9
654 32. CALYMPERACEAE Kindberg William D. Reese† Plants small to medium-sized, mostly erect [prostrate and with ascending branches], in tufts, cushions, or gregarious, rarely single or in tufts of a few; dark green to yellowish brown stems. Stems simple or forked; central strand absent; rhizoids brown to red or dark purple, scanty to numerous, sometimes abundant and conspicuous; axillary hairs 2 to many per axil, proximal cells short and colored, or undifferentiated. Leaves often slightly to strongly dimorphic, mostly variously contorted when dry, straight and erect-ascending when moist, teniolae (intramarginal files of differentiated cells) present (in some Calymperes) or absent; proximal portion of leaves commonly sheathing, including usually conspicuous areas of smooth, enlarged, hyaline, internally and externally porose cells (cancellinae); distal portion of leaves oblong to ligulate, lanceolate, acuminate, or narrowly long-linear; margins of distal lamina mostly thickened and toothed, sometimes bordered entirely or in part with elongate hyaline cells; costa single, strong, percurrent to excurrent, in cross section showing a median row of guide cells with abaxial and adaxial bands of stereid cells (the ad- and abaxial bands of cells rarely thin-walled and parenchyma- like); medial cells of distal lamina mostly isodiametric, rarely transversely elongate, mostly variously papillose. Specialized asexual reproduction common by seriate-multicellular, fusiform- clavate to filiform gemmae, borne mostly adaxially on apices of often highly modified leaves. Sexual condition mostly dioicous, rarely monoicous; perigonia axillary, gemmiform, with highly reduced leaves; perichaetia terminal but quickly overtopped by innovations and then appearing lateral, leaves few, scarcely different from cauline leaves. Seta single, yellow to red, mostly elongate, smooth, erect. Capsule erect, mostly exserted, rarely immersed, yellowish to brown, mostly cylindric, smooth; stomata scanty, on neck, phaneropore; annulus differentiated in several rows of cells; operculum rostrate; peristome single, often vestigial or absent, teeth 16, variously papillose on external surface, mostly smooth on internal surface, with weak transverse bars. Calyptra cucullate and deciduous or rarely conic-mitrate, sometimes clasping the seta below the capsule and persistent and the spores then escaping through vertical fissures in the calyptra, naked, smooth or papillose, sometimes plicate. Spores spherical, mostly granular-papillose. Genera 3, species ca. 150 (2 genera, 11 species in the flora): worldwide, mostly tropical and subtropical regions. The Calymperaceae are defined by clasping leaf bases with cancellinae, absence of stem central strand, usually thickened and toothed leaf margins, mainly tropical-subtropical range, corticolous

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Page 1: 32. CALYMPERACEAE Kindberg · 2009. 10. 21. · 654 32. CALYMPERACEAE Kindberg William D. Reese† Plants small to medium-sized, mostly erect [prostrate and with ascending branches],

654

32. CALYMPERACEAE Kindberg

William D. Reese†

Plants small to medium-sized, mostly erect [prostrate and with ascending branches], in tufts,cushions, or gregarious, rarely single or in tufts of a few; dark green to yellowish brown stems.Stems simple or forked; central strand absent; rhizoids brown to red or dark purple, scanty tonumerous, sometimes abundant and conspicuous; axillary hairs 2 to many per axil, proximalcells short and colored, or undifferentiated. Leaves often slightly to strongly dimorphic, mostlyvariously contorted when dry, straight and erect-ascending when moist, teniolae (intramarginalfiles of differentiated cells) present (in some Calymperes) or absent; proximal portion of leavescommonly sheathing, including usually conspicuous areas of smooth, enlarged, hyaline, internallyand externally porose cells (cancellinae); distal portion of leaves oblong to ligulate, lanceolate,acuminate, or narrowly long-linear; margins of distal lamina mostly thickened and toothed,sometimes bordered entirely or in part with elongate hyaline cells; costa single, strong, percurrentto excurrent, in cross section showing a median row of guide cells with abaxial and adaxialbands of stereid cells (the ad- and abaxial bands of cells rarely thin-walled and parenchyma-like); medial cells of distal lamina mostly isodiametric, rarely transversely elongate, mostlyvariously papillose. Specialized asexual reproduction common by seriate-multicellular, fusiform-clavate to filiform gemmae, borne mostly adaxially on apices of often highly modified leaves.Sexual condition mostly dioicous, rarely monoicous; perigonia axillary, gemmiform, with highlyreduced leaves; perichaetia terminal but quickly overtopped by innovations and then appearinglateral, leaves few, scarcely different from cauline leaves. Seta single, yellow to red, mostlyelongate, smooth, erect. Capsule erect, mostly exserted, rarely immersed, yellowish to brown,mostly cylindric, smooth; stomata scanty, on neck, phaneropore; annulus differentiated in severalrows of cells; operculum rostrate; peristome single, often vestigial or absent, teeth 16, variouslypapillose on external surface, mostly smooth on internal surface, with weak transverse bars.Calyptra cucullate and deciduous or rarely conic-mitrate, sometimes clasping the seta belowthe capsule and persistent and the spores then escaping through vertical fissures in the calyptra,naked, smooth or papillose, sometimes plicate. Spores spherical, mostly granular-papillose.

Genera 3, species ca. 150 (2 genera, 11 species in the flora): worldwide, mostly tropical andsubtropical regions.

The Calymperaceae are defined by clasping leaf bases with cancellinae, absence of stem centralstrand, usually thickened and toothed leaf margins, mainly tropical-subtropical range, corticolous

rzander
Text Box
Flora of North America, Volume 27, 2007
Page 2: 32. CALYMPERACEAE Kindberg · 2009. 10. 21. · 654 32. CALYMPERACEAE Kindberg William D. Reese† Plants small to medium-sized, mostly erect [prostrate and with ascending branches],

CALYMPERACEAE 655·habitat, and common production of uniseriate leaf-tip gemmae. Plants of the family arespecialized for water capture and retention by their sheathing leaf bases, and the large, empty,porose cells of their cancellinae. The habitat is typically on bark but many taxa occur primarilyon decaying wood, rock, and soil. Our taxa occur almost exclusively in the southeastern UnitedStates.

SELECTED REFERENCES Reese, W. D. 1993. Calymperaceae. In: Organization for Flora Neotropica. 1968+. Flora Neotropica.98+ nos. New York. No. 58. Reese, W. D. and R. H. Zander. 1988. Comparison of Calymperaceae with Pottiaceae. Bryologist91: 18–20.

1. Plants with sporophytes.2. Calyptra deciduous, cucullate, not clasping the seta, without vertical fissures . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Syrrhopodon (in part), p. 6552. Calyptra persistent, enclosing the capsule and clasping the seta proximally, with vertical

fissures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. Calymperes (in part), p. 6591. Plants without sporophytes.

3. Leaves bordered entirely or in part with elongate hyaline cells . . . . . . . 1. Syrrhopodon (in part), p. 6553. Leaf margins without elongate hyaline cells.

4. Teniolae (intramarginal files of differentiated cells) present at leaf shoulders . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. Calymperes (in part), p. 659

4. Teniolae absent.5. Leaves 4–8 mm; margins coarsely toothed and multistratose distally; gemmae

adaxial on leaf apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Syrrhopodon (in part), p. 6555. Leaves 1–3 mm; margins entire and 1- to 2-stratose distally; gemmae borne

circling apex of excurrent costa . . . . . . . . . . . . . . . . . . . . . . 2. Calymperes (C. tenerum), p. 659

1. SYRRHOPODON Schwägrichen, Sp. Musc. Frond. Suppl. 2(12): 110. 1824 · [Greek

syrrepo, to close the eye, and odon, tooth, alluding to narrow, connivent, horizontal

peristome teeth of some species closing capsule mouth upon drying]

Plants small to medium-sized, gregarious to tufted or caespitose, green to yellowish brown,sometimes with pink tinge. Stems erect. Leaves sometimes dimorphic with vegetative leavesmostly grading into gemmiferous leaves, teniolae absent (teniola-like features rarely present ina few taxa); margins of distal lamina mostly thickened and toothed, rarely 1-stratose, in manytaxa regularly bordered entirely or in part with elongate hyaline cells; costa showing two bandsof stereids in cross section; medial leaf cells isodiametric. Specialized asexual reproduction bygemmae common on often modified leaves, borne on apices or adaxially and sometimes abaxiallyalong costa of distal lamina, clavate-fusiform or filamentous. Sexual condition mostly dioicous,rarely apparently monoicous. Seta single, rarely 2–3 per perichaetium. Capsule mostly exserted,sometimes immersed, mostly cylindric; peristome present or absent. Calyptra deciduous, mostlycucullate, rarely conic-mitrate.

Species ca. 80–90 (6 in the flora): worldwide except Antarctica, mostly tropical and subtropicalregions.

Syrrhopodon is characterized by its erect stems, leaves bordered with hyaline cells or withthickened and toothed margins, cancellinae, and mostly cucullate calyptra. It differs consistentlyfrom Calymperes in the latter character. Gemmae are not as uniformly present in species ofSyrrhopodon as in Calymperes.

SELECTED REFERENCE Reese, W. D. 1987. World ranges, implications for patterns of historical dispersal and speciation, andcomments on phylogeny of Syrrhopodon (Calymperaceae). Mem. New York Bot. Gard. 45: 426–445.

Syrrhopodon

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656 ·CALYMPERACEAE

1. Leaves without border of elongate hyaline cells . . . . . . . . . . . . . . . . . . . . . . . . . 6. Syrrhopodon incompletus1. Leaves bordered entirely or in part with elongate hyaline cells.

2. Leaves regularly conspicuously toothed at shoulders . . . . . . . . . . . . . . . . . . . . . . 1. Syrrhopodon texanus2. Leaves entire or irregularly and inconspicuously toothed at shoulders.

3. Leaves completely bordered all around with hyaline cells.4. Distal lamina more or less crispate when dry, folded when moist . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. Syrrhopodon gaudichaudii4. Distal lamina straight or somewhat contorted when dry but never crispate,

plane when moist . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. Syrrhopodon prolifer3. Leaves incompletely bordered; hyaline cells absent on portions of the margins.

5. Distal lamina ligulate; cancellinae rounded distally . . . . . . . . . . . . . . . 4. Syrrhopodon ligulatus5. Distal lamina acuminate; cancellinae narrowly acute . . . . . . . . . . . . 5. Syrrhopodon parasiticus

1. Syrrhopodon texanus Sullivant, Musc. Hepat. U.S.,103. 1856

Plants gregarious, tufted, to deeplycaespitose, dark green to brownishwhen dry, to 3 cm but mostlymuch shorter. Leaves dimorphic,the vegetative 3–4 mm, looselycontorted when dry; distal laminanot reflexed, plane when moist,broadly linear, apex obtuse;margins bordered nearly to apex

with hyaline cells, coarsely toothed-ciliate at shouldersand irregularly so to apex; medial cells distinct, 5 × 5–7µm, bulging papillose adaxially, 1-papillose abaxially;

cancellinae rounded or ending in acute angles distally;gemmiferous leaves terminal and in zones along stem,mostly rigidly erect wet and dry, consisting mostly of costawith narrow wings of lamina. Gemmae clavate-fusiform,smooth, abundant, forming pale spheres all around onexcurrent tip of costa. Seta single, 15 mm. Capsuleexserted, 2–5 mm; peristome teeth 120 µm, blunt,segmented.

Mature spores present Oct–Mar. Tree trunks andbases, stumps, rotted logs, soil on banks, and sandstonein forests; low to moderate elevations (0–600 m); Ala.,Ark., Del., Fla., Ga., Ill., Ky., La., Md., Miss., Mo., N.Y.,N.C., Ohio, S.C., Tenn., Tex., Va.; Mexico (Nuevo León).

SYRRHOPODON

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Syrrhopodon

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CALYMPERACEAE 657·The bordered, coarsely toothed leaf margins and

narrow, rigidly erect gemmiferous leaves makeSyrrhopodon texanus easy to recognize. Thegemmiferous leaves arise annually at the stem tips; oldplants may have several zones of gemmiferous leavesalong the stem representing annual increments, theseseparated by zones of vegetative leaves. In larger plants,proximal portions of the stems may be felted withrhizoids. Populations of this moss from interior states inthe flora area are female or apparently asexual; maleplants and sporophytes are known only from a relativelyfew sites along the Atlantic and Gulf coasts, and from asingle station in northwestern Alabama (Marion County).

SELECTED REFERENCE Reese, W. D. 1984b. Reproductivity, fertilityand range of Syrrhopodon texanus Sull., a North American endemic.Bryologist 87: 217–222.

2. Syrrhopodon gaudichaudii Montagne, Ann. Sci.Nat., Bot., sér. 2, 2: 376. 1834

Plants caespitose, dark green tobrownish, to 1 cm. Leavesmonomorphic, 2–3 mm, crispate-contorted when dry; distal laminareflexed at shoulders, usuallyfolded when wet, broadly linear-acuminate, apex obtuse; marginsbordered all around with elongatehyaline cells, mostly toothed only

at apex but rarely with denticulations at shoulders; medialcells obscure because of papillosity, 7.5–8.5 µm, withstout, multifid papillae ad- and abaxially; cancellinaerounded distally. Gemmae clavate-fusiform, smooth,sparse, on adaxial apex of leaf.

Not producing sporophytes in flora area. Bark; lowelevations (ca. 30 m); Fla.; Mexico; West Indies; CentralAmerica; South America; Africa; Pacific Islands (Hawaii).

Syrrhopodon gaudichaudii is widespread and oftenabundant elsewhere in its range but is known in the floraarea only from a few collections from HighlandsHammock, Highlands County. It is similar in a generalway to S. prolifer but differs in its leaves contorted-crispate when dry and folded when moist. The leaves ofS. prolifer are usually flexuous or straight when dry andlie flat on the microscope slide when wet.

3. Syrrhopodon prolifer Schwägrichen, Sp. Musc. Frond.Suppl. 2(1,2): 99. 1824

Varieties 3 (1 in the flora): se United States; Mexico;West Indies; Central America; South America; Asia;Africa; Australia.

SELECTED REFERENCE Orbán, S. and W. D. Reese. 1990. Syrrhopodonprolifer (Musci: Calymperaceae): A world view. Bryologist 93: 438–444.

3a. Syrrhopodon prolifer var. scaber (Mitten) W. D.Reese in Organization for Flora Neotropica, Fl.Neotrop. 58: 13. 1993

Syrrhopodon scaber Mitten, J. Linn.Soc., Bot. 12: 119. 1869

Plants tufted or in loose to densesods, pale glaucous green whendry, 0.5–1 cm. Leaves mono-morphic, 2–3 mm, erect andstraight or flexuous when dry;distal lamina not or only slightlyreflexed at shoulders, plane when

wet, narrowly linear, apex rounded; margins borderedall around with elongate hyaline cells, toothed to spinoseonly at apex; medial cells obscure due to papillosity, 7–9µm, with bulging multifid papillae ad- and abaxially;cancellinae ending in acute angles distally. Gemmaeclavate, smooth, very sparse, adaxial and sometimesabaxial on leaf apex.

Not producing sporophytes in flora area. Sandy soil,roof of sandstone cave, humid forests in ravines, alongbases of bluffs; low elevations (0–100 m); Ala., Fla., La.;Mexico; West Indies; Central America; South America.

Syrrhopodon prolifer var. scaber is easy to recognizeby its pale color, grass-like aspect of the colonies due tothe narrow erect leaves, and leaves bordered all aroundwith elongate hyaline cells and spinose-toothed at apices.

4. Syrrhopodon ligulatus Montagne, Syll. Gen. Sp.Crypt., 47. 1856

Calymperes crispum (Austin) Austin;Syrrhopodon crispus Austin

Plants gregarious to caespitose,dark green to brownish green, to0.5 cm. Leaves monomorphic, 2mm, crispate when dry; distallamina not reflexed at shoulders,often folded-falcate when wet,ligulate, apex rounded or retuse;

margins bordered at shoulders and irregularly distallywith hyaline cells, entire or irregularly dentate atshoulders, entire distally except for projecting papillosecells; medial cells obscure due to papillosity, 5.5–6.5 µm,with bulging simple or multifid papillae ad- and abaxially;cancellinae rounded distally. Gemmae clavate, smoothto slightly papillose, sparse, subterminal on adaxialsurface of leaf apex.

Not producing sporophytes in flora area. Fissures oftree bark, sandstone, humid forests along rivers and inswamp forests; low elevations (0 m); Fla., Ga., La., Miss.;Mexico; West Indies; Central America; South America.

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658 ·CALYMPERACEAE

Syrrhopodon ligulatus is not often collected becauseof its small size and obscure habitat, which is mostly onbark fissures. The short-ligulate bluntly tipped leaves,crispate when dry, and with restricted hyaline border,make this species easy to recognize.

5. Syrrhopodon parasiticus (Bridel) Bescherelle, Ann.Sci. Nat., Bot., sér. 8. 1: 298. 1895

Bryum parasiticum Bridel, Muscol.Recent. 2(3): 54. 1803; Calymperesfiligera Austin; Calymperopsisparasitica (Bridel) Brotherus;Syrrhopodon filigerus (Austin)R. S. Williams

Plants solitary, gregarious, ortufted, dark green to brownish, to2 cm but mostly much shorter.

Leaves dimorphic; vegetative 2–4 mm, loosely contortedwhen dry; distal lamina somewhat reflexed aboveshoulders, often somewhat folded when wet, oblong tobroadly lanceolate, apex acute; margins mostly borderedentirely or in part with hyaline cells at least on someleaves, entire; medial cells distinct, 8–10 µm, bulgingadaxially, smooth to 1-papillose abaxially; cancellinaeending in acute angles distally; gemmiferous leaves oftenaggregated into splash-cup comae at stem tips, tightlyappressed when dry, spreading-ascending when wet,oblong-deltoid, acute. Gemmae filamentous, smooth,abundant, adaxial along costa of distal lamina. Seta singleor 2–3 per perichaetium, 2–3 mm. Capsule emergent toexserted, 1–1.5 mm; peristome vestigial.

Capsules mature Mar. Twigs and bark of shrubs andsmall trees, humid forests; low elevations (0–100 m); Ala.,Fla., Ga., La., Miss., Tex.; Mexico; West Indies; CentralAmerica; South America; Asia; Africa; Australia.

In dry plants of Syrrhopodon parasiticus, the abaxialsurface of the costa is often very conspicuous, an aid toidentification in the field. In the northern portion of itsrange plants are mostly small and inconspicuous, but insouthern Florida the plants are often robust, tufted, andconspicuous. Plants with sporophytes are restricted tosouth of Highlands County, Florida. Robust plants insouthern Florida often have teniola-like intramarginalfetures in the proximal parts of the leaves.

SELECTED REFERENCE Reese, W. D. 1959. Syrrhopodon parasiticusin the southern United States. Bryologist 62: 182–186.

6. Syrrhopodon incompletus Schwägrichen, Sp. Musc.Frond. Suppl. 2(1,2): 119. 1824

Syrrhopodon floridanus Sullivant

Plants gregarious to denselycaespitose, dark green to brownishwhen dry, mostly 1–2 cm. Leavesmonomorphic, mostly 3–5 mm,often secund to circinate at stemtips when dry; distal lamina notreflexed at shoulders, plane orchanneled when wet, lanceolate to

linear-lanceolate, apex broadly acute; margins absenthyaline cells, thickened and with two rows of teethdistally; medial cells distinct, 7.5–8.5 µm, bulgingadaxially, plane to minutely 1-papillose abaxially;cancellinae rounded distally. Gemmae fusiform,papillose, sparse, terminal and subterminal on adaxialapex of leaf. Seta single, 7–12 mm. Capsule well exserted,1.5–2.5 mm; peristome absent or rudimentary.

Capsules mature year around. Tree trunks, especiallypalms, logs, and stumps, low and tall forests; lowelevations (0–50 m); Ala., Fla., Ga., La., Miss., N.C.,S.C., Va.; Mexico; West Indies; Bermuda; CentralAmerica; South America; c, e tropical Africa.

Fertile specimens of Syrrhopodon incompletus areknown in the flora area only from Florida. This is aweedy moss of low humid forests. It is the only one ofits genus in the flora area that lacks hyaline cells on theleaf margins. Syrrhopodon incompletus is representedin the flora area by the var. incompletus. H. A. Crumand L. E. Anderson (1981) noted that attribution of thismoss to New York is probably incorrect. According toits label, a specimen (DUKE) was collected in 1935 atAmityville, on Long Island, by M. L. Wickes. Thespecimen consists of three plants, two of them withsporophytes. It is very unlikely that this southern mosswould occur in New York, let alone produce sporophytesthere.

SELECTED REFERENCE Reese, W. D. and S. Orbán. 1986. Syrrhopodonincompletus Schwaegr. var. incompletus (Musci: Calymperaceae) disjunctbetween the Americas and Africa. Bryologist 89: 213–214.

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Syrrhopodon

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CALYMPERACEAE 659·

2. CALYMPERES Swartz in F. Weber, Tab. Calyptr. Operc., 3. 1814 · [Greek kalymma,

covering, and peiro, pierce, alluding to fissured calyptra]

Plants small to medium-sized, mostly tufted, green to yellowish brown, rarely with pink to purpletinge. Stems erect. Leaves often dimorphic with vegetative leaves mostly grading intogemmiferous leaves, teniolate or teniolae absent; margins of distal lamina without elongatehyaline cells, mostly thickened and toothed, rarely 1-stratose and entire; costa mostly showingtwo bands of stereid cells or stereids rarely absent; medial cells of leaves isodiametric [rarelytransversely elongate]. Specialized asexual reproduction common, by fusiform-clavate gemmaeborne on apices of often modified leaves. Sexual condition dioicous. Seta single. Capsulemostly well-exserted, sometimes more or less immersed, cylindric; peristome absent. Calyptrapersistent, enclosing the capsule, with vertical slits distally, spirally plicate when dry.

Species ca. 40 (5 in the flora): worldwide except Antarctica, mostly tropical and subtropicalregions.

Calymperes is easy to recognize when fruiting because of its unique, persistent calyptra; noneof our species, however, bears sporophytes in the flora area. The apex of the calyptra grips theoperculum by its rostrum and under dry conditions lifts the operculum away from the mouthof the capsule, allowing the spores to escape through the gaping fissures in the calyptra. Whenthe sporophyte is wet, the mouth of the capsule is sealed by the operculum, and the fissures inthe calyptra are closed. Sterile specimens of teniolate species of Calymperes are easy to recognizebecause of their characteristic teniolae. Our only nonteniolate Calymperes—C. tenerum—isusually easily recognizable by the excurrent costa of the gemmiferous leaves bearing gemmaeall around. Fertile specimens of Calymperes are not known from the flora area.

SYRRHOPODON ° CALYMPERES

Calymperes

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660 ·CALYMPERACEAE

SELECTED REFERENCE Reese, W. D. 1987b. Calymperes (Musci: Calymperaceae): World ranges, implications for patterns ofhistorical dispersal and speciation, and comments on phylogeny. Brittonia 39: 225–237.

1. Leaves without teniolae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. Calymperes tenerum1. Leaves with teniolae.

2. Distal cells of cancellinae prominently mammillose adaxially.3. Gemmae borne all around excurrent tip of costa; costa in cross section showing

well-defined abaxial and adaxial bands of stereid cells . . . . . . . . . . . . . . . . . . 2. Calymperes erosum3. Gemmae only adaxial on tip of costa; costa in cross section absent well-defined

bands of steroids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. Calymperes pallidum2. Distal cells of cancellinae plane.

4. Cancellinae forming acute angles with costa distally; teniolae usually 4 or morecells in from margin at shoulders . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Calymperes afzelii

4. Cancellinae forming very broad angles with costa distally; teniolae 2–3 cells infrom margin at leaf shoulders . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. Calymperes palisotii

1. Calymperes afzelii Swartz, Jahrb. Gewächsk. 1: 3,plate 1. 1818

Calymperes brittoniae Bescherelle;C. donnellii Austin

Plants gregarious or tufted, dark-green to yellowish brown,uncinate-curved at tips when dry,to 10 mm. Leaves dimorphic, thevegetative 3–4 mm; distal laminalanceolate; margins thickened andtoothed distally; costa in cross

section showing ad- and abaxial bands of stereid cells;medial cells obscure, 5–6 µm, bulging adaxially, minutelypapillose abaxially; teniolae prominent at leaf shouldersand above; cancellinae ending in acute angles distally,adaxial cells smooth distally; gemmiferous leaves withconstricted apices consisting of the costa with narrowlyrevolute marginal laminae, bearing gemmae only onadaxial surface of apex.

Not producing sporophytes in flora area. Tree trunks,bases, exposed roots, and rotted wood, hammocks andforests; low elevations (0 m); Fla.; Mexico; West Indies;Central America; South America; Asia; Africa; IndianOcean Islands; Pacific Islands; Australia.

In the flora area, Calymperes afzelii occurs sparinglyin peninsular Florida, but it is weedy in other parts of itswide range. It is somewhat similar to C. erosum buteasily distinguished by the smooth distal adaxial cells ofits cancellinae, which are distally acute, and by its gemmaeborne only on the adaxial surface of the leaf apex.

2. Calymperes erosum Müller Hal., Linnaea 21: 182.1848

Calymperes emersum Müller Hal.

Plants gregarious or tufted, greento brownish, often somewhatuncinate-curved at tips when dry,to 5 mm. Leaves dimorphic, thevegetative 2–3 mm; distal laminabroadly to narrowly lanceolate;margins somewhat thickened andtoothed distally; costa in cross

section showing ad- and abxaxial bands of stereid cells;medial cells obscure or distinct, 5–6 µm, minutelypapillose ad- and abaxially; teniolae usually distinct atleaf shoulders and beyond but sometimes weak orinterrupted; cancellinae usually broadly rounded distally,adaxial distal cells distinctly mammillose; gemmiferousleaves with apex of costa excurrent, bearing gemmae allaround on apex of costa.

Not producing sporophytes in the flora area. Treebark, bases, and logs, hammocks and forests; lowelevations (0 m); Fla.; Mexico; West Indies; CentralAmerica; South America; Europe; Asia; Africa; AtlanticIslands; Indian Ocean Islands; Pacific Islands; Australia.

Calymperes erosum occurs rarely in peninsular Florida,but it is common in other parts of its range outside of theflora area. It may superficially resemble C. afzelii butcan be easily distinguished by the mammillose distaladaxial cells of its cancellinae, and by bearing gemmaeall around on the excurrent apex of the costa ofgemmiferous leaves. The distal ends of the cancellinaeare usually rounded, and the plants often have a pinkishtinge.

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Calymperes

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CALYMPERACEAE 661·

3. Calymperes palisotii Schwägrichen, Sp. Musc. Frond.Suppl. 1(2): 334. 1816

Calymperes palisotii subsp. richardii(Müller Hal.) S. R. Edwards;C. richardii Müller Hal.

Plants gregarious or tufted,brownish green, uncinate-curvedat tips when dry, to ca. 10 mm butmostly shorter. Leaves dimorphic;vegetative 2–3 mm; distal laminaoblong to oblong-lanceolate;

margins slightly thickened distally, entire; costa in crosssection showing ad- and abaxial bands of stereid cells;medial cells distinct, 7–9 µm, bulging adaxially, smoothto slightly bulging or minutely 1-papillose abaxially;teniolae usually distinct at leaf shoulders and above,sometimes weak or interrupted; cancellinae ending inbroad angles distally, often appearing truncate, adaxialcells smooth distally; gemmiferous leaves with apicesconstricted and often with narrow revolute marginallaminae, bearing gemmae only on adaxial surface in ausually conspicuous hairbrush-like tuft.

Not producing sporophytes in flora area. Tree trunks

and logs in hammocks and forests; low elevations (0 m);Fla.; Mexico; West Indies; Central America; SouthAmerica; Asia; Africa; Pacific Islands.

In the flora area, Calymperes palisotii is rather frequentin peninsular Florida, especially in the southern part,including some of the upper Keys. Its broad, more orless distally truncate cancellinae and gemmae in ahairbrush-like tuft on the adaxial surface of the leaf apexmake it easy to recognize. Calymperes palisotii isrepresented in the flora area by the subsp. palisotii.

4. Calymperes tenerum Müller Hal., Linnaea 37: 174.1872

Calymperes nashii R. S. Williams; C.tenerum subsp. edamense Fleischer

Plants gregarious or tufted, palegreen to brownish, straight orcurved at tips when dry, to 5 mmor more but mostly much shorter.Leaves dimorphic; vegetative 1–3mm; distal lamina oblong tobroadly linear-lanceolate; margins

1-stratose or slightly thickened distally, entire; costa incross section showing ad- and abaxial bands of stereidcells; medial cells distinct, 7–8 µm, slightly bulging

CALYMPERES ° OCTOBLEPHARUM

Calymperes

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662 ·CALYMPERACEAE

adaxially, smooth or minutely papillose abaxially;teniolae absent; cancellinae ending in broad anglesdistally, adaxial cells smooth distally; gemmiferous leaveswith apex of costa excurrent, bearing gemmae in palegolfball-like spheres all around on costa apex.

Not producing sporophytes in flora area. Tree trunksand logs in hammock vegetation; low elevations (0 m);Fla.; Mexico; West Indies (Grand Bahama, Haiti, PuertoRico); South America (s Brazil); Asia; Africa; IndianOcean Islands; Pacific Islands; Australia.

Calymperes tenerum is very rare in the flora area,where it is known only from a few specimens fromsouthern peninsular Florida. Its absence of teniolae,narrow cancellinae, and gemmae in golfball-like spheresmake it distinctive. This species has a very widedistribution in the tropical parts of the world and is verycommon in the paleotropics; it is extremely scarce in theNeotropics, suggesting that it is an introduction there.

5. Calymperes pallidum Mitten, Philos. Trans. RoyalSoc. London 168(extra vol.): 388. 1879

Plants gregarious and tufted, palegreen, erect when dry, to 7 mm.Leaves slightly dimorphic;vegetative 2–2.5 mm; distal laminaoblong-lanceolate; margins 1-stratose to slightly thickeneddistally, irregularly serrate; costain cross section absent well-defined stereid cells; medial cells

distinct, 8–10 µm, bulging and 1- to pluripapilloseadaxially, bulging to 1-papillose abaxially; teniolae

distinct at leaf shoulders and above; cancellinae roundedor ending in acute angles distally, adaxial distal cellsmammillose at least in part; gemmiferous leaves withnarrowed apices, bearing gemmae mostly adaxially butalso abaxially on apices.

Not producing sporophytes in flora area. Tree bark;low elevations (0 m); Fla.; West Indies; South America;Africa; Asia.

Calymperes pallidum is known in the flora area onlyfrom one collection from Collier County, southernFlorida. The species is very widespread but almost alwaysrare in tropical regions around the world. Its mostdistinctive feature is the absence of well-defined stereidcells in the costa in contrast to our other species ofCalymperes, which uniformly show two distinct bandsof stereids in the costa. Calymperes erosum is similar toC. pallidum in having teniolae and with distal cells ofthe cancellinae mammillose, but its costa has strong bandsof stereids. In C. pallidum the apices of the vegetativeleaves are often rounded and sometimes almost cucullate.

Calymperes