FIELDIANA Botany Published by Field
New
Series,
Museum
of Natural History
No. 12
FERNS AND FERN ALLIES OF GUATEMALA PART III MARSILEACEAE, SALVINIACEAE, AND THE FERN ALLIES (INCLUDING A COMPREHENSIVE INDEX TO PARTS I, II, AND III) ROBERT
G.
STOLZE
Custodian, Pteridophyte Herbarium Field Museum of Natural History
The Genus Lycopodium BENJAMIN 0LLGAARD
by:
Botanical Institute
University of Aarhus,
The Genus R.
Denmark
Isoetes with:
JAMES HICKEY
University of Connecticut, Starrs
Accepted for publication July
August
31, 1983
Publication 1349
16,
1982
CONTENTS INTRODUCTION
1
ACKNOWLEDGMENTS
3 4 4 7 7
Marsileaceae Marsilea Salviniaceae Azolla Salvinia
11
Psilotaceae
13 14 16 16 20 21 21 25 28 44 44 62 63 68
Psilotum
Equisetaceae Equisetum
Lycopodiaceae Lycopodium
Subgenus Cernuistachys Subgenus Lycopodium Subgenus Selago Selaginellaceae Selaginella
Isoetaceae Isoetes
COMPREHENSIVE INDEX TO PARTS
I,
II,
LIST 1.
Marsilea:
M.
mexicana;
M.
AND
III
OF ILLUSTRATIONS 5 9 12
berteroi
filiculoides; A. microphylla 3. Salvinia: 5. minima; S. auriculata 2. Azolla:
A.
4. Psilotum: P.
nudum;
P.
complanatum
5.
Equisetum: E. hyemale var.
6.
Lycopodium subgenus Cernuistachys: L. carolinianum; L. cernuum Lycopodium subgenus Lycopodium: L. clavatum; L. thyoides Lycopodium subgenus Selago: L. mexicanum; L. taxifolium; L. myrsinites
7. 8.
affine; E.
myriochaetum; E. giganteum
9. Selaginella: S. umbrosa; S. steyermarkii; S. galeottii; S. pallescens var. pallescens 10. Isoetes: I. montezumae; I. panamettsis; I. cubana
15 17 23 26 31 48 65
FIELDIANA:
62
BOTANY
mm
maturity; sporophylls 1-1.5 long, carinate, usually spreading at maturity; megaspores white; microspores pale orange.
This and
S.
faucium were described at the
same time by Baker and were
said to
differ principally in size of plants and width of branches. The Peruvian S. tarabroad. The potensis is a smaller, more delicate plant, with branches about 3
mm
mm
faucium is much larger and coarser, with larger branches 4 to 6 broad. Alston (1955) combined the two, and rightly so, for other than size and
Mexican
S.
texture, there are
no
significant differences.
The coarser plants
are found in south-
ern Mexico and Guatemala. Other than one or two specimens, the South American plants are the delicate variants represented by the type. Selaginella umbrosa Lemaire ex Hieron., in Engl. & Prantl, Die Naturlichen Pflanzenfamilien 1(4): 683. 1901. S. lemairei Hieron., Hedwigja 58: 287. 1917, nom. provis. S. erythropus var.
major Spring,
Mem Acad. Roy. Sci.
Belgique 24: 156. 1850.
In forests, thickets or wooded ravines, on banks of lakes or streams, rarely on bases of trees or on rocky slopes, sea level to 650 m; Alta Verapaz; Izabal; Peten; San Marcos. British Honduras; Honduras; Costa Rica; Panama; Jamaica; Grenada;
Barbados; Tobago; Colombia; Venezuela; Brazil; Peru. Plants commonly terrestrial, erect, heterophyllous, but homophyllous as to the stem below the leafy portion; stems solitary or remote, never caespitose, unbranched in the 2 proximal Va- /}, deep reddish nearly throughout; rhizophores few, basal, produced from the lower surface of the stem; lateral leaves 1-3 mm long, older ones turning reddish, approximate to contiguous, or slightly imbricate on the branches, obliquely ovate or lanceolate, falcate, acute, margins essentially entire, but ciliate in the proximal half, at base broadly rounded acroscopically, truncate to geniculate basiscopically; axillary leaves ovate to broadly lanceolate, acute, rounded at base, the margins entire to serrulate, but ciliate in the proximal portion; median leaves 0.7-2.7 long, oblique-ovate, acute or (occasionally) acuminate, subcordate at base or unequally biauriculate, the margin entire to remotely serrulate or setose, but with 3-4 cilia at base; strobili to ca. 1 cm long, sharply
mm
tetragonous; sporophylls ca. 1 bright orange.
This
is
Spring. In
mm long, sharply carinate; megaspores white; microspores
most respects to the South American S. haematodes (Kze.) two are often confused where their ranges overlap in Panama.
similar in fact,
the
The most obvious
difference
is
in the lateral leaves, with those of S. umbrosa rather
conspicuously ciliate in the proximal portion and sharply truncate or geniculate at base basiscopically. Lateral leaves of S. haematodes are essentially naked along the
margin and are rounded
at the base.
ISOETACEAE ROBERT G. STOLZE Field
Museum R.
of Natural History
JAMES HICKEY
University of Connecticut, Storrs Plants heterosporous, small, herbaceous, perennial, aquatic, amphibious, or terrestrial;
corms subterranean or epiterranean, rounded or elongated horizontally or vertically, 2- to many-lobed, the lobes often difficult to discern; roots numerous, dichotomously branched, produced within a single or several root grooves (fossae) which are continuous along the basal or lateral surfaces of the conn or are confined to a single side of the corm (as in older specimens of /. andicola [Amstutz] Gomez); leaves quill-like, (2-) 5-40 (-128), imbricate,
STOLZE: FERNS OF
GUATEMALA
63
spiral or rarely distichous, erect to recurved, with an expanded, winglike margin, triangular or rounded in cross section, with a single, unbranched midvein, surrounded by 4 transversely septate air channels and 0-6 or more peripheral groups of fibrous supporting cells; stomata numerous to lacking; ligule small, delicate, broadly to narrowly triangular, ephemeral, arising from a pit on the adaxial leaf surface just above the sporangium, frequently with a fleshy labium inserted between the ligule and sporangium which partially or completely covers the ligular pit; sporangia adaxial, borne singly at or near the base of the leaf, covered by or partially covered by the membranaceous velum; megaspores globose-tetrahedral, 200-800 (-1,000) /im in diameter, variously ornamented, developing
into endosporic gametophytes which produce several archegonia; microspores ellipsoid, 20-60 fim long, variously ornamented, developing into endosporic gametophytes which produce but a single antheridium.
This is a monotypic family containing the single, cosmopolitan genus Isoetes, with 175-200 species. A second genus, Stylites, from Peru and Bolivia, has often been recognized and considered distinct based on its unusual corm morphology,
and sporangial position. These characters, while unusual, are sufficiently discrete to merit generic distinction. The unusual vertically elongated corm of Stylites appears to be nothing more than an adaptation to root production, not,
however,
where the surrounding substrate is continually and rapidly The unusual root production in adult plants of Stylites, where roots are
a cushion-plant habitat increasing.
at the apex of the single, laterally positioned fossa is likewise a modification to this unusual habitat, and young plants have been found in which the fossa is continuous around the base of the corm and extends up each side.
produced only
Finally, other taxa in the
sporangia located
fairly
genus
Isoetes (e.g.,
/.
high up on the adaxial
andina Spruce ex Hook.) have
leaf surface.
ISOETES Linnaeus REFERENCE: N. E. Pfeiffer, Monograph of the Isoetaceae, Ann. Missouri Bot. Card. 9: 79-233. 1922. Characters are those of the family.
The "quillwort" genus,
Isoetes,
can be mistaken for no other group of ferns or
fern allies; but species are often difficult to delineate, even with the aid of a compound microscope. The problems are greater still in the study of dried specimens. Most of the so-called "diagnostic" characters used in the past are inconsistent and/or misunderstood. Pfeiffer (1922)
and others attempted
to base a classi-
features as shape and size of the ligule, number of corm lobes, presence or absence of stomata, and number of fibrous peripheral bundles. Most of these are difficult to assess, especially in dried specimens, and the nature of many such characters is dependent upon the age of the plant or leaf or the fication partially
on such
habitat (whether the plant
is fully
submersed,
partially so, or
completely
ter-
restrial).
To add to the taxonomic difficulties, there is the problem of paucity of specimens, especially in more remote areas of the Tropics. Plants are easily overlooked, or are passed over by collectors as clumps of sedges or rushes. Additional efforts to seek out these plants must be made, especially in the Tropics, but more intelligent collecting
and data recording are necessary as
well.
Attempts
to collect
plants within a particular habitat should include individuals of varying degrees of maturity, as well as those from the center and at the periphery of the colony, and
those of varying degrees of submersion. Furthermore, individual plants should be tagged to record these three pieces of data.
FIELDIANA:
64
BOTANY
occurs along fresh or tidal shores, in or at margins of lakes, ponds, and and wet meadows, alpine bogs, and rarely in low forests or
Isoetes
streams, in ditches
rocky slopes. Aquatic taxa are frequently evergreen, whereas terrestrial species often lose all their leaves and can remain dormant for one or several years. Two species have been collected in Guatemala,
perhaps a.
to
be expected
Megaspores the proximal
a.
in
and a
third,
from
British
Honduras,
Guatemala.
bacillate, the projections and distal surfaces
much
taller
than broad, and evenly distributed on
Megaspores smooth, verrucate or pustulate, the protuberances as
tall
/. panamensis. as or shorter than
broad. b.
Megaspores pustulate, the protuberances smooth, rounded, distinct on both proximal and distal surfaces, evenly distributed and well developed, even near the
b.
Megaspores smooth or verrucate, the protuberances when present poorly developed and then rounded or well developed and then apically muricate, the protuberances 7. montezumae. becoming smaller and less distinct near the equatorial ridge
7.
equatorial ridge
Isoetes
Thus
cubana Engelm. ex Baker,
far
known only from
J.
cubana.
Bot. 18: 110. 1880.
the type location in
Cuba and two
locations in British
Honduras.
Corm rounded, 3(?)-lobed, 0.4-2 cm in diameter; fossa continuous around corm; leaves broad at mid-length, triangular in cross section, 10-40, these 10-46 cm long, 0.5-1.75 with 3 darker, shallow, lateral ridges continuing to the apex; differentiated leaf border broad at the sporangium, gradually chartaceous, brown to barely transparent, 1-2 tapering into the leaf 1.5-6 cm above the base; sporangia elliptic, 5-9 long, 3-4 broad along the lateral edges of the broad, tan, unspotted; velum incomplete, 0.3-0.6 sporangium, 0.6-0.7 deep along the upper edge of the sporangium; ligule not apparent, eroding to below the labium; labium transversely ovate, 0.4-1.2 high, 1.1-3 = 351) /*m in broad, tan, margins entire to laciniate; megaspores white or gray, 270-470 (x diameter, pustulate, with 20-50 or more projections on the distal surface, each of these 30-70 /tm in diameter, as high as or shorter than broad, occasionally merging laterally to
mm
mm
mm
mm
mm
mm
mm
mm
short, meandriform markings, equatorial and proximal ridges straight, thick and = 29) /Ltm = 23) /nm wide, exospore prominent; microspores gray, 25-35 (x long, 20-30 (x 1-4 yum thick, sparsely to densely echinate (smooth in immature spores).
form
This species is apparently aquatic or semiaquatic, based on label data, the flexuous nature of the leaves in the British Honduras material, and the dense covering of diatoms on the leaves of the Cuban collections. Although the British Honduras material does not possess aborted scalelike leaves, this may be an artifact of collecting, and more ter can be fully assessed.
specimens need
to
be examined before
this charac-
Honduras and Cuban material is very close morphologically, but some of the megaspore and microspore characters. The megaspore pustules in the British Honduras material are generally more numerous, more crowded, and tend to be somewhat shallower than those in the Cuban material. In the latter, there are fewer than 40 pustules on the distal surface, while in the former, there are generally more than 50. Although the microspores of the Cuban material cannot be completely characterized due to their immature nature, it is The
does
British
differ in
obvious from measurements of the most mature spores found that they are somewhat longer (x = 30 /j,m) than those from British Honduras (x = 28 /xm) and less densely ornamented.
FIG. 10. Isoetes.
d-e,
a-c,
/.
montezumae:
panamensis: d, leaf base, x 38. g, /. cubana, megaspore, x /.
x 5; HO.
a, e,
x 1/2; b, leaf base, x 6; c, scale leaves, x 5. megaspore, x 94. f, /. montezumae, megaspore,
habit,
65
FIELDIANA:
66
montezumae A. A.
Isoetes
BOTANY
Eaton, Fern Bull.
5:
25. 1897.
In shallow ponds, wet depressions, and meadows, 2,000-3,500 m; Huehuetenango; San Marcos. Mexico (type from the state of Mexico: damp soil, plains near Flor de Maria, Aug. 28, 1890, Pringle 3459).
Corm rounded
to slightly elongated laterally, 2-lobed; fossa continuous
corm; leaves 7-21, these 4-23
around the
mm
cm
broad at mid-length, stiff, upright to long, 0.4-1.3 slightly recurved, triangular in cross section with 3 green, inconspicuous lateral ridges continuing upward to near the apex, leaf tip abruptly tapering to a blunt, rounded apex, differentiated leaf border chartaceous, light brown, nearly transparent, 0.4-2 broad at
mm
mm
the sporangium, gradually tapering into the leaf 1.5-4.5 above the base; scale leaves persistent, black to dark brown, oblong to transversely oblong, 2-2.5 long, 2-4 broad, with mucronate apex, 0.5-1 long; sporangia elliptic to circular, 2.5-4.5
mm
mm
mm mm
mm broad, tan, lightly spotted to unspotted; velum absent, or incomplete, mm broad along the upper edge of the sporangium; ligule occasionally represented by a tan, triangular to ovate remnant, 1-1.25 mm long, 1-1.6 mm broad; labium very small, barely covering the ligular pit, 0.2-0.3 mm high, 0.6-0.8 mm broad, tan, entire; megalong, 2-3.5
0.2-0.3
=
510) /am in diameter, smooth to inconspicuously and more crowded near the equatorial ridges, each protuberance 20-30 /urn high, 20-30 /^.m broad, equatorial and proximal ridges straight, = 39) /tm = 31) pirn broad, distinct; microspores light gray, 34-45 (x long, 27.5-35 (x exospore 3.25 ;u,m thick, minutely granular, often with larger tubercles superimposed on
spores white, not shiny, 350-660
(x
pusticulate, the protuberances smaller
the granular surface. is found throughout central and southern Mexico from Jalisco Oaxaca and Chiapas. Its southern limit appears to be in central Guatemala, though it should be expected in Honduras and El Salvador. The plant is quite seasonal, as evidenced by its habitat and the persistent scale leaves which apparently protect the apical meristem during drought conditions. The description given above is based primarily on Guatemalan material. This material differs, however, in several respects from the more typical Mexican collections. In general, the Guatemalan plants are much smaller, have fewer leaves, and have much smaller sporangia. The megaspore ornamentation is also much less well developed. Typically, /. montezumae has distinct tubercles on both proximal and distal surfaces, though, like the Guatemalan material, there is a tendency
Isoetes
montezumae
and Mexico
to
to
show numerous but
well-developed protuberances near the equatorial
less
ridge. However, even within Mexico, there is a tremendous amount of variation in many of these characters, and it seems reasonable at this time to assign the
Guatemalan Isoetes /.
paciftca
collections to
/.
montezumae.
panamensis Maxon
Svenson, Amer. Fern
south of Engabao,
Chanduy
& Morton, J.
Ann. Missouri
34: 123. 1944. (type
Hills,
Bot. Card. 26: 272. 1939. from Ecuador, Prov. Guayas,
Mar. 22/23, 1941, Svenson 11002.)
boggy swamps, and depressions, 950-1,500 m. As yet, only from two localities in Guatemala: Chiquimula and Jutiapa, and from one locality each in Costa Rica, Panama (type from Prov. Panama, vicinity of Bejuco, Aug. 7, 1938, Woodson, Allen & Seibert 1685), Ecuador, In seasonal rain pools,
this species is
known
and Peru.
Corm rounded, 3-lobed, 0.7-3 cm in diameter; fossa continuous around corm; leaves broad at mid-length, stiff, upright, triangular in 9-128, these 17-28 cm long, 0.5-2 cross section, with three, darker, shallow, lateral ridges continuing to the apex; persistent scale leaves not present, leaf tip finely tapering to a thin, pointed apex, stomates present, broad at the spodifferentiated leaf border chartaceous, tan, nearly transparent, 2-4
mm
mm
STOLZE: FERNS OF
GUATEMALA
67
rangium, gradually tapering into the leaf 3.5-10 cm above the base; sporangia obovate, 13 broad along broad, tan, unspotted; velum virtually absent, 0.2-0.5 long, 5-7 the lateral edges of the sporangium, absent along the upper edge of the sporangium; ligule usually absent, occasionally represented by a blackened, triangular fragment, 3-4 broad, its base partially or completely covered by the labium; labium high, 4-4.5 broad, whitened, margins laciniate; megatransversely elliptic, 1-2 high, 4.5 = 451) /xm in diameter, bacillate, the spores white, 350-580 (x projections up to 70 ptm in height, ca. 35 pirn in diameter; equatorial ridge thin, often undulate, proximal ridges thin, = = 26) pirn broad, exdistinct; microspores ash-gray, 27-40 (x 34) /xm long, 20-33 (x ospore ca. 2.5 jum thick, smooth to sparsely echinate.
mm
mm
mm
mm
mm
mm
Isoetes
Svenson
mm
panamensis grows near sea level from Peru to Costa Rica. According to (1944), the Ecuador material was collected in pools which are dry for part
of the year; such habitats are probably common throughout the range of this species, though label data is scanty. The spotty distribution of the species will undoubtedly be filled in as more intensive collections are made along the west
America and northern South America. The type descriptions of /. panametisis and /. pacifica coincide in most characters. The greater leaf width reported for /. panamensis is most likely a reflection of the more vigorous nature of this collection, which may be a result of lower water stress in that locality. The only significant discrepancy in the two descriptions is in the reports of microspore "diameter." Maxon and Morton report a diameter of 25 fj.m for I. panamensis, whereas Svenson gives a size range of 36-40 /u,m for the microspores of /. pacifica. Given the fact that the spores are ellipsoid, it is possible that Svenson's measurements were of microspore length, while those of Maxon and Morton were of width. Recent measurements of spores from both types indicate that the spores are quite comparable in both length and width. coasts of Central
Gomez has described a new species, /. savannarum, from Costa which strongly resembles /. panamensis. The descriptions of these two plants are sufficiently similar to warrant a close examination of the two before relationships can be postulated. Recently, L. D.
Rica,
COMPREHENSIVE INDEX TO PARTS This index covers
have appeared
I,
II,
AND III
three parts of the pteridophyte flora of Guatemala which and consists of 719 pages and numerous plates.
all
in Fieldiana: Botany
The numbers following an entry refer first to the part number of the flora and second to the page(s) where the entry occurs. The index includes all accepted names (in roman), synonyms (italics ), common English names (roman), and vernacular names (italics). Entries (in roman) are also included for medicinal or local uses. Hyphenated words and multiple words are alphabetized letter by reader, also
letter.
Names
of persons,
when
considered useful to the
have been indexed. Page numbers of descriptions are
page numbers
of illustrations are in
italics.
INDEX Achomanes,
huacsaro 2-225
section 1-78
Acrospermum maxonii 2-243
japurense 2-296
subgenus 3-25 Acrostichum 2-11 acuminatum 2-116 affine 2-215 amygdalifolium 2-216 araneosum 2-231
latifolium 2-226 lindenii 2-226
Acrostachys,
aureum 2-11,
22,
mathewsii 2-227
mexicanum 2-220 mulleri 2-231 muscosum 2-229 nicotianaefolium 2-116 nitidum 2-220 nodosum 1-16 osmundaceum 2-357 ovatum 2-236 paleaceum 2-229 pallidum 2-237 peltatum 2-341 petiolatum 2-230 piloselloides 2-231 pilosum 2-231 polypodioides 2-416 portoricense 2-119 pringlei 2-227 pumilum 2-226 rampans 2-233 recognitum 2-234 reptans 2-369 revolutum 2-233 rubiginosum 2-237 rufum 2-267 sanctum 2-485 schiedei 2-230 schmitzii 2-230 serrulatum 2-264
13
auricomum 2-217 bernoullii 2-116 blepharodes 2-220 bonariense 2-319 calomelanos 2-344 cervinum 2-356 citrifolium 2-42 cladorrhizans 2-119 crinitum 2-218, 220 daneaefolium 2-22, 13, 14 decoratum 2-219 dombeyanum 2-219 ebeneum 2-344 engelii 2-220 erinaceum 2-220 excelsum 2-13 eximium 2-221 fitnbriatum 2-220 flaccidum 2-234 glaucum 2-221 gratum 2-222 guatemalense 2-222 herminieri 2-224 hirtum 2-229 68
in boldface;
STOLZE: INDEX Hemionitis dealbata 2-347
2-440 2-440
79
lindenii
1-70
lanceolata
lineare 1-62, 65
lineata
maxonii 1-64, 65 var. angustius 1-66 microcarpum 1-64 microcarpum 1-64 myriocarpum 1-58, 67 pannosum 1-68 plumieri 1-70
palmata 2-273, 275 pedata 2-120 pinnatifida 2-274, 275 Hetniphlebium krausii 1-83 membranaceum 1-84 petersii 1-85 reptans 1-88 sphenoides 1-86 Hentitelia costaricensis 1-126 decurrens 1-106 guatemalensis 1-106 /wdda 1-106 mexicana 1-106 multiflora 1-111 nigricans 1-111 Hemitdia, subgenus Cnemidaria 1-105 Heteropteris lanceolata 2-334 Hickey, R. James 3-3, 62 Hicriopteris bancroftii 1-46 Hippochaete, subgenus 3-16, 19 Histiopteris 2-276 incisa2-276, 277 Holly ferns 2-164 Horsetail 3-16 Huperzia crassa 3-34 dichotoma 3-36 hartivegiana 3-36
3-37 3-39 passerinoides 3-42 lini/olia
polyanthos 1-57, 58, 67, 68 pulchellum 1-62, 68 schiedeanum 1-60 sieberi 1-60, 69 spinulosum 1-63 tablaziense 1-56 trapezoidale 1-60 tricophyllum 1-70 undulatum 1-59 ivercklei 1-69 Hymenophyllum, subgenus 1-53 Hymenostachys diversifrons 1-79 Hypoderris adnata 2-351 heteroneuroides 2-351 marginalis 2-351 seemannii 2-351
Hypolepis 1-102; 2-173, 276, 278, 340 hostilis 2-280 mexicana 2-280 nigrescens 2-278, 279 radiata 2-140 repens 2-279, 280
myrsinites
reflexa
Isoetaceae 3-62
3-41 3-42
Isoetes
taxifolia
Hymenasplenium, Asplenium section 2-51, 182
Hymenodium 2-219 crinitum 2-218
montezumae pacifica
Hymenophyllaceae 1-51
Hymenophyllum 1-52, 53 abruprum 1-55, 58 asplenioides 1-56 brevifrons 1-55, 56, 58 brevisripes 1-58 caudatellum 1-61
1-64 contortum 1-58,
3-63
andina 3-63 cubana 3-64, 65
ciliatum
59, 71
crassipetiolarum 1-59, 60, 70
crispum 1-60
3-65, 66 3-66, 67
panamensis 3-65, savannarum 3-67 Ithycaulon 2-458 inaequale 2-459
66, 67
Jamesonia 2-238, 280, 313 alstonii 2-281,282 goudotii 2-283 hispidula 2-240 Juniperus standleyi 2-203
ectocarpon 1-60, 61, 63 elegans 1-61, 65 elegantissimum 1-65
elegantulum 1-62, 69 fragile 1-62
Kamcha 3-27 Kam chax 3-27 Kidney medicine 2-377; 3-19
fucoides 1-57, 61, 63
Kolo macho ujekiej 3-18
hirsurum 1-64 1-62 interruprum 1-70 lanatum 1-57, 64
Lagarta 2-482
intercalatum
Lady
fern 2-98, 99