Bay HONDURAS. ATLANTIDA: Lancetilla, ca. 10 mi SE
of Utria, Apr. 1848, Seeman s.n. (holotype K!). of Tela, 3 Aug. 1977, Croat 42663 (MO).
NICARAGUA. ZELAYA: Bahía de Bluefields, Río can differentiated by the characteristics used in
Escondido, 30 Mar. 1949, Molina 2024 (US), 3.6 km SE the key.
Cerro San Isidro, Río Kama, Río Escondido, 6 Mar.
1966, Proctor 27004 (NY); SE Cerro San Isidro, 9 Mar.
1966, Proctor et al. 27063 (NY). MATAGALPA:
Jinotega, between Las Camelias and La Salvadora,
18. Campyloneurum cubense Fée, Gen. Filic. 259.
along small tributary od Río Jigüina, 31 Oct. 1979, 1852. Type. Cuba, 1843-1844, Linden 1912
Stevens & Grijalva 15323 (MO, UC). (holotype RB, isotypes BM!, K, L, photo BM!).
COSTA RICA. ALAJUELA: Upala, Dos Ríos, Río Polypodium vexatum D. C. Eaton, Mem. Amer.
Cucaracha, 4 Nov. 1987, Herrera 1114 (MO). Acad. Arts n.s. 8: 199. 1860. Nomen Novum for
PUNTARENAS: Parque Nacional Corcovado, Monkey Campyloneurum cubense.
Woods, 9 Jun. 1988, Kernan 575 (MO). Polypodium cubense (Fée) Christ, Bot. Jahrb.
PANAMA. BOCAS DEL TORO: Herrera, 10 km W of Syst. 24: 131. 1897. Nom. Illeg. Non
Las Minas on road to El Toro, 24 Jan. 1981, Sytsma &
Polypodium cubense Fée 1852.
D'Arcy 3254 (MO). CANAL ZONE: pipeline road, 4-6
mi N of Gamboa, 24 Sep. 1981, Knapp 1272 (MO); over Campyloneurum fasciale var. gracile T Moore,
Río Masambi Grande, 1 km NW of Summit Garden, 20 Index Filic. 224. 1861.
Oct. 1973, Nee 7497 (US); along Río Mendosa, 8 km Stem creeping, not pruinose, 1-4 (-5) mm
NW of Gamboa, 16 Apr. 1974, Nee & Smith 11364 (US). wide. Stem scales brown in mass, 4 mm long,
DARIEN: Isthmus, Seemann s.n. (US). Near upper (1.5-) 2-2.5 mm wide, ovate, clathrate, apices
Juan Díaz river, Killip 2851 (US). short acuminate, central cells usually oblong, cell
CUBA. PINAR DEL RIO: La Plata, Sierra del Rosario, walls 12.5-15 (-20) µm thick, along the major axes
Jan. 1957, Bro. Alain 6100 (US); Bahía Honda, N of Pan
of the scale, marginal cells usually isodiametric,
de Guajaibón, Aug. 1968, Bisse 9637 (HAJB);
mountains N of San Diego de los Baños, 11 Apr. 1900,
cell walls 5-7.5 (-10) µm wide, transversal to the
Palmer & Riley 511 (BM). ORIENTE: Florida Blanca, major axes of the scale. Phyllopodia 0.5-1 mm
Apr. 1947, Bro. Clement 5230 (US); Sierra de Nipe, on long, 1.5-2 (-3) mm wide, 2-3 mm apart. Leaves
Río Piloto, 20 Jul. 1914, Ekman 2063 (S); Bayate, Cayo 23-60 (-90) cm long; petiole stramineous, (1-) 3-15
del Rey, 6 Sep. 1914, Ekman 2757 (S); Sierra de Nipe, ad (-20) cm long, ranurate adaxially, indument
Río Piloto, 9 Jun. 1915, Ekman 5979 (S); Farallones of La sometimes of scattered caducous scales less than
Perla, N of Jaguey, Yateras, 2 May 1907, Maxon 4377 1 mm long; lamina narrowly obovate, bases
(US). El Palenque, Mar. 1889, Eggers 4854 (F, S).
attenuate, apex acuminate, (0.6-) 1-2 (-3.5) cm
Without locality: 1849, Rugel 22 (BM); 19 Aug. 1889,
wide, herbaceous-chartaceous, margin sinuate,
Eggers 4854 (B, F); 1859-1860, Wright 802 (MO, S).
JAMAICA. PORTLAND: Seamen's valley, 14 Feb. plane, indument of simple bicellular hairs,
1920, Maxon & Killip 5 (F); Mansfield, near Bath, 2 May disperse and caducous abaxially, stomata
1903, Maxon 1820 (US); Hartford, near Priestman's polocytic; costa prominent, primary veins
river, 9 Jun. 1904, Maxon 2554 (US); Ginger river, May inconspicuous or slightly prominulous adaxially,
1884, Syme s.n. (BM). ST. THOMAS: Corn Puss Gap,
prominulous abaxially, flexuosous, 50-65o
27 Jun. 1954, Wilson & Webster 466 (GH).
divergent from the costa, 3-5 (-6) mm apart,
DOMINICAN REPUBLIC. Pacificador: Villa Riva, 11-
19 Jan. 1912, Abbott 570 (US). Samaná: Santo Domingo, secondary transversal veins forming 2-4 (-6)
Cordillera Central, 27 Jun. 1930, Ekman 15450 (S). areoles between the costa and margin, areoles
TRINIDAD. Without locality, Jenman 205 (NY). entire or divided, 1-2 excurrent veinlets in each
VENEZUELA. PORTUGUESA: dist. Araure, Ríos secondary areole; sori subterminal, paraphyses
Bocoy and Riecito, Ortega & Aymard 1801 (UC). not seen; sporangia 14-17 cells of the bow, spores
ECUADOR. LOS RIOS: surroundings of Montalvo, ca. (45-)50-60 µm long, 30-40 µm wide. Figs. 10 c;
40 km E of Babahoyo, 30 Mar.-2 Apr. 1973, Holm- 36.
Nielsen et al. 2827 (AAU). Without exact locality:
Eggers 14373 (F); 6 Sep. 1896, Eggers 15306 (F).
This species is known from Cuba, Jamaica,
and Haiti, where it grows between 100 m and
Campyloneurum costatum is characterized by
600 m elevation.
lanceolate or elliptical-lanceolate leaves, with
inconspicuous or slightly prominulous veins.
REPRESENTATIVE SPECIMENS: CUBA.
It is closely related to C. xalapense, and they GUANTANAMO: Las Ninfas, Dec. 1917, Hioram 1443
(S). LA HABANA: Lomas de Tapaste, 4 Feb. 1973, and Polypodium laevigatum Cav. (a name that he
León 3528 (S). CIENFUEGOS: Cumanayagua, Las used probably for P. lapathifolium Poiret, a
Vegas, Cafetal de Buenos Aires, 29 Oct. 1985, Berazaín synonym of C. repens), suggesting a probable
et al. 57968 (HAJB). LAS VILLAS: Valley of the Río
hybrid origin for C. cubense.
Los Negros, 10-14 km beyond Siguanea, around El
Junco, Jul. 1950, Hawkes 2082 (UC); Trinidad
During this study, the pattern of venation of
Mountains, Loma Ventana, Aug. 1941, Howard 6482 Campyloneurum cubense were compared with
(GH); above San Blas, 4 Dec. 1928, Jack 6769 (US); San those of C. angustifolium and C. repens. In C.
Blas, La Sierra, 4 Mar. 1929, Jack 6966 (F); Sierra cubense, primary veins form narrow angles with
Gavilán, 9-10 Nov. 1941, Morton 4002 (GH, MO). the costa and divided primary areoles, as in C.
ORIENTE: Sierra Maestra, Loma del Gato, 1923, angustifolium, although undivided areoles, as in
Clement 779 (S); Loma del Gato, El Cobre, Aug. 1924, C. repens are more common. Other characters,
Clement 1387 (BM); Jaguey, Eggers 4921 (F); Loma del such as pattern of stomata, spore number and
Jaguey, Mar. 1889, Eggers 4942 (B, S); Bayate, 13 Jul.
morphology were also examined in C. cubense.
1914, Ekman 1969 (S); Cayo del Rey, in the Cañón de
Canapu, 6 Nov. 1914, Ekman 2756 (S); S of Jaguey, Specimens of C. cubense showed the epidermis
Yateras, Apr. 1907, Maxon 4161 (BM, S); summit of El with some irregular or abortive stomata. Spores
Yunque, near Baracoa, 30-31 Jan. 1902, Pollard & in herbarium specimens of C. cubense are very
Palmer 176 (F); gorge Río Yamuri, 7-9 Dec. 1910, Shafer scarce, and usually collapsed; the number of
7862 (GH); Monte Verde, 13 Feb. 1911, Shafer 8704 spores in each sporangia, however, were 64, as in
(MO); Monte Verde, Jan.-Jul. 1859, Wright 801 (BM, normal meiosis.
MO, S); Wright 1020 (B, MO, S, US). PINAR DEL RIO: This information collectively may suggest
Baños San Vicente, 12-16 Sep. 1910, Britton et al. 7354
hybridization. A hybrid origin cannot be
(F). Mountains near El Guama, 11 Mar. 1900, Palmer &
Riley 254 (BM, US); San Diego de los Baños, 1 Feb.
discarded, especially considering one of the
1917, Palmer s.n. (US). Without exact locality: w.d., parental species to be C. angustifolium, but at the
Eggers 4740 (B, F); Finca Las Prendas, 30 Dec. 1920, same time there is not still cytological evidence
Hioram & Maurel 4124 (US); 1860-1864, Wright 1829 for solving the origin of this species.
(MO).
JAMAICA. Gordontown, 11 Sep. 1906, Moore s.n. (BM);
Mansfield, near Bath, 2 May 1903, Maxon 1832 (US). 19. Campyloneurum decurrens (Raddi) C. Presl,
Saint Thomas: Cuna Cuna Pass, Mar. 1895, Gilbert s.n.
Tent. Pterid. 190. 1836.
(MO); vic. of House Hill, 15 Jun. 1926, Maxon 9226 (S,
Polypodium decurrens Raddi, Syn. Fil. Bras. 287.
UC); Cuna Cuna Gap and vicinity, 19 Jun. 1926, Maxon
9390 (F, GH, US); Arntully, 19 Mar. 1963, Proctor 23346 1819. Type. Brazil, Raddi s.n. (holotype not
(BM); along track between House Hill and Cuna Cuna, seen).
26 Dec. 1969, Proctor 31145 (F). Venegas Hill, 12-13 Aspidium pentaphyllum Willd., Sp. Pl. 5: 216-217.
Apr. 1909, Watt 130 (S, US). Without exact locality, 1810. Type: Plumier, Fil. tab. 114. Non
w.d., Swartz s.n. (BM). Polypodium pentaphyllum Baker, 1891.
HAITI. Gorge Crete-a-Piquants, Port au Prince, 14 Feb. Cyrtophlebium decurrens (Raddi) J. Sm., J. Bot. 4:
1927, Ekman 7604 (S); Massif de la Holle, Jeremie, 58. 1841.
between Maffrand and Maron, 11 Jul. 1928, Ekman
Stem short-creeping, not pruinose,
10292 (S); Massif de la Holle, Dame Marie, 14 Aug.
1928, Ekman 10519 (S); Riviere Glace, 4 Aug. 1945, stramineous, 8-10 (-15) mm wide. Stem scales
Holdridge 2090 (US); vicinity of Mission Fonds adpressed, light brown in mass, 3-4 mm long,
Varettes, 17 Apr.-4 May 1920, Leonard 3762 (US), 2.5-3 mm wide, ovate, bases auriculate, apices
Leonard 4031 (US); vicinity of Furcy, 26 May-15 Jun. acute, clathrate, the cells broadly oblong.
1920, Leonard 4616 (GH). Phyllopodia 5-6 mm long, 5-7 mm wide, 8-10
mm apart. Leaves 1-pinnate, 75-100 cm long;
Campyloneurum cubense is characterized by its petiole brown stramineous, usually more than
lamina narrowly obovate, primary veins with a half the lenght of the lamina, ranurate adaxially,
50-65o divergence angle from the costa, and by convex abaxially; 5-10 pinnae on each side of the
ovate-lanceolate stem scales. Christ (1897) rachis, pinnae narrowly lanceolate, bases
mentioned the intermediate characters of this assymmetrically attenuate, apices acuminate,
species compared with those of C. angustifolium upper pinna decurrent on the rachis, basal
pinnae sessile or very shortly petiolulate, 17-32 magnificum, but it differs from the latter by its
cm long, 2.5-3.2 cm wide, herbaceous- small size, narrowly lanceolate pinnae. Both
chartaceous, margins cartilaginous, slightly species may represent an ancient lineage in the
ondulate, indument not seen, stomata polocytic; genus.
venation areolate, costulae prominent, primary
veins prominent and usually stramineous, 55o
divergent from the costulae, transverse veinlets 20. Campyloneurum densifolium (Hieron.)
prominulous, forming 5-7 primary areoles Lellinger, Amer. Fern J. 78: 19. 1988.
between the costula and margin, excurrent Polypodium angustifolium var. amphostenon
veinlets 2-3 in each undivided non-costal areole, (Kunze) Baker f. densifolium Hieron., Bot.
costal areole with one excurrent veinlet, simple Jahrb. Syst. 34: 532. 1904. Lectoype (chosen by
or furcate; sori terminal, paraphyses not seen, Lellinger, Amer. Fern J. 78: 19-20. 1988):
spores 40-45 µm long, 20-25 µm wide. Figs. 9; Ecuador, Azuay, near Las Yerbas Buenas,
11; 13; 35. Lehmann 5723 (US!, isolectotype B!, F!; photo
of US, AAU!).
This species is found in Colombia, Venezuela, Stem creeping, black, sometimes pruinose, 3-5
Martinica, and southern Brazil, where it grows in (-7) mm wide. Stem scales brown, light brown in
forests between 200-950 m elevation. mass, 4.5-7 mm long, 2.5-3 mm wide, ovate or
broadly ovate, bases auriculate, apices
REPRESENTATIVE SPECIMENS: COLOMBIA. acuminate, subadpressed, slightly clathrate, the
ANTIOQUIA: Río Verde, 14 Jul. 1889, Kalbreyer s.n. (S). cells oblong or ovate, cells along the main axes or
MAGDALENA: Sierra Nevada de Santa Marta, 1898- irregularlly disposed at the apex, cell walls
1899, H. Smith 2456 (F). slightly diffuse, 15-17 µm thick, scales sometimes
VENEZUELA. Colonia Tovar, 1854-1855, Fendler 231 with marginal hairs. Phyllopodia 1-2 mm long,
(F).
(1-) 2-3 mm wide, 4-10 mm apart. Leaves 30-70
MARTINICA. Piton Marcel, entre la montagne Pelee
cm long; petiole stramineous or dark
et le Pricheur, Jul. 1885, Duss 1568 (B, US).
BRAZIL. ESPIRITO SANTO: Santa Bárbara de stramineous, 2-14 cm long, slightly ranurate
Caparaó, 5 Dec. 1929, Mexia 4093a (UC). MINAS adaxially, plane convex abaxially; laminae linear-
GERAIS: Viçosa, 22 Jul. 1930, Mexia 4892 (B, S, UC). lanceolate or narrowly lanceolate, bases and
RIO DE JANEIRO: Corcovado, Apr. 1913, Brade 6461 apices attenuate, 1-3 (-5) cm wide, chartaceous or
(UC); Itatiaia, Apr. 1913, Brade 6461 (UC); Itatiaia, 4-10 subcoriaceous, margins cartilaginous, slightly
Jun. 1913, Brak 6461 (S); Itatiaia, 23 Jul. 1902, Dusen 743 sinuate, sometimes revolute, stomata polocytic,
(S); Tijuca, 3 Sep. 1904, Dusen 5144 (S); Rio de Janeiro, indument of bicellular, simple hairs, scattered
Gaudichaud 2a (F); Nova Friburgo, 22 Sep. 1947, Leite
abaxially; costa prominent, primary veins
4204 (F); Organ mountains, Jul. 1871, Luessen 1237 (F);
Corcovado, 1911, Lutzelburg 363 (UC); Corcovado, 17 prominulous on different degree on both sides of
Jul. 1873, Mosen 63 (S); Corcovado, 10 Sep. 1874, Mosen the lamina, inconspicuous, partially stramineous
2683 (B, S); trail between Sylvestre and Paineiras, 14 or darker when dry, 50-60o (-65)o divergent from
Apr. 1929, Smith 2251 (S, UC). SÃO PAULO: Alto da the costa, straight or slightly sinuate, 4-7 mm
Serra, Wacket 134 (S); São Paulo, 25 Dec. 1873, Mosen apart, secondary veins inconspicuous or very
2224 (S); Alto da Serra, 1905, Wacket 134 (UC).
slightly prominulous, forming 2-4 primary
PARANA: Porto de Cima, 24 Jul. 1914, Jonsson 717a
(S).
areoles between the costa and margin,
symmetrically divided or rarely entire, excurrent
Campyloneurum decurrens is characterized by veinlets 1-2 in each secondary areole; sori medial
its pinnate leaves with more than 5 pairs of or subterminal, paraphyses not seen, spores (50-)
narrowly lanceolate pinna. The known 57-60 µm long, (35-) 40-42 µm wide. Figs. 37, 38.
distribution of this species appears to be
restricted to the presently scarce Atlantic humid This species is found from Central America to
forests of South America. Bolivia. It grows in open areas, among rocks,
It is closely related to Campyloneurum above 2000 m elevation.
REPRESENTATIVE SPECIMENS: GUATEMALA. USM). NAPO: Papallacta, Harling et al. 10329 (F, GB);
Huehuetenango: 5 mi S of San Juan Ixcoy, Breedlove SE of Playón de San Francisco, on the slopes of cerro
8531 (US). EL PROGRESO: between Calera and Mirador, Holm-Nielsen et al 29831 (AAU, USM); road
summit of volcan Siglo, 21 Jan. 1942, Steyermark 43067 San Miguel Salcedo-Puerto Nuevo, 54 km from San
(F). Miguel, Øllgaard & Balslev 9820 (AAU). BOLIVAR:
COSTA RICA. HEREDIA: forest of Río Vueltas, Gómez Cerro Negro, Parroquia Chillanes, Acosta Solís 6793 (F).
2297 (F); between Sacramento and Laguna de Barba, TUNGURAHUA: Runtún caserío, ca. 3-4 km from
12 Apr. 1975, Utley & Utley 2031 (F). LIMON: Baños, Lugo 1222 (AAU, GB). CHIMBORAZO:
Cordillera Talamanca, Cerro Kamuk massif, between between San Andrés and Cuatro Esquinas, Fagerlind &
Cerro Dudu and Cerro Apri, Davidse et al. 25897 (MO); Wibom 883 (S). AZUAY: above Sayaus, E of Cuenca,
unnamed cordillera between the Río Terbi and the Río Correll E339 (S); Cuenca, Paramo Quinoas, Harling
Sinú, Davidse et al. 29070 (MO). SAN JOSE: Cerro de la 1481 (S); Paramo de Matanga, km 25 on road Sigsig-
Muerte, N 5 km, 30 Aug. 1983, Saiki 83 (F); 15-18 km Gualaquiza, Holm-Nielsen et al. 29526 (AAU).
SE of Empalme, 17-26 Mar. 1973, Stolze 1517 (AAU, F, PERU. CAJAMARCA: 25 km from Cajamarca to
UC). PUNTARENAS: Cantón de Buenos Aires, Bambamarca, 25 Mar. 1960, Correll & Smith 859 (GH);
Ujarrás, 13 Oct. 1989, Herrera 3673 (F). environs of Huancabamba, Las Huaringas, 20 Feb.
PANAMA. CHIRIQUI: shoulder of El Barú, above 1981, Davis & Turner 707 (F, GH); Cajamarca, summit
Bajo Grande, Folsom et al. 2140 (MO); 2 km S of Questa between Cajamarca and San Juan, Gutte & Müller 8915
Piedra, along Concepción, Folsom 3966 (UC); vicinity (USM); Celendín, canyon of the Río Marañón, above
of cerro Punta, above Guadalupe, McPherson 9399 Balsas, Hutchison & Wright 5282 (F, UC); San Miguel,
(MO). cerro Quillón, Agua Blanca, 5 Jul. 1986, Mostacero et al.
CUBA. ORIENTE: alround Alto del Olimpo, López- 1287 (F); Contumazá, Pampa de la Sal, 27 Jun. 1983,
Figueiras 405 (US). Sagástegui et al. 10747 (F). LA LIBERTAD: Otuzco,
HAITI. Massif de la Selle, Grand-Gosier, slope of M. Llaguen, Shilte, López 1560 (GH); Otuzco, cerro
Commissaires, Ekman 6862 (C, S); Massif de la Holle, Chologday, 19 May 1957, Sagástegui 78 (GH); Santiago
Torbec, Ekman 7504 (S); Massif de la Selle, Croix des de Chuco, Chota, Motil-Shorey, 12 Jun. 1984,
Bouquets, Ekman 7631 (S, US). Sagástegui et al. 11699 (F, MO); Santiago de Chuco,
COLOMBIA. ANTIOQUIA: Mun. Helmeira, Acosta- Santiago-Shorey road, 26 km from Santiago, D.N.
Arteaga 880a (AAU); Mun. Caldas, trail La Corrala, Smith 2329 (USM). SAN MARTIN: Mariscal Cáceres,
Morales et al. 5869 (F). CUNDINAMARCA: northern P. N. Río Abiseo, Chochos valley, Young 3800 (USM);
end of sabana near Suba, Cuatrecasas & Jaramillo 25942 Young 2619 (USM); Chochos forest, Young 4847 (USM);
(US); Cogua-San Cayetano, 9 May 1967, Murillo et al. forest patch C9 above timberline, Chochos valley,
1052 (F, US). VALLE: Cordillera Central, Río Young 2586 (F, USM); forest patch C10, Young 2534 (F,
Bugalagrande, Cuchilla de Barragán, 15, 16, 24 Apr. USM); C15, Young 2461 (USM), Young 2462 (F, USM);
1946, Cuatrecasas 20797 (F). CAUCA: Mun. Puracé, forest patch C17, Young 3655 (USM); entre El Mirador
Parque Nacional de Puracé, near Laguna San Rafael, 6 y Puerta del Monte, Young & León 4443 (USM); Puerta
Oct. 1984, Lozano et al. 4682 (F). del Monte, P6 forest patch, Young 1986 (F, USM);
VENEZUELA. MERIDA: 25 km from Mérida, along El Young 1765 (USM); forest patch P10, Young 1883 (F,
Valle road, Breteler 4663 (NY, US). DISTRITO USM); forest patch P12, Young 2056 (USM). ANCASH:
FEDERAL: arriba de Galipán, Williams 12386 (F, S). Bolognesi, Tinya, Río Fortaleza valley, 28 Apr. 1956,
Colonia Tovar, Fendler 226 (US). Cerrate 2585 (GH, USM). HUANUCO: Mito, Bryan 196
ECUADOR. CARCHI: road Julio Andrade-Palestina, (F); Bryan 365 (F). LIMA: Huarochirí, Río Blanco,
Holm-Nielsen et al. 29704 (AAU, F, USM); Maldonado- Asplund 11299 (S, US); 23 Jul-14 Aug 1922, Macbride &
Tulcan road, 5 Oct. 1981, Werling & Leth-Nissen 253 Featherstone 1919 (F); Huarochirí, Viso, Goodspeed et al.
(AAU, F). PICHINCHA: Cerro Pasochoa, Balslev 2783 11547 (US). PASCO: 95 km S from Huánuco, Polylepis
(AAU); Los Alpes, N of Cordillera Occidental, Acosta- forest, on road to Cerro de Pasco, Gentry et al. 37489
Solís 7094 (F); Quebrada Sombría, betweenMagdalena (F). JUNIN: Tarma, Incatacuna, Tarmatambo-Acolla,
and Chillogallo Firmin 454 (S); northern slopes of Constance & Tovar 2348 (UC); Jauja, laguna de Paca,
Cerro Corazón, 2-4 km of Aloag, Holm-Nielsen 18023 Gracey et al.6 (USM); entre Tarma y San Ramón, 35 km
(AAU, USM); road Chillogallo-San Juan-Chiriboga- from Tarma, Gentry et al. 39778 (F, MO, USM); Yauli,
Empalme, 21 Feb. 1986, Zak 897 (F, MO); between Llocllapampa, 22 Oct. 1966, Saavedra 6317 (GH); La
Quito and Santo Domingo de los Colorados, Alisal, 19 Merced, Aug. 1947, Soukup 3400 (F). APURIMAC:
Sep. 1989, Zogg & Gassner 13035 (QCA). COTOPAXI: Abancay, Andahuaylas, Alvarado s.n. (USM); quebrada
Quevedo-Latacunga road, Holm-Nielsen et al. 3262 of Juccuchic-chupan, on trail Andahuaylas-
(AAU, F, GB, MO); road Pilaló-Zumbagua, 10 km Chincheros, West 3723 (UC). CUSCO: Chincheros,
above Pilaló, Holm-Nielsen & Quintana 24647 (AAU, Antakillpa, Davis et al. 1649 (F, USM); Cusco, 1 Sep.
1914, Rose & Rose 19062 (US). lamina margins cartilaginous, usually revolute,
BOLIVIA. LA PAZ: Sur Yungas, San Felipe, Asplund indument of inconspicuous hairs, scattered
1782 (S); Omasuyos, Isla del Sol, Yumani, Asplund 3583 abaxially, stomata polocytic; costa prominent,
(BM, S, US); vicinity of La Paz, Bang 140 (UC, US);
sometimes with scales similar to those on the
Murillo, valle de Zongo, Santa Rosa, Beck 1098 (LPB);
Camacho, Puerto Acosta, 6 km hacia La Paz, Beck 7685
stem, primary veins inconspicuous, 1-2 areoles
(F, LPB); Nord Yungas, Unduavi, Buchtien 78 (F, S); between the costa and margin, excurrent veinlet
Murillo, Río Zongo valley, below dam at Lago Zongo, one per areole; sori medial, paraphyses not seen;
Solomon 8386 (LPB, UC); Nor Yungas, 0.9 km W of spores 45-55 µm long, 30-40 µm wide. Fig. 29.
Chuspipata, Solomon 9650 (LPB); Murillo, Valle del Río
Zongo, 8 Nov. 1987, Solomon 17272 (MO). This species is found from Mexico to
Guatemala, Nicaragua, where it grows mainly as
Campyloneurum densifolium is characterized by an epiphyte, rarely on rocks, between 500 m and
its adpresed ovate stem scales, these being 2900 m elevation.
usually persistent and light brown in color.
It resembles C. fallax from Brazil, from which REPRESENTATIVE SPECIMENS: MEXICO.
it differs by decurrent leaf bases, and stem scales SINALOA: 5 Km N of El Palmito, 28 Oct. 1973,
with acuminate apex. BothC. densifolium and C. Breedlove 35732 (MO). JALISCO: Sierra de Manantlán,
fallax are disjunct in distribution. Cuautitlan, 19 Jan. 1975, Diaz-Luna 5598 (UC).
Campyloneurum densifolium is closely related to MICHOACAN: vic. Morelia, Campanario, 14 Sep.
1911, Arsene 5612 (MO); Uruapan, Tancitaro, 14 Nov.
C. amphostenon, from which it is easily
1940, Hinton et al. 15685 (MO, US); Tancitaro, Uruapan,
distinguished by stem scale characters (Fig. 37), 17 Oct. 1940, Hinton 15543 (MO); Tancitaro, 21 Jul.
which are in the latter lanceolate and with 1941, Leavenworth & Hoogstraal 1095 (F, GH, MO).
spreading apices. VERACRUZ: Mun. Jalisco, Taxolo, 8 Jun. 1983, Barnett
Campyloneurum densifolium belongs to the et al. 85a (MO); Mun. Acajete, NW of Mazatepec, 9 Jun.
Campyloneurum amphostenon group. 1983, Barnett t al. 101 (MO); Córdoba, Aug. 1936,
Matuda 208 (MO); Cerro del Aguila, 13 km N of
Altotonga, 28 Jun. 1980, Nee & Hansen 18570 (F); 1 km
NE of San Antonio Ixtatetla, 27 Apr. 1983, Nee & Taylor
21. Campyloneurum ensifolium (Willd.) J. Sm., Cat.
26818 (F) Huayacocotla, 26 Jan. 1984, Nee 29080 (F).
Cult. Ferns 12. 1857. CHIAPAS: SE of Cerro Baul, 16 km NW of Rizo de
Polypodium ensifolium Willd., Sp. Pl. 5: 152. 1810. Oro, 3 Nov. 1971, Breedlove & Smith 21793 (F); finca
Type. Probably Mexico (as Peru Lima), Née Irlanda, May 1914, Purpus 7238 (BM, F, MO). Without
s.n. (Herb. Willd. 19610) (holotype, B!; photo locality: Tocuila, 1869, Hahn 19 (B).
BM!, USM!). GUATEMALA. ALTA VERAPAZ: San Juan
Goniophlebium ensifolium (Willd.) Brackenridge in Chameles-Cobán, 19 Aug. 1973, Dary-Rivera 254 (F).
Wilkes, U. S. Explor. Exped. 33. 1854. SAN MARCOS: 12 Mar. 1940, Steyermark 37600 (F).
Polypodium angustifolium var. ensifolium Hicken, QUEZALTENANGO: slopes of volcán Zunil, at Aguas
Amargas, 17 Febr. 1939, Standley 65432 (F); below
Revista Mus. La Plata, Secc. Bot. 5: 271. 1908.
Santa María de Jesús, 11 Mar. 1939, Standley 68419 (F);
Campyloneurum angustifolium var. ensifolium between Finca Pirineos and Patzulín, 9 Feb. 1941,
(Hicken) Farwell, Amer. Midl. Naturalist 2: Standley 86975 (F, UC), Standley 87028 (F), Standley
296. 1931. 87095 (F); along quebrada San Gerónimo, lower south-
Stem creeping, pruinose, 2-4 mm wide. Stem facing slopes of Volcán Santa María, 1-2 Jan. 1940,
scales brown, 3-4 mm long, 1-1.5 mm wide, Steyermark 33446 (F). SOLOLA: San Lucas, 10 Jun.
scales ovate, bases auriculate, apices acuminate, 1948, Williams 14343 (BM, F, US).
rarely obtuse, clathrate, the cells roundish, cell HUEHUETENANGO: near El Reposo, about 8 km
from Mexican frontier, 14-18 Dec. 1972, L.O. Williams et
walls 7.5-12 µm thick. Phyllopodia 1-2 mm long,
al. 41388 (AAU, F) CHIMALTENANGO:
1.5-2 mm wide, 1-4 mm apart. Leaves 16-60 cm Chimaltenango, Johnston 1270 (F). SACATEPEQUEZ:
long; petiole stramineous, 0.5-3 cm long, slightly just above Barranco Hondo, 11 Mar. 1941, Standley
ranurate on the adaxial side, convex abaxially, 88933 (F); lower slopes of Volcán de Fuego, SW of
glabrate; lamina linear, bases and apices Alotenango, 16 Jan. 1974, L.O. Williams & Williams
attenuate, 0.4-1 (-1.5) cm wide, chartaceous, 43521 (F). JALAPA: 8 km N of Jalapa, 13 Nov. 1940,
Cutler 4320 (MO, US). SUCHITEPEQUEZ: S lower peltate, apices acuminate, cells oblong along the
slopes of Volcán Zunil, E of Pueblo Nuevo, 1 Feb. main axes of the scale, central cell walls 10-17 µm
1940, Steyermark 35361 (F). ESCUINTLA: Monte Rey, thick, marginal cell walls 7.5 µm thick.
El Zapote, 27 Apr. 1937, Muenscher 12119 (F); between
Phyllopodia 0-1 mm long, 1-1.5 mm wide, 8-20
Río Jute and Río Pantaleón, on road between Escuintla
and Santa Lucia, 24 Jan. 1939, Standley 63479 (F).
mm apart. Leaves 10-30 cm long; petiole
SANTA ROSA: Jumaytepequez, Aug. 1892, J. D. Smith stramineous or brownish, 3-7 cm long (1/3-1/5
4087 (B); along road SE of Barbarena, 21 Nov. 1940, of the total lenght of the lamina), indument not
Standley 77870 (F, UC); near Cuilapilla, 23 Nov. 1940, seen, slight ranurate adaxially, convex abaxially;
Standely 78056 (F); near El Molino, 26 Nov. 1940, lamina lanceolate to narrow lanceolate, bases
Standley 78500 (F); Volcán Tecuamburro, N of attenuate or subcuneate, apices long acuminate,
Chiquimulilla, 20 Dec. 1939, Steyermark 33153 (F); up 1.5-2.5 (-4) cm wide, herbaceous or herbaceous-
Loma Bandera Shac, lower S facing slopes of Volcán chartaceous, margins cartilaginous, repand or
Tajumulco, 9 Mar. 1940, Steyermark 37351 (F). Finca
slightly sinuate, indument of scatered hairs on
Naranjo, Chicacao, 14 Mar. 1947, Brenckle 47-94 (F);
Coatepeque, 15 Mar. 1944, Varrelman s.n. (F) the lamina and caducous scales on the costa;
HONDURAS. EL PARAISO: Quebrada Tapahuasca, stomata polo or copolocytic; costa prominent on
14 Aug. 1964, Molina 14664 (F). both sides of the lamina, 1.5 mm long 0.5 mm
EL SALVADOR. AHAUACHAPAN: Laguna Verde, 1 wide, similar to those at the stem, primary veins
Mar. 1979, Seiler 969 (F, NY, UC); near Ataco, 19 Jan. inconspicuous or slightly prominulous on both
1947, Standley & Padilla 2640 (F). SONSONATE: Los sides of the lamina, 3-5 mm apart, flexuous, 50-
Pedregales de San Isidro, 19 May 1977, Seiler 3 (F). LA
LIBERTAD: Mun. Antiguo Cuscatlan, 13 Jul. 1989, 70o divergent from the costa, transverse
Villacorta & Martinez 310 (MO). SAN SALVADOR: secondary veins forming 3-4 primary areoles
vicinity of San Salvador, 30 Mar.-24 Apr. 1922, Standley between the costa and margin, usually entire,
21824 (MO), Standley 22660 (MO, S, US), Standley 22672 excurrent veinlets 1(-2), sometimes forming
(F); near Apulo, near lake Ilopango, 19 Jun. 1949, L.O. secondary areoles at the margin of the lamina;
Williams & Molina 16740 (BM, F, MO). SAN MIGUEL: sori subterminal, usually one row between
Volcán San Miguel, Las Placitas, 5 May 1979, Salgado secondary veins, paraphyses not seen, spores
67 (F)
(50-) 60-70 µm long, 40-50 µm wide. Figs. 5 d, e;
NICARAGUA. ESTELI: Salto de Estanzuela, ca. 5 km
sur de Esteli, 29 Sep. 1980, Guzman et al. 1215 (MO).
17 d; 36.
Campyloneurum ensifolium is characterized by This species is known from Costa Rica,

its small, adpressed stem scales, with roundish Panama and southwestern Colombia, where it
cells. grows as an epiphyte between 1300 m and 2500
Although the label of the type collection m elevation.
mentioned Obrajillo, Peru, as the type locality, I
agree with Lellinger (1988) that the Nee's
ALAJUELA: from Vara Blanca to La Concordia,
specimen was mislabeled. between Poas and Barba volcanoes, 23 Jul. 1923,
Maxon & Harvey 8488 (BM); property of R. Gonzales,
28 Jun. 1875, Polakowsky 201 (BM); Palmira, region of
22. Campyloneurum falcoideum (Kuhn ex Hieron.) Zarcero, 30 Aug. 1937, A. Smith F-48 (F); Palmira, 13
M. Meyer ex Lellinger, Proceed. Biol. Soc. Jan. 1938, A. Smith H82 (F). HEREDIA: Carpintera, 10
Wash. 89: 708. 1977. Apr. 1908, A.C. Brade & A. Brade 19 (S, US); N of
Polypodium falcoideum Kuhn ex Hieron. Bot. Heredia, ca. 1 km beyond Porrosati, 18 Aug. 1970,
Lellinger & White 1675 (US); Puerto Viejo, on the
Jahrb. Syst. 34: 533. 1904. Lectotype (chosen
Sarapiquí River, 2 Mar. 1956, Stork 4841 (NY, US).
by Lellinger, Proceed. Biol. Soc. Washington PUNTARENAS: upper Río Burú, 19 Aug. 1983, Gómez
89: 708. 1977). Costa Rica: Río Sucio, 17 Mar. et al. 21491 (MO), Gómez et al. 21691 (MO, UC). SAN
1882, Lehmann 1741 (B!, BM!, K!, US!). JOSE: Tablazo, 4 Mar. 1908, A.C. Brade & A. Brade 44
Stem long-creeping, not pruinose, 1-1.5 (-2) (BM, NY); ca. 15 km N of Tres Ríos, ca. 4 km N of
mm wide. Stem scales 4-7 mm long, usually 1 Cascajal, 2 Aug. 1970, Lellinger & White 1375 (F, US);
mm wide, narrowly ovate, slightly falcate, Río Parrita Chiquita, 5 km N of Santa María de Dota,
10 Oct. 1976, Lent 3912 (F, NY); near Quebradillas, 24 on both sides, margin cartilaginous, slightly
Dec. 1925, Standley 42922 (BM); upper slopes of Cerro sinuate; stomata polocytic; costa prominent,
Daser, Azulillo, 6 km W of rt. 4, 5 km S of Aserri, 19 primary veins very slightly prominulous on the
Mar. 1973, Stolze 1418 (AAU, F, UC). LIMON:
Cordillera de Talamanca, Atlantic slope, canyon of Río ventral side, slightly flexuosus, 45-50o divergent
Siní, 15 Sep. 1984, Davidse & Herrera 29159 (MO). with the costa, (4-) 5-7 mm apart, secondary
Without exactly locality: Cerro del Gallito, 26 Jun. veins inconspicuous, 4-5 areoles between the
1926, Valerio A95 (US). costa and margin, usually 2-3 veinlets, entire or
PANAMA. CHIRIQUI: valley of the Río Caldera, from furcate; sori medial or subterminal, paraphyses
El Boquete to the Cordillera, 6 Feb. 1918, Killip 5076 not seen, spores (55-) 72-87 µm long, 40-50 µm
(BM); 2 mi W of Cerro Punta, 22 Jan. 1968, McDaniel
wide. Figs. 7 c; 14 d; 17 c; 38.
10222 (GH); valley of the upper Río Chiriquí Viejo, vic.
of Monte Lirio, 27 Jun.-13 Jul. 1935, Seibert 284 (GH);
dist. Bugaba, Santa Clara, Cerro Pando, 28 Feb. 1985, This species is known from southern Brazil,
Werff & Herrera 7243 (UC). where it grows between 1000 m and 2000 m
COLOMBIA: Putumayo, above Sibundoroy, 4 May elevation.
1939, Alston 8372 (MO).
REPRESENTATIVE SPECIMENS: BRAZIL. RIO DE
Campyloneurum falcoideum is easily recognized JANEIRO: Serra do Itatiaia, n. d, Brade 6551 (NY, S);
by the falcate stem scales, remote leaves, and Serra dos Orgãos, Pedra Assú 30 Jul. 1940, Brade 16501
(BM, MO, NY, S, US); Itatiaia, Pousada Nova, 21 Mar.
venation of undivided primary areoles with one
1943, Brade 17327 (MO); Mun. Resende, Itatiaia
included free veinlet. It belongs to the C. National Park, S face of Mt. Itatiaia, below Macieiras,
sphenodes group. Eiten & Eiten 7483 (US); Campo do Itatiaia, 13 May
1906, Luederwaldt s.n. (S); vicinity of Itatiaia, 26-30 Jul.
1915, Rose & Russell 20589 (US). SÃO PAULO: Serra
23. Campyloneurum fallax Fée, Crypt. vasc. Brésil da Bocaina, 21 Apr. 1951, Brade 20676 (MO); Campos
1: 114. t. 35. f.2. 1869. Type. Brazil: Rio de do Jordão, 5-20 Feb. 1937, Campos Porto 3213 (F, BM);
Janeiro, Serra dos Orgãos, n.d., Glaziou 2819 São José do Barreiro, Serra da Bocaina, 29 May 1958,
Handro 785 (US); Campos do Jordão, Sep. 1945, Leite
(S!, RB!, probably isotype K!, photo BM!). In
3632 (MO, UC). PARANA: banks of Rio da Santa,
the label of the specimen at Kew, the date is between Curitiba and Joinville, 5 Jan. 1974, Conrad &
October 1871. Non Polypodium fallax Schlecht, Dietrich 2052 (UC). Without locality: 1870, White s.n.
1830. (BM).
Polypodium longipetiolatum Brade, Rodriguesia 3:
115. 1937. Nomen novum for Campyloneurum Campyloneurum fallax belongs to the C.
fallax Fée. amphostenon group, which consists os species
Stem long-creeping, no pruinose, 3-5 mm distributed in the Andes and montane areas of
wide. Stem scales light brown in mass, 5-7 mm Mexico and Central America. Campyloneurum
long, 2.5-3.5 mm wide, broadly ovate, adpressed, fallax resembles C. densifolium, but it differs from
persistent, slightly clathrate, the cells oblong or it by broader stem scales, and by its oblong,
broadly oblong, central cells along the main axes isodiametric cells of the stem scales (Fig. 7 c).
of the scale, cell walls 12-16 µm thick, marginal Campyloneurum fallax is the only disjunct species
cells transverse or with irregular disposition, cell in the group, and it may have evolved from an
walls usually 4-8 early isolation, since the Brazilian shield is
(-10) µm thick. Phyllopodia 4-5 mm long, 2.5-3 geologically older than the Andes.
mm wide, 7-10 mm apart. Leaves 25-45 (-75) cm
long; petiole stramineous or dark stramineous,
(5-) 8-14 (-16) cm long, slightly ranurate on the 24. Campyloneurum fasciale (Willd.) C. Presl, Tent.
ventral side, convex on the dorsal, indument of Pterid. 190. 1836.
caducous scales; lamina narrowly lanceolate, Polypodium fasciale Willd., Sp. Pl. 5: 156. 1810.
bases attenuate or subcuneate, rarely cuneate, Type. Caribe, Humboldt 426 (Herb. Willd.
apices acuminate, 1.5-4.5 cm wide, chartaceous- 19632) (holotype, B! ; isotype, P).
subcoriaceous, rarely chartaceous, usually bright
Polypodium serpentinum Christ, Bull. Herb. Boiss. BELIZE: 1907, Peck 636 (F).
2, 6: 51. 1906. Type. Costa Rica: Navarro, HONDURAS. ATLANTIDA: Lancetilla valley, near
Wercklé s.n. (holotype, P!, photo of P, BM!). Tela, 6 Dec. 1927-20 Mar. 1928, Standley 54150 (F, US).
COSTA RICA. ALAJUELA: Santiago, 25 Apr. 1901,
Campyloneurum serpentinum (Christ) Ching,
Amz. 14200 (F); colinas de San Pedro de San Ramón, 27
Sunyatsenia 5: 263. 1940. May 1925, Brenes 4234 (F); San Miguel de San Ramón,
Stem long-creeping, not pruinose, brown, 21 Jul. 1934, Brenes 19258 (F); San Isidro de San Ramón,
greenish-stramineous or black, (1-) 2-3 mm wide. 22 Oct. 1986, Herrera 98 (MO); Buena Vista, San Carlos,
Stem scales brown or dark brown in mass, (1.5-) 21 Jul. 1963, Jimenez 918 (F); Buena Vista, San Carlos,
2-3 mm long, 0.8-1 mm wide, narrowly oblong, 14 Aug. 1964, Jimenez 2295 (F); Santiago, near San
usually pseudopeltate, apices acuminate, Ramón, 21 Apr. 1913, Tonduz 17572 (BM, F, NY, S,
clathrate, cell walls sometimes diffuse, cells UC). LIMON: cerro Colonel, E of Laguna Danto, 15-
oblong, along the main axes of the scale, cell 20 Sep. 1986, Stevens & Montiel 24622 (F, MO). SAN
JOSE: Río Negro, cerro La Cangreja, 20 Jun. 1986,
walls 8-10 µm thick, without differentiate
Chacón & Chacón 1959 (MO); Zona Protectora La
margins. Phyllopodia 0.5-1 mm long, 1-2.5 mm Cangreja, along Quebrada Grande, ca. 2 km NNE of
wide, 0.7-20 mm apart. Leaves 20-40 (-50) cm Mastatal de Puriscal, 23 Jul. 1988, Grayum 8642 (MO);
long; petiole stramineous or brownish, 1-2 cm vicinity of El General, Jul. 1936, Skutch 2663 (GH, NY,
long, slightly ranurate adaxially or slightly S, US). CARTAGO: 2 km W of Orosi, 16 Jan. 1977, Lent
convex abaxially; lamina lanceolate or narrowly 4056 (F, NY) PUNTARENAS: Reserva Forestal Golfo
lanceolate, bases attenuate, apices acuminate, (1-) Dulce, Península Osa, Rancho Quemado, ca. 15 km W
2-5 cm wide, herbaceous-chartaceous, indument of Rincón, 30 May 1988, Hammel et al. 16891 (MO);
slope of Río Java, 20 Nov. 1986, Hennipman et al. 7069
of bicellular, simple hairs, inconspicuous;
(MO); San Vito de Java, 24 Jul. 1972, McAlpin 1474 (F).
stomata polocytic; costa prominent, primary PANAMA. CHIRIQUI: El Boquete, 5 Feb. 1918,
veins prominulous on both sides of the lamina, Cornman 804 (MO, UC, W); around El Boquete, 17
stramineous or darker than the leaf tissue, 70-80o Mar. 1918, Cornman 1151 (W); on NW side of Cerro
divergent from the costa, slightly flexuosous, 5-7 Pando, 21 Jul. 1971, Croat 15933 (AAU, MO); E of
mm apart, secondary veins slightly prominulous Canas Gordas, near Costarican border, 26 Feb. 1973,
Croat 22350 (MO); Ojo de Agua, vicinity of Santa
on only darker than the leaf tissue, forming (4-)
Clara, 17 Jun. 1987, Croat 66296 (MO); Las Lagunas, 18
5-10 primary areoles between the costa and Mar. 1983, Hamilton & Stockwell 3588 (MO, UC); 7.5 mi
margin, areoles entire, usually with 2 free from bridge over Río Chiriqui Viejo, on road to Río
excurrent veinlets; sori medial or subterminal, Sereno, 7 Apr. 1979, Hammel et al. 6855 (MO); valley of
paraphyses not seen, spores 47-50 µm long, 32 Río Caldera, 5-19 Feb. 1918, Killip 5039 (MO); km 3 on
µm wide. Fig. 39. La Unión road, 23 May 1971, Proctor 32029 (MO).
BOCAS DEL TORO: 10-15 mi S from Changuinola
This species is known from Mexico, Costa river, 18 Dec. 1966, Lewis et al. 990 (MO), Lewis et al. 994
Rica and Panama to Bolivia. It grows as an (MO, UC). VERAGUAS: along base of Cerro Tute, 10
Sep. 1982, Hamilton et al. 1310 (MO). COCLE: El Cope
epiphyte in forests, between 100 m and 2000
on Pacific side, 16 Oct. 1979, Antonio 2152 (MO); above
(-2500) m elevation. El Valle, 13 Aug. 1972, Gentry 5667 (MO); 9 km from El
Valle, 12 May 1973, Kennedy et al. 3221 (MO).
REPRESENTATIVE SPECIMENS: MEXICO. PANAMA: 16 km above Pan-Am highway, on road
VERACRUZ: Mun. Hidalgotitlán, NE Hnos. Cedillos from El Llano to Carti-Tupile, 13 Feb. 1973, Kennedy et
camp, 13 Feb. 1974, Dorantes et al. 2453 (F, MO). al. 2451 (F, MO); summit of cerro Campana, 31 Mar.
OAXACA: Santa María Chimalapa, Río Milagro, 4 1969, Porter et al. 4920 (MO). DARIEN: Serranía del
Sep. 1985, Hernandez 1465 (MO). CHIAPAS: 6-8 km N Piré, above Cana gold nime, between Río Cana and
of Ocosingo, along road to Bachajón, 9 Nov. 1971, Río Escucha, 27 Jul. 1976, Croat 37781 (MO); Serranía
Breedlove & A.R. Smith 22151 (F, NY); 45 km N of del Darién, top of cerro Mali, 17 Jan. 1975, Gentry &
Ocozocoautla, above lake of Malpaso, 31 Jan. 1973, Mori 13685 (MO); W ridge of cerro Mali, 23 Jan. 1975,
Breedlove 32816 (F); Mun. Ocosingo, 1 km SE of Monte Gentry & Mori 13838 (MO); S slope of west peak of
Líbano, 8 Aug. 1954, Dressler 1617 (GH, NY, US); 3 km cerro Tacarcuna, 28 Jan. 1975, Gentry & Mori 13967
S de Lacanja-Chanzayab, 11 Aug. 1984, Martinez 6955 (MO); trail from cerro Mali to Río Pucuro, 20 Jul. 1976,
(MO) Finca Mexiquito, Jun. 1913, Purpus 6751 (F, NY, Gentry et al. 16838 (MO); E slope of Cerro Sapo, 3 Fb.
S, UC). 1978, Hammel 1303 (MO); 6 km S from gold mining
camp at Cana, W to Alturas de Nique, 20 Apr. 1980, Campana, Tarapoto, Aug. 1856, Spruce 4647 (BM).
Lellinger 1969 (MO, US); Pirre massif, Alturas de LORETO: Santa Rosa, Yurimaguas, 1-5 Sep. 1929, Killip
Nique, 3 Mar. 1988, McPherson 12204b (MO); along & Smith 28927 (S, US). MADRE DE DIOS: Río Manu,
ridge trail from Cana up the Cerro Pirre, 3 May 1990, Cocha Cashu , 14 Aug. 1978, Foster & Terborgh 6621 (F);
Moran 5045 (F); Río Tuquesa, at lower Tuquesa mining Cocha Cashu, 10 Sep. 1986, Nuñez 6070 (F, NY).
camp, 4 Jul. 1975, Mori 6943 (MO); Parque Nacional BOLIVIA. LA PAZ: Larecaja, Consata, 14 Dec. 1981,
Darien, slopes of cerro Tacarcuma, 27 Oct. 1987, Nevers Beck 4917 (F); Murillo, 44 km below Lago Zongo dam,
et al. 8532 (MO). vic. of Cahua hydroelectric plant, 12-15 Sep. 1983,
WINDWARD ISLANDS. ST. VINCENT: valley of N Solomon 10853 (MO). COCHABAMBA: Carrasco,
fork of Cumbrland river, 2-3 May 1947, Morton 5542 junction of Río Leche with Río Isarsama, 5 May 1977,
(MO). Beck 1633 (F, LPB).
COLOMBIA. CHOCO: Hoya del Río San Juan,
Quebrada La Sierpe, 1 Apr. 1979, Forero & Jaramillo Campyloneurum fasciale is characterized by
4454 (MO). VALLE: Pacífic basin, Río Cajambre, 21-30 clathrate narrowly ovate scales, without
Apr. 1944, Cuatrecasas 17180 (F). META: Sierra de la
differentiated margins. It is closely related to C.
Macarena, between Río Guejar and Sansa, 29 Aug.
1950, Idrobo 521 (AAU, US).
fuscosquamatum, from which it differs by clear
VENEZUELA. LARA/YARACUY: Sierra de Aroa, 10- cell lumen in the stem scales. Campyloneurum
13 mi NW of Urachiche, NW from Urachiche to fasciale belongs to the C. repens group.
Duaca, 16 Nov. 1982, A.R. Smith et al. 1343 (MO).
Parque Nacional Aragua, 2 Dec. 1938, L.Williams 10801
(F). TACHIRA: 35 km SSE of San Cristobal, La 25. Campyloneurum fuscosquamatum Lellinger,
Buenaña, 6-12 km W of Quebrada Colorada, 20-21 Amer. Fern J. 78: 21. 1988. Type. Peru:
Mar. 1981, Liesner & Gonzalez 10876 (MO, UC). Huánuco, Tingo María on the Río Huallaga, 2
BOLIVAR: 5 km S of El Paujil, Río Samay, affluent of
Nov. 1938, Stork & Horton 9452 (holotype US!;
Icabaru, 21 Oct. 1985, Liesner & Holst 18843 (MO).
FRENCH GUIANA. Saul: Monts La Fume, 13 Oct. isotypes F!, GH, UC!, photo of US at F!,
1982, Boom & Mori 2016 (CAY, F); camp Tigre, 4.5 km USM!).
N of Saut Mais , 16 Jan. 1980, Cremers 6121 (CAY). Stem green, geenish-stramineous to black, not
ECUADOR. CARCHI: trail from Rafael Quindís finca pruinose, 2-3 mm wide. Stem scales dark borwn,
to Río Verde, 26 Nov. 1987, Hoover & Wormley 1679 caducous, persistent only at the growing areas,
(MO), Hoover & Wormley 1710 (MO). MANABI: narrowly ovate or linear, 2.5-3 (-3.5) mm long,
Chone-Santo Domingo road, near Río La Morena, 15 0.5-0.8 mm wide, the cells narrowly oblong along
km NNE of Flavio Alfaro, 7 May 1980, Harling &
the main axes of the scale, usually not
Andersson 18911 (AAU, F, GB, QCA). PICHINCHA:
Tinalanadia, 9.6 km E of Santo Domingo, 3 Apr. 1983,
differentiate at the margins, cell walls (8-) 12-16
Croat 55708 (MO); road La Unión del Toachi, San (-20) µm thick, lumen yellowish or obliterate.
Francisco, 19 Mar. 1985, Harling & Andersson 23146 Phyllopodia 0.5-1 mm long, 1.5-2 mm wide, 7-10
(QCA). NAPO: San Pablo de los Secoyas, 6 Aug. 1980, (-20) mm apart. Leaves 20-40 cm long; petiole
Brandbyge et al. 32545 (AAU, QCA); Río Aguarico, 0.5-5 cm long, stramineous, sometimes darker
Tangoy, 29 Aug. 1979, Holm-Nielsen et al. 20145 (AAU, abaxially; lamina narrowly obovate or narrowly
QCA); Nuevo Rocafuerte, 27 Feb. 1981, Jaramillo & elliptic, bases attenuate or narrowly cuneate,
Coello 4414 (AAU, QCA); San Pablo at Río Aguarico, 5
apices acuminate, 2-5 (-8) cm wide, herbaceous-
May 1984, Laegaard 52083 (QCA); Añangu, Parque
chartaceous, margins cartilaginous, slightly
Nacional Yasuní, 30 May-21 Jun. 1982, Øllgaard et al.
39007 (AAU, QCA), Øllgaard et al. 39110 (AAU); revolute or plane, indument of inconspicous,
Cabañas Aliniahui, S of Río Napo, 29 Sep. 1989, Zogg simple, multicellular hairs, 80-96 m long,
& Gassner 13127a (QCA). ZAMORA-CHINCHIPE: scattered abaxially; costa prominent, indument
new road Loja-Zamora, 15 Feb. 1991, Øllgaard & Moran of scales similar to those at the stem, primary
98831 (QCA). veins prominent or prominulous, stramineous,
PERU. AMAZONAS: Bagua, 12 km E of La Peca,
(60o-) 70-75o (-80o) divergent from the costa,
Barbour 2487 (MO, UC), Barbour 2495 (F, UC); Serranía
de Bagua, 14 Jun. 1978, Gentry et al. 22930 (F, MO, UC); straight or slightly sinuate, 4-6 mm apart,
Bongará, SW of Pomacocha, Wurdack 844 (F, NY, UC, transverse secondary veins forming 6-10 primary
US, USM); Sipabamba, Shillac, 5 May 1981, Young & areoles between the costa and margin, excurrent
Eisenberg 352a (MO, NY, UC). SAN MARTIN: Monte veinlets 2 (-3), irregular marginal areoles present
with 0-1 (-4) excurrent veinlets; stomata Huallaga, below Yurimaguas, 1-5 Sep. 1929, Killip &
copolocytic or polocytic; sori subterminal or Smith 28854 (F, US); San Antonio, Río Itaya, 18 Sep.
medial, paraphyses not seen, spores 56 µm long, 1929, Killip & Smith 29372 (NY, US); above Pongo de
Manseriche, left bank of Río Santiago, 23 Nov. 1931,
32 µm wide. Figs. 3 b, c; 14 g; 17 e; 40.
Mexia 6141a (MO, UC, US); creek Inche, 3 Jan. 1932,
Mexia 6371a (UC, US). HUANUCO: Fundo Chela,
This species is known from Colombia to Sinchono, 3 Aug. 1948, Aguilar 945 (F, USM); Tingo
Bolivia, where it grows as an epiphyte in María, 30 Oct. 1949-19 Feb. 1950, Allard 22329 (US);
lowland forests between 100 m and 1500 m Tingo María, 4 Oct. 1972, Croat 21034 (US); Tingo
elevation. María-Pucallpa, 1971, Ellenberg 3849 (GH); abajo de la
Divisoria, cerca Sinchono, 21 Jul. 1948, Ferreyra 1101 (F,
REPRESENTATIVE SPECIMENS: COLOMBIA. USM); Huánuco, Tingo María, 24 Sep. 1954, Ferreyra
META: Cordillera La Macarena, trail between Río 10276 (GH, USM); Pachitea, Codo de Pozuzo, 22 Oct.
Guejar and Guapayita, 20-28 Dec. 1950, Idrobo & 1982, Foster 9399 (F, USM); La Divisoria, Tingo María-
Schultes 817 (GH, US); Sierra de La Macarena, Caño Pucallpa road, near Loreto border, 29 Mar. 1977,
Estrada, 15 Dec. 1949, Philipson & Idrobo 1753 (BM, Gentry et al. 18821 (F, MO); Churubamba, Balsa-Playa,
US). 12 Sep. 1936, Mexia 8176 (BM, F, GB, GH, MO, NY, S,
ECUADOR. NAPO: Añangu, S bank of Río Napo, 95 UC, US); near confluence of Río Cayumba with Río
km downstream from Coca, 19 Jun.-14 Jul. 1985, Huallaga, 10 Oct. 1936, Mexia 8271 (BM, F, GB, GH,
Balslev et al. 60573 (QCA); 2 km downstream from MO, NY, S, UC. US); above Río Cayumba, 19 Oct.
Puerto Aguarico, 6 Apr. 1980, Brandbyge et al. 30478 1936, Mexia 8312 (UC); Pachitea, Honoria, Bosque
(AAU, QCA); San Pablo de los Secoyas, 11 Aug. 1980, Nacional Iparía, 14 Mar. 1967, Schunke V. 1762 (F, GH,
Brandbyge & Asanza 32792 (AAU, QCA); Río Aguarico, USM); Tingo María on Río Huallaga, 19 Oct. 1938,
San Pablo de los Secoyas, 13 Feb. 1980, Holm-Nielsen et Stork & Horton 9452 (F, MO, UC); Huamalíes, Supte
al. 21061 (AAU, QCA); Añangu, Río Napo, 16-27 Apr. River, N of Tingo María, 2 Nov. 1938, Stork & Horton
1983, Lawesson et al. 39459 (AAU); trail from Santa 9568 (F, GH, UC, US). PASCO: Puerto Bermudez, 14-
Cecilia up Río Aguarico, 29 Mar. 1972, Mac Bryde & 17 Jul. 1929, Killip & Smith 26444 (US); Pozuzo, 20-22
Dwyer 1347 (MO, QCA). PICHINCHA: Cantón Santo Jun. 1923, Macbride 4585 (F, US); Oxapampa, vicinity of
Domingo, 12 km E of Patricia Pilar, 23 Aug. 1978, Chequitavo, Gran Pajonal, 26 Sep. 1983, D.N. Smith
Dodson et al. 7181 (AAU, F, MO); station ENDESA, km 5625 (UC). JUNIN: Pichis trail, Yapas, 28-29 Jun. 1929,
113 along Quito-Puerto Quito, 16-17 Nov. 1989, Luteyn Killip & Smith 25606 (F, NY, US); Perené, 1960, Kunkel
& Borchsenius 13362 (QCA); ENDESA, km 113 via 618 (GH); Chanchamayo valley, 1924-1927, C. Schunke
Quito-Puerto Quito, 23-24 Apr. 1983, Rodriguez et al. 123 (US); above San Ramón, 1923, C. Schunke 166 (GH,
113 (QCA), 25 Feb. 1984, Rodriguez 293 (QCA). US); La Merced, 1909, Schunke 25 (S, UC);
ZAMORA-CHINCHIPE: N side of Río Palanda with Chanchamayo, Feb. 1939, Soukup 1103 (F).
Río Zumba, 30 Jan. 1985, Harling & Andersson 21286 AYACUCHO: Río Apurímac, valley near Kimpitiriki,
(QCA). 10-11 May 1929, Killip & Smith 22861 (F, US); La Mar,
PERU. SAN MARTIN: Mariscal Cáceres, Madre Mía, along Río Catute, 2 km NW of Santa Rosa, 8 Sep. 1976,
16 Mar. 1977, Boeke & Ramirez 1329 (NY, UC); San Wasshausen & Encarnación 621 (MO, US, USM).
Martín, km 28 of Tarapoto-Yurimaguas road, 17 Aug. CUSCO: La Convención, 4 km NE from the Hacienda
1986, Knapp 8040 (MO, USM), Knapp & Mallet 8393 Luisiana, 31 Jul. 1968, Dudley 11501 (GH, US).
(NY); Tarapoto, Alto Pucayacu, Montes 47 (F); MADRE DE DIOS: Tambopata Reserve Zone, 13 Mar.
Campanillas, trail to Achiras, SW from Sión, 23 Oct. 1988, Bell & Wiser 88-314 (F); Parque Nacional Manu,
1969, Schunke V. 3556 (F, MO, US, USM); Mariscal Cocha Cashu Biological Station, Foster P-84-17 (MO);
Cáceres, Tocache Nuevo, Schunke V. 3590 (UC, US); Parque Nacional Manu, Cocha Cashu, 20 Aug. 1976,
San Roque, Williams 7255 (F, US); Tarapoto, 4 mi E of Foster & Augsburger 3282 (F); Tambopata, Río Piedras,
Tarapoto, Woytkowski 35235 (MO, S, UC). LORETO: 17 Jan. 1967, Vargas 18666 (GH).
Maynas, near Brilla Nueva, Boro indian village, on BOLIVIA. LA PAZ: Nor Yungas, Polo Polo bei
upper Río Yaguasyacu, 8 Nov. 1977, Gentry & Revilla Coroico, Oct.-Nov. 1912, Buchtien 3535 (UC); Buchtien
20412 (MO); Maynas, Quebrada Tamshiyacu, E of 3536 (F, S, US); 6 km N (below) Consata, 15 Dec. 1981,
Tamshiyacu, 19 Mar. 1979, Gentry et al. 25837 (F, MO, Solomon et al. 6575 (MO). BENI: Ballivian, Río
US); Maynas, Yanamano, Explorama Tourist Camp, on Colorado, 21 Jun. 1989, Fay & Fay 2083 (F, MO);
Río Amazonas, between Indiana and mouth of Río Ballivian, Río Colorado, 22 Jun. 1989, Fay & Fay 2092
Napo, 26 Jul. 1980, Gentry et al. 29028 (MO); (F), 24 Jun. 1989, Fay & Fay 2130 (F). Ballivian, lower
Yurimaguas, lower Río Huallaga, 23 Aug.-7 Sep. 1929, slopes of serranía Pilón Lajas, 14.3 km N of the bridge
Killip & Smith 27668 (F, US); Santa Rosa, lower Río over the Río Quiquibey, Solomon 13893 (MO); Río
Chaparé, Mamoré, Aug. 1926, Werdermann 2169 (MO).

SANTA CRUZ: Ichilo, Parque Nacional Amboró, This species is known only from Colombia
along Río Saguayo, 18 Jan. 1988, Nee & Saldias 36839 and Peru, where it grows in forests between 2200
(MO); Parque Nacional Amboró, ca. 15 km SE up the
m and 2400 m elevation.
Río Pitasama, from the Río Surutú, Solomon & Urcullo
14090 (MO, US); Sara, bosques del Río Surutu, 1 Oct.
1925, Steinbach 7246 (F, S, UC). COCHABAMBA: EXAMINED SPECIMENS: PERU. AMAZONAS:
Carrasco, Cantón Chirquioma, Isarsama camp, 3 May Bongará, WSW of Pomacocha, Wurdack 868 (US).
1979, Beck 1569 (LPB); Antahuacana, N von
Cochabamba, Jun. 1909, Buchtien 2154 (S, UC, US);
Buchtien 2155 (S, US). 27. Campyloneurum lorentzii (Hieron.) Ching,
Sunyatsenia 5: 263. 1940.
Campyloneurum fuscosquamatum is Polypodium lorentzii Hieron., Bot. Jahrb. Syst. 22:
characterized by its dark brown, narrowly ovate 406. 1896. Lectotype (chosen by Sota, Opera
or linear stem scales. It is closely related to C. Lilloana 5: 102. 1960). Argentina: Tucumán,
fasciale, from which it might be derived. They Tafi, 4 Apr. 1872, Lorentz 781 (B!, photo BM!).
can be differentiated by the characters provided Stem creeping, dark stramineous, pruinose,
in the key. 4-6 mm wide. Stem scales light brown in mass,
Campyloneurum fuscosquamatum belongs to the 3-4 mm long, 2.5-3 mm wide, ovate, bases
C. repens group. auriculate, apices obtuse or ending in a
multicellular hair, scales slightly clathrate, the
cells broadly oblong, cell walls diffuse, central
26. Campyloneurum inflatum Lellinger, Amer. cell walls 20 µm thick, marginal cell walls 5 µm
Fern J. 78: 22. 1988. Type. Colombia: Cauca, thick. Phyllopodia 2.5-4 mm long, 2.5-3 mm
W slope of Cerro Munchique, Pérez-Arbelaez wide, 0.6-10 mm apart. Leaves 50-75 cm long;
& Cuatrecasas 6244 (holotype US!; isotypes petiole dark stramineous, 5-27 cm long, ranurate
COL, F!; photo of US, USM!). adaxially, convex abaxially, indument of
Stem long creeping, green stramineous or caducous scales similar to those on the stem;
black, not pruinose, 2-3 mm wide. Stem scales lamina narrowly lanceolate, bases attenuate,
light brown in mass, 4-5 mm long, 0.9-1 mm apices long acuminate, 2.5-4.5 cm wide,
wide, lanceolate, bases auriculate, apices herbaceous-chartaceous, margins cartilaginous,
acuminate, clathrate, the cells oblong. slightly undulate, indument of inconspicuous,
Phyllopodia 1-2 mm long, 2.5-3 mm wide, 30-40 bicellular, simple hairs; stomata polocytic; costa
mm apart. Leaves 30-55 cm long; petiole prominent, primary veins prominulous,
stramineous to dark stramineous, 9-21 cm long; stramineous, 50-60o divergent from the costa, 5-
lamina elliptic, bases acute, apices caudate, 6- 7 mm apart, secondary veins slightly
10.5 cm wide, subcoriaceous, margins strongly prominulous, same color as the leaf tissue,
cartilaginous, slightly revolute, indument of transverse secondary veins forming (3-) 4-5
scattered bicellular, simple hairs; hypostomatic, primary areoles between the costa and margin,
stomata polocytic; costa prominent, indument of primary areoles symmetrically divided, (0-) 1-2
caducous scales, primary veins slightly free excurrent veinlets in each secondary areole,
prominulous adaxially, prominent abaxially, sometimes free excurrent veinlets at the margin;
slightly flexuous, usually stramineous, 65-70o sori medial, paraphyses not seen, spores 50-65
divergent from the costa, 7-8 mm apart, µm long, 32-40 µm wide. Fig. 36.
secondary veins forming 6-7 primary areoles
between the costa and margin, 2 (-4) free This species is found in Bolivia and northern
excurrent veinlets in each primary areole, entire, Argentina, where it grows between 1400 m and
sometimes secondary areoles close to the 3000 m elevation, usually as a terrestrial.
margin; sori subterminal on the excurrent
veinlets, paraphyses absent; spores 67-76 µm REPRESENTATIVE SPECIMENS: BOLIVIA. LA PAZ:
long, 37-42 µm wide. Fig. 40. Prov. Murillo, Valle de Zongo, Santa Rosa, 3 km hacia
La Paz, 8 Apr. 1979, Beck 1099 (F); Nor Yungas, along hairs, scattered abaxially; stomata polocytic;
road between Unduavi and Chulumani, ca. 5 km costa prominent, primary veins prominulous on
beyond Aceromarca, 25 Nov. 1980, Croat 51465 (LPB, both sides of the lamina, stramineous, slightly
UC).
ARGENTINA. CATAMARCA: Río Potrero, 10 Feb. flexuosous, 65-70o divergent, 6-7 mm apart,
1949, Brücher & Brücher s.n. (S); Rodeo, 30 May 1910, secondary veins forming 6-10 primary areoles
Castillón s.n. (GH, US); Ambato-Rodeo, 20 May 1910, between the costa and margin, primary areoles
Castillón s.n. (NY, S, UC). Andalgalá, Esquina Grande, entire, 2 (-3) free excurrent veinlets; sori
3 May 1915, Jörgensen 1497 (UC). TUCUMAN: road subterminal, paraphyses filamentosous,
from Concepción to Andalgalá (Catamarca), 16 Oct. branched, same length as sporangia, spores 60-
1989, Kramer et al. 10708 (F); Tafí, Tafí del Valle-Los
66 µm long, 42 µm wide. Fig. 41.
Nogales, 6 May 1947, Meyer 12110 (W); Tafí, Quebrada
de las Sosa, Los Nogales, 12 Nov. 1952, Petersen &
Hjerting 609 (BM); Chicligasta, Quebrada de Cochuna, This species is known from Colombia and
17 Oct. 1957, Sota s.n. (S); Monteros, Piedra Labrada Venezuela, where it grows as an epiphyte
and Puerto del Pino, 24 Apr. 1949, Verworst 482 (US, between 1500 m and 2500 m elevation.
W); Tafí, Taficillo, 13 Mar. 1928, Venturi 5891 (BM, F,
MO). JUJUY: road to Lozano a Tiraxi, 3 Nov. 1974, REPRESENTATIVE SPECIMENS: COLOMBIA.
Schinini et al. 10210 (UC). CUNDINAMARCA: Salto de Tequendama, 8 Mar.
1939, Alston 7405 (MO); Cordillera Oriental, WNW of
Campyloneurum lorentzii is closely related to Bogotá, 31 Dec. 1944, Little & Little 9151 (F, US).
C. densifolium. The former has stem scales with Sebastopol, 3 Sep. 1944, Little & Little 8603 (F).
obtuse apices, while the latter has scales with VENEZUELA. TACHIRA. Valencia, Pico de Vela,
Buena Vista, 11 Nov. 1945, Charpin & Jacquemont 13137
acuminate or acute apices.
(F); Junín, S slopes of Cerro San Isidro, Davidse &
González 22222 (MO); trail leading to summit of
Páramo de Tama, 29 Jan. 1978, Luteyn et al. 5352 (F,
28. Campyloneurum macrosorum Fée, Gen. Fil. 96. UC). YARACUY: Sierra de Aroa, 9 km W of San
1854-1857. Type. Colombia: Ocaña, Schlim Felipe, 4 Apr. 1980, Liesner & González 10010 (MO).
440 (holotype, not found; isotypes, B!, K!,
photo of K, BM!). Campyloneurum macrosorum is characterized
Polypodium lindigii Mett., Ann. Sc. Nat. 5: 257. by lanceolate, long petiolate leaves, with black
1864. Cited Syntypes: Colombia, Bogotá, San stems, and by persistent, subadpressed,
Antonio, Lindig 172 (B!, BM!, K!); La Vega, marginally differentiated stem scales.
Lindig 338 (B!, BM!, K!); Ocaña, Schlim 440 (B!, Among the synonyms is included Polypodium
K!, photo of K, BM!); Schlim 724 pp (not seen). lindigii Mett., which was based on material from
Lectotype (chosen here) Colombia: Bogotá, Colombia, among them the type collection of
San Antonio, Lindig 172 (B!, BM!, K!). Campyloneurum macrosorum Fée. During this
Campyloneurum lindigii (Mett.) Ching, study three of the four syntypes were studied;
Sunyatsenia 5: 263. 1940. they clearly belong to this taxa as defined today.
Stem long-creeping, green turning black or This species can be differentiated from C.
stramineous, 2-4 mm wide. Stem scales light repens by having persistent, subclathrate stem
brown in mass, persistent, subadpressed, 3-4 scales and a thick stem. In addition, cells of the
mm long, 1 mm wide, lanceolate, bases slightly stem scale run along the axis, and the margin is
auriculate, apices acuminate, slightly clathrate, as broad as the center part of the scale.
the cells oblong, cell walls diffuse. Phyllopodia Campyloneurum macrosorum belongs to the C.
1-2 mm long, 1.5-3 mm wide, 25-30 mm apart. repens group.
Leaves 30-50 cm long; petiole dark stramineous,
4.5-22 cm long, ranurate adaxially; lamina
lanceolate or narrowly lanceolate, bases 29. Campyloneurum magnificum T. Moore, Ind. Fil.
narrowly cuneate, apices acuminate, 5.5-7 cm 226. 1861. Type. Venezuela, Fendler 410
wide, margins cartilaginous, slightly sinuate to (holotype, K!; isotypes, B!, F!, MO!, NY!).
undulate, indument of inconspicuous, bicellular Figs. 1 c; 41.
Polypodium fendleri D. C. Eaton, Mem. Amer. VENEZUELA. YARACUY: Sierra de Aroa, Cerro
Acad. Arts n.s. 8: 199. 1860. Type. Venezuela, Tigre, 10 km of Aroa,30 Mar. 1980, Liesner & González
Fendler 410 (holotype MO!, isortypes B!, F!, 9722 (MO); Liesner & Gonzalez 9738 (MO). LARA-
YARACUY: dist. Urdaneta and Bolívar, 16-17 Feb.
K!, NY!). Non Campyloneurum fendleri T.
1985, Ortega & R. F. Smith 2455 (MO). FALCON:
Moore, 1861. Sierra de San Luis, above Santa María, 10 Jan. 1979,
Polypodium decurrens Raddi var. fendleri (D. C. Werff et al. 126 (MO, UC).
Eaton) Hook., Sp. Fil. 5: 43. 1864. ECUADOR. PICHINCHA: Cantón Quito, Reserva
Campyloneurum fendleri (D. C. Eaton) J. Smith, Maquipucuna, 13 Sep. 1989, Webster & Addison 27527
Hist. Fil. 97. 1875. Nom. superfl. (QCA); San Miguel de los Colorados, Oct. 1893, Sodiro
Campyloneurum juglandifolium Fée, Crypt. Vasc. s.n. (UC). PASTAZA: Baños (Jivaría), Stübel 983 (B).
Br. 1. 1869. Type. Brazil: Rio de Janeiro, PERU. SAN MARTIN: Tarapoto, 4 mi E of Tarapoto,
Glaziou 1750 (holotype, P!). Woytkowski 35218 (MO, UC). PASCO: Oxapampa,
Gran Pajonal, trail to Shumahuani from Chequitavo,
Polypodium magnificum (T. Moore) Hieron., Bot.
24 Sep. 1983, D.N. Smith 5212 (MO). JUNIN:
Jahrb. Syst. 34: 535. 1904. Chanchamayo, Schunke 22 (F); Schunke 683 (F); Schunke
Stem creeping, not pruinose, 10-15 mm wide. 913 (F); Soukup 1089 (F).
Stem scales light brown in mass, 4-6 mm long, 2 BOLIVIA. LA PAZ: Polo-Polo, Coroico, Oct. Nov.
mm wide, ovate, clathrate, the cells oblong, 1912, Buchtien 3490 (F, S, US); Larecaja, 6 km N of
along main axes of the scale, cell walls 10-20 µm Consata, 15 Dec. 1981, Solomon et al. 6574 (MO).
thick. Phyllopodia 2-7 mm long, 5-9 mm wide,
5-10 mm apart. Leaves 1.5-3 m long; petiole dark Campyloneurum magnificum is one of two
stramineous, 40-95 cm long, ranurate adaxially; species with pinnate leaves; it is characterized by
lamina 1-pinnate, 4-7 pair of pinnae, pinnae its elliptic pinnae with caudate apices. During
elliptic, base attenuate, apices caudate, 5-11 cm this study all southern Brazilian specimens,
wide, herbaceous-chartaceous, subsessil, margin except the Glaziou specimen (Glaziou 1750),
cartilaginous, sinuate, indument of scattered, belong to C. decurrens and not to this taxa, which
simple branched hairs, mainly along veins; occurs in more continental areas in South
stomata polocytic; costula prominent, primary America.
veins prominent, stramineous or darker than the Together with Campyloneurum decurrens form
leaf tissue, 55-60o divergent from the costula, 8- the C. magnificum group, that may represent an
10 mm apart, secondary transversal veins ancient lineage in the genus.
forming 8-12 (-15) primary areoles, primary
areoles entire, 3-4 free excurrent veinlets in each
areole, simple or furcate; sori terminal, 30. Campyloneurum minus Fée, Gen. Fil. 258.
paraphyses not seen, spores 35-45 µm long, 22-27 1852. Type. America Austral, Glaziou s.n.
µm wide. (holotype, n.s.; isotype RB).
Polypodium herbaceum Christ in Schwacke, Pl.
This species is known from Panamá, Nov. Mineiras 2: 22. 1900. Type. Brazil: Rio
Trinidad, Colombia, Venezuela to Bolivia and de Janeiro, Paineiras, Schwacke 5384 (holotype
Brazil. It grows in shady forested areas, from P!; isotypes BM!).
sea level to 2100 m. Campyloneurum herbaceum (Christ) Ching,
REPRESENTATIVE SPECIMENS: PANAMA. Darien: Stem long creeping, green turning black,
cerro Pirre, 10-20 Jul. 1977, Folsom 4534 (MO). dark stramineous, not pruinose, 1-2 mm wide.
COLOMBIA. DEL VALLE: Western Cordillera, Río Stem scales dark brown in mass, 2-2-5 mm long,
Digua, Quebrada de San Juan, below Queramal, 8 1.2-1.5 mm wide, ovate, slightly clathrate,
Nov. 1946, Cuatrecasas 22735 (F, S); Mun. Calima, slightly bullate, bases auriculate, apices acute or
Campo Alegre, Alto Calima, 29 Aug. 1981, Silverstone obtuse, the cells oblong or ovate, irregularly
539 (MO). Cordillera Occidental, Quebrada del Río arranged, cell walls 8-10 µm thick. Phyllopodia
Blanco, Dec. 1942, Cuatrecasas 13667 (F, UC); San
0.5 mm long, 0.5-2 mm wide, 2-7 mm apart.
Antonio, Bogotá, Lindig 307 (B, F). Cerro Pelado,
Stübel 1252 (B). Leaves 15-40 cm long; petiole stramineous or
dark stramineous, 0.7-8 cm long, slightly 21 Dec. 1980, Abrell 32 (MO). Cordillera de Altos, 7
ranurate adaxially, convex abaxially; laminae Aug. 1902, Fiebrig 13a (B, F). Cordillera Mbatobi, Apr.
narrowly lanceolate, bases and apices attenuate, 1881, Balansa 2882 (B). Without locality. 1885-1895,
Hassler 421 (K).
1-2.5 (-4.5) cm wide, herbaceous-chartaceous,
margins cartilaginous, slightly sinuate or
Campyloneurum minus is characterized by its
undulate, indument of adaxially scattered
slightly bullate, ovate lanceolate stem scales. It
bicellular, simple hairs; stomata polocytic; costa
belongs to the C. repens group.
prominent, ranurate adaxially, primary veins
prominulous, stramineous or darker in color
than the leaf tissue, slightly flexuous, 60-65o 31. Campyloneurum nitidissimum (Mett. in Triana
divergent from the costa, 5-7 mm apart, et Planchon) Ching, Sunyatsenia 5: 263. 1940.
secondary veins inconspicuous, forming (3-) 5-7 Polypodium nitidissimum Mett. in Triana et
primary areoles between costa and margin, Planchon, Ann. Sc. Nat. n.s. 5: 258. 1864.
areoles undivided, 2 free excurrent veinlets in Type. Colombia: San Antonio, Jan. 1861,
each areole, margins sometimes with free Lindig 363 (holotype B!; isotypes BM!, K!,
excurrent veinlets; sori medial or subterminal, US!).
paraphyses not found, spores 40-45 µm long, 25 Stem long-creeping, not pruinose, 5-10 mm
µm wide. Figs. 17 f; 39. wide. Stem scales dark brown in mass, linear or
narrowly ovate, (5-) 7-15 mm long, (0.5-) 1-1.5
This species occurs in Brazil, Argentina and mm wide, scales non-clathrate, the cells oblong,
Paraguay, where it grows between 100 m and marginal cell walls 8 µm thick, central cell walls
1800 m elevation. 12-16 µm thick, sometimes with hairs on the
margins 5-7 m long, cell lumen dark yellow.
Phyllopodia 0.5-1 mm long, 4-6 mm wide, 5-7
REPRESENTATIVE SPECIMENS: BRAZIL. MINAS
mm apart. Leaves entire; petiole with or without
GERAIS: Carangola, trail Araponga to fazenda de
Grama, 27 Jan. 1930, Mexia 4243 (B, F, MO, UC). RIO
wings, slightly ranurate on the adaxial side,
DE JANEIRO: Guanabara, Paineiras, near Pedra de convex on the abaxial side, glabrate; lamina
Beijo, 15 Nov. 1965, Carauta 285 (F); Rio de Janeiro, narrowly lanceolate or elliptic-lanceolate,
1873, Mosén 66 (B); Guanabara, 14 Feb. 1945, Occhioni chartaceous or coriaceous, margins
20 (F); Parque Nacional Serra dos Orgãos, 6 Aug. 1961, cartilaginous, plane or scarcely revolute,
Pabst 5653a (B, F); vic. Paineiras, Corcovado, 14 Nov. sinuous, bases attenuate or abruptly cuneate
1928, Smith 1213 (GH); Serra dos Orgãos, Vauthier 605 then decurrent on the petiole, indument of
(F). SÃO PAULO: km 78 on road 116, between
bicellular hairs, entire, inconspicuous, scattered;
Curitiba and São Paulo, 5 Jan. 1974, Conrad & Dietrich
stomata polocytic, rarely anomocytic; costa
1980 (MO); São Paulo, Serra da Cantaeira, 18 Jul. 1960,
Eiten et al. 2162 (F); Caraguatuba, 3.5 km NNW of prominent, slightly ranurate adaxially, plane or
Caraguatuba, 20 May 1961, Eiten & Eiten 2840 (F); angulate abaxially, primary veins prominent,
Cunha, 11-14 Feb. 1981, Filho 528 (MO), secondary veins forming 7-15 primary areoles,
Paranapiacaba, 4 Oct. 1956, Handro 634 (US); Serra da primary areoles asymmetrically divided, 3-4
Bocaina, Casa do Peixe, 8 Feb. 1959, Pabst 4702 (B); 11 simple or furcate excurrent veinlets in each
Feb. 1959, Pabst 4786 (B). SANTA CATARINA: Mun. primary areole, 1-2 veinlets connected to the
Florianópolis, Rio Tavares, 13 Mar. 1952, Smith & Reitz transverse veins forming asymmetrical
6181 (MO, US). PARANÁ: Iguaçú falls, 16 Sep. 1976,
secondary areoles; sori 2-4 rows between
Davis & Shepherd 60593 (F); Mun. Morretes, Estação
Marumbi, 23 Nov. 1984, Kummrow & Hatschbach 2529 primary veins, sori subterminal or compital;
(UC); Estação Marumbi, 1 Feb. 1986, Kummrow & paraphyses not seen.
Cordeiro 2704 (UC).
ARGENTINA. MISIONES; Iguazu, Osten 7263 (S). Campyloneurum nitidissimum grows mainly as
PARAGUAY. ITAPUA: El Tirol, 19.5 km NNE of a terrestrial plant in disturbed forests. It belongs
Encarnación, 16 Oct. 1981, Foster 81-31 (UC); San to the Campyloneurum brevifolium group.
Bernandino, 10 Oct. 1893, Lindman 2191 (F, S). Morphological variation within this taxon is
PARAGUARI: Parque Nacional Ybicui, pond Guaraní,
found in lamina size and in the division of the Polypodium nitidissimum Mett. var. latius (latior)
primary areoles. These variations appear to be Rosenst., Repert. Spec. Nov. Regni Veg. 12:
continuous, and they may be a response to 474. 1913. Type. Bolivia: Yungas
environmental conditions, since those specimens septentrionalis, Polo Polo, prope Coroico, 900
growing in open areas have thicker and m, Buchtien 3526 (holotype not located;
narrower leaves compared to those growing in isotypes F!, S!, US!).
forests. Based on the available information, two Stem 5-10 mm wide, stem scales (5-) 7-15 mm
varieties are recognized, although future studies long, (0.5-)1-1.5 mm wide. Leaves 60-110 cm
may allow the recognition of two different long; petiole dark stramineous, 2-7 cm long, with
species. The typical variety is characterized by wings 1.5-4 mm wide, lamina lanceolate or
narrow lanceolate leaves, and with primary elliptic-lanceolate, bases abruptly cuneate or
veins more than 70o divergent from the costa. attenuate, then decurrent on the petiole, apices
The variety latius has more elliptical lanceolate usually caudate, (4.5-) 6.5-13.5 cm wide,
herbaceous-chartaceous or chartacea; primary
leaves, and primary veins 70o or less divergent
from the costa. veins (60°-) 65-70° divergent from the costa, (3-)
5-7 (-10) mm apart, secondary transverse veins
- Lamina herbaceous-chartaceous, usually more forming 9-15 areoles between the costa and
than 5 cm wide. var. latius margin; spores 50-60 µm long, 35-40 µm wide.
- Lamina subcoriaceous, usually less than 5 cm Figs. 6 a; 42.
wide. var. nitidissimum
This variety is found in Colombia, Ecuador,
31a. Campyloneurum nitidissimum var. Peru, and Bolivia. On abundant humus, usually
nitidissimum as a terrestrial plant, although sometimes it has
Stem 10 mm wide, stem scales 10-14 mm been found as a climber, in forested areas
long, 1-1.5 mm wide. Leaves 60-90 cm long; between 100 m and 2000 m elevation.
petiole dark stramineous, 17-25 cm long; lamina
narrowly lanceolate, bases attenuate, apices REPRESENTATIVE SPECIMENS: COLOMBIA.
TOLIMA: Alto del Consuelo, Honda, Jul. 1923, Ariste
acuminate, 4-6 cm wide, subcoriaceous; primary
997 (F, GH). CUNDINAMARCA: near bridge San
veins 75-80o divergent from the costa, 4-9 mm Antonio de Tena, 10 May 1940, Cuatrecasas 8270 (F).
apart, secondary transverse veins forming 7-12 CAUCA: Central cordillera, basin of Río Palo,
primary areoles between the costa and margin; betweenTacueyó and La Tolda, 19 Dec. 1944,
spores 58-60 µm long, 35-40 µm wide. Fig. 42. Cuatrecasas 19499 (F, GH, US).
ECUADOR. NAPO: Rio Waui Si Ayá, a northern
tributary to Río Aguarico, 8 Aug. 1980, Brandbyge et al.
The typical variety is known from Colombia,
32661 (AAU); Río Yasuní, Garza Cocha, 12 Apr. 1983,
Peru and Bolivia, where it grows in open areas
Lawesson et al. 43536 (AAU). PASTAZA: Río Bufeo,
above 1500 m elevation. northern tributary of Río Bobonaza, 19 Jul. 1980,
Øllgaard et al. 34779 (AAU, QCA). PICHINCHA:
REPRESENTATIVE SPECIMENS: COLOMBIA. between Nono and Nanegalito, NW of Quito, 4 Sep.
CAUCA: Cordillera Central, western side, Hoya del 1976, Croat 38838 (UC). ZAMORA-CHINCHIPE:
Río Palo, 19 Dec 1944, Cuatrecasas 19499 (F, GH, MO, Shaime at junction of Río Nangaritza. 7 Dec. 1990,
US). Øllgaard 98440 (QCA).
PERU. HUANUCO: Muña, Macbride 4040 (F, GH, US). PERU. HUANUCO: near confluence of Río Cayumba
Without locality: Peruvian Andes, Ruiz 12 (B). with río Huallaga, 10 Oct. 1936, Mexia 8272 (BM, F, S,
BOLIVIA. LA PAZ: Murillo, 44 km below Lago Zongo UC, US). PASCO: Oxapampa, canyon of
dam, vic. of Cahua hydroelectric plant, 12-15 Sep. Huancabamba, above Quebrada Honda, 17 Aug. 1985,
1983, Solomon 10758 (MO). León 667 (F, USM). JUNIN: Pichis trail, between
Miriatiriani and Yessup, 28 Jun.-8 Jul. 1929, Killip &
31b. Campyloneurum nitidissimum var. latius Smith 26221 (US); road to Tarma, before Carpapata, 1
(Rosenst.) B. León, Fieldiana Bot. n.s. 1993: Oct. 1982, León 335 (USM), León 340 (F, GH, USM);
in press. Yaupe, 2 Jul. 1961, Woytkowski 6401 (MO, US); Manto
and Yaupi, 11 Jul. 1961, Woytkowski 6542 (MO); long, ranurate adaxially, with sparse bicellular,
Yunguy, 14 Jul. 1961, Woytkowski 6592 (MO); simple hairs; lamina lanceolate or narrowly
Yucapata, 17 Jul. 1961, Woytkowski 6657 (MO). lanceolate, bases and apices attenuate, 2-7 cm
MADRE DE DIOS: Manu, Atalaya, Hacienda
wide, margins cartilaginous, repand or slightly
Amazonía, 2-3 km W of village, 12 Dec. 1983, Foster &
Waechter 7453 (F). PUNO: San Gavan, Lechler 2374 (B).
undulate; costa prominent, primary veins
BOLIVIA. LA PAZ: Nor Yungas, Polo-Polo, Coroico, prominulous or prominent, stramineous on both
Oct-Nov. 1912, Buchtien 3384 (MO); Larecaja, 6-10 km sides of the lamina, 60-65o divergent from the
E of Consata, along new road, 15 Dec. 1981, Sperling et costa, secondary veins, slightly prominulous,
al. 5451 (MO); Larecaja, 6 km N below Consata, 15 sometimes stramineous, forming (4-) 5-7 primary
Dec. 1981, Solomon 6579 (MO).
areoles between the costa and margin, entire or
symmetrically divided, 1-2 free excurrent
Campyloneurum nitidissimum var. latior has
veinlets on each secondary areole; sori medial or
been often misidentified in herbaria as C.
subterminal, paraphyses dendritic, shorter than
coarctatum (Kunze) Fée, but it has a much wider
the sporangia, 11-16 µm long, spores 64 µm long,
stem, with closely spaced phyllopodia, and
40 µm wide. Chromosome number: n=37 (as C.
leaves more than 40 cm long.
phyllitidis by Smith & Foster, 1984). Figs. 17 g;
40.
32. Campyloneurum nitidum (Kaulf.) C. Presl,
This species is known from the southern part
Tent. Pterid. 190. 1836.
of South America, from southern Bolivia to
Polypodium nitidum Kaulf., Enum. Fil. 92. 1824.
Argentina, and southern Brazil, where it grows
Type. Brazil: Chamisso s.n. (B!).
as a terrestrial between 500 m and 1100 m
Campyloneurum leuconeuron Fée, Crypt. Vasc.
elevation.
Brésil 113. 1869. Type. Brazil: Glaziou 1001
(holotype, P!; isotype RB!). REPRESENTATIVE SPECIMENS: BOLIVIA.
Polypodium phyllitidis f. minus (minor) Hieron., COCHABAMBA: Cavernas del Repechón, Parque
Bot. Jahrb. Syst. 22: 405. 1896. Type. Nacional Carrasco, 9 Oct 1996, Kessler 8336 (LPB).
Uruguay: Misiones, Paggi at Río Alto BRAZIL. RIO GRANDE DO SUL: Pelotas, 12 May
Uruguay, Aug. 1887, Niederlein 1947 1959, Brauner 74 (F); Pelotas, 22 May 1959, Costa-Saco
(holotype, B!). 1248 (F); Rio Grande do Sul, Jul. 1940, Eugenio &
Polypodium phyllitidis f. majus (major ) Hieron. ex Leopoldo 1499 (F); São Leopoldo, 10 Aug. 1930, Flach
936 (BM, MO); Porto Alegre, 15 Oct. 1982, Lindman 499
Hicken, Rev. Mus. La Plata 15: 272. 1908.
(S); 1865, Page s.n. (F); 1897, Reineck & Czermack 35 (F);
Cyted Syntypes. Argentina: Misiones, Ruins Esmeralda, Estação Ecológica de Aracuri, 23 Aug.
at Candelaria, 20 Feb. 1883, Niederlein s.n. 1981, Waechter 1843 (F); Caxias do Sul, Conceição, 24
(B!); El Primer Misionero, von Hernandez, Oct. 1987, Wasum et al. 3422 (F). RIO DE JANEIRO:
Puck and Fernández, Niederlein 237 (B!). Macaé, Serra do Frade, perto do Rio São João, 18 Oct.
Lectotype (chosen here) Argentina: Misiones, 1970, Carauta 1217 (F); Serra dos Orgãos, entre a
Arroyo Ñacanguazú, 12 Feb. 1883, Niederlein Barragem e o Abrigo no. 1, 28 Mar. 1971, Carauta 1331
(B!) (F); Parque Nacional Serra dos Orgãos, 6 Aug. 1961,
Pabst 5662 (B, F); Serra dos Orgãos, 9 Aug. 1964, Pabst
Campyloneurum majus (major) (Hieron. ex
8139 (F); Mun. Petrópolis, road from Araras to Vale de
Hicken) Lellinger, Amer. Fern J. 78: 26. 1988.
Videiras, 22 Apr. 1980, Plowman & Martinelli 10168 (F);
Stem short-creeping, black or stramineous, Serra dos Orgãos, Teresópolis, 31 Jan. 1983, Simonis &
not pruinose, 3-4 mm wide. Stem scales brown Martinelli 25 (UC); SE side of Itatiáia, Riberão Campo
in mass, adpresed or subadpressed, 1.8-2 mm Belo, 31 Oct. 1965, Tryon & Tryon 6619 (F). SÃO
long, 1-2 mm wide, broadly ovate, bases PAULO: km 78 of road 116, between Curitiba and São
auriculate, apices obtuse, slightly clathrate, cells Paulo, 5 Jan. 1974, Conrad & Dietrich 1980 (MO);
broadly oblong, cell walls 4-8 µm thick. Ipanema, 29 Sep. 1925, Freine & Azevedo 153 (UC);
Phyllopodia 2-3.5 mm long, 1.5-4 mm wide, 2-6 Campos do Jordão, Jun. 1947, Leite 3387 (MO), Leite
3492 (MO); Estação Biológica Alto da Serra, 14 Feb.
mm apart. Leaves 25-75 cm long; petiole
1929, L.B. Smith 1891 (GH, US). PARANA: banks of
stramineous or dark stramineous, 1.7-8 (-9.5) cm the Rio do Santa, between Curitiba and Joinville, 5 Jan.
1974, Conrad & Dietrich 2052 (MO); Jaguariaíva, 26 Jun. belongs to the C. phyllitidis group. This species
1910, Dusén 10057 (BM, S, US); Jacareí, 18 Jul. 1914, has been confused with C. phyllitidis, but it is
Dusén 15311 (F); Jaguariahyba, 28 Feb. 1915, Dusén smaller, with a narrow stem and with an
17354 (F, UC); Mun. Lapa, Rio Passa Dois, 30 Sep.
attenuate leaf apex. In Campyloneurum nitidum
1969, Hatschbach 22255 (UC); Mun. Balsa Nova, Serra
Santa Ana, 1 Nov. 1969, Hatschbach 22780 (UC); Mun.
there are two forms, one has very narrow leaves
Ortiguera, Rio do Barreiro, 7 Aug. 1970, Hatschbach with the divergent angle of the primary veins
24526 (UC); Mun. Catanduvas, 10 Oct. 1974, less than 55o and they belong to the
Hatschbach & Pelanda 35134 (MO); Jacareí, 20 Aug. "leuconeuron" leaf type, and the second has
1914, Jönsson 97a (F); Mun. Tijucas do Sul, Campina, 46 broader leaves with the divergent angle of the
km S de Curitiba, 14 Feb. 1978, Krapovickas & Cristóbal
33640 (MO); Mun. Quedas do Iguaçú, 20 Aug. 1974, primary veins more than 60o and it corresponds
Kummrow & Golte 603 (UC); Mun. Lapa, São Carlos, 13 to the "nitidum" type; however they are not
Aug. 1982, Oliveira 625 (UC); Mun. Quatro Barras, Rio recognized as different species because of the
Taquarí, Damata, 25 Aug. 1982, Oliveira 657 (UC); presence of many intermediate specimens.
Mun. São José dos Pinhais, 4 Sep. 1986, Silva & Silva
182 (UC). SANTA CATARINA: Oct. 1904, Haerchen
s.n. (Rosenst. 115, S); Frinvihe, 1 Jul. 1901, Schmalz 8 (F,
33. Campyloneurum oellgaardii B. León, sp. nov.
S); Joinville, 17 Jul. 1901, Schmalz 49 (F, MO); Joinville,
ined. Type. Ecuador, Carchi, Cerro
30 Jul. 1901, Schmalz 76 (F, MO).
ARGENTINA. CORRIENTES: Santo Tomé, estancia Golondrinas, 0o52'N, 78o07'W, 21 Dec. 1987,
Garruchos, 8 Feb. 1972, Krapovickas et al. 21365 (QCA). Hoover 2211 (holotype MO, isotype QCA).
MISIONES: Dep. Libertador General San Martin, Species cum habitu Campyloneurum inflatum,
Gruta 3 de Mayo, 12 Jan. 1970, Krapovickas & Cristóbal a qua differt rhizomate longe repente, 5 mm
15635 (MO, UC); Dep. San Pedro, 20 Jul. 1957, Montes crasso, atrofusco, dense paleaceo, squamis
27453 (UC); San Pedro, 80 km E of El Dorado, 22 Jan.
adpressis, brunneolis, foliis amplissimis.
1973, Schinini & Fernandez 5971 (F).
URUGUAY. TACAUREMBO: Gruta Helechos, 28 Stem long-creeping, black, not pruinose, 5
Sep. 1928, Herter 1231 (MO, S, UC); Tacuarembo, 24 mm wide. Stem scales brown in mass, broadly
Aug. 1907, Herter 3538 (B), Herter 3739 (B); Gruta de ovate, adpressed, 3-4 mm long, 2-2.5 mm wide,
los Cuervos, Herter 10225 (B). TREINTA Y TRES: bases auriculate, apices acuminate, margins
Quebrada de los Cuervos, Serranías de Yerbal, Apr. dentate, scales slightly clathrate, the cells oblong
1936, Legrand 716 (F). or broadly oblong, cell walls 6-9 µm wide, walls
PARAGUAY. AMAMABAY: Cerro Corá, 3 Sep. 1978, of central cells brown dark, walls of marginal
Herbst s.n. (F). ALTO PARANA: 2.8 km W de Puerto
cells yellowish or brownish, cell lumina
Stroessner, Krapovickas 13403 (MO). ITAPUA: Hotel El
Tirol, 19.5 km by road NNE Encarnación, 8 Sep. 1978,
transparent. Phyllopodia 10-20 mm apart.
M. Foster 78-(2)-17 (UC), 15 Oct. 1979, Foster 79-58 Leaves erect, 100 cm long, petiole 30-40 cm long,
(MO). MISIONES: Santiago, Estancia La Soledad, 13 dark stramineous; lamina elliptic, 21 cm wide,
Dec. 1969, Pedersen-Myndel 9548 (UC). CAAGAZU: herbaceous-chartaceous, base cuneate, apex
Guayaquí, 22 May 1987, Zardini et al. 2458 (MO). acuminate, margins cartilaginous, sinuate, leaves
ALTO PARAGUAY: San Pedro, 13 Sep.1959, Woolston puberulous, indument of inconspicuous,
1118 (UC). GUAIRA: Cordillera Ybytyruzú, cerro bicellular glandular hairs, scattered abaxially;
Peró, 23 Jul. 1989, Zardini & Velasquez 13903 (MO).
stomata polocytic rarely copolocytic; costa
CAAZAPA: Tavai, 29 Oct. 1988, Basualdo 1688 (MO).
prominent, slightly angular abaxially; primary
Without locality. Cordillera de Altos, 7 Aug. 1902,
Fiebrig 13 (B, F); Río Apa, und Río Aquidaban, 1908- veins prominent, 75o divergent from the costa,
1909, Fiebrig 5079 (BM); Tobaty, Sep. 1903, Hassler 6283 straight, lighter in color than the adjacente
(BM, S, UC); Caaguazuensis, 1905, Hassler 9077 (BM, S, tissue, 7-9 mm apart, secondary veins slightly
UC); Sapucay, Aug. 1913, Hassler 12241 (BM, GH, MO, prominulous on both sides of the lamina,
S, UC); 1931, Jorgensen 4602 (MO, S, US). transverse secondary veins forming 19 primary
areoles between the costa and margin, primary
Campyloneurum nitidum is characterized by areoles undivided, with 2-3 excurrent veinlets,
narrow lanceolate leaves, attenuate at both ends, sometimes with one recurrent veinlet in the
and by stem scales with obtuse apices. It areoles close to the margin, veinlets entire, free.
Sori subapical on the excurrent veinlet;

paraphyses and spores not seen. Fig. 50. This species is distributed from Colombia
and Venezuela to Bolivia, where it grows as an
This species is found in northern Ecuador, at epiphyte mostly between 1500 m and 2500 m
1200 m. It is known only from the type elevation.
collection.
REPRESENTATIVE SPECIMENS: COLOMBIA. SUR
Campyloneurum oellgaardii is closely related to DE SANTANDER: along highway between Pamplona
C. inflatum; they can be distinguished because and Bucaramanga, Mun. Tona, Corregimiento
Corcova, Vereda La Mariana, 5 May 1983, Croat
the former has a 5 mm wide stem; adpressed,
56507a (MO). VALLE: San Antonio, 18 May 1939,
broad lanceolate stem scales with Alston 8609 (MO). CALDAS: Cordillera Occidental,
undifferentiated margins; and elliptic lanceolate Pueblo Rico, 21 Dec. 1945, Sneidern 5258 (F), 11 Jan.
leaves reaching 1.5 m long. The latter species 1946, Sneidern 5437 (F, S). VALLE: Cordillera
has a 2-3 mm wide stem; spreading, lanceolate Occidental, al sur de las Brisas, 27 Oct. 1946,
stem scales with differentiated margins; and Cuatrecasas 22663 (F). CAUCA: Munchique, 21 Apr.
elliptic leaves less than 50 cm long. 1939, Alston 8175 (MO); Carpinterías, between cerros
This species is named in honor of B. Munchique and Altamira, 15 Jul. 1939, Cuatrecasas &
Øllgaard, who with his work on the Ecuadorean Perez-Arbelaez 6136 (F) . Cerro Gualcala, above Piedra
Ancha, Lehmann 5012 (F).
plants, especially with the pteridophytes, is
VENEZUELA. TACHIRA: along Quebrada Agua
contributing to the knowledge of that rich flora. Azul, S of El Reposo, 14 km SE of Delicias, 22-23 Jul.
1979, Steyermark & Liesner 118424 (MO). Los Venados,
Caracas, Aug. 1939, Elias 21 (F).
34. Campyloneurum ophiocaulon (Klotzsch) Fée, ECUADOR. CARCHI: Chical, 16 Nov. 1983, Barfod et
Gen. Fil. 258. 1852. al. 48644 (QCA); Valle de Maldonado, km 60 on road
Polypodium ophiocaulon Klotzsch, Linnaea 20: Tulcán-Maldonado, 18 May 1973, Holm-Nielsen et al.
401. 1847. Type. Peru: Junín, Tarma, Dombey 5775 (F, MO). NAPO: 3.5 km NW of Borja, 20 Sep.
1980, Holm-Nielsen et al. 26325 (QCA); Francisco de
41 (holotype, B!; photo of B, BM!).
Orellana, near Cañón de los Monos, 1987, Zak &
Stem dark stramineous, black, not pruinose, Jaramillo 3604 (F, MO), Zak & Jaramillo 3607 (F, MO);
(2-) 2.5-4 mm wide. Stem scales light brown in Baeza path, 1 km SW of the village,20 Oct. 1976,
mass, 3-4 mm long, 2 mm wide, ovate, bases Øllgaard & Balslev 10238 (AAU, USM); road Baeza-
peltate, apices obtuse, scales clathrate, the cells Tena, in the pass S of Río Salado, 8 Aug. 1980, Øllgaard
oblong, basal and marginal cells irregular et al. 35766 (AAU, QCA), Øllgaard et al. 35786 (AAU,
arranged, central cells along main axes of the QCA); Cerro Huacamayos, on road Baeza-Tena, 9-10
scale. Phyllopodia 1-2 mm long, 2-3 mm wide, Aug. 1980, Øllgaard et al. 35921 (F, QCA, UC).
PICHINCHA: Nono-Nanegalito, N of Cerro
10-20 mm apart. Leaves 30-50 cm long; petiole
Pichincha, 9 May 1982, Balslev & Boom 2500 (QCA)
dark stramineous, 2.5-4.5 cm long; lamina broad
along road Nono and Nanegal, NW of Quito, 11-12
lanceolate, obovate or narrow lanceolate, bases km NW of Nono, 4 Sep. 1976, Croat 38818 (MO); km 17
attenuate, apices acuminate, 3.5-7 cm wide, Nono-Tandapaya, road along Río Alambi, 14 May
chartaceous, margins cartilaginous, undulate, 1981, Dodson et al. 10793 (F, MO); Estación Científica
indument of scattered, inconspicuous, bicellular Guajalito, along road Chiriboga-El Tránsito, 10 Jun.
hairs; stomata not seen; costa prominent, 1990, Øllgaard 90410 (QCA). TUNGURAHUA:
Colonia Mexico, 4 km from Río Topo, 5 Mar. 1969,
primary veins 65-80o divergent from the costa,
Lugo 647 (F). PASTAZA: ca. 5 km E of Mera, 30 Ju;.
stramineous, prominulous, straight, 6-7 mm 1980, Øllgaard et al. 35581 (AAU, QCA). SANTIAGO-
apart, secondary veins slightly prominulous, ZAMORA: between La Esperanza and Santa Ana, 15
inconspicuous, transverse veins forming 8-11 Feb. 1944, Acosta-Solís 7436 (F); W side of Río
primary areoles between the costa and margin, Valladolid, 15 Oct. 1943, Steyermark 54722a (F).
primary areoles undivided, 2 free excurrent Without locality: Mille s.n. (MO).
veinlets in each areole; sori medial, paraphyses PERU. CAJAMARCA: Cutervo, San Andrés de
not seen, spores 50 µm long, 35 µm wide. Fig. Cutervo, above Saucedal, Chorro Blanco, 3 Aug. 1988,
Diaz & Osores 2963 (MO). AMAZONAS: Bagua, ca. 20
43.
km E of la Peca, 22 Jul. 1978, Barbour 2812 (AAU, F, 1 Nov. 1979, Foster 79-145 (MO). Without locality:
MO, UC); Bagua, Cordillera Colán, SE of La Peca, 17 Bang 2395 (BM, F); Río Tocorani, Jul. 1911, Herzog 2310
Oct. 1978, Barbour 4188 (F, MO). SAN MARTIN: (B, S, UC).
Venceremos, near Amazonas border, km 291 on Rioja-
Pomacocha road, 12 Feb. 1984, Gentry et al. 45488 Campyloneurum ophiocaulon is closely related
(MO); Rioja, Pedro Ruiz-Moyobamba road, km 390 to C. repens, but it differs by the ovate lanceolate
Venceremos, 29-31 Jul. 1983, D.N. Smith 4498 (F, MO,
scales with obtuse apices. The
NY, UC). HUANUCO: Pampayacu, 28 Jan. 1927,
Kanehira 119 (GH, US); Cushi, 19-23 Jun. 1923,
studied specimens of Campyloneurum ophiocaulon
Macbride 4841 (F); La Divisoria, ca. 25 km NE of Tingo are mostly distributed above 1500 m elevation.
María, 4 Jul. 1984, Moran & Fernandez 3693 (MO, UC); However, two specimens (Zak & Jaramillo 3604
Leoncio Prado, dist. Hermilio Valdizán, La Divisoria, and Zak & Jaramillo 3607) came from 250 m
from Pumahuasi to La Cumbre, 26 Jun. 1978, Plowman elevation, a very unusual distribution for the
& Schunke 7414 (F); Leoncio Pardo, km 35 on road species.
between Tingo María and Pucallpa, 3 Jun. 1981,
Sullivan & Young 1147 (F, MO). PASCO: Oxapampa,
Ulcumanu, SW of Oxapampa, road to María Teresa
35. Campyloneurum oxypholis (Maxon) Ching,
and Llaupi, 31 Dec. 1983, Foster et al. 7682 (MO);
Oxapampa, 31 Jan. 1983, León 495 (F, USM); Sunyatsenia 5: 263. 1940.
Oxapampa, 5 km SE of Oxapampa, D.N. Smith 2914 (F, Polypodium oxypholis Maxon, J. Wash. Acad. Sci.
MO). JUNIN: Villa Amoretti, 1960, Kanehira 524 (GH); 14: 140. 1924. Type. Haiti: Morne de Brouet,
in the area of Pichita Caluga, 1-3 May 1957, Walden 26 near Furcy, 13 Jun. 1920, Leonard 4782
(BM). UCAYALI: La Divisoria, ca. 25 km NE of Tingo (holotype US!, photo of US, BM!, F!; isotypes
María, Moran & Fernández 3693 (MO). CUSCO: valle B!, BM!, C!).
de Accobamba, Aug. 1922, Bues 866 (US); 28 May Stem creeping, 2-2.5 mm wide. Stem scales
1915, Cook & Gilbert 951 (US); Quillabamba, Abra de
brown in mass, spreading, 5-6 mm long,
Málaga, 8-9 Mar. 1971, Ellenberg 4739 (LPB); ca. 20 km
of Pampa Hermosa, 25 Jul. 1978, Ellenberg 9122 (LPB);
narrowly ovate, bases auriculate, apices
valle de San Miguel,near bridge Media Naranja, 20 acuminate, clathrate, the cells oblong, with
Jul. 1928, Herrera 2050 (BM, C, US); Machu Picchu, yellowish lumina, cell walls 14-24 µm thick.
Oct. 1931, Herrera 3298 (US); Paucartambo, trail below Phyllopodia 1.5 mm high, 5-7 mm apart. Leaves
Buenos Aires, km 136 road Acjanaco-Pilcopata, 16 Feb. 30-40 cm long; petiole stramineous, 2-5 cm long;
1990, León 2183 (F, USM), 17 Feb. 1990, León 2195 (F, lamina lanceolate, bases and apices attenuate, 2-
USM); Urubamba, above the first waterfall of the Río 3 cm wide, herbaceous-chartaceous, margins
Mandor, 2.5 km from Machu Picchu, 5 Jun. 1982,
sinuate, indument and stomata not seen; costa
Peyton & Peyton 452 (MO); La Convención,
prominent, primary veins slightly prominulous
Huayopata, 3 km from village of Incatambo, 31 Jul.
1982, Peyton & Peyton 844 (GH, MO); Urubamba, on both surfaces of the lamina, 45-50o divergent
Machu Picchu, hillside called Puncuyoj, 10 km SW of from the costa, secondary veins immersed,
Incatambo, 3 Oct. 1982, Peyton & Peyton 1365 (GH, darker than the leaf tissue, forming 4-5 primary
MO); La Convención, Amaibamba, 23 Nov. 1950, areoles between the costa and margin, primary
Vargas 9804 (UC); La Convención, Potrero, Garavito,
areoles with 1-3 included veinlets, undivided or
13-14 May 1960, Vargas 13182 (GH).
symmetrically divided; sori medial, paraphyses
BOLIVIA. LA PAZ: Sud Yungas, Huancané, 8 Mar.
1980, Beck 3065 (F, LPB), 9 Mar. 1980, Beck 3114 (F, not seen, spores 85-90 µm long, 50 µm wide.
LPB); Nor Yungas, Coroico-Yolosa, 1 Apr. 1982, Beck Fig. 43.
7537 (F); Nor Yungas, Polo Polo, Coroico, 1912,
Buchtin 3534 (F); Sud Yungas, La Paz-Chulumani road, This species is found in Haiti.
12 km E of Chuspipata, 1 Aug. 1989, Fay & Fay 2473
(LPB); Nor Yungas, 14.4 km NE Chuspipata, 21 Oct. EXAMINED SPECIMENS: HAITI. Morne des
1982, Solomon 8627 (MO, UC); Murillo, 30.5 km N of Commissaires, Massif de la Selle, 4 Sep. 1926, Ekman
dam at Lago Zongo, 16-17 Dec. 1982, Solomon 9103 6884 (B, S, US).
(MO); Carrasco, Serranía de Bella Vista, 31 Oct. 1984,
Solomon & Nee 12620 (MO); Nor Yungas, 12.8 km NE Campyloneurum oxypholis is a rare species,
de Chuspipata, 11 Nov. 1987, Solomon 17362 (MO,
closely related to C. vulpinum. It belongs to the
NY). COCHABAMBA: Chapare, road to San Onofre,
Campyloneurum vulpinum group. Plowman et al. 3941 (S); Río Panteor, SW of Borja, 22
Sep. 1980, Holm-Nielsen et al. 26769 (AAU, QCA); 1 km
SW of Baeza,20 Oct. 1976, Øllgaard & Balslev 10239
(AAU); road Baeza-Lago Agrio, ca. 114 km from Lago
36. Campyloneurum pascoense R. Tryon & A.
Agrio, 8 Aug. 1980, Øllgaard et al. 35778 (AAU, QCA),
Tryon, Rhodora 84: 125. 1982. Type. Peru: Øllgaard et al. 35788 (AAU, QCA). PICHINCHA: road
Pasco (as Junín), Oxapampa, Soukup 2340 Quito-Chiriboga-Empalme, km 50, sector Zapadores,
(holotype GH!). 21 Jul. 1987, Zak & Jaramillo 2199 (F). ZAMORA-
Stem short-creeping, black or dark CHINCHIPE: above Valladolid,,on road to Yangana, 1
stramineous, not pruinose, 10-30 mm wide. Stem feb. 1985, Harling & Andersson 21397 (QCA). Without
scales brown in mass, 8-10 mm long, 4 mm wide, locality: Baños, Rio de Machai, Stübel 863 (B).
broadly ovate to ovate, pseudopeltate, bases PERU. CAJAMARCA: Chota, Huambos, 10 Sep. 1956,
auriculate, apices acuminate, the cells oblong, Soukup 4488 (US). AMAZONAS: Bagua, Colán, SE of
La Peca, 17 Oct. 1978, Barbour 4187 (F). SAN MARTIN:
central cell walls 8-16 µm thick, marginal cell
Mariscal Cáceres, Parque Nacional Río Abiseo, Río
walls 4-5 µm thick. Phyllopodia 6-10 mm long, 8- Montecristo, Gran Pajatén ruins, 13 Aug. 1986, Young
10 mm wide, 20 or more mm apart. Leaves 1.50-2 4323 (NY). HUANUCO: Huacahi, near Muña, 20 May-
m long; petiole brownish, 8-14 (-75) cm long, 1 Jun. 1923, Macbride 4082 (F, US); Huánuco, km 468 on
ranurate adaxially; lamina lanceolate, bases Lima-Tingo María road, above San Miguel Chinchao, 2
attenuate or narrow cuneate, apices long Jun. 1981, Young & Sullivan 619 (MO, NY, UC). PASCO:
acuminate, 10-20 cm wide, chartaceous, margins Oxapampa, Cordillera Chanachaga, road over
cartilaginous, sinuate, indument not seen; shoulder of Cerro Pajonal, 9 Oct. 1982, Foster & Smith
9071 (F, MO); Oxapampa, camino a Quebrada San
hypostomatic, stomata polocytic; costa
Alberto, 12 Aug. 1985, León 636 (F, GH, USM); 5 km SE
prominent, sometimes with scattered scales of Oxapampa, 24 Dec. 1983, D.N. Smith 5361 (F, MO,
abaxially, similar to those on the stem, primary UC). AYACUCHO: San Miguel, Urubamba valley, 10
veins prominent, 70-75o divergent from the costa, Jul. 1915, Cook & Gilbert 1756 (US). CUSCO: Machu
stramineous, straight, (6-) 9-11 mm apart, Picchu station, 5 Nov. 1957, Hutchison 1760a (UC);
secondary veins prominulous, stramineous, Urubamba, Jan. 1936, Soukup 173 (F).
BOLIVIA. LA PAZ: Coroico, 1912, Buchtien 3526 (F);
forming 12-20 primary areoles between costa and
Murillo, Zongo valley, 0.6 km up valley from Sainani,
margin, costal areole with 1 free excurrent 5-6 Aug. 1990, Fay & Fay 2907 (MO); Murillo, 27.4 km
veinlet, simple or furcate; non-costal primary below N dam at Lago Zongo, 16 Mar. 1984, Solomon et
areole assymmetrically divided with 3-7 al. 11919 (MO, USM).
excurrent veinlets, 1-2 veinlets forming 2-4
assymmetrical secondary areoles, secondary Campyloneurum pascoense is characterized by
areoles with 1-3 simple or furcate veinlets, these large leaves, more than 1 m long, and also by its
veinlets forming tertiary areoles; sori subterminal prominulous secondary veins and the presence of
or compital, paraphyses simple, unbranched, secondary and tertiary areoles.
spores (50-) 60-65 (-70) µm long, (30-) 35-40 µm Campyloneurum pascoense is closely related to
wide. Figs. 14 k; 18 b; 19 b; 44. C. brevifolium and C. tucumanense. It can be
differentiated from C. brevifolium by the
This species is distributed from Colombia and prominence of its tertiary veins and the relatively
Ecuador to Bolivia, where it grows between 1500 large size of its leaves. It differs from C.
m and 2500 m elvation, usually as a terrestrial in tucumanense, by having a chartaceous leaf texture.
disturbed forests.
REPRESENTATIVE SPECIMENS: COLOMBIA. 37. Campyloneurum phyllitidis (L.) C. Presl, Tent.

SANTANDER: Jan. 1878, Kalbreyer 451 (B). Pterid. 190. 1836.
ECUADOR. GUAYAS-CAÑAR-CHIMBORAZO-
Polypodium phyllitidis L. Sp. Pl. 1083. 1753.
BOLIVAR: near village of Bucay, 8-15 Jun. 1945, Camp
E-3688 (F, UC, US). NAPO: road Baeza-Tena, 8 km Lectotype (chosen by Proctor, in R. A.
from Baeza, 28 Oct. 1976, Balslev & Madsen 10420 Howard 2: 341. 1977): Plumier t.38, Descr. Pl.
(AAU); Cantón Quijos, 28 mi E of Baeza, 29 Jul. 1974, Amer. 1693.
Polypodium comosum L., Sp. Pl. 1084. 1753. Type. subterminal or terminal, paraphyses not seen,
Plumier t.131, Descr. Pl. Amer. 1693. spores (57-) 65-70 µm long, 40-50 µm wide.
Polypodium conjugatum Poiret in Lam., Encycl. 5: Chromosome number 2n=74. Figs. 18 c, d; 20 b;
516. 1804. Type. Herb. Jussieu 1071 (holotype, 45.
P; photo of P, BM!, C!).
Cyrtophlebium phyllitidis (L.) J. Sm., J. Bot. This species is distributed from southern
(Hooker) 4: 58. 1841. United States (Florida), Central America, the
Polypodium phyllitidis var. linneanum Hook., Sp. Caribbean Islands, Colombia to Bolivia and
Fil. 5: 38. 1864. Type. Plumier t. 130, 131, central Brazil, where it grows as a terrestrial or
Descr. Pl. Amer. 1693. epiphyte between 100m and 2500 m elevation.
Polypodium phyllitidis var. swartziana Griseb., Fl. Common names: "ba sú tape" (cayapa).
Br. West Ind. 702. 1864. Type. Plumier t. 130,
131, Descr. Pl. Amer. 1693. REPRESENTATIVE SPECIMENS: U.S.A. FLORIDA:
Polypodium phyllitidis var. elongata Hieron., Bot. Stuart and vicinity, Arch Creek, near Natural Bridge,
Jahrb. Syst. 34: 534. 1904. Cited Syntype. 20 Feb. 1917, Atwood s.n. (MU); Palm Beach, 10 Apr.
1897, Curtiss 867 (S); Indian River Narrows, Curtiss
Ecuador: Azuay, Cordillera Oriental de
3668 (BM, NY); Collier County, Big Cypress, vic. of
Cuenca, Cerro Yanghuan, near Pindilic, Falkahatchee, 7 Jun. 1966, Lakela & Almeda 29969 (US);
Lehmann 7679 (B!, F!, K!, US!). Lectotype Redlands district, Hattie Bauer Hammock, 2 Feb. 1930,
(chosen here) Colombia: Tolima, Río Paez, Moldenke 569 (NY, S); Aiken, Key Largo, 30-31 Mar.
Lehmann 5721 (B!, F!, K!, US!). 1898, Pollard et al. 204 (BM, MU, NY); Dade County,
Stem black, green turning black, not pruinose, Nixon Lewis Hammock, 16 Mar. 1915, Small & Mosier
(4-) 6-15 mm wide. Stem scales brown in mass, 5885 (GH, MO, NY, S); Merritt's Isalnd hammock,
(4-) 6-8 mm long, (1.5-) 2-3 mm wide, ovate or Little River Prairie Miami, 8 Jul. 1915, Small et al. 6936
trinagular-ovate, bases auriculate or shortly (MU); 7 May 1904, Tracy 9139 (BM, F, US).
MEXICO. SAN LUIS POTOSI: near banks of
auriculate, apices acuminate, the cells oblong,
Moctezuma, 25 May 1939, Frye & Frye 2657 (UC);
central cells along main axes of the scale, mountains along road to Jalpan, 2 mi W of Xilitla, 25
marginal cells irregularly arranged, central cell Mar. 1961, King 4284 (UC). MICHOACAN: Dist.
walls 15 µm wide, marginal cell walls 8-10 µm Coalcoman, Huizontla, 9 Aug. 1941, Hinton 15969 (F,
thick. Phyllopodia 1-3 mm long, 1.5-4 mm wide, UC); Aquila and Coahuayana, Jan. 1942, Hinton 16269
2-5 (-7) mm apart. Leaves (25-) 60-150 cm long; (UC). HIDALGO: dist. Huejutla, woods Tehuetlan, 30
petiole stramineous or brownish, 0.7-4 (-6) cm May 1947, Moore 3038 (UC). VERACRUZ: E of Playa
long, ranurate adaxially, convex abaxially; lamina Vicente, 21 Sep. 1973, Mickel 7227 (UC). OAXACA:
Santa María Chimalapa, al N de Santa María, 2 Aug.
obovate or lanceolate, bases attenuate, apices
1984, Hernández 284 (F); Santa María Chimalapa, Piedra
acuminate or subcaudate, (2.7-) 5-16 cm wide, Blanca (Popotzá), E of Santa María, 15 Dec. 1984,
chartaceous or subcoriacous, margins Hernández 728 (MO); Chiltepec, Sierra Juarez, Tuxtepec,
cartilaginous, slightly sinuate, plane or slightly 30 May 1966, Martinez 873 (F). TABASCO: Mun.
revolute, indument of abaxially scattered, simple, Tacotalpa, NW de Tapijulapa, 30 May 1982, Cowan et
bicellular hairs; hypostomatic, stomata polocytic; al. 3540 (UC); San Isidro, Balancan, 7-11 Jun. 1939,
costa prominent, sometimes with scales similar to Matuda 3368 (F). CHIAPAS: Esperanza, Escuintla, 23
those on the stem, primary veins prominent or Feb. 1948, Matuda 17632 (F). YUCATAN: 1895, Armour
38 (F); Yuxpeña, Campeche, 30 Jan. 1932, Lundell 1268
prominulous on both sides of the lamina,
(F, US); Yucatan, Schott 781 (F).
stramineous or dark stramineous, straight,
BELIZE: Yucatan Peninsule, Maskall, Dec. 1933, Gentle
(55o-) 65-75o divergent from the costa, (5-) 6-10 1108 (F, GH, S).
mm apart, secondary veins slightly prominulous, HONDURAS. FRANCISCO MORAZAN: Quebrada
same color as the leaf tissue, transversal veins Hierba Buena, 15 km NE of Tegucigalpa, 24 Sep. 1983,
forming (6-) 8-16 primary areoles between the Calderón 55 (UC). EL PARAISO: drainage of the Río
Yeguare, Quebrada Dantas, 10 km N of Yuscarán, 12
costa and margin, 3 (-4) free excurrent veinlets in
Mar. 1950, L.O. Williams 17220 (F). ATLANTIDA:
each non costal primary areoles, usually one
about 15 mi E of Ceiba, 21 Jul. 1938, Yuncker et al. 8575
veinlet connected with the transversal vein (GH). San Pedro Sula, 30 May 1888, Thieme 6 (UC).
forming isodiametric secondary areoles; sori
GUATEMALA. PETEN: Puerto Chimino, Laguna valley, 6 km SW of Jéremie, 6 May 1941, Bartlett 17290
Petexbatún, 20 km S of Sayaxchá, Oct-Dec. 1989, Zomer (GH, UC).
95 (F), 26 Apr. 1990. Zomer 210 (F). SUCHITEPEQUEZ: DOMINICAN REPUBLIC. MACORIS: Consuelo, along
near Santo Domingo, S of Mazatenango, 5 Mar. 1941, the Macoris river, 22-24 Nov. 1909, Taylor 262 (F).
Standley 88877 (F). Without locality: Finca Panama, 26 SANTO DOMINGO: Llano costero, banks of Río
Mar. 1947, Brenckle 47-188 (F). Ozama, 30 Apr. 1929, Ekman 12342 (F, S, US); Santo
COSTA RICA. ALAJUELA: W of San Ramón, ca. 1 km Domingo, 24 Jan. 1899, Millspaugh 813 (F).
S of Socorro, 25 Jul. 1970, Lellinger & White 1317 (F); Río PUERTO RICO. NW of Utuado, 18 Sep. 1941, Blomquist
San Rafael, Cantón Aguas Zarcas, 8 Feb. 1965, L.O. 11805 (F); Vega Alta, 15 Oct. 1941, Blomquist 11991
Williams et al. 29103 (F). PUNTARENAS: about 4 mi W (UC); near The Caves, Aguas Buenas, 12 Oct. 1941,
of Rincón de Osa, 4-7 Jun. 1968, Burger & Stolze 5404 (F, Blomquist 12015 (UC); vic. of Catano, 1 Apr. 1922,
US), Burger & Stolze 5605 (F). LIMON: between Britton et al. 6990 (F, US); near Mayaguez, 30 Jan. 1900,
Siquerres and the Río Pacuare, 20-22 Dec. 1969, Burger Heller 4438 (F); on the Adjuntas road, 7 mi from Ponce,
& Liesner 6904 (F, GH, NY). SAN JOSE: basin of El 2 Dec. 1902, Heller 6177 (F, US); Río Abajo forest, 2 Apr.
General, Jul.-Ago. 1943, Skutch & Barrantes 5161 (UC); 1985, Luteyn & Lebrón-Luteyn 11486 (UC); Toro Negro
along Río Convento, El General valley, 31 Jan. 1965, recreation area, 3 Jan. 1978, Tullis s.n. (MU).
L.O. Williams et al. 28722 (F). CARTAGO: 22 km N of LEEWARD ISLANDS. ANTIGUA: St. Mary,
Turrialba, 23 Jul. 1983, Givens 3230 (F). ISLA DEL downstream from Walling's dam, 12 jun. 1974, Holland
COCO: Twin mountains, Mar. 1970, Gomez 3353 (F); 8 (F). DOMINICA: bank of the Mantipo river, 21 Jul.
Mar. 1940, Valerio 1102 (F). 1938, Hodge 34 (UC). MONTSERRAT: Canan river, 7
PANAMA. VERAGUAS: Islas Contreras, Isla Feb. 1907, Shafer 451 (F, US); in the mountains, 13 Feb.
Brincaneo, 20 Jul. 1984, Churchill 5739 (MO). COLON: 1907, Shafer 756 (F). VIRGIN ISLANDS. ST. CROIX:
trail to lago Gatún, 17 May 1975, Salazar 13 (MO). mountain Eagle, 31 Jan. 1896, Ricksecker 251 (F, MU,
CANAL ZONE: Barro Colorado Island, 7 Nov. 1972, UC).
Kennedy 1905 (F); Barro Colorado Island, 9 Nov. 1931, WINDWARD ISLANDS. ST. VINCENT: St. Patrick,
Shattuck 359 (F); Parque Nacional Soberanía, 20 Mar. upper Rutland river, 15 Jan. 1962, Cooley 8155 (F, UC,
1980, Vasquez 165 (F), 4 Jun. 1980, Vasquez 232 (MO, US); .
UC). DARIEN: vic. of gold mine at Cana, 26 Jul. 1976, BAHAMAS. GRAND BAHAMA: Coppice, Baruetts
Croat 37588 (MO). Without locality: 1860, Eaton 40 (F). Point, 5-13 Feb. 1905, Britton & Millspaugh 2636 (F).
CUBA. PINAR DEL RIO: source of Río Taco-Taco, GREAT ABACO: along Great Abaco highway, ca. 22
Sierra de los Organos, 18 Nov. 1941, Morton 4339 (UC); mi SE of Marsh Harbour, 12 Mar. 1975, Correll & Meyer
vic. of Sumidero, 2-4 Aug. 1912, Shafer 13501 (F); 44541 (F); Vanilla Hummock, edge of Marsh Harbour,
Guanajay, 19 Sep. 1907, Wilson 1773 (F). LAS VILLAS: 15 Jun. 1981, Correll & Wasshausen 52030 (F). ANDROS:
Arroyo Cimarrón, 5 Mar. 1910, Britton & Britton 5095 Conch Sound, 12 May 1890, Northrop & Northrop 566 (F,
(US); Cieneguita, 6 Jul. 1895, Combs 285 (F); El Junco, GH); Corpice, near Staniard Creek, 1-3 Feb. 1910, Small
above Siguanea, in San Juan mountains, 1-20 Jul. 1950, & Carter 8869 (F). NEW PROVIDENCE: Midenhead
Howard et al. 175 (GH, UC); above San Blas, 9-10 Nov. Coppice, 12-24 Mar. 1907, Britton 6544 (F, US).
1941, Morton 4105 (UC). ORIENTE: SE summit of CROOKED ISLANDS: Salt Hope, 9-23 Jan. 1906, Brace
Yunque de Baracoa, 28 Feb. 1979, Bisse et al. 40158 4734 (F).
(HAJB); slopes and summit of El Yunque, near COLOMBIA. CHOCO: Mun. Río Sucio, near to
Baracoa, 30-31 Jan. 1902, Pollard & Palmer 184 (F, US); campamento Tilupo, Forero et al. 1688 (MO); Hoya del
vic. of Baracoa, 1-7 Feb. 1902, Pollard et al. 243 (F, US); Río San Juan, near to Palestina, 23 Mar. 1979, Forero et
La Perla, 9 Feb. 1911, Shafer 8554 (F); Monteverde, 1859, al. 3797 (MO); 26 Mar. 1979, Forero et al. 4059 (MO),
Wright 1021 (UC). San Antonio, Mar. 1906, Hitchcok s.n. Forero et al. 4066 (MO), 5 Apr. 1979, Forero et al. 4681
(F). (MO); area of Baudó, 11 Feb.-29 Mar. 1967, Fuchs &
JAMAICA. ST. ANDREW: about 2 mi NE of Kingston, Zanella 22351 (F); Chocó, Nov. 1857, Schott 10 (F).
on road to Newcastle, 15 Jun. 1963, Crosby et al. 162 (F, ANTIOQUIA: 1934, Daniel 212 (F); Río Samana, Cord.
UC). ST. ANN: hill on E side of station TL, 8 mi S of Central, 2 km above Argelia, 29 May-1 Jun. 1944, Ewan
Brown's town, 2 Aug. 1977, Goodfriend s.n. (F). Point 15773 (UC); carretera Mutatá-Pavarando, before bridge
Antonio, 28 May 1891, Metcalf s.n. (MU); Port Antonio, Río Sucio, 3 Mar. 1987, Fonnegra et al. 1684 (MO);
Dec. 1898-Mar. 1899, Millspaugh 999 (F). ST. Anori, Providencia hydroelectric station, 6 Jun. 1971,
ELIZABETH: Cooks Bottom, N of Ipswich, 31 Mar. Soejarto 2901 (F). CALDAS: Santa Cecilia, Cord.
1920, Maxon & Killip 1447 (F). Port Moraut, 1890-91, Occidental, Tatamá, 17 Apr. 1945, Sneidern 5128 (F).
Rothrock 428 (F). CUNDINAMARCA: Salto de Tequendama, 1-3 Oct.
HAITI. DU NORD: vic. of Dondon, 8 Jan. 1926, Leonard 1938, Cuatrecasas 146 (F); Río Magdalena, Quebrada de
8681 (UC); at source of Rivière Tissier, Grand'Anse los Ñeques, 7 Feb. 1962, Murillo et al. 579 (F, US).
VALLE: Mun. Zarzal, between La Paila and Zarzal, 17 river, Saut Pararé, 11 Feb. 1969, Oldeman 3016 (CAY);
Nov. 1986, Silverstone et al. 2590 (MO). Saul. L'eaux Claires, 22 Jun. 1988, Windisch 5283 (F).
VENEZUELA. TACHIRA: along road between San ECUADOR. ESMERALDAS: Río Cayapa, Zapallo
Cristóbal and Delicias, 11 km N of Delicias, 10 Aug. Grande, 1-2 Aug. 1982, Kvist & Asanza 40806 (QCA).
1982, Croat 54993 (MO). APURE: dist. Páez, selva de CARCHI: around Maldonado, W of Tulcan, 5 Sep.
Cutifí, between Catufí on the Río Cutufí and the Río 1981, Balslev 1993 (F, QCA). IMBABURA: between El
Sanare, 8-12 Nov. 1982, Davidse & González 21760 (MO); Pajón and Cachaco, 2 Jun. 1949, Acosta-Solís 12704 (F).
25 km by car E of El Nula, 2 Jul. 1983, Werff & Gonzalez NAPO: Sucumbios, Reserva Faunística Cuyabeno, 6
4742 (MO). PORTUGUESA: 50 km WNW of Apr. 1989, Balslev 84890 (QCA); laguna Cuyabeno, 21
Guanare,15 Mar. 1982, Liesner et al. 12804 (MO); ESE of Aug. 1981, Brandbyge et al. 33840 (QCA), 22 Aug. 1981,
Paraiso de Chabasquén, 5 Nov. 1982, Smith et al. 1008 Brandbyge et al. 33930 (AAU, QCA); Parque Nacional
(MO). MIRANDA: dist. Páez, cerro Riberón, between Yasuní, Pozo Amo 2, 9-13 Jan. 1988, Cerón & Coello
Río Guapo and Río Chiquito, 1-2 Jun. 1977, Davidse & 3300 (QCA); at Río Aguarico, 20-21 Jan. 1984, Laegaard
Gonzalez 13580 (MO). BOLIVAR: 7 km E of Hato de 51544 (QCA); Río Yasuní, Garza Cocha, 8 Apr. 1983,
Nuria, E of Miamo, 14 Jan. 1961, Steyermark 88474 (F, Lawesson et al. 43344 (AAU, QCA); Lago Agrio, 3.5-4.8
GH). TERRITORIO FEDERAL AMAZONAS: 25 May km E of Río Conejo, 1 Apr. 1972, Mac Bryde & Dwyer
1975, Berry 701 (MO); Alto Orinoco, 18 Aug. 1951, 1407 (QCA); Reserva Biológica Jatun Sacha, 8 km ESE
Croizat 524 (F). TERRITORIO FEDERAL DELTA of Puerto Misahualli, 23 Jun. 1986, Miller et al. 2179
AMACURO: Pedernales, Caño Simoina, W of Isla (MO); Añangu, Parque Nacional Yasuní, 30 May-21
Cocuinma, 8 Oct. 1977, Steyermark et al. 11348 (MO). Jun. 1982, Øllgaard et al. 38908 (AAU, QCA).
ISLA MARGARITA: San Juan mountain, 27 Jul. 1903, PICHINCHA: vía Santo Domingo-Quinindé, 8 Sep.
Johnston 151 (F); El Valle, 30 Jul. 1901, Miller 165 (BM, 1949, Acosta-Solís 13850 (F); road Cotocollao-
F). Pumdupamba-Nono-Nanegalito, 6 May 1980, Jaramillo
TRINIDAD. St. Amés, 21 Apr. 1924, Broadway 5279 (F). et al. 2405 (QCA). LOS RIOS: Jauneche forest, km 70
TOBAGO. Calder Hall, 27 Jun. 1910, Broadway 4650 (F, Quevedo-Palenque road, vía Mocachi, 14 Jul. 1979,
US); Providence road, 27 Apr. 1910, Broadway 3597 Dodson et al. 7988 (MO). COTOPAXI: Quevedo-
(BM, F, S). Latacunga road, along Río Pilaló, 6 Apr. 1973, Holm-
GUYANA. Amakura river, NW district, 23-30 Mar. Nielsen et al. 3098 (AAU, F, QCA). TUNGURAHUA:
1923, Cruz 3549 (F, MO, UC); Essequibo river, Jun. Baños, over the river to Baños, 2 Nov. 1981, Madsen et
1923, Persaud 356 (F); bassin of Essequibo river, near al. 36475 (AAU). CHIMBORAZO: 20 Aug. 1943,
mouth of Onoro creek, 15-24 Dec. 1937, A.C. Smith 2724 Acosta-Solís 5466 (F). PASTAZA: Ceilán, 6 Jun. 1980,
(F). Brandbyge & Asanza 31633 (QCA); Río Villano, 24 Mar.
SURINAME: station Victoria, Dec. 1843, Kappler 1386 1980, Holm-Nielsen et al. 22662 (AAU, QCA); 3-4 km of
(F, S); Jodensavanne-Mapane Kreek area, 15 Dec. 1953, Puyopungu, 28 Sep. 1976, Lugo 5043 (F, GB); Río
Lindeman 5258 (MO); Table mountain, base S Bobonaza, near outlet into Río Pastaza, between
escarpment Arrowhead Basin, 26 Aug. 1944, Maguire Destacamento Cabo Pozo and La Boca, 21 Jul. 1981,
24495 (F); Nassau mountains, Marowijne river, 3 Jan. Øllgaard et al. 34932b (AAU, QCA). LOJA: Loja, 31 May
1955, Maguire et al. 39088 (UC). 1946, Espinoza 457 (GH, LOJA). GALAPAGOS
FRENCH GUIANA. Saint Jean du Maroni, 26 Mar. ISLAND. Santa Cruz: 17 Apr. 1974, Adsersen & Adsersen
1914, Benoist 1004 (F). SAUL: Mont La Fumée, 2 Sep. 78 (QCA); 5 mi N Academy Bay, 3 Mar. 1953, Bowman
1982, Boom & Mori 1594 (CAY). Saint Laurent, km 10 98 (UC).
road Paul Isnard, 5 Nov. 1981, Billiet & Jadin 1309 PERU. PIURA: Huancabamba, Canchaque-
(CAY). Ytany island, Haut Maroni, 20 Nov. 1977, Huancabamba, km 16-25 from Canchaque, 17 Apr.
Cremers 5098 (CAY); Cayenne, island 17 May 1979, 1987, Díaz & Baldeón 2394 (MO, NY, USM);
Cremers 5665 (CAY); 45 km SE from Aul, 31 Aug. 1980, Huancabamba, W side just below summit of Abra
Cremers 6503 (CAY); Riv. Mana, Ilots du Saut, 22 Jul. Porculla, between Olmos and Río Marañón, 26 Sep.
1981, Cremers 7290 (CAY); Baboune river, affluent of 1957, Hutchison 1386 (UC); Ayabaca, around Ayabaca,
Mana river, 29 Jul. 1981, Cremers 7358 (CAY); region 9 Sep. 1976, Sagástegui & Cabanillas 8703 (MO). LA
Paul Isnard, SW Citron, 7 Feb. 1983, Cremers 7879 LIBERTAD: Otuzco, Chuquizongo, 5 Jun. 1958, López et
(CAY); Salut, Ile Royale, 21 Feb. 1985, Cremers 8444 al. 2630 (GH). SAN MARTIN: Mariscal Cáceres, dist.
(CAY); Haut Oyapock, Saut Cambrouze, 18 Jul. 1975, Campanilla, Mashuyacu, 12 Aug. 1970, Schunke V. 4225
Granville 2482 (CAY); 12 km E fromSaul, 10 Jan. 1980, (F); Mariscal Cáceres, Tocache Nuevo, Pushurumbo, E
Granville 3252 (CAY); 18 km S from Saul, 4 Apr. 1983, of Puente Palo Blanco, 24 Dec. 1972, Schunke V. 5792
Granville 5542 (CAY); Haute Camopi, N of Mont (F); Alto Río Huallaga, Dec. 1929, L. Williams 5621 (F);
Belvedere, 2 Dec. 1984, Granville 7106 (CAY); St. Tarapoto, 14 Feb. 1947, Woytkowski 35070 (MO, UC).
Joseph, 6 Feb. 1967, Oldeman 2502 (CAY); Arataye LORETO: Maynas, Colonia Río Zumun, near Río
Yahuas-Yacu, 19 Mar. 1980, Barrier 1959 (USM); Río Mar. 1971, Prance et al. 11216 (F, MO, S); vic. of Uaicá
Ampiaco, 24 Sep. 1972, Croat 20731 (F, MO, UC); airstrip, 3 Dec. 1973, Prance 20008 (F). ACRE: Rio
Varadero de Mazán, 27 Sep. 1972, Croat 20807 (UC); Juruá-Mirim, near Lucania, 14 May 1971, Maas et al.
Río Ampiyacu, Brillo Nuevo-Río Yaguasyacu, 13 Mar. 12932 (F, S); Cruzeiro do Sul, Rio Juruá and Rio Moa,
1981, Davis et al. 900 (F, GH); Alto Amazonas, Río 28 Apr. 1971, Prance et al. 12622 (F). PARÁ : Lageira
Pastaza, 1 Aug. 1979, Diaz et al. 1324 (F, MO); airstrip on Rio Maicuru, 1 Aug. 1981, Strudwick et al.
Yanamono, Explorama tourist camp, trail to Río Napo, 3979 (F); Sete Varas airstrip on Rio Curua, 4 Aug. 1981,
19 Feb. 1981, Gentry et al. 31530 (MO, UC, USM); dist. Strudwick et al. 4077 (F), 6 Aug. 1981, Strudwick et al.
Indiana, quebrada Yanayacu, below Bombonaje, 27 4263 (F), 8 Aug. 1981, Strudwick et al. 4377 (F). GOIAS:
May 1973, McDaniel & Rimachi 17347 (GH); above Córrego Itaquera, ca. 30 km N of Formosa, 2 May 1966,
Pongo de Manseriche, left bank of Río Santiago, 22 Irwin et al. 15586 (F, US); Córrego Itaquera, 2 May 1966,
Dec. 1931, Mexia 6324 (BM, F, GH, MO, NY, S, UC, US); Irwin et al. 15596 (US). DISTRITO FEDERAL: Córrego
Maynas, dist. Pebas, Río Ampiyacu, 19 Jul. 1976, Revilla Vicente Pires, near Taguatinga, 8 Sep. 1965, Irwin et al.
862 (F, MO, NY, UC); Maynas, Iquitos, 17 Feb. 1968, 8111 (F); Planalto, ca. 15 km W of Brasilia, Riacho
Simpson & Schunke 679 (F). HUANUCO: Tingo María, Vicente Pires, 12 Jul. 1966, Irwin et al. 18169 (F, MO,
30 Oct. 1959-19 Feb. 1950, Allard 22332 (US); Muña, 23 US).
May-4 Jun. 1923, Bryan 549c (F); near carretera
marginal de la selva, near Puente Colombia, between Campyloneurum phyllitidis is characterized by
Río Mayo and Río Huallaga, 10 Jul. 1970, McDaniel the prominence of its secondary veins and its
13846 (GH); Pachitea, dist. Honoria, Bosque Nacional
symmetrically divided primary areoles.
Iparia, río Pachitea, near Miel de Abeja camp, 26 Sep.
1967, Schunke V. 2181 (F, GH, NY, US); Mariscal
This species has been confused with C.
Cáceres, Tocache Nuevo, trail to Pushurumbo, 7-8 km brevifolium and C. nitidum. However, C. phyllitidis
E de puente Palo Blanco, 24 Dec. 1972, Schunke V. 5792 differs from C. brevifolium by having
(F, NY, US); Tingo María, 9 Sep. 1956, Tryon & Tryon symmetrically divided primary areoles, and
5290 (GH, US). PASCO: Oxapampa, Quebrada therefore they represent different lineages within
Castillo, sobre el Río Omaiz, afluente del Chuchurras, the genus. From C. nitidum, it differs by having
12 Jun. 1987, León & Young 1083 (F, USM); Oxapampa, leaves predominantly with caudate or acuminate
5 km SE de Oxapampa, 9-11 Dec. 1982, D.N. Smith 2917 apices and acuminate stem scales.
(F, MO); Oxapampa, Palcazú valley, near the
Campyloneurum phyllitidis is a widespread
confluence of Río Palcazú and Río Iscosacin, 23 Apr.
1983, D.N. Smith 3874 (UC). JUNIN: Puente species and several phenotypes occur along its
Paucartambo to La Merced, 30 Jan. 1983, Gentry et al. range of distribution. Two cytotypes have been
39804 (F, MO); Satipo, Alto Quimiriqui, 30 Aug. 1982, reported in this taxa, and this fact may correlate
León 278 (AAU, F, USM); Chanchamayo, La Merced- with some of the morphological variation.
Puente Paucartambo road, 3 km from La Merced, 17
May 1983, D.N. Smith 4051 (MO, UC). CUSCO: La
Convención, Rosario Mayo, 10 Aug. 1968, Chavez 137 38. Campyloneurum repens (Aublet) C. Presl, Tent.
(GH); La Convención, Cocalpampa, Chaullay, 29-30
Pterid. 190. 1836.
Dec. 1986, Nuñez et al. 6772 (MO). MADRE DE DIOS:
Tambopata, Tambopata Natural Reserve, 14 Apr. 1980,
Polypodium repens Aublet, Hist. Pl. Guiane 2: 962.
Barbour 4768 (F, MO), Barbour 4967 (MO); Parque 1775. Lectotype (chosen by Proctor,in R. A.
Nacional Manu, Cocha Cashu Biological Station, 8 Howard 2: 340, 1977): Plumier t.134, Traité
Nov. 1984, Foster P-84-68 (MO, UC); Parque Nacional Foug. Amér. 117. 1705.
Manu, Cocha Cashu, 28 Sep. 1979, Foster et al. 7065 (F); Polypodium lapathifolium Poiret, Encycl. 5: 514.
Tambopata, 39 km SW of Puerto Maldonado, 10 Oct. 1804. Type. America Meridionale, Jussieu s.n.
1985, S.F. Smith et al. 669 (F, MO, NY). (holotype, P, photo of P, BM!, S!).
BOLIVIA. LA PAZ: Antahuacana, Jun. 1909, Buchtien
Campyloneurum lapathifolium (Poiret) Ching,
2167 (UC); Polo-Polo, Coroico, Oct-Nov. 1912, Buchtien
3533 (F, S); Casanare, Tipuani-Tali, 6 Oct. 1922,
Buchtien 7078 (MO); Nor Yungas, 32.1 km S of Polypodium caespitosum Link, Hort. berol. 2. 91.
Caranavi, 26 Mar. 1982, Solomon 7368 (UC, MO). 1833. Type: Cultivated, ex Hort. Loddiges
BRAZIL. RORAIMA: Serra dos Surucus, 17 Feb. 1969, (holotype B!).
Prance et al. 9974 (AAU, F); Posto Mucajaí, Rio Mucajaí, Campyloneurum caespitosum (Link) Link, Fil. spec.
vic. of Mucajaí airstrip, 14 Mar. 1971, Prance et al. 10996 125. 1841.
(F, S), 18 Mar. 1971, Prance et al. 11086 (F, S, US), 25 Campyloneurum crispum Fée, Gen. Filic. 259. 1852.
Type. Brazil: Martius 303 (holotype, not Williams & Molina 11338 (F).
found; isotypes, BM!, MO!). GUATEMALA. PETEN: Dolores, on Santo Toribio
Stem dark stramineous, black, not pruinose, 1- road, 20 Apr. 1961, Contreras 2138 (MO); between Finca
Yalpemech and Chinajá, 28 Mar. 1942, Steyermark 45437
3 (-4) mm wide. Stem scales brown or light
(F). IZABAL: along trail between Dartmouth and
brown, 3-4 mm long, 1-1.5 mm wide, ovate, the Morales towards lago Izabal, 7 Apr. 1940, Steyermark
cells oblong, cell walls 5-7 µm thick, marginal cell 39065 (F). SOLOLA: S facing slopes of volcán Atitlán,
walls lighter than the central cell ones. 20 Jun. 1942, Steyermark 47892 (F).
Phyllopodia 1 mm long, 2-2.5 mm wide, 0.7-15 NICARAGUA. RIO SAN JUAN: Río San Juan, 4 Dec.
mm apart. Leaves 20-60 cm long; petiole 0.5-7 (- 1982, Araquistain 3416 (MO). BOACO: cerro
13) cm long, stramineous; lamina lanceolate or Mombacito, 4 km NW de Camoapa, 1 Feb. 1979,
obovate lanceolate, sometimes narrowly Grijalva & Araquistan 42 (MO).
lanceolate, bases attenuate, apices acuminate or COSTA RICA. PUNTARENAS: 4 mi W of Rincón de
Osa, 4-7 Jun. 1968, Burger & Stolze 5405 (F, NY), Burger
caudate, (2.5-) 3-8 cm wide, chartaceous, margins
& Stolze 5536 (F); about 5 km W of Rincón de Osa, 24-
cartilaginous, slightly sinuate, usually plane, 30 Mar. 1973, Burger & Gentry 9014 (F, NY); foothills of
indument of scarce bicellular glandular hairs, the Cordillera de Talamanca, N of Las Alturas, 28 Aug.
scattered abaxially; hypostomatic, stomata 1983, Davidse 24149 (MO, UC); ca. 10 mi SE of Rincón
polocytic or copolocytic; costa prominent, slightly de Osa, along road to Pacific, 18 Jul. 1967, Evans &
ranuarte adaxially, angular or convex abaxially, Bowers 2806 (MO); Santiago, near San Ramón, 21 Apr.
primary veins prominent or prominulous, 1913, Tonduz 17572 (BM, F, NY, S, UC). CARTAGO:
stramineous or darker than the leaf tissue, Río Chitaria, E of Buenavista, 20 km from Turrialba, 24
Aug. 1983, Saiki 58 (F).
slightly flexuous or straight, (65o-) 70-75o PANAMA. CHIRIQUI: Burica Peninsula, 10-11 mi W
divergent from the costa, (3-) 5-7 mm apart, of Puerto Armuelles, in vic. of San Bartolo, 19 Feb.
secondary transversal veins forming (4-) 6-12 1973, Croat 21983 (MO); 25 km W of El Hato del
primary areoles between the costa and margin, 2 Volcán, 19 Oct. 1980, Maas & Dressler 4939 (MO); km 3
(-4) free excurrent veinlets in each non costal on La Unión road NW of volcano, 23 May 1971, Proctor
areole; sori medial or subterminal, spores 50-70 32029 (F). PANAMA: at headwaters of Río Indio,
slopes of Cerro Jefe, 2 Nov. 1979, Antonio 2433 (MO), 18
µm long, 30-40 µm wide. Figs. 1 b; 46.
Dec. 1980, Antonio 3229 (MO); over Río Guanche, 19
Jan. 1980, Antonio 3344 (MO); Río Maje, 20 Apr. 1976,
This species is distributed from Mexico, Croat 34415 (MO); 20 km above Pan. Am. highway, on
Central America, Greater Antilles to Bolivia and trail from El Llano to Carti-Tupile, 22 Feb. 1973,
central Brazil, where it grows mostly as an Kennedy 2563a (MO); Altos de Pacora, 4 Feb. 1979,
epiphyte between 100 m and 2000 m elevation. Windisch 2199 (F). CANAL ZONE: Limbo Hunt Club
road, 1 Nov. 1972, Kennedy & Andrews 1878 (MO);
Common name: "Shan tape" (colorado). Parque Nacional Soberanía, trail to oilline, 17 May
1980, Vásquez 192 (MO). COCLE: vic. of El Valle, 14
May 1939, Allen 1799 (F, MO); near La Mesa, 11 Feb.
REPRESENTATIVE SPECIMENS: MEXICO.
1971, Croat 13346 (F, MO, NY); road to Coclosito, 12 mi
OAXACA: 23.5 mi S of road to Jesus Carranza, 28 Jul.
from Llano Grande, 9 Dec. 1983, Churchill et al. 4019
1966, Cruden 1113 (UC); dist. Ixtlán, 29 km S of Valle
(MO, UC); El Valle, 11 May 1977, Folsom 3110 (MO,
Nacional, 80 km N of Ixtlán de Juárez, 13 Aug. 1971,
UC); 7 km N of Llano Grande on road to Coclesito, 19
Mickel 6374 (UC). CHIAPAS: Mun. Las Margaritas, at
Apr. 1978, Hammel 2511 (MO); foot of cerro Pilón,
confluence of the Río Ixcán with the Río Lacantum (Río
above El Valle de Antón, 28 Mar. 1969, Porter et al. 4615
Jatatí), 14 Mar. 1973, Breedlove & McClintock 34063 (F,
(MO). HERRERA: 18 km W of Las Minas, trail to top
MO); between Palenque and Bonampak, SW of
of Alto Higo, 5 Aug. 1978, Hammel 4230 (MO); between
Palenque, 5 Jul. 1977, Croat 40209 (MO).
Las Minas and El Toro, near village of Chepo, 24 Jan.
BELIZE. CAYO: Vaca Plateau, Blue Hole Camp, 6 Aug.
1987, McPherson 10290 (MO). LOS SANTOS: trail
1980, Whitefoord 2044 (BM, MO). TOLEDO: vicinity of
between Jobero and Río Pedregal, 29 Apr. 1976, Croat
San Jose, 6.7 mi N of Columbia, 13 Jun. 1973, Croat
34517 (MO); from El Cortezo to Arenas, 27 Oct. 1978,
24462 (MO). Maya mountains, vicinity Cockscomb
Hammel 5367 (MO). DARIEN: N Punta Guayabo
mountains, 8 Jun. 1930, Schipp 527 (BM, F, NY, S, UC).
Grande, 21 Apr. 1980, Antonio & Hahn 4321 (MO);
HONDURAS. CORTES: Santa Cruz de Yojoa, 26 Oct.
Parque Nacional Darién, at N base of cerro Pirre, 8 Oct.
1933, Edwards 706 (F); Agua Azul, 27 Dec. 1946, L.O.
1987, Hammel et al. 16153 (MO); RENARE hut in Darien
National Park, 5 Aug. 1986, McDonagh et al. 444 (MO). from Cerro Neblina camp, 12 Apr. 1984, Gentry & Stein
JAMAICA. PORTLAND: gorge of the Stony river, 46563 (MO); Río Negro, 1.5 km E of Cerro de la
below junction of the Macungo river, 24 Jul. 1967, Neblina, 2-3 Dec. 1984, Liesner 17473 (MO); Río Negro,
Proctor 28322 (F, MO). Dollwood, 6 Aug. 1898, Harris Cerro de la Neblina, valley N base of Pico Cardona, 21-
7273 (F); foothills of John Crow mountains, E of 24 Mar. 1984, Liesner & Stannard 16873 (F, MO).
Seamen's valley, 18 Feb. 1920, Maxon & Killip 235 (F); DISTRITO FEDERAL: Colonia Tovar, 1854-55, Fendler
lower eastern slopes of Mount Diabolo, 29 Feb. 1920, 229 (F); 6 km NE of Colonia Tovar, before turnoff to El
Maxon & Killip 543 (F); road from Silver Hill Gap to Limón on road to Chichiriviche, 7 Apr. 1982, Liesner &
Hardwar Gap, 19 Mar. 1920, Maxon & Killip 1216 (F, Medina 13546 (MO). Upper Orinoco region, La
GH); trail from Morces gap to Vinegar hill, 21 Mar. Mantequilla, 22 Sep. 1951, Croizat 710 (F, MO); Santa
1920, Maxon & Killip 1321 (F); Vinegar Hill road, 2 Jun. Marta, H.H. Smith 1040 (MO).
1910, York 115 (MO); Petit Bordel, 9 Jan. 1890, Eggers GUYANA. Upper Rupununi river, near Dadanawa, 2
6871 (F, S). Without locality, 1895, Moore s.n. (MO); Jun. 1922, De la Cruz 1449 (F, MO); New River, 1/4
Mart. 2874 (MO). mile S of camp 3, 5 Oct. 1952, Guppy 6376 (BM, F);
WINDWARD ISLANDS. MARTINIQUE: 1869, Barima-Waini region, 3 mi W of Eclipse Falls, below
Belanger s.n. (F); 1868, Husnot s.n. (F). Wanamaparu, 2 Aug. 1986, Pipoly & Lall 8172 (F),
COLOMBIA. ANTIOQUIA: Mun. San Luis, Cañón del Pipoly & Lall 8183 (F).
río Claro, 2 Sep. 1984, Cogollo 1887 (MO). VALLE: SURINAM. Haute crique, Waamahpann, 25 Jul. 1972,
Western Cordillera, hoya del Río Anchicayá, 20 Dec. de Granville 966 (CAY); Tumac Humac, between
1942, Cuatrecasas 13745 (F); Río Cajambre, 21-30 Apr. Kouaipann and Palouloui, 1 Aug. 1972, Granville 1067
1944, Cuatrecasas 17111a (F, S); western cordillera, (CAY); Tafelberg, Arrowhead Basin, 26 Aug. 1944,
Hoya del Río Cali, trail to Miralindo, 31 Oct. 1944, Maguire 24507 (F, MO, NY); Nassau mountains,
Cuatrecasas 18439 (F); along road between Cali and Marowijne river, 7 Jan. 1955, Maguire et al. 39195 (NY);
Buenaventura at km 18.5 on road from Finca Santa Tumuc Humuc Mountains, source of Litani river, 7
Elena, 27 Aug. 1976, Croat 38504 (MO). CAUCA: valle Nov. 1937, Rombouts 879 (MO, US).
del Cauca, May 1948, Dryander 2931 (F); Río Naya, near FRENCH GUIANA: Arataye river, Sauts Parare, 5 Feb.
El Pastico, 23 Feb. 1983, Gentry & Juncosa 40624 (UC). 1981, Barrier & Feuillet 2524 (CAY); 45 km SE of Saul,
META: Mun. La Macarena, sobre el Río Guayabero, 11 summit of Tabulaire, 19 Aug. 1980, Cremers 6337
Aug. 1988, Callejas & Marulanda 7066 (MO). CHOCO: (CAY), 21 Aug. 1980, Cremers 6383 (CAY); riviere
Mun. Río Sucio, Alto del Limón, 4 Jun. 1976, Forero et Mana, Saut Dalles, 18 Jul. 1981, Cremers 7220 (CAY);
al. 1811 (MO); Hoya del Río San Juan, La Sierpe, 1 Apr. Mont Gauthiot, Yaroupi, 20 Apr. 1970, Granville 420
1979, Forero & Jaramillo 4454 (MO); Mun. San José del (CAY); Monts Galbao, 10 May 1973, Granville 1594
Palmar, Hoya del Río Torito, 1 Mar. 1980, Forero et al. (CAY).
6433 (MO); 4 Mar. 1980, Forero et al. 6619 (MO), Forero ECUADOR. CARCHI: valle de Maldonado, km 60 on
et al. 6683 (MO); Hoya del Río Atrato, Beté, 5 Apr. 1982, road Tulcán-Maldonado, 18 May 1973, Holm-Nielsen et
Forero et al. 8889 (MO); area of Baudó, 11 Feb.-29 Mar. al. 5775 (AAU, F, USM). ESMERALDAS: Río San
1967, Fuchs & Zanella 22253 (F). Mutatá, road to Bajirá, Miguel, upstream from San Miguel de Cayapas, 1 Sep.
Nuevo Oriente, 19 Mar. 1983, Brand & Ascanio 270 1980, Holm-Nielsen et al. 25465 (AAU, USM).
(MO). SANTANDER: between Pamplona and IMBABURA: Collapi, 4 Jun. 1949, Acosta-Solís 12795 (F).
Bucaramanga, 5 May 1983, Croat 56507 (UC). NAPO: Río Eno al NE de Shushufindi, 11 Apr. 1982,
PUTUMAYO: Puerto Porvenir, above Puerto Ospina, Balslev 2326 (AAU, QCA); San Pablo de los Secoyas, on
22 Nov. 1940, Cuatrecasas 10755 (F). CAQUETA: the path to Shushufindi, WSW of the village, 6 Aug.
Eastern cordillera, Sucre, 4 Apr. 1940, Cuatrecasas 9074 1980, Brandbyge et al. 32545 (UC), 7 Aug. 1980,
(F). Brandbyge et al. 32633 (AAU, QCA); Río Wai si ayá, 6
VENEZUELA. LARA: Moran, between Agua Amarilla km upriver from San Pablo, 10 Aug. 1980, Brandbyge &
and Santo Domingo, 24 Oct. 1987, Rivero & Diaz 1316 Asnza 32739 (AAU, QCA); Reserva Biológica Jatun
(MO). MIRANDA: Paéz, Fila La Tigra, Quebrada San Sacha, 8 km E de Misahuali, 21-25 May 1987, Cerón
Juan, 2-7 Sep. 1977, Ortega & González 386 (MO), 1436 (MO); along road from Tena, past Muyuna, ca. 5.7
Ortega & González 405 (MO). ANZOATEGUI: along km W of Tena, 1 May 1984, Croat 58853 (MO); Río
Río León, by Quebrada Danta, 20 Feb. 1945, Steyermark Yasuní, 80 km upriver from Rocafuerte, 21 Sep. 1977,
61033 (F, MO). BOLIVAR: Caño Pablo, tributary of Río Foster 3794 (F); Río Yasuní, Charapillo, 26 Aug. 1979,
Caura, ca. 6 km ESE of Las Pavas, 8 May 1982, Liesner Holm-Nielsen et al. 19954 (AAU, QCA); Río Cuyabeno,
& Morillo 13946 (MO); Salto del Para, Medio Caura, 3 near to Río Aguarico, 20 Feb. 1980, Holm-Nielsen et al.
Mar. 1939, L. Williams 11361 (F). TERRITORIO 21609 (AAU, QCA); San Pablo de los Secoyas, 4 Jul.
FEDERAL AMAZONAS: Sierra Parima, headwaters of 1980, Jaramillo & Coello 2793 (AAU, QCA), Jaramillo &
Río Siapa, 11-23 Mar. 1946, Cardona 1321 (F); trail S Coello 2821 (AAU, QCA); Río Yasuní, Garza Cocha, 10
Apr. 1983, Lawesson et al. 43381 (AAU, QCA); Río Rimachi 22536 (NY); Maynas, Iquitos, 26 Jun. 1984,
Yasuní, upstream from Garza Cocha, 11 Apr. 1983, Moran 3649 (F, MO, UC); Gamitanacocha, Río Mazan,
Lawesson et al. 43439 (QCA); Parque Nacional Yasuní, Feb. 1935, Schunke s.n. (US, USM); 9 Feb. 1935, Schunke
Anango, 15 Jul. 1982, Luteyn et al. 8698 (F); below 205 (F, GH, NY, S, UC, US). PASCO: Prov. Oxapampa,
Puerto Misahualli, 18-30 May 1985, Palacios et al. 407 Palcazú valley, Iscozacin, 10 Jan. 1984, Foster et al. 7833
(QCA). PICHINCHA: road Nono-Nanegalito, N de (F); Río Alto Iscozacín, Ozuz to Río Pescado, 12 May
Cerro Pichincha, 9 May 1982, Balslev & Boom 2500 1985, Foster & d'Achille 10113 (F); Prov. Oxapampa,
(AAU); Tinalandia, 9.6 km E of Santo Domingo de los Quebrada Castilla, Amuesha community on Río
Colorados, S of highway to Alaog and Quito, 3 Apr. Omaiz, 10 Jun. 1987, León & Young 1065 (MO, USM).
1983, Croat 55708 (MO); Centinela, cantón Santo JUNIN: Pichis trail, Santa Rosa, 6,7 Jul. 1929, Kiilip &
Domingo, 23 Aug. 1978, Dodson et al. 7181 (F); Smith 26176 (US); E of Satipo, Aug. 1940, Ridoutt s.n.
Congoma Grande, at km 23 on the Santo Domingo- (US). AYACUCHO: La Mar, between Tambo San
Puerto Limón road, 4 Jun. 1982, Kvist & Holm-Nielsen Miguel, Ayna and the Hacienda Luisiana, 20 Aug.
40095 (QCA), 7 Jun. 1982, Kvist & Holm-Nielsen 40132 1968, Dudley 11908 (GH). CUSCO: La Convención,
(QCA), 21 Jul. 1982, Kvist 40672 (QCA). COTOPAXI: along Río Klause, 15 Jun. 1968, Dudley 10181 (GH);
Tenefuerte, Río Pilaló, Latacunga, 7 Feb. 1982, Dodson Urubamba, Machu Picchu, on the Río Mandor, 2.5 km
& Gentry 12204 (MO); Tenefuerte, km 52-54 Quevedo- from Machu Picchu, 2 Jun. 1982, Peyton & Peyton 379
Latacunga, 9 Apr. 1984, Dodson & Thurston 14228 (MO); (MO); Quispicanchis, Quincemil, May 1951, Vargas
road Quevedo-El Corazón, 9 km ENE of Moraspungo, 10090 (UC); Paucartambo, Hacienda Villa Carmen, 19
13 May 1980, Harling & Andersson 19044 (F, GB, QCA). Jul. 1963, Vargas 14679 (GH); Quispicanchis, Punkiri, 14
PASTAZA: Curaray, SE of the airstrip, 20 Mar. 1980, May 1964, Vargas 15414 (GH). MADRE DE DIOS:
Holm-Nielsen et al. 22242 (AAU, QCA), 22 Mar. 1980, Tambopata, SSW of Puerto Maldonado, Tambopata
Holm-Nielsen et al. 22511 (AAU, QCA); Río Papayacu at Natural Reserve, 19 Apr. 1980, Barbour 4853 (F, MO),
Río Curaray, 23 Mar. 1980, Holm-Nielsen et al. 22595 10 May 1980, Barbour 5216 (F, MO); Prov. Manu,
(AAU, QCA); Río Bobonaza, between Cachitama and Atalaya, vic. Hacienda Amazonía, 12 Dec. 1983, Foster
the outlet of Río Bufeo, 19 Jul. 1980, Øllgaard et al. 34718 & Wachter 7428 (F); on trail between lodge and Río La
(AAU, QCA), Øllgaard et al. 34808 (AAU, QCA); Torre, 2 Jun. 1986, Funk et al. 8379 (US); Tambopata
between destacamento Chiriboga and Apachi Entza, 24 Wildlife Reserve, 30 km S of Puerto Maldonado, 2 Dec.
Jul. 1980, Øllgaard et al. 35190 (AAU, QCA). 1984, H.J. Young & Stratton 330 (MO); Tambopata,
MORONA-SANTIAGO: between La Esperanza and Explorer's Inn, 39 km SW of Puerto Maldonado, 24 Jan.
Santa Ana, Huamboya, 15 Feb. 1944, Acosta-Solís 7436 1989, S.F. Smith et al. 1576 (US); Tambopata Reserve,
(F); Guaruma, km 38 carretera Saloya, 20 Aug. 1945, junction of Río La Torre and Río Tambopata, 16 Mar.
Acosta-Solís 11010 (F); Taisha, Río Panguientza, 21 Jun. 1981, Young 121 (MO, NY, UC).
1980, Brandbyge & Asanza 32175 (AAU, QCA); BOLIVIA. PANDO: Manuripi, 21 km from Puerto
Pumpuentza, 1 km E of village, 28 Jun. 1980, Brandbyge Rico, towards Conquista, 28 Jan. 1983, Fernández Casas
& Asanza 32353 (AAU, QCA); road Limón (General & Susanna 8513 (MO); Nicolas Suarez, SW of Cobija, on
Plaza)-Macas, ca. km 20 from Limón, 26 Mar. 1974, Río Naraueda, 1 Aug. 1982, Sperling & King 6460 (MO).
Harling & Andersson 12909 (GB, MO); Misión LA PAZ: Larecaja, Consata, 7 km hacia Mapiri, 14 Dec.
Bomboiza, 23 Apr. 1973, Holm-Nielsen et al. 4288 (AAU, 1971, Beck 4917 (LPB); Antahuacana, Jun. 1909, Buchtien
F, MO). ZAMORA-CHINCHIPE: Río Nangaritza, 2156 (BM, S, UC); Polo Polo bei Coroico, 1912, Buchtien
Colina Salada, 8 Dec. 1990. Øllgaard 98482 (QCA), 3536 (F, S, US); Murillo, 44 km below lago Zongo dam,
Øllgaard 98457 (QCA). 12-15 Sep. 1983, Solomon 10853 (MO).
PERU. AMAZONAS: Río Cenepa, Kayamas creek, 5 BRAZIL. ACRE: 25-30 km NW of Rio Branco, on road
km N of confluence of Huampani and Cenepa, 4 Dec. to Sena Madureira, 25 Feb. 1978, Anderson 12123 (F).
1972, Berlin 451 (MO); Puerto Nazareth, 22 Dec. 1970, TERRITORIO DO RORAIMA: vic. Auaris, 6 Feb. 1969,
Ellenberg 3490 (LPB); Nazareth, 14 Sep. 1912, Osgood 26 Prance et al. 9657 (AAU, F); Serra dos Surucus, 6 Feb.
(F), Osgood 27 (F). SAN MARTIN: San Martín, Caserío 1971, Prance et al. 13516 (AAU, F). PARÁ: Marabá, 1
El Progreso, on Tarapoto-Yurimaguas road,25 Sep. Jun. 1982, Secco et al. 404 (F); Serra dos Carajás, 10 Jun.
1986, Knapp & Mallet 8415 (MO, NY); Mariscal Cáceres, 1982, Sperling et al. 6068 (F). RONDONIA: Rio dos
Tocache Nuevo, Puerto Pizana, 14 Jun. 1974, Schunke V. Pacaás Novos, 22 Mar. 1978, Anderson 12235 (F).
6952 (F, MO, UC). LORETO: near Tuta Pishco on Río
Napo, 16 Sep. 1972, Croat 20293 (MO); Maynas, Campyloneurum repens is characterized by its
Quebrada Yanamono, Río Amazonas, 15 Nov. 1979, acuminate stem scales with differentiated
Gentry & Jaramillo 28084 (MO); Maynas, dist. Iquitos, 11
margins. It is closely related to C. ophiocaulon,
Aug. 1978, Hickok 612 (GH); Nauta, Quebrada de
Sapira, caserío Florida, 26 May 1979, McDaniel &
from which it differs by the obtuse stem scales.
24 Sep. 1908, Dusén 6607 (BM, S); Río Uruguay, 31 May

39. Campyloneurum rigidum J. Sm., Cult. Ferns 13. 1911, Dusén 11782 (S); Porto da Cima, 30 Jul. 1912,
1857. Type. Cultivated, from Tropical Dusén 14168 (S); Jacareí, 11 Nov. 1915, Dusén 17327
(BM, GH, S); Paranaguá, Praia de Mendanha, 13 Aug.
America, Herb. J. Smith s.n. (probably type
1961, Hatschbach 8205 (US). SANTA CATARINA:
collection, BM!). Santa Catarina, Gaudichaud 268 (F); Hammonia, Jun.
Polypodium rigidum (J. Sm.) Lowe, Ferns 2, t.37a. 1911, Luederwaldt 634 (BM); Joinville, 1906, Müller 398
1858. Nom. nud. Not Polypodium rigidum (BM); Brusque, 30 Aug. 1947, Reitz 1839 (S); Morro da
Aublet, 1775; Hoffman, 1795; Hooker & Lagoa, Ilha de Santa Catarina, 2 Aug. 1945, Hno. Rohr
Greville, 1829. 325 (US); Joinville, 15 Jul. 1901, Schmalz 38 (F).
Stem creeping, dark stramineous, not
pruinose, 2-4 mm wide. Stem scales dark brown Campyloneurum rigidum is characterized by
in mass, 2-3 mm long, 1-1.5 mm wide, ovate, the having light-green shiny leaves and adpressed
cells oblong, central cell walls 10-15 µm thick, stem scales. It belongs to the C. amphostenon
marginal cell walls 5 µm wide. Phyllopodia 0.5-1 group.
mm long, 1-2 mm wide, 2-4 mm apart. Leaves
30-55 cm long; petiole green stramineous, (2.5-)
3.5-6 (-7) cm long, slightly ranurate adaxially, 40. Campyloneurum solutum (Klotzsch) Fée, Gen.
convex abaxially, rarely with spreading scales, Filic. 258. 1852.
similar to those on the stem; lamina linear- Polypodium solutum Klotzsch, Linnaea 20: 399.
lanceolate or narrowly lanceolate, bases and 1847. Lectotype (chosen here) Colombia,
apices attenuate, (1-) 1.5-2.5 cm wide, coriaceous, Moritz 309 (B, BM!). Syntype. Without
green yellowish shining on both sides of the locality: 1843, Hartweg 1493 (B!, isolectotypes
lamina, margin cartilaginous, repand, indument BM!, LD!).
of inconspicuous, scarce, bicellular, simple Polypodium nodosum Klotzsch, Linnaea 20: 400.
glandular hairs scattered abaxially, 1847. Type. Colombia, Páramo de la Culata,
hypostomatic, stomata not seen; costa prominent, Moritz 310 (holotype B!, isotypes, BM!).
slightly ranurate adaxially, convex Campyloneurum nodosum (Klotzsch) Fée, Gen.
or slightly angulate abaxially, primary veins Filic. 258. 1852.
inconspicuous, secondary veins forming 2-3 Campyloneurum jamesoni Fée, Gen. Filic. 259.
primary areoles between the costa and margin, 1852. Type. Ecuador: Pichincha, Quito,
primary areoles symmetrically divided, each Jameson s.n. (holotype,probably P, isotypes
secondary areole with one free excurrent veinlet; BM!).
sori subterminal, paraphyses and spores not seen. Polypodium angustifolium f. remotifolium Hieron.,
Fig. 47. Bot. Jahrb. Syst. 34: 531. 1904. Type:
Colombia, Tolima, Mt. Ruiz, May 1882,
This species is known from southern Brazil, Schmidtchen (B); and Cauca, Páramo de las
where it grows as a terrestrial between 50 m and Delicias, Lehmann 4439 (B, isosyntype US!, F!).
800 m elevation. Campyloneurum remotifolium (Hieron.) Lellinger,
Amer. Fern J. 78: 26. 1988.
REPRESENTATIVE SPECIMENS: BRAZIL. RIO DE Stem long-creeping, black, usually pruinose,
JANEIRO: Rio de Janeiro, Glaziou 2073 (S); Rio de 1-3 mm wide. Stem scales shiny dark brown in
Janeiro, Sellow 1815 (BM). SÃO PAULO: Ribeira, May mass, caducous and spreading, (3-) 4-7 mm long,
1911, Brade 5099 (S); Morro das Pedras, 1922, Brade s.n. 1-2 (-2.5) mm wide, narrowly ovate,
(US); Feb. 1928, Brade s.n. (UC); Capivary, Mar. 1897,
pseudopeltate, bases auriculate or short
Gerdes 38 (UC); Tietê, 23 Dec. 1906, Gerdes 60 (S);
Campinas, 27 May 1905, Heiner 484 (S); Santos, 20 Mar. auriculate, apices acuminate, slightly clathrate,
1875, Mosén 3745 (BM, S); Toledo, 18 Feb. 1903, Ulricht without differentiate margins, except at the base,
24 (S); Santos, Guarujá, Jan. 1907, Usteri 22087 (BM); the cells narrowly oblong, along the main axes of
Alto da Serra, Wacket 139 (S). PARANÁ Mun. the scale, cell walls 15 µm thick, cellular lumen
Antonina, Serrinha, 30 Nov. 1983, Callejas t al. 1814 yellowish ot translucent. Phyllopodia 1-2 mm
(MO); Morretes, 21 Apr. 1904, Dusén 4458 (S); Jacareí, long, 1.5-2 mm wide, 5-10 mm apart. Leaves 15-
40 (-60) cm long; petiole stramineous or dark (AAU, UC, USM). along Río Topo, SE of Aucacocha,
stramineous, 4.5-15 cm long; lamina narrowly 18 May 1982, Øllgaard & Holm-Nielsen 38784 (AAU,
lanceolate, bases and apices attenuate, 0.7-2.5 (-3) USM). PICHINCHA: Chaparro de Sebritana, 9 Jul.
1944, Acosta-Solís 8359 (F); N slope of Volcán
cm wide, chartaceous, margins cartilaginous,
Pichincha, 24 Jan. 1981, Balslev et al. 1743 (AAU, NY,
slight or strongly revolute, indument not seen, UC); road to Yanacocha, NW of Cerro Pichincha, 3
stomata polocytic; costa prominent, primary Oct. 1981, Balslev 2045 (QCA); 30 Sep. 1982, Balslev &
veins usually slight prominulous abaxially, more Steere 3264 (QCA); Volcán Cayambe, 3 Jul. 1980, Holm-
or less concolorous with the adjacent tissue, Nielsen & Øllgaard 24333 (AAU, UC); Volcán Iliniza, 14
slightly flexuosous, secondary transverse veins Aug. 1980, Holm-Nielsen et al. 25018 (AAU, F); Volcán
forming 2-3 primary areoles between the costa Atacazo, W slope 17 km from San Juan, 25 Aug. 1980,
and margin, each primary areole symmetrical Holm-Nielsen & Asanza 25143 (AAU); Volcán
divided, with one free excurrent veinlet; sori Pichincha, Cerro Ventanillas, 9-10 Jan. 1984, Laegaard
51048 (QCA); Páramo de Guamani, 5 km W of Paso de
medial or subterminal, paraphyses not seen,
la Virgen, 8 Feb. 1984, Laegaard 51368 (QCA); Páramo
spores 55-60 µm long, 35-40 µm wide. Fig. 47. de Guamani, N of road Pifo-Papallacta, W of the pass,
10 Nov. 1990, Øllgaard 98237 (QCA); Páramo de
This species is distributed in Colombia, Guamani, 16 Jan. 1981, Proctor 38745 (QCA); NW of
Venezuela, Ecuador, and Perú. It grows in open Volcán Atacazo, 30 Nov. 1985, Zak 730 (F, QCA); NW
grassland areas, and high montane forest; of Volcán Pichincha, 21 Mar. 1987, Zak 1843 (F).
usually terrestrial, among rocks, usually COTOPAXI: Parque Nacional Cotopaxi, al lado W de
between 3000 m and 4500 m elevation. loma Ingapirca, 2 Oct. 1982, Balslev 3337 (QCA),
Balslev & Muñoz 3398 (AAU); trail from El Corazón to
Facundo Vela, 1-3 km from El Corazón, 17 May 1980,
REPRESENTATIVE SPECIMENS: COLOMBIA:
Harling & Andersson 19194 (AAU, GB); Latacunga-
CALDAS: Cordillera Central, Río Otún, towards
Quevedo road, between Pujilí and Zumbagua, 26 May
Nevado de Santa Isabel, 24 Nov. 1946, Cuatrecasas
1979, Lojtnant et al. 13695 (AAU, GB, QCA).
23146 (F, S, US); road between Manizales andHotel
BOLIVAR: road above Balzapamba, 2 May 1942,
Termales del Ruiz, 8 Jun. 1966, Forero et al. 523 (F);
Haught 3299 (S). TUNGURAHUA: Páramo of Miniza,
road Termales-Refugio, 22 Oct. 1961, Murillo 464 (F);
7 Apr. 1939, Penland & Summers 351 (F).
vic. Termales, 20 km SE of Manizales, 22 Oct. 1961,
CHIMBORAZO: road Riobamba-Penipe-Bayuschi-
Tryon & Tryon 6131 (S, UC, US). CUNDINAMARCA:
Llurarllacu, 18 Feb. 1987, Jaramillo 9419 (QCA); Cerro
Macizo de Bogotá, páramo de Palacio, El Tablón, 14
Chiguazo, 24 Sep. 1968, Lugo 474 (F, GB); 10 km NE of
Dec. 1959, Cuatrecasas 25651 (US). VALLE: Cordillera
Alao, at Cuspipaccha, 6 May 1982, Øllgaard et al. 38139
Occidental, Los Farallones, Alto del Buey, 11,12 Oct.
(AAU); road 10 km NE of Alao at Cuspicaccha, 6 May
1944, Cuatrecasas 17969 (F); Cordillera Central, Río
1982, Øllgaard et al. 38153 (AAU, QCA). AZUAY:
Tulua, Quebrada de Las Vegas, 21-23 Mar. 1946,
Páramo El Cajas, W of Sayausí and Cuenca, 4 Jan.
Cuatrecasas 20385 (F)
1981, Balslev 1443 (AAU, NY, UC); Páramo de Cajas,
VENEZUELA. MERIDA: Páramo de los Leones (La
27 Dec. 1976, Boeke & Loyola 635 (UC); 5-6 km above
Lagunita, La Cañada Grande), W de Mucurubá, 31
Angas, 5 Mar. 1985, Harling & Andersson 22750 (QCA);
May 1930, Gehriger 141 (F); dist. Rangel, Cuenca de la
between Molleturo and Quinoas, 15 Jun. 1943,
Quebrada Las Escaleras, Páramo de Minugú, 10 km
Steyermark 53106 (F). LOJA: Alamor-Celica road, 2-3
SE de San Rafael de Mucuhíes, 16-17 May 1972, Ruiz-
km S of Río Alamor, 5 Apr. 1980, Harling & Andersson
Terán 7239 (UC); Cerro de Turumiquire, 1925, Tate 192
17939 (AAU, GB); W side of Laguna Parcacocha,18
(US).
Mar. 1979, Lojtnant & Molau 11149 (AAU, GB, QCA).
ECUADOR. CARCHI: Páramo El Angel, road El
Without locality. Mar. 1906, Rimbach 2 (S, US, UC);
Angel-Tulcán, 15 May 1973, Holm-Nielsen et al. 5480
Gualea, 1888, Sodiro s.n. (UC).
(AAU, F, GB, UC); base of Volcán Chiles, km 34-36 on
PERU. CAJAMARCA: Sánchez V. 438 (AAU).
road Tulcán-Páramo Maldonado, 19 May 1973, Holm-
Nielsen et al. 5912 (AAU, F, GB, UC). IMBABURA:
Lago San Marcos Cayambe, 28 Nov. 1961, Cazalet & Campyloneurum solutum is characterized by its
Pennington 5374 (UC); Laguna Mojanda, Laguna spreading narrowly ovate stem scales, which are
Negra, 22 Sep. 1990, Øllgaard 98197 (QCA). NAPO: N dark brown and shiny, with 15-20 narrowly
side of cerro Sumaco, loma NW of campsite, 28 Apr. oblong cells at the middle part of the scale, and
1979, Holm-Nielsen et al. 17398 (AAU, QCA); road with a yellowish cellular lumen.
Quito-Baeza, 17 Jul. 1976, Øllgaard & Balslev 8022 Campyloneurum solutum grows in open areas,
and it usually has small narrowly lanceolate transverse tertiary veins inconcpicuous or
leaves, less than 35 cm long, rarely reaching 60 slightly prominulous, primary areoles
cm (Cuatrecasas 17969). undivided, 2 (-3) free excurrent veinlets in each
Campyloneurum solutum and C. asplundii areole; sori subterminal or medial, paraphyses
might be confused because both have shiny dark not seen, spores 50-57 µm long, 30-37 µm wide.
brown scales, and the latter grows up to 3500 m Figs. 3 a; 48.
elevation. But the leaves of C. solutum are
usually narrowly lanceolate and the stem is This species is known from Costa Rica to
pruinose with well-spaced leaves. Peru, where it grows mostly as an epiphyte
Campyloneurum solutum belongs to the C. between 200 m and 1800 m elevation.
amphostenon group.
ALAJUELA: Reserva Río San Lorenzo, below Fila
41. Campyloneurum sphenodes (Kunze ex Klotzsch) Volcán Muerte, 14-17 Jul. 1983, Barringer & Pérez-
García 3782 (F); La Palma de San Ramón, 24 Nov.
Fée, Gen. Filic. 258. 1852.
1923, Brenes 3968 (F), cataratas de San Ramón, 23 Feb.
Polypodium sphenodes Kunze ex Klotzsch, Linnaea 1931, Brenes 13472 (F); los Angeles de San Ramón, 21
20: 402. 1847. Type. Venezuela, Moritz 304 Jul. 1932, Brenes 16138 (F); upper drainage of the Río
(holotype B!; isotypes, BM!, K!). Peñas Blancas below the Monteverde Nature Reserve,
Polypodium wercklei Christ, Bull. Herb. Boissier 2. 25-26 Feb. 1977, Burger et al. 10725 (AAU, F);
5: 7. 1905. Lectotype (chosen by Lellinger, Cordillera de Tilarán, between San Ramón and Bajo
Amer. Fern J. 78: 28. 1988). Costa Rica: Wercklé Rodriguez, 26 Sep. 1987, Croat 68038 (MO); Río
s.n. (P, not seen). Sarapiquí at bridge on road to Colonia Virgen del
Campyloneurum wercklei (Christ) Lellinger, Amer. Socorro, 1 Oct. 1987, Croat 68337 (MO); Cordillera de
Tilarán, 5 May 1976, Dryer 93 (F); Reserva
Fern J. 78: 28. 1988.
Monteverde, Cordillera de Tilarán, 1 Jun. 1976, Dryer
Stem long-creeping, green turning black or 113 (F, MO); 7 mi N of San Ramón, 27 Jul. 1967, Evans
dark stramineous, (1-) 2-3 (-4) mm wide. Stem & Bowers 2972 (MO); Monteverde Reserve, Peñas
scales light brown or brown in mass, sub- Blancas, 13 Jun. 1986, Haber et al. 5143 (MO);
adpressed, 2-3 mm long, 0.8-1 mm wide, Monteverde Reserve, on the Atlantic slope, 14 Jun.
narrowly ovate, bases peltate or slightly 1986, Hammel et al. 15409 (MO); Reserva Forestal de
auriculate, apices acuminate, clathrate or rarely San Ramón, Quebrada Cacical, 2 May 1987, Herrera et
slightly clathrate, concolorous or bicolorous, al. 595 (MO, UC); NW of Zarcero, ca. 2 km of Zapote,
on road to Santa Elena, 27 Jul. 1970, Lellinger & White
margins differentiated at the bases or along the
1357 (MO, US); 25 km NNW of San Ramón, on way to
scale, the cells narrowly oblong, some times San Lorenzo, 24 Apr 1983, Liesner & Judziewicz 14765
irregularly arranged, cell lumina transparent, (MO), Liesner & Judziewicz 14829 (MO); Vara Blanca de
cell walls 10 µm thick. Phyllopodia 1-2 mm Sarapiquí, Jul-Sep. 1937, Skutch 3139 (F, MO); la Peña
long, 1-2 mm wide, 7-25 mm apart. Leaves 25-50 de Zarcero, Cantón Alfaro Ruiz, 11 May 1938, A. Smith
(-65) cm long; petiole stramineous or dark 570 (F); ca. 1 km SE of La Balsa de San Ramón, 3 Feb.
stramineous, 5-20 cm long, ranurate adaxially, 1986, A.R. Smith et al. 2295 (UC); Río San Rafael,
convex abaxially; lamina narrowly elliptic or cantón Aguas Zarcas, 8 Feb. 1965, L.O. Williams et al.
29062 (F). HEREDIA: NW slope of Volcán Barba, 2
lanceolate, bases narrowly cuneate, short
km NE of los Cartagos, 16 Mar. 1986, Grayum &
attenuate, apices acuminate or subcaudate, (2-)
Yatskievych 6638 (MO, UC); La Selva, along Río Viejo,
4-6 (-9) cm wide, herbaceous-chartaceous, 1 Feb. 1988, Hennipman et al. 7152 (MO); near the Río
margin cartilaginous, slightly undulate, Segundo, 2 km SW of Cerro Chompipe, 12 Jul. 1970,
indument of scarce bicellular hairs; stomata Lellinger & White 1110 (F); base of the Cerro Zurquí, 8
polocytic or copolocytic; costa prominent, Jul. 1973, Lent 3567 (F); road between San Rafael and
primary veins prominulous on both sides of the Río Las Vueltas, 4 Sep. 1979, Stevens 13972 (F, MO).
lamina, stramineous or darker than the leaf ALAJUELA-HEREDIA: Vara Blanca, N slope of
Central Cordillera, Poas-Barba, 12 Jul. 1940, Chrysler
tissue, (60o-) 65-70o divergent from the costa, 5-7 5048 (MO); Vara Blanca de Sarapiquí, N slope of
mm apart, secondary veins forming 7-12 Central cordillera, Jul.-Sep. 1937, Skutch 3139 (F, MO,
primary areoles between the costa and margin, NY, S). LIMON: path beyond Río Sucio, May 1984,
Gómez et al. 22753 (MO). PUNTARENAS: W side of Continental divide, 6 Feb. 1984, Churchill et al. 4661
Fila Gamba, ca. 6 km from Golfito airport, 6 Mar. 1985, (MO, UC); Fortuna dam area to N of reservoir near
Croat & Grayum 59921 (MO); Monteverde reserve, Quebrada Bonito, 30 Jul. 1984, Churchill 5799 (MO); El
along Río Peñas Blancas, 14 Jun. 1986, Hammel et al. Boquete, Dexter trail, 7 Feb. 1918, Cornman 865 (MO);
15409 (MO); Monteverde Reserve, Pacific slope, 5 vic. of Planes de Hornito, beyond Gualaca, 28 Nov.
Nov. 1986, Haber & Bello 6199 (MO); Monteverde, 19 1979, Croat 48858 (MO); along road between Fortuna
Jun. 1979, Koptur SK-150 (UC). SAN JOSE: valley of lake and Chiriquí Grande, 8 Mar. 1985, Croat &
the Río Hondura, below La Palma, NE of San Grayum 59977 (AAU, MO, UC); Boquete district, Bajo
Jerónimo, 15 May 1968, Burger & Stolze 4867 (F); La Chorro, 11 Jan. 1938, Davidson 106 (F); Cerro
Palma area, NE of San Jerónimo, above La Hondura, Horqueta, 1 Jul. 1968, Dwyer & Lallathin 8763 (MO); La
15 Sep. 1978, Burger & Antonio 11063 (F); Virgen del Fortuna hydroelectric project, 20 Mar. 1978, Hammel
Socorro-Río Sarapiquí-Cariblanco, 31 Aug. 1983, 2035 (MO); Palo Alto, 4.5 mi NE of Boquete, along E
Chacón & Herrera 1224 (MO); on western ascent of fork of Palo Alto river, 26 May 1979, Hammel 7510
Cerro de La Muerte, 27 Feb. 1976, Croat 32849 (MO); 1 (MO); S slopes of Cerro Pate Macho, along Río Palo
mi beyond divide between San Isidro del General and Alto, 11 Nov. 1981, Knapp et al. 2028 (MO); near
Dominical, 22 May 1976, Croat 35277 (MO); 10 km N Fortuna dam, along Quebrada de Arena, 6 Dec. 1985,
of San Rafael de Heredia, on Volcán Barba, 30 Jul. McPherson 7802 (MO); N of San Felix at Chiriquí-Bocas
1967, Lellinger 791 (MO); ca. 15 km N of Tres Ríos, ca. 4 del Toro border, 4 May 1975, Mori & Kallunki 5860
km N of Cascajal, 2 Aug. 1970, Lellinger & White 1393 (NY, US); Cerro Colorado, 50 km N of San Felix, 17
(US); above Río Cascajal, 3 km NE of Cascajal, 26 Sep. Aug. 1975, Mori & Dressler 7808 (MO); along trail
1971, Lent 2176 (F); above Río Hondura, 10 Mar. 1973, between N fork of Río Palo Alto and Cerro Pate
Lent 3229 (F); Bajo La Hondura area, 4 Jul. 1972, Macho, 6 Feb. 1986, A.R. Smith 2311 (MO, UC); along
McAlpin et al. 1186 (F); Pan Am hwy. above San Isidro Río Caldera, ca. 3.5 km NW of Bajo Mono, 8 Feb. 1986,
de General, 17 Nov. 1973, McAlpin 2383 (MO); Parque A.R. Smith 2460 et al. (MO, UC); SE slopes and summit
Nacional Bravillo Carrillo, 19 Jul. 1983, Moran 3281 (F); of Cerro Pate Macho, 4 km NE of Boquete, 26 May
along the road to La Hondura, 8 Apr. 1956, Scamman 1981, Sytsma et al. 4886 (MO), 27 May 1981, Sytsma et
& Holdridge 8074 (GH); vic. of El General, Aug. 1936, al. 4998 (MO). VERAGUAS: NW of Santa Fé, 20 Dec.
Skutch 2840 (MO, NY, S); along unnamed N fork of 1974, Mori et al. 3958 (MO). BOCAS DEL TORO: 10-15
Río Zurquí, Cordillera Central, 18 Jan. 1986, A.R. Smith mi S from mouth of Changuinola river, 18 Dec. 1966,
1696 (MO); above La Hondura valley, La Palma area, Lewis et al. 994 (UC). COCLE: La Mesa region, N of
23 Mar. 1973, Stolze 1434 (F, UC), Stolze 1446 (MO); S Cerro Gaital, 2 Jul. 1978, Hammel 3862 (MO). DARIEN:
slopes of Cerro Zurquí, 5 km N of San Luis Norte, 28 S of El Real on trail up Cerro Pirre, 29 Mar. 1985,
Mar.-4Apr. 1973, Stolze 1561 (UC); Alto de La Palma, McPherson 7036 (MO, UC); on ridge of cerro Pirre, 14
ca. 5 km N of San Jerónimo, 18 Aug. 1975, Utley & Sep. 1989, McPherson 14082 (F).
Utley 2900 (F). CARTAGO: Carpintera, 10 Apr. 1908, COLOMBIA. CAUCA: near Cerro Munchique, 5 Nov.
Brade & Brade 45 (NY, S, UC); 10 km S of Tapantí, 1968, Espinal & Ramos 3201 (MO). CHOCO: Mun. Río
above the Río Grande de Orosí, 10-24 Jun. 1968, Burger Sucio, Alto Limón, 4 Jun. 1978, Forero et al. 1812 (MO).
& Stolze 5633 (F, MO); Tapantí, near banks of Río Without locality: Triana 99 (W).
Grande de Orosí, 24 Jun. 1968, Burger & Stolze 6089 (F); VENEZUELA. FALCON: Sierra San Luis, near Hotel
along tributary of Quebrada Casa Blanca Tapantí, 6 Parador, 25 Aug. 1978, Wingfield & Werff 6565 (MO).
Aug. 1984, Grayum & Sleeper 3680 (MO, UC); Tapantí, TRUJILLO: near Vitú, Cerro El Zamuro, Quebrada El
valley of Río Reventazón, 18 Mar. 1956, Scamman & Limón, 23 Nov. 1984, Ortega & Werff 2292 (MO, UC).
Holdridge 8073 (GH); above the Río Grande de Orosí, LARA: dist. Morán (Andrés Eloy Blanco), 4 mi SE of
25 Mar. 1973, Stolze 1479 (F); Cerro Carpintera, 7 mi W Sanaré, Parque Nacional Yacambú, 13 Nov. 1982, A.R.
of Cartago, 7 Mar. 1928, Stork 1186 (UC, US); Naranjo, Smith et al. 1195 (MO).
S of Cartago, 4 May 1928, Stork 1822 (UC). ECUADOR. CARCHI: Maldonado, km 60 on road
CARTAGO-SAN JOSE: NW of Cartago, 3 Apr. 1928, Tulcán-Maldonado, 18 May 1973, Holm-Nielsen et al.
Stork 1360 (UC); Cerro Carpintera, 23 May 1928, Stork 5782 (AAU, F, MO, UC); above San Marcos de los
2135 (UC). Coaiqueres, on trail towards Gualpí Bajo, 7 Feb. 1985,
PANAMA. CHIRIQUI: Cerro Colorado, near Øllgaard et al. 57533 (AAU). PICHINCHA: road El
Continental divide, 26 Jul. 1979, Antonio 1463 (MO); Paraíso-Saguangual, 3 km from El Paraíso, 2 May
road to Fortuna dam, N of Gualaca, 22 Nov. 1979, 1982, Øllgaard et al. 37798 (AAU); along road SE from
Antonio 2766 (MO); road between Gualaca and the La Aurora, passing through La Reforma, 24 Jul. 1990,
Fortune, NW of los Planes de Hornito, 10 Apr. 1980, Øllgaard 98060 (AAU). BOLIVAR: road Chillanes-
Antonio 4171 (MO); Cerro Horqueta, 24 Jul. 1966, Blum Bucay, 29 Aug. 1987, Zak & Jaramillo 2571 (F, MO).
& Dwyer 2615 (MO); Fortuna dam area, at the CHIMBORAZO: Sibambe, Hacienda La Carmela, 18
Aug. 1943, Acosta-Solís 5389 (F); valley of the Río light brown in mass, persistent, 2-4.5 mm long,
Chanchan, about 5 km N of Huigra, 19-28 May 1945, 1-1.5 mm wide, ovate, pseudpeltate, bases
Camp E-3381 (F, MO, UC). AZUAY: between Cruz auriculate, apices acuminate, slightly clathrate,
Pampa and Loma de Canela, in region of the Río
the cells oblong, lumen yellow, cell walls 5-9 µm
Sadacray, 12 Jun. 1943, Steyermark 52959 (F, US).
SANTIAGO-ZAMORA: between Río Sordo and La
thick. Phyllopodia 0.5 mm long, 1 mm wide, 10-
Esperanza, road to Huamboya, 13 Feb. 1944, Acosta- 25 mm apart. Leaves 15-25 cm long; petiole
Solís 7318 (F). ZAMORA-CHINCHIPE: río stramineous, (2.5-) 8-10 cm long; lamina elliptic-
Nangaritza, Colina Salada, c. 2 km of Destacamento lanceolate, base cuneate, apices caudate, 2-5 cm
Shaime, 8 Dec. 1990, Øllgaard 98482 (AAU). wide, chartaceous or subcoriaceous, indument of
PERU. PIURA: Canchaque, near Chorro Blanco, 4 scattered simple hairs and deciduous scales;
Nov. 1985, Ramirez & Lamas s.n. (USM). stomata polocytic, 49-61 µm long, 67 µm wide;
CAJAMARCA: Santa Cruz, dist. Catache, upper Río costa prominent, primary veins inconspicuous or
Zaña, ca. 5 km above Monte Seco, 2-4 May 1987, Dillon
et al. 4901 (F). AMAZONAS: Bagua, Cordillera Colán, slight prominulous adaxially, 50-55o divergent
SE of La Peca, 7 Oct. 1978, Barbour 3893 (MO). from the costa, 5-7 mm apart, secondary
HUANUCO: La Divisoria, near Ucayali border, 29 transverse veins forming 5-6 areoles between the
Mar. 1977, Gentry et al. 18821 (MO). PASCO: Pozuzo, costa and margin, primary areoles usually
20-22 Jun. 1923, Macbride 4581 (F, US); Oxapampa, 4-5 undivided,
km N of Mallampampa, 22 Jan. 1984, D.N. Smith & (1-) 2 free excurrent veinlet in each areole,
Canne 5802 (MO). JUNIN: E of Quimirí bridge, near
sometimes marginal areoles without excurrent
La Merced, 1-3 Jun. 1929, Killip & Smith 23896 (F, GH,
US); Colonia Perené, 14-22 Jun. 1929, Killip & Smith veinlets; sori subterminal, paraphyses not seen;
24917 (F, NY, US). AYACUCHO: Aina, between spores 55-60 µm long, 30-36 µm wide. Fig. 49.
Huanta and Río Apurímac, 7-17 May 1929, Killip &
Smith 22720 (NY, US). This species is known from Costa Rica and
Ecuador, where it grows pendent in calacareous
Campyloneurum sphenodes is recognized here rocks, in low montane or lowland forests, in very
in a broad sense. It is defined by its stem scales shady places at 700-1500 m elevation.
with differentiated margins along the length of
the scale or at its bases. Stem scales are EXAMINED SPECIMENS: COSTA RICA. SAN JOSE:
concolorous or bicolorous, depending on the above the Río La Hondura, 28 Jan. 1979, Montgomery
development of the margin. Stem color is & Huttleston 79-102 (US). CARTAGO: 12 km S of
Turrialba, 4 km SE of Pejibaye, along Río Gato, 16-17
variable in herbarium material and in this study
Apr. 1983, Liesner 14426 (MO, UC); vic. of Pejivalle, 7-8
therefore this character is not considered to be Feb. 1926, Standley & Valerio 47108 (US). HEREDIA:
taxonomically important. road between San Rafael and Río Vueltas, 4 Sep. 1979,
Campyloneurum sphenodes might be confused Stevens 13972 (AAU, F).
with C. coarctatum. But stem scales in C. ECUADOR. MORONA-SANTIAGO: km 35 of road
sphenodes are adpressed, lanceolate or narrowly Gualaquiza-Limón, 4 Mar. 1992, Øllgaard 98 (AAU, F,
lanceolate, and usually more than 0.8 mm wide, QCA).
with differentiated margins. Campyloneurum Campyloneurum sublucidum has yellowish,
sphenodes is closely related to C. sublucidum, from persistent stem scales, and its leaves are
which can be differen-tiated by the characters chartaceous subcoriaceous with a glossy surface.
used in the key. This species belongs to the C. sphenodes group.
42. Campyloneurum sublucidum (Christ) Ching, 43. Campyloneurum tenuipes Maxon, Contr. U.S.
Sunyatsenia 5: 263. 1940. Natl. Herb. 13: 7. 1909. Type. Guatemala:
Polypodium sublucidum Christ, Bull. Herb. Alta Verapaz, near Cobán, Sep. 1907, von
Boissier sér. 2, 7: 261. 1907. Type. Costa Rica, Türckheim II 1952 (holotype, US, photo of US,
Wercklé 17051 (holotype, P!, isotype BM!). BM!).
Stem long creeping, stramineous, dark Polypodium tenuipes (Maxon) C. Christ., Index
brown, not pruinose, 2-3 mm wide. Stem scales Filic. Suppl. 63. 1913.
Stem dark stramineous or black, not Mexicanos, at the E base of Cerro Tres Picos, 4 May
pruinose, 4-7 mm wide. Stem scales dark brown 1972, Breedlove 25053 (MO); Mun. Ocozocautla de
or black in mass, 6-10 mm long, 2-2.5 mm wide, Espinosa, Cerro del Ocote, 30 km NW of Ocozocautla,
14 Oct. 1972, Breedlove 28894 (F); Mun. San Andrés
narrowly ovate, pseudopeltate, bases auriculate,
Larrinzar, near the summit of Chuchil Ton, NE of
apices acuminate, clathrate, the cells oblong, Bochil, 1 May 1973, Breedlove 34590 (MO); between
along the main axes of the scale, cell walls (5-) Rizo de Oro and Cerro Baul, 9.2-10.4 mi N of Rizo de
12.5-17.5 µm thick. Phyllopodia 1-2 mm long, Oro, 15 Feb. 1979, Croat 47627 (MO); 1864-1870,
1.5-2.5 mm wide, 2-4 mm apart. Leaves 40-80 cm Ghiesbreght 292 (BM); Motozintla, Mt. Boquerón, 1
long; petiole brown stramineous, 8-18 cm long; May 1945, Matuda 15343 (F); San Juan de Panamá,
lamina narrowly lanceolate or lanceolate, bases Escuintla, 28 Jul. 1948, Matuda 18051 (F, US); Libertad,
attenuate or narrowly cuneate, apices usually Acocayagua, 3 Jul. 1948, Matuda 18145 (F); Simojovel,
caudate, 5-8 cm wide, chartaceous, margins Tierra Caliente, 1900, Munch 9 (US); Mun. Unión
Juarez, Volcán Tacaná, near trail to Talquián, 4 Feb.
slightly cartilaginous, repand or sinuate,
1987, Martínez et al. 19148 (F).
indument of scarce bicellular hairs; stomata GUATEMALA. ALTA VERAPAZ: 14 mi E of Cobán,
polocytic and/or copolocytic, rarely anomocytic; 18 Jul. 1977, Croat 41467 (MO); along Río Carchá,
costa prominent, slightly ranurate adaxially, between Cobán and San Pedro Carchá, 26, 27 Mar.
angular abaxially, primary veins prominent or 1941, Standley 90037 (F); along Río Frío, about 8 km
prominulous on both surfaces of the lamina, below Tactic, 1 Apr. 1941, Standley 90834 (F); 1-8 km
NW of Cobán, 4 Jan. 1973, L.O. Williams et al. 42001
stramineous, slightly flexuosous, (55o-) 60-70o
(AAU, F). SAN MARCOS: 10 mi S of San Marcos, 13
divergent from the costa, 5-6 mm apart, Jul. 1977, Croat 41011 (MO); 6 mi SW of town
secondary veins inconspicuous or slightly Tajumulco, NW slopes of Volcán Tajumulco, 26 Feb.
prominulous, same color as the leaf tissue, 1940, Steyermark 36670 (F); Sierra Madre Mountains,
transverse veins forming 6-9 primary areoles between San Rafael Pie de la Cuesta and Palo Gordo,
between the costa and margin, primary non- 10-18 Dec. 1963, L.O. Williams et al. 25807 (F, NY).
costal areoles usually symmetrically divided, 1 QUEZALTENANGO: slopes of Volcán Zunil, at and
free excurrent veinlet in each secondary areole, above Aguas Amargas, 17 Feb. 1939, Standley 65424
(F); Aguas Amargas 14 Jan. 1941, Standley 83288I(nr;
marginal areoles sometimes irregularly divided;
region of Boxantin, SE of San Martin Chile Verde, 16
sori subterminal or supramedial, paraphyses not Jan. 1941, Standley 83834 (F, US), along Río Samalá,
seen, spores 60-70 µm long, near Santa María de Jesús, 25 Jan. 1941, Standley 84586
35-40 µm wide. Chromosome number 2n=74. (UC); along old road betwen Finca Pirineos and
Fig. 24. Palzulin, 9 Feb. 1941, Standley 87148 (F); on SE slopes
of Volcán Santa María, 18 Jan. 1940, Steyermark 34372
This species is distributed from Mexico to (F). SOLOLÁ: Sierra Madre Mountains, near Nahuala,
Guatemala and Honduras, where it grows 17 Dec. 1962, L.O. Williams et al. 23211 (F, US).
HONDURAS. FRANCISCO MORAZAN: Montaña La
mostly as terrestrial between 1200 m and 2400 m
Tigra, 30 km NE of Tegucigalpa, 5 Jun. 1977, Alduvía et
elevation.
al. 266 (MO); Montaña La Tigra, 17 Apr. 1983, Guerra
152 (MO); 20 km fromTegucigalpa, 5 Jun. 1977, Ochoa
REPRESENTATIVE SPECIMENS: MEXICO. 61 (MO); 20 km from Tegucigalpa, 5 Jun. 1977, Rubio
MICHOACAN: Galena, Río de las Selvas, 3 Jan. 1938, 94 (MO); Valle de Angeles, 20.8 km NE of
Hinton et al. 11177 (US). VERACRUZ: Alto Lucero, La Tegucigalpa, Pinares, Soihet 66 (UC).
Piedra Cuata, between Plan de las Hayas and Rancho
Nuevo, 7 Apr. 1981, Castillo & Vásquez 1354 (F);
Campyloneurum tenuipes is characterized by
Teocillo Cañón, just before Teocillo, 11 Aug. 1966,
Knobloch 2195 (US). OAXACA: above San Gabriel, conspicuously petiolated leaves and linear
Mickel s.n. (UC); Pochutla, 185 km S of Oaxaca, 60 km lanceolate stem scales. It belongs to the
N of Pochutla, 29 Sep. 1970, Mickel & Leonard 5097 C.phyllitidis species group.
(UC); dist. Juquila, 96 km S of Sola de Vega, 33 km N
of San Gabriel, 9 Aug. 1971, Mickel 6050 (UC).
CHIAPAS: Mun. Berriozabal, 13 km N of Berriozabal, 44. Campyloneurum tucumanense (Hieron.) Ching,
2 Nov. 1971, Breedlove & A.R. Smith 21686 (F, NY); Sunyatsenia 5: 263. 1940.
Mun. Villa Corzo, SW of Colonia Agrónomos
Polypodium tucumanense Hieron., Bot. Jahrb. Syst.
22: 405. 1896. Type. Argentina, Tucumán, pascoense. Future cytological studies could reveal
Quebrada Monteros, 5 Apr. 1872, Lorentz 304 their affinities. The main difference to recognize
(holotype, B!; isotype. CORD, n.v.) C. tucumanense as a different species from C.
Terrestrial or sometimes epipetric. Stem dark pascoense is the remarkable herbaceous texture of
stramineous, not pruinose, 8-25 mm wide, scales the leaf in the former species.
brown, 5-6 mm long, (1.5-) 2-2.5 mm wide,
lanceolate, ovate lanceolate, bases auriculate,
apices acuminate, clathrate, cells oblong, 45. Campyloneurum vulpinum (Lindman) Ching,
marginal cells irregularly arranged, central cells Sunyatsenia 5: 263. 1940.
along main axes of the scale, cell walls 15 µm Polypodium vulpinum Lindman, Ark. Bot. 1: 245.
wide. Phyllopodia 5-6 mm long, 6 mm wide, 2-5 1903. Nom. nov. for Polypodium laevigatum
mm apart. Leaves (80-) 95-120 cm long; petiole Cav. var. crispatum C. Christ.
green stramineous, stramineous, 12-20 cm long, Polypodium laevigatum Cav. var. crispatum C.
ranurate adaxially, convex abaxially, with scales Christ., Bot. Tiddskr. 25: 79. 1903. Type.
at the base similar to those at the stem; laminae Brazil, Minas Geraes, Serra de Caldas, 25 Oct.
lanceolate, bases attenuate, apices subcaudate or 1873, Mosén 2220 (holotype, S!; isotypes BM!,
acuminate, US!).
7-9.5 (-14) cm wide, membranaceous or Epiphyte. Stem long creeping, not pruinose,
herbaceous, margins sinuate, cartilaginous, 1-2 mm wide, scales ferrugineous, slightly
indument of inconspicuous, scarce bicellular clathrate, 4-7 mm long, 0.6-1.2 mm wide, linear
hairs, stomata not seen; costa prominent, lanceolate, bases peltate, rarely slightly
primary veins prominent, stramineous, 65-70o auriculate, apices acuminate, cells oblong along
divergent from the costa, 5-8 mm apart, the main axes of the scale. Phyllopodia 0.5-1.5
secondary veins slightly prominulous, slightly mm long, 0.5-1 mm wide, 7-15 mm apart. Leaves
stramineous or darker than the leaf tissue, 15-45 cm long; petiole stramineous to dark
transverse veins forming 12 primary areoles stramineous, 4-12 cm long; laminae linear
between the costa and margin, primary areoles lanceolate or narrowly lanceolate, bases
usually asymmetrical divided in 2-3 secondary attenuate, apices acuminate, 1.5-3 cm wide,
areoles, 3-4 excurrent veinlets in each primary herbaceous, margins slightly cartilaginous,
areole, simple or furcate, ; sori medial or sinuate or undulate, indument formed by scarce
subterminal, 2-3 rows between secondary veins, bicellular hairs; stomata polocytic; costa
paraphyses not seen, spores 52 µm long, 35 µm prominent, primary veins inconspicuous or
wide. Chromosome number unknown. Fig. 44. slightly prominulous on both sides of the
lamina, same color as the leaf tissue, 30-40o
Campyloneurum tucumanense is known only divergent from the costa, 4-7 mm apart, primary
from Bolivia and Argentina. It grows between veins forming 2-4 primary areoles between the
1000 m and 1500 m elevation, usually as a costa and margin, primary areole usually
terrestrial in shady habitats. isodiametric divided; sori subterminal,
paraphyses dendritic, spores (50-) 60-70 (-90)
REPRESENTATIVE SPECIMENS: BOLIVIA. LA PAZ: µm long, (30-) 35-40 (-50) µm wide. Figs. 3d; 7 b;
Murillo, Valle de Zongo, Cahua, 7 Apr. 1979, Beck 1219 16 b; 49.
(F). ARGENTINA. TUCUMAN: Quebrada Montero, 5
Apr. 1872, Lorentz 780 (F); Tafí, Rodeo Aspero, 14 Apr.
This species is known from Haiti, and the
1926, Schreiter 4344 (GH); Quebrada de los Sosa, 2 Sep.
1957, Sota 1672 (S, US). MISIONES: Libertador General
Dominican Republic, and from Ecuador to
San Martín, Gruta 3 de Mayo, 9 Nov. 1974, Krapovickas Bolivia and central Brazil, where it grows above
& Cristóbal 26638 (MO). 1000 m elevation, mostly as an epiphyte.
This species belongs to the Campyloneurum REPRESENTATIVE SPECIMENS: HAITI. Massif de

brevifolium group. It is closely related to C. la Selle, Morne Trauchant, 13 Sep. 1924, Ekman 1892
(BM, F, S); Morne des Commissaires, Grand Gosier, at
ravine Fanchon, 4 Sep. 1926, Ekman 6885 (S); Massif de Vareschi, Flora de Venezuela 1, 2: 950. 1968.
la Holle, M. Columette, 26 Nov. 1926, Ekman 7324 (S); Type: Venezuela, Bolívar, Cerro Pijiguao,
Massif de la Selle, Croix des Bouquets, Badeau, 4 Mar. Sierra Suapure, Wurdack 41130. Nomen
1927, Ekman 7781a (F, S), Ekman 7781b (S); Morne des
nudum.
Commissiares, near Petite Source, 17 Apr. 1932,
Stem long creeping, not pruinose, 2-3 mm
Holdridge 1134 (US); Morne des Commissaires, Jul.
1942, Holdridge 1368 (GH, UC); vic. of Furcy, 26 May- wide. Stem scales light brown in mass, 3-4 mm
15 Jun. 1920, Leonard 4638 (BM), Leonard 4778 (F); long, 1-1.3 mm wide, lanceolate, bases
Ouest Départment, summit of Morne Guimby, above auriculate, apices obtuse or rarely acute, the cells
Morne des Commissaires, 16 Sep. 1955, Proctor 10812 oblong, cell walls 10 µm thick. Phyllopodia 1-1.5
(US). mm long, 1.5-2 mm wide, 5 mm apart. Leaves
DOMINICAN REPUBLIC. BARAHONA: Polo, 26 19-41 cm long; petiole dark stramineous or
Feb.-12 Mar. 1922, Abbott 1798 (S, US), Abbott 1805 stramineous, 4-8 cm long; lamina lanceolate, 4.5-
(BM, GH); Montiada Nueva, 21 Aug. 1946, Howard &
8 cm wide, bases cuneate then long decurrent,
Howard 8548 (US).
ECUADOR. TUNGURAHUA: Montaña Woma, 11 apices acuminate to slight caudate; costa
km E of Baños, 3 Jun 1968, Holm-Nielsen & Jeppesen 298 prominent, primary veins slight prominulous,
(AAU, GH). NAPO: 4 km NW of Borja, 20 Sep. 1980, darker than the leaf tissue, 60-65o divergent
Holm-Nielsen et al. 26365 (AAU); Baeza, 1 km S of the from the costa, 5-7 mm apart, secondary veins
village, 20 Oct. 1976, Øllgaard & Balslev 10211 (AAU, inconspicuous, forming 7-8 primary areoles
NY, UC, USM). MORONA-SANTIAGO: Pachicutza,
between the costa and margin, 3-4 excurrent
km 140 on road Loja-Gualaquiza, 26-27 Apr. 1973,
Holm-Nielsen et al. 4616 (AAU).
veinlets in each primary areole, simple or
PERU. CAJAMARCA: Santa Cruz, Catache, upper Río furcate, central veinlet generally anastomosed
Zaña valley ca. 5 km above Monteseco on path to with the transverse veins forming assymetric
Chorro Blanco, 16-18 Mar. 1986, Dillon et al. 4358 (F), secondary areoles; sori subterminal or medial,
Dillon et al. 4428 (F). AMAZONAS: Bagua, 12 km E of paraphyses and spores not seen. Figs. 5 a-c; 44.
La Peca, 23 Jun. 1978, Barbour 2487a (UC); 12 km E of
La Peca, 29 Jun. 1978, Barbour 2585 (F, UC). This species is known from Venezuela, where
HUANUCO: Muña, 23 May-4 Jun. 1923, Bryan 421 (F);
it has been found between 90 m and 500 m
Muña, 8 Mar. 1959, Woytkowski 5218 (GH); below Río
elevation.
Santo Domingo, Macbride 4208 (F). PASCO:
Oxapampa, S of Oxapampa, 1 Feb. 1983, León 506
(USM). JUNIN: Yucapata, 17 Jul. 1961, Woytkowski EXAMINED SPECIMEN: VENEZUELA.
6658 (US). AYACUCHO: Ccarrapa, between Huanta TERRITORIO FEDERAL AMAZONAS: Atures, 23 km
and río Apurímac, 5-17 May 1929, Killip & Smith 22401 NE of Puerto Ayacucho and 10 km E of the highway,
(US). 17-19 Apr. 1978, Davidse & Huber 15306 (MO).
BOLIVIA. COCHABAMBA: Espíritu Santo, NE von
Cochabamba, Jun. 1909, Buchtien 2165 (BM, US), Campyloneurum wurdackii belongs to the C.
Buchtien 2208 (S, US). Santa Bárbara, 30 Aug. 1902, phyllitidis group. It is characterized by the
R.S. Williams 1055 (US). closely spaced leaves, light brown and persistent
stem scales, and undivided primary areoles.
Campyloneurum vulpinum is easily
distinguished by its persistent ferrugineous stem
scales. 47. Campyloneurum xalapense Fée, Gen. Filic. 258.
1852. Type. Mexico, Veracruz, Xalapa, Jun.-
46. Campyloneurum wurdackii B. León, Ann. Oct. 1840, Galeotti 6273 (holotype, probably P ;
Missouri Bot. Gard. 77: 212-214. 1990. Type: isotype, K!).
Venezuela, Bolívar, Cerro Pijiguao, Sierra Campyloneurum caudatum Fée, Mém. Foug. 8: 96.
Suapure, E slopes of Cerro Pijiguao (N end of 1857. Type: Mexico, Cordoba et Huatusco,
Serranía Suapure), 19 Jan. 1956, Wurdack & 1853, Schaffner 176 (holotype, P; isotype K!).
Monachino 41303 (holotype, MO!; isotypes Polypodium xalapense (Fée) Christ, Bull. Soc. Roy.
US!). Bot. Belgique 35: 231. 1896.
Polypodium repens Aublet var. spathulatum Polypodium phyllitidis L. f. multipunctatum Christ,
Bull. Herb. Boissier 2, 5: 7. 1905. Type: Costa Siera Madre Oriental, 4 km NE of Villa Juárez, 27 Jun.
Rica, Navarro, 1903, Wercklé 174 (holotype, P!, 1969, Marcks & Marcks 820 (BM, UC). VERACRUZ:
isotype US, photo of US, BM!). near Jalapa, Hacienda Concepción, 11 Sep. 1906,
Barnes et al. 87 (F); Cerro San Martín, Calzada 426 (GH);
Polypodium multipunctatum (Christ) Christ, Bull.
near Fortín above plant Cervecería Moctezuma, 26
Herb. Boissier 2: 51-52. 1906. Jun. 1977, Croat 39386 (MO), Croat 39413 (MO); El
Polypodium weatherbyanum Seymour, Phytologia Mirador, 21 km E of Huatusco, 23 Aug. 1977, Croat
31: 171. 1975. Nomen novum for 44010 (MO, UC); along highway 125 to Huatusco, 15
Campyloneurum caudatum. Jan. 1987, Croat & Hannon 63103 (MO); Salto del Gato,
Campyloneurum multipunctatum (Christ) along Río Sedeño, about 3 km NE of Xalapa, 31 Dec.
Lellinger, Proc. Biol. Soc. Wash. 89: 708. 1977. 1973, Dorante et al. 780 (GH); Cordoba, Finck 55 (UC),
Stem long creeping, dark stramineous, not Fink 84 (MO); La Luz, Cordoba, 13 Oct. 1882, Kerber
pruinose, (2-) 4-5 (-10) mm wide. Stem scales (3- s.n. (BM); 7.2 km E of Tebanca, 7.2 km E of E side of
Lago Catamaco, 2.6 km W of Bastonal lumber camp,
) 4-6 mm long, 0.75-2.5 mm wide, narrowly
15 Jan. 1981, Nee & Schatz 19947 (F); Cordoba, 6 Apr.
ovate or ovate, bases auriculate, apices 1910, Orcutt 3212 (BM, MO); near Orizaba, 29 Jan.
acuminate, sometimes obtuse, cell walls 12-24 1895, Pringle 6082 (BM, GH, MO, NY, S, UC);
µm thick, basal margins of the scale with hair- Zacuapán, Nov, 1906, Purpus 2163 (F, MO, NY);
like teeth 48-56 µm long. Phyllopodia 1-2 mm Zacuapán, Nov. 1906, Purpus 2163 (F); Barranca de
long, 2-2.5 mm wide, 2-7 mm apart. Leaves 30- Tenampa, Apr. 1934, Purpus 16479 (F); Zacuapán, Jan.
85 cm long; petiole stramineous or dark 1908, Purpus s.n. (UC); Copatepec, Dec. 1943, Sánchez
stramineous, 2-21 cm long, indument of scales 10 (US); Barranca de la Concepción, near Jalapa, 24
Dec. 1984, C. Smith 2008 (F, GH); Mun. Perote, deroute
similar to those on the stem; lamina narrowly
to Magueyitos, 25 Aug. 1975, Vásquez 2131 (UC); vic.
lanceolate or narrowly oblong, bases attenuate, of El Ejido de Tepetlampa, El Palmar, Zongolica, 6
apices acuminate, long acuminate or subcaudate, Jun. 1944, Vera 3010 (US). OAXACA: along road
1.7-5 cm wide, chartaceous or subcoriaceous, between Teotitlán to Chichotla, 23 Feb. 1979, Croat
margins cartilaginous, undulate, indument of 48412 (MO); dist. Villa Alta, 25 Jul. 1962, Mickel 964
scarce, simple, bicellular hairs, 80-96 µm long; (NY, UC, US), 27 Jul. 1962, Mickel 1015 (NY, US).
stomata polocytic or copolocytic; costa CHIAPAS: Mun. Motozintla de Mendoza, 45-50 km
prominent on both sides of the lamina, plane or NE of Huixtla, along road to Motozintla, 17 Nov. 1971,
Breedlove & Smith 22658 (F, NY); Selva Negra, 10 km
slight ranurate adaxially, angled abaxially,
above Rayón Mezcalapa, Breedlove 26079 (MO); Mun.
primary veins slight prominulous adaxially,
Cintalapa, 3 km E of Francisco Madero, NE of
inconspicuous abaxially, same color as the leaf Cintalapa, 4 Oct. 1974, Breedlove 38039 (MO); above El
tissue or prominulous and stramineous at its Rosario, 8 mi S of Motozintla, 10 Jul. 1977, Croat 40731
origin, (60o-) 65-70o (-75o) divergent from the (MO), Croat 40732 (MO); along road between
costa, 4-8 mm apart, transverse secondary veins Motozintla and Siltepec, 26-30 mi N of Motozintla, 12
Feb. 1979, Croat 47460 (MO); Mun. Las Margaritas, 12
forming 4-7 primary areoles between the costa
km E of Tziscao, 16 Nov. 1984, Davidse et al. 29866
and margin, primary areoles usually
(MO); Mun. Ocosingo, near Laguna Ocotal Grande, ca.
isodiametric divided, 2-4 excurrent veinlets in 25-30 km SE of Monte Líbano, 8 Aug. 1954, Dressler
each primary areole; sori subterminal or medial, 1618 (GH, NY, US); Ocozocautla, El Ocote, 14 Feb.
paraphyses not seen, spores 60-64 µm long, 35- 1986, Palacios-Ríos 2780 (UC). Without locality: Aug.
45 µm wide. Chromosome number 2n=74. Figs. 1855, Botteri 5 (BM); 1857, Muller s.n. (BM).
11; 14 f; 18 e; 50. BELIZE. TOLEDO: Edwards road beyond Columbia,
27 Apr. 1948, Gentle 6515 (F, S, US).
This species is distributed from Mexico to GUATEMALA. ALTA VERAPAZ: between San
Pedro Carcha and Campur, 20 Mar. 1970, Harmon &
Costa Rica. It grows between 1000 m and 2500
Fuentes 2181 (MO); Montaña Ixocuvain, W of
m elevation, mostly in open areas or disturbed Cubilguitz, 12 Mar. 1942, Steyermark 44975 (F). SAN
forests. MARCOS: Río Mopá, below Rodeo, 14 Mar. 1939,
Standley 68764 (F). QUEZALTENANGO: Aguas
REPRESENTATIVE SPECIMENS: MEXICO. Amargas, W slopes of Volcán Zunil, 14 Jan. 1941,
HIDALGO: Molango, between Calnali and Standley 83353 (F, UC); El Pocito, S of San Martín Chile
Huazalingo, 39 May 1947, Moore 3019 (UC). PUEBLA: Verde, on road to Colomba, 27 Jan. 1941, Standley
85012 (F, US), above Mujuliá, between San Martín road from Santa Cruz to Vista de Mar, 22-26 Jul. 1985,
Chile Verde and Colomba, 1 feb. 1941, Standley 85620 Gómez & Herrera 23667 (MO); Jul. 1857, Hoffman 894
(F). SOLOLA: Atitlán, 16 Feb. 1906, Kellerman 5779 (S); 58 km from San José, Poas, Saiki 109 (F); basin of El
(US); Volcán Atitlán, 11 Jun. 1942, Steyermark 47378 General, Jul.-Aug. 1943, Skutch & Barrantes 5161 (MO).
(F). CHIMALTENANGO: Volcán Pacayca, 16 Feb. CARTAGO: 2 km N of Orosí, 3 Jul. 1967, Mickel 2283
1947, Brenckle 4737 (F); 8 km S of Acatenango, 2 Sep. (NY, US); Navarro valley, 6-8 mi SW of Cartago, 7
1972, Madison 672 (GH); region of Los Ositos, above Apr. 1928, Stork 1402 (NY, UC); 3 km SE of Cartago, 10
Las Calderas, 16 Dec. 1940, Standley 80182 (UC). Aug. 1967, Taylor 4259 (MO).
JALAPA: Volcán Jumay, N of Jalapa, 1 Dec. 1939,
Steyermark 32450 (F). ZACAPA: Sierra de las Minas, Campyloneurum xalapense is characterized by
between Cerro de Monosand Monte Virgen, 17 Jan. narrowly lanceolate or oblong lanceolate leaves
1942, Steyermark 42848 (F). SUCHITEPEQUEZ: S
and by 4-7 primary areoles between margin and
slopes of Volcán Zunil, vic. Finca Las Nubes, along
Quebrada Chita, E of Pueblo Nuevo, 2 Feb. 1940,
costa.
Steyermark 35433 (F). CHIMALTENANGO: 8 km S of It is recognized here in a broad sense,
Acatenango, 2 Sep. 1972, Madison 672 (GH); region of including populations with a wide range of
Los Positos, above Las Calderas, 16 Dec. 1940, Standley morphological variation in size and shape of
80182 (F, UC). ESCUINTLA: between Río Jute and Río stem scales from linear lanceolate scales with
Pantaleón, on road between Escuintla and Santa bases less than 1 mm wide to lanceolate scales
Lucía, 24 Jan. 1939, Standley 63496 (F). SANTA ROSA: with bases more than 2 mm wide. All this range
near El Molino, 26 Nov. 1940, Standley 78511 (F). of variation can be found within an individual
HONDURAS. COMAYAGUA: Cerro Azul-Meambar,
specimen (for example Croat 40732). Moreover,
8 Aug. 1974, Horwath 81 (F). CORTES: N side of Lake
Yojoa, 10 Apr. 1951, Morton 7643 (US). SANTA at any point of its distributional range, a similar
BARBARA: 10 km W de Lago Yojoa, 28-30 Apr. 1973, amount of variation can be found in stem scale
Clewell & Hazlett 3796 (MO, US). OCOTEPEQUE: size and shape. For these reasons stem scale
Cordillera Merendón, vic. of El Portillo, 2 Sep. 1975, characteristics do not have taxonomic value for
Molina 31007 (F). discerning different taxa within C. xalapense.
EL SALVADOR. SANTA ANA: Cerro Monte Cristo, Stem width, leaf texture, and pattern of
ca. 14 mi NE of Metapan, 31 Jul. 1977, Croat 42399 venation are also variable in most specimens
(MO, UC); Montecristo, 23 May 1963, Molina & Molina
examined. Some of that variation was used to
12658 (F, NY, US); Parque Nacional Montecristo, 24
Sep. 1988, Pfeiffer 48 (MO); Montecristo, 11 Oct. 1978,
distinguish Campyloneurum multipunctatum as
Seiler 670 (F, NY, UC); Laguna Las Ninfas, 8 Feb. 1979, different from C. xalapense (cf. Lellinger, 1988).
Seiler 907 (F, NY). CHALATENANGO: E slope of Los However, for the vast majority of specimens
Esesmiles, 14 Mar. 1942, Tucker 1047 (UC, US). SAN studied these character states are not associated
SALVADOR: Volcán San Salvador, 3 Feb. 1946, (e.g. Steyermark 44975, Arsene s.n, Standley 85012).
Carlson 505 (F, UC); Santa Tecla, Cantón Las Victorias,
1941, García 109 (UC). SAN VICENTE: Volcán San
Vicente, 7-8 Mar. 1922, Standley 21610 (MO); Volcán
NOMINA DUBIA
San Vicente, 26 Apr. 1978, Seiler 324 (F); Seiler 325 (F).
1. Polypodium calaguala Ruiz, Mem. c tab. 1805.
COSTA RICA. GUANACASTE: upper slopes of
Cerro San José de Líbano, 15 Feb. 1930, Dodge et al. Moore (1861), included this name as a
6461 (US); along Río San Juan, W slopes of Volcán synonym of Campyloneurum angustifolium.
Tenorio, 25 Jan. 1985, Grayum et al. 4963 (MO); Rincón Attempts to localize the type specimen were
de la Vieja National Park, ridge SE of Quebrada unsuccessful. For this reason it is not attributed
Zopilote, 24 Jan. 1986, A.R. Smith et al. 1927 (UC), 26 here to any particular species.
Jan. 1986, A.R. Smith et al. 1982 (UC). ALAJUELA: San
Isidro de San Ramón, 21 Oct. 1986, Herrera 73 (MO). 2. Polypodium gladiatum Vellozo, Fl. Flum. 11,
HEREDIA: Barba, at Volcán Barba, 18 Apr. 1953,
t.59. 1827.
Scamman 7247 (US). PUNTARENAS: Las Cruces
Tropical Botanical Garden, 6 mi S of San Vito de Java, The illustration appears to be that of
Aug. 1974, Herb. Tropical Studies 885 (US). SAN JOSE: Campyloneurum nitidum. However, I have been
Tablazo, 17 Sep. 1908, Brade & Brade 243 (BM, NY, S); unable to locate the type specimen.
La Palma, 1909, Brade & Brade 243a (S); Llanuras de
San Carlos, Apr.-May 1910, Brade & Brade 710 (S, UC); 3. Campyloneurum lanciforme (J. S. Presl) C. Presl,
Tent. Pterid. 190.1836. as a synonym of Campyloneurum falcoideum.

Polypodium lanciforme J.S. Presl, Deliciae Although the figures in the original publication
Pragensis 164. 1822. Type: Brazil, Rio de clearly show a specimen of Campyloneurum, here
Janeiro, Sellow s.n. they are not attributed to any particular species
According to the description, this name refers since leaf morphology, especially among
to a Brazilian species with narrowly lanceolate narrowly lanceolate does not help to distinguish
leaves. The description applies either to species. Attempts to localize the type specimen
Campyloneurum minus or C. nitidum. were unsuccessful; according to Pablo Sánchez,
curator of CR (pers. comm.) no Gómez types are
4. Campyloneurum loreum (Kaulfuss ex Kunze) kept there.
Fée, Mém Foug. 8. 129. 1857.
Polypodium loreum Kaulfuss ex Kunze, Flora 1839. 9. Campyloneurum sieberianum C. Presl, Tent.
There were no seen types or descriptions, and Pterid. 190. 1836. Tab. 7. Fig. 17.
therefore it is not included in any species. According to the pattern of venation shown
in Presl (1836) this could be a synonym for a
5. Polypodium medicinale Rojas, Bull. Acad. Int. species within the Campyloneurum brevifolium
Géogr. Bot. 28: 156. 1918. group.
According to Morton (1973), the type
specimen is deposited at the herbarium in 10. Polypodium schnittspahnii Christ, Bull. Herb.
Asunción, Paraguay, and "by the process of Boissier 6: 836. 1898. Type: Andes, ?Moritz
elimination" was determined to be a synonym s.n..
for P. phyllitidis. However, since C. phyllitidis In the original description, the stem scales are
does not occur in Paraguay, it is probable that described as "ovato-lanceolatis" and atrofuscous
this name is a synonym for Campyloneurum on a pruinose glaucous stem (farina glauca).
nitidum. These characters, together with the pattern of
venation, may apply to the Campyloneurum
6. Campyloneurum moritzianum Fée, Gen. Fil. 258. angustifolium group. Lellinger (1988) suggested
1852. Type. Venezuela, Caracas, Moritz 3. that this name might be an earlier epithet for C.
Non Polypodium Link. falcoideum. However, based on the available
I have not seen the type specimen, which is evidence it is not included in any species.
deposited at the herbarium of Rio de Janeiro
(Windisch, 1982), but attempts to get a loan were
not successful. T. Moore (1861) included this
name as a synonym of C. phyllitidis. However, EXCLUDED TAXA
after reading the description, I can only be sure 1. Campyloneurum laevigatum (Cav.) C. Presl,
that it belongs to the group of C. phyllitidis. Tent. Pterid. 190. 1836. =Polypodium
laevigatum Cav. Descr. Pl. 244. 1802.
7. Campyloneurum polyanthum C. Presl, Tent.
Pterid. 190. 1836. Tab. 7. Fig 16. 2. Campyloneurum decumanum (Willd.) Fée, Crypt.
Presl mentioned Polypodium polyanthum as the Vasc. Brésil 1. 115. 1869. =Polypodium
basionym of this name. According to the pattern decumanum Willd., Sp. Pl. 5: 170. 1810.
of venation shown in Presl (1836), this name
could be a synonym for one of the species in the 3. Campyloneurum oligophlebium (Kunze) Fée,
C. phyllitidis species group. Gen. Fil. 258. 1852. =Polypodium oligophlebium
Kunze, Linnaea 23: 73. 1850. Type. N. Holl. et
8. Polypodium rodriguezianum L.D. Gómez, Rev. Tasmannia, Houtteau 1848.
Biol. Trop. 17: 107, f. 5-6. 1970. Type: Costa
Rica, Cartago, Cerro Carpintera, Gómez pt.C-
2063.
This name was considered by Lellinger (1988)
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Nac. Autónoma de México, México.
Aspidium pentaphyllum Willd., 58.
Blechnum, 8.
Calceolaria, 16.
Campyloneuron, 3.
Campyloneurum abruptum (Lindman) Ching, (1),
6, 7, 8, 13, 14, 17, 20, 22, 24, 27-28.
— acrocarpon Fée, (2), 4, 5, 11, 12, 14, 16, 17, 19,
20, 22, 24, 25, 26, 28-29.
— aglaolepis (Alston) Sota, 4, 5, 6, 11, 16, 18, 21,
27, 29-30, 43.
— amphostenon (Kunze ex Klotzsch) Fée (4), 1, 2,
4, 5, 6, 7, 9, 10, 11, 12, 14, 15, 17, 18, 19, 20, 21,
22, 25, 27, 30-36, 43, 47, 53, 56, 59, 61, 63, 84,
85.
var. amphostenon (4a), 31-35.
var. irregulare (Lellinger) B. León (4b), 35-36.
— anetioides (Christ) L. D. Gómez (5), 3, 4, 5, 6, 7,
8, 9, 10, 11, 12, 13, 14, 15, 19, 21, 23, 26, 36-37,
46.
— angustifolium (Sw.) Fée (6), 1, 2, 4, 5, 7, 8, 9, 10,
11, 12, 13, 14, 15, 16, 17, 18, 21, 22, 27, 28, 30,
37-43, 44, 47, 52, 58, 93, 94.
var. amphostenon (Kunze) Farwell, 32, 38.
var. ensifolium (Hicken) Farwell, 62.
— angustipaleatum (Alston) Meyer ex Lellinger
(7), 5, 18, 21, 22, 28, 43-44.
— aphanophlebium (Kunze) T. Moore, (8), 2, 6, 7,
9, 10, 11, 12, 13, 15, 18, 20, 22, 23, 26, 38, 44-46.
— asplundii (Christ) Ching (9), 4, 5, 6, 16, 21, 27,
36, 46-47, 87.
— austrobrasilianum (Alston) Sota (10), 14, 17, 18,
21, 22, 28, 44, 47.
— brevifolium (Lodd. ex Link) Link (11), 4, 8, 10,
11, 13, 14, 15, 17, 18, 20, 21, 22, 25, 38, 47-51,
72, 78, 82, 92, 96.
— caespitosum (Link) Link, 80.
— caudatum Fée, 10, 94.
— centrobrasilianum Lellinger (12), 6, 14, 19, 21,
22, 27, 44, 52-53.
— chlorolepis Alston (13), 6, 10, 21, 27, 52-53.
— chrysopodum (Klotzsch) Fée (14), 6, 7, 11, 14,
16, 19, 20, 22, 26, 38, 53.
— coarctatum (Kunze) Fée (15), 4, 5, 6, 7, 14, 15,
20, 22, 25, 26, 53-55, 73, 90.
— cochense (Hieron.) Ching (16), 4, 6, 7, 14, 16,
18, 21, 22, 24, 55-56.
— cooperi Lellinger, 11, 32.
— costatum (Kunze) C. Presl (17), 4, 5, 10, 12, 14,
15, 18, 24, 26, 56-57 . var. nitidissimum (31a), 1, 9, 10, 70.
— crispum Fée, 82. var. latius (Rosenst.) B. León (31b), 71.
— cubense Fée (18), 8, 10, 14, 15, 22, 26, 27, 57-58. — nitidum (Kaulf.) C. Presl, (32), 5, 6, 8, 9, 11, 12,
— decumanum (Willd.) Fée, 97. 13, 14, 17, 19, 20, 22, 26, 27, 71, 82, 93.
— decurrens (Raddi) C. Presl (19), 3, 7, 8, 9, 10, — nodosum Klotzsch, 86.
13, 14, 19, 21, 23, 58-59, 69. — occultum (Christ) L. D. Gómez (8), 9, 11, 45.
— densifolium (Hieron.) Lellinger (20), 2, 4, 5, 14, — oellgaardii B. León (33), 7, 13, 16, 19, 20, 22, 24,
15, 18, 21, 22, 24, 25, 27, 57, 59-61, 65, 69. 25, 75-76.
— difforme (Lodd.) T. Moore, 38. –– oligophlebium (Kunze) Fée, 94.
— ensifolium (Willd.) J. Sm. (21), 6, 15, 18, 21, 28, — ophiocaulon (Klotzsch) Fée (34), 4, 6, 14, 16, 17,
44, 61-62. 20, 22, 25, 76-77, 85.
— falcoideum (Kuhn ex Hieron.) Meyer ex — oxypholis (Maxon) Ching (35), 14, 15, 19, 22,
Lellinger (22), 4, 5, 7, 11, 13, 15, 19, 22, 25, 26, 27, 73-75.
38, 62-63, 96. — pascoense R. M. Tryon & A. F. Tryon (36), 4, 5,
— fallax Fée (23), 6, 14, 17, 18, 19, 21, 25, 62, 63. 7, 9, 10, 11, 13, 14, 16, 21, 22, 25, 52, 75, 91.
— fasciale (Willd.) C. Presl (24), 4, 5, 15, 20, 22, — phyllitidis (L.) C. Presl (37), 1, 4, 7, 8, 10, 12, 13,
26, 64-65, 68. 14, 15, 17, 18, 19, 21, 22, 23, 24, 26, 29, 52, 75,
var. gracile T. Moore, 59. 76-80, 92, 93.
— fendleri (D. C. Eaton) J. Sm., 70. var. costatum (Kunze) Farwell, 56.
— fendleri T. Moore, 54. var. latum (T. Moore) Farwell, 49.
— fuscosquamatum Lellinger (25), 2, 5, 6, 16, 17, — pittieri (Christ) Ching, 32.
20, 22, 26, 65-66. — polyanthum C. Presl, 93.
— heterolepis (Rosenst.) Lellinger, 52. — remotifolium (Hieron.) Lellinger, 84.
— herbaceum (Christ) Ching, 71. — repens (Aublet) C. Presl (38), 1, 3, 4, 5, 6, 7, 8,
— immersum J. Sm., 57. 9, 10, 12, 15, 17, 20, 21, 22, 23, 25, 30, 38, 55,
–– irregulare Lellinger, 36. 60, 68, 70, 72, 77, 80-83.
— inflatum Lellinger (26), 6, 7, 13, 19, 22, 26, 67, — rigidum J. Sm. (39), 8, 4, 17, 19, 21, 22, 27, 83-
75. 84.
— jamesonii Fée, 86. — serpentinum (Christ) Ching, 65.
— juglandifolium Fée, 70. — sieberianum C. Presl, 94.
— laevigatum Cav., 97. — solutum (Klotzsch) Fée (40), 4, 5, 6, 16, 21, 25,
–– lanciforme (J. S. Presl) C. Presl, 96. 27, 84-85.
— lapathifolium (Poiret) Ching, 80 — sphenodes (Kunze ex Klotzsch) Fée (41), 2, 4, 5,
— latum T. Moore, 49, 52. 9, 10, 15, 20, 23, 25, 56, 64, 85-87.
— leuconeuron Fée, 74. — sublucidum (Christ) Ching (42), 7, 8, 13, 15, 19,
–– leucorhizon (Kunze ex Klotzsch) Fée, 32. 20, 25, 87-88.
— lindigii (Mett.) Ching, 70. –– taeniosum (Willd.) Fée, 38.
— lorentzii (Hieron.) Ching (27), 4, 5, 9, 14, 16, 17, — tenuipes Maxon (43), 8, 12, 15, 17, 20, 22, 24,
18, 19, 21, 24, 25, 67. 26, 88-89.
–– loreum (Kaulfus ex Kunze) Fée, 93. — trichiatum (Rosenst.) Ching, 45.
— macrosorum Fée (28), 6, 11, 13, 14, 16, 17, 19, — tucumanense (Hieron.) Ching (44), 2, 5, 8, 10,
20, 25, 68. 13, 14, 16, 17, 19, 21, 22, 25, 52, 78, 89-90.
— magnificum T. Moore (29), 3, 6, 7, 10, 13, 15, 18, — vulpinum (Lindman) Ching (45), 4, 5, 6, 10, 11,
19, 20, 22, 24, 68-69. 12, 14, 15, 18, 21, 23, 25, 77, 90.
–– majus (major) (Hieron.) Lellinger, 74. — wacketii Lellinger, 29.
— minus Fée (30), 5, 12, 17, 19, 20, 26, 69-70, 93. — wercklei (Christ) Lellinger, 88.
–– multipunctatum (Christ) Lellinger, 94, 95. — wurdackii B. León (46), 6, 14, 16, 17, 19, 20, 22,
— nitidissimum (Mett.) Ching (31), 5, 6, 8, 16, 19, 26, 90-91.
21, 22, 24, 52, 70-71. — xalapense Fée (47), 6, 8, 10, 12, 13, 15, 18, 21,
var. abruptum (Lindman) B. León, 28. 22, 24, 26, 27, 57, 58, 91-93.
Cyrtophlebium, 23. — comosum L., 78.

— angustifolium (Sw.) J. Sm., 38. — conjugatum Poiret, 78.
— costatum (Kunze) J. Sm., 56. — costale Jenm. , 56.
— decurrens (Raddi) J. Sm., 60. — costatum Kunze, 56.
— difforme Lodd., 38. — crassifolium L.
— phyllitidis (L.) J. Sm., 78. f. angustissimum Rosenst., 36.
Fuchsia sec. Fuchsia, 16. — cubense (Fée) Christ, 59.
Goniophlebium — decumanum Willd., 94.
— angustifolium (Sw.) Brackenridge, 38. — decurrens Raddi, 58.
–– ensifolium (Willd.) Brackenridge, 62. var. fendleri (D. C. Eaton) Hook., 68.
Grammitis — difforme (Lodd.) Kunze, 38.
— angustifolia (Sw.) Heward, 38. — ensifolium Willd., 62.
Hyalotricha , 23. — falcoideum Kuhn ex Hieron., 64.
— anetioides, 23, 37. — fallax Schlecht, 64.
Hyalotrichopteris , 1, 3, 23, 38. — fasciale Willd., 65.
— anetioides , 3, 23, 37. — fendleri D. C. Eaton, 70.
Marginaria, 3. — gladiatum Vellozo, 93.
— angustifolia (Sw.) C. Presl, 38. — herbaceum Christ, 71.
Microgramma, 4, 6, 11, 12, 21 — laevigatum Cav., 94.
— ciliata, 11 var. crispatum Christ, 92.
Microsorum, 8, 9. –– lanciforme J. S. Presl, 93.
Niphidium , 4, 5, 6, 11, 12, 21. –– lapathifolium Poiret, 82.
Pecluma, 9. — latum (T. Moore) T. Moore, 13, 49.
Platycerium, 9. — leuconeuron
Pleopeltis, 4, 5, 12, 21. var. latifolia Rosenst. , 32.
Polybotrya, 16. –– leucorhizon Kunze ex Klotzsch, 32, 36, 48.
Polypodiaceae, 1, 4, 9, 13, 20, 22. — lindigii Mett., 11, 70.
Polypodium, 3, 5, 6, 9, 12. — longipetiolatum Brade, 63.
–– aglaolepis Alston, 30. — lorentzii Hieron., 69.
— amphostenon Kunze ex Klotzsch, 31. — loreum Kaulfuss ex Kunze, 94.
— anetioides Christ, 37. — magnificum (T. Moore) Hieron., 70.
— angustifolium Sw., 38. — medicinale Rojas, 96.
f. remotifolium Hieron., 84. — multipunctatum (Christ) Christ, 95.
var. amphostenon (Kunze ex Klotzsch) Baker, — nitidissimum Mett., 72.
31. var. latius (latior) Rosenst., 73.
f. densifolium Hieron., 61. — nitidum Kaulfuss, 74.
var. ensifolium (Hicken) Farwell, 62. — nodosum Klotzsch, 86.
var. gramineum Sodiro, 38. — occultum Christ, 45.
var. heterolepis Rosenst., 53. — oligophlebium (Kunze) Fée, 97.
var. monstruosum Mett., 7, 56. — ophiocaulon Klotzsch, 76.
— angustipaleatum Alston, 45. — oxypholis Maxon, 77.
–– aphanophlebium Kunze, 45. — phyllitidis L., 78.
— asplundii C. Chr., 47. f. latum (T. Moore) Proctor, 49.
— austrobrasilianum Alston, 47. f. majus (major) Hieron., 74.
— brevifolium Lodd. ex Link, 47. f. minus (minor) Hieron., 74.
— caespitosum Link, 82. f. multipunctatum Christ, 91.
— calaguala Ruiz, 93. var. elongata Hieron., 78.
— chrysopodum Klotzsch, 54. var. lata (T. Moore) Kaulfuss, 49.
— coarctatum Kunze, 55. var. linneanum Hook., 78.
–– cochense Hieron., 56. var. swartziana Griseb., 78.
— pittieri Christ, 32.

— poloense Rosenst., 32.
— repens Aublet, 82.
var. abruptum Lindman, 28.
var. spathulatum Vareschi, 90.
— rigidum Aublet, 86.
— rigidum (J. Sm.) Lowe, 86.
— rodriguezianum L. D. Gómez, 93.
— schnittspahnii Christ, 94.
— serpentinum Christ, 90.
–– taeniosum Willd., 38.
— solutum Klotzsch, 84.
— sphenodes Kunze ex Klotzsch, 85.
— sublucidum Christ, 87.
— tenuipes (Maxon) Christ, 91.
— trichiatum Rosenst., 45.
— tucumanense Hieron., 89.
— vexatum D. C. Eaton, 10, 59.
— vulpinum Lindman, 92.
— weatherbyanum Seymour, 94.
— wercklei Christ, 88.
— xalapense (Fée) Christ, 91.
Polypodium subg. Cyrtophlebium, 3, 23.
Pyrrosia, 3, 4, 5, 11, 21.
Quercus-Liquidambar, 15.
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BLen 1993 ATaxonomicRevCampyloneurum Thesis

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A Taxonomic Revision of the Fern Genus Campyloneurum (Polypodiaceae)

Thesis · January 1993

Ferns and lycophytes of Peru: rarity and endemism View project

The GRIN-Taxonomy Crop Wild Relative Inventory View project

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Nordlandsvej 68, DK 8240 Aarhus, Danmark

Afhandling indleveret til det Naturvidenskabelige Fakultet ved Aarhus

Universitet til bedømmelse for Ph.D. graden

Vejleder: Lektor Benjamin Øllgaard

to my husband, Kenneth R. Young.

Danish summary - Dansk sammenfatning viii-xii

II. MATERIALS AND METHODS 1-2

III. TAXONOMIC HISTORY 3

IV. MORPHOLOGY 4-12

V. CYTOLOGY AND BIOCHEMISTRY 12-13

VI. ECOLOGY AND GEOGRAPHICAL DISTRIBUTION 13-18

VII. CONSERVATION AND USES 18-19

VIII. INFRAGENERIC CLASSIFICATION AND PHYLOGENY 19-23

Nomina Dubia 93-94

X. CITED REFERENCES 95-99

XI. LIST OF TAXA 100-102

Figure 1. Stem morphology in Campyloneurum.

Figure 30. Distribution of Campyloneurum angustifolium.

Table 1. Variablity of morphological characters in Campyloneurum.

og pinnate blade. Definitionen af Campyloneurum forblev uændret indtil Tryon &

henført til slægten Hyalotrichopteris.

Nærværende afhandling har til formål at afklare slægtens afgrænsning, og at

af morfoloiske karakterer og belyse deres indbyrdes slægtskab.

Herbariorum Part I. 8th ed.).

Slægtens morfologi gennemgås, og visse kritiske karakterer, såsom

behåring, sporangier og parafyser, beskrives for første gang (Kap. IV).

Slægten deler mange karakterer med andre slægter i Polypodiaceae.

Stænglen (rhizomet) hos Campyloneurum er krybende, med skæl og

dorsiventralt arrangerede rødder og blade. Rhizomskæl har spillet en vigtig

rolle for mange Polypodiace-slægters taxonomi, fordi de leverer mange

konstante bygningstræk. Dette gælder også Campyloneurum, hvor materiale

form og størrelse; de er i reglen anbragt i to rækker langs stænglen, og falder

af ved et distinkt løsningslag. Bladpladen er hel hos de fleste arter; kun C.

decurrens og C. magnificum har fjersnitdelte bladplader. Med undtagelse af

en enkelt art, Campyloneurum aphanophlebium, beskrives arterne som glatte.

affaldende skæl. Fuldt udvoksede blades nervation kan inddeles i 4 typer,

der danner grundlag for en inddeling i artsgrupper (Kap. VIII).

Sporehushobene er afrundede eller elliptiske, uden indusier, og 1-2 (-3) mm

i diameter. Sori sidder på bladundersiden, på afrundede eller elliptiske

områder i niveau med bladoverfladen, oftest i to rækker mellem

primærnerverne, men undertiden tre eller flere hos C. brevifolium gruppen,

og undertiden i en enkelt række hos C. anetioides og C. falcoideum.

Forekomsten af parafyser, organer der kan findes blandt sporangierne, blev

sporangierne, og tyndere end sporangiestilkene; der findes to typer: simple

og dendritiske. Sporerne er ellipsoidale, de aborterede dog mere afrundede

og kollaberede, 40—100 x 30—60 µm; deres overflade er forsynet med

spredte sfæriske legemer eller granuli under under et tyndt perispor.

Exosporet er mere eller mindre vortet.

Campyloneurum er tidligere anset for at stå nærmest de andre

neotropiske slægter Microgramma og Niphidium, men anses her for at stå

særlig nær Pleopeltis (Kap. IV).

De fleste af arterne hører til i skov-vegetation (Kap. VI), især

stedsegrøn skov, men C. xalapense findes i delvis løvfældende skov i

Mexico. De fleste arter forekommer på mange forskellige mikrohabitater på

forstyrrede eller uforstyrrede skov-arealer. Seksogtyve arter hører til på

skyggede og fugtige mikrohabitater i sluttet skov. De fleste af disse arter er