Kew Bulletin

, Volume 65, Issue 2, pp 137–188

Megalastrum (Dryopteridaceae) in Central America

Authors

    • The New York Botanical Garden
  • Jefferson Prado
    • Instituto de Botânica
Article

DOI: 10.1007/s12225-010-9214-3

Cite this article as:
Moran, R.C. & Prado, J. Kew Bull (2010) 65: 137. doi:10.1007/s12225-010-9214-3

Summary

Twenty-one species of Megalastrum are here recognised for Central America. A dichotomous indented key is given to identify the species, and each species is provided with a description, complete synonymy, discussion, specimens examined, and illustrations. Eight new species are described: Megalastrum apicale R. C. Moran & J. Prado, M. costipubens R. C. Moran & J. Prado, M. glabrum R. C. Moran & J. Prado, M. gompholepis R. C. Moran & J. Prado, M. intermedium R. C. Moran & J. Prado, M. longiglandulosum R. C. Moran & J. Prado, M. mexicanum R. C. Moran & J. Prado, and M. sparsipilosum R. C. Moran & J. Prado. Two new combinations are made: M. galeottii (M. Martens) R. C. Moran & J. Prado and M. heydei (C. Chr.) R. C. Moran & J. Prado. The central mountains of Costa Rica and Panama are the most species-rich region in Central America, harbouring 15 species, 10 of which are endemic.

Key words

Fernsfloristicssystematicstaxonomy

Introduction

Megalastrum Holttum is a genus of medium- to large-sized ferns found terrestrially (rarely saxicolous) in wet forests. The genus consists of about 90 species, all of which are neotropical except for 3 species in the Africa-Madagascar region (Rouhan & Moran 2011), and 6 on islands in the southern Pacific, Atlantic and Indian Oceans. The best character to distinguish the genus is venation: as one goes distally along the pinna rachis, the basal basiscopic lobe of the pinnules gradually becomes adnate and decurrent to the pinna rachis (Fig. 3D). This adnate lobe is nourished by a vein that arises from the pinna rachis, not the costule. Among ferns, this character combination is unique. Other helpful characters to distinguish the genus, found in most but not all species, are ciliate laminar margins (Fig. 6C) and veins that end well before the margins in prominent hydathodes (Fig. 3N). The rachis of the lamina, pinna rachises, and costules are typically pubescent adaxially, the hairs often coarse, whitish, and multiseptate. Most species have perispores that are either spiny or cristate with parallel ridges (see the over 250 images of Megalastrum spores available at www.plantsystematics.org).

The above characteristics (except those of the spores) were first pointed out by Christensen (1913, 1920), who treated the species now called Megalastrum as the “Group of Dryopteris subincisa” of Dryopteris subgen. Ctenitis. He assigned about 30 species to the group, citing 7 of the species as occurring in Central America. Smith & Moran (1987) cited 12 taxa for Central America and later accepted 9 in their treatment of the genus for Flora Mesoamerica (Smith & Moran 1995).

The purpose of the present study is to review the species of Megalastrum in Central America, examine all relevant types, and provide an up-to-date taxonomic treatment. This study is one in a series of papers treating the genus in large geographic regions. So far we have treatments of the 18 species of Megalastrum in Brazil, Paraguay, and Uruguay (Moran et al.2009a) and the 9 species in the West Indies (Moran et al.2009b). Other papers in this series will treat separately the remaining species of the genus; namely, those that occur in Africa-Madagascar-La Réunion (Rouhan & Moran 2011), tropical South America, and Chile and the southern oceanic islands (Sundue et al.2010). Between any two of these regions there is little or no overlap of species.

Methods

Herbarium specimens were borrowed from 24 herbaria (see Acknowledgments). To show the variation in laminar cutting, silhouettes were prepared from herbarium specimens for all species. To do this, digital images were taken of lamina dissection (usually of the basal pinnae), and the images were then adjusted in PhotoShop to provide a white background and a black lamina. In the specimens examined section, geographic coordinates often had to be estimated because this information was not provided on the labels. The estimates are given in brackets for the herbarium specimens cited below.

Results

In total, the herbarium specimens studied represent 665 separate gatherings. We recognise 21 species, without any varieties or subspecies, and in Appendix I give a synopsis of their distributions in the Central America. In Central America the most species-rich region is the Talamanca mountains of Costa Rica and Panama, which harbour 15 species, 10 of which are endemic. A list of collectors and their numbers is given in Appendix II. The 21 species of Megalastrum in Central America are keyed out and described below (Figs 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15 and 16; Maps 1, 2 and 3).
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Fig. 1

Details of three species of Megalastrum. A – EM. gompholepis (Luteyn & Almeda 3471, F). A lamina division, basal and suprabasal pinnae; B abaxial surface of pinnules and rachises; C scale from rachis of lamina; D adaxial surface of pinna rachis and pinnules; E adaxial surface of pinna rachis; F – JM. squamosum (Herrera 8717, K); F petiole, densely scaly; G petiole scale; H abaxial surface of pinnules and rachises; J adaxial surface of pinna rachis and pinnules; K – QM. apicale (Lellinger & White 1480, US); K abaxial surface of pinnules; L detail of abaxial surface showing scattered sessile, spherical glands; M indusiate sorus; N segment with sori restricted toward apex; P adaxial surface of pinnules; Q adaxial surface of the midrib of the pinnule. drawn by haruto fukuda.

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Fig. 2

Basal pinnae of two species of Megalastrum. AM. dentatum (Grayum 7093, MO); BM. apicale (Herrera et al. 9094, K).

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Fig. 3

Indument details of three species of Megalastrum. A – FM. atrogriseum (Skog et al. 4055, US). A petiole base; B petiole scale; C adaxial surface of lamina; D, E abaxial surface of lamina; F hairs from lamina tissue between the veins; the one on the left is capitate-glandular; G – JM. glabrum (Palmer 125, NY). G adaxial surface of lamina; J, H abaxial surface of lamina; K – NM. mexicanum (Sinaca C. 1154, MO). K abaxial surface of lamina; L scale from rachis of the leaf; M hair from abaxial surface lamina; N adaxial surface of lamina. drawn by haruto fukuda.

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Fig. 4

Lamina division of four species of Megalastrum. AM. glabrum, basal pinna (Sundue et al. 1737, NY); BM. atrogriseum, leaf (Maxon & Harvey 8321, US); CM. mexicanum, basal pinna (Sinaca C. 1154, MO); DM. squamosum, basal pinna, note densely scaly rachis of the lamina (Herrera & Solís 2454, MO).

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Fig. 5

Indument characters of Megalastrum atrogriseum.A abaxial surface of rachis, costa, and pinnule; B rachis scale; C abaxial surface of lamina showing mixed glandular and non-glandular hairs; D gland-tipped and non-glandular hairs; E adaxial surface of rachis, costa, and pinnule; F adaxial surfaces of rachis and costa showing dense pubescence. All from Croat 15683 (MO). drawn by haruto fukuda.

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Fig. 6

Indument details of three species of Megalastrum. A – FM. galeottii (Wercklé s.n., NY). A abaxial surface of lamina; B scale from rachis of leaf; C abaxial surface of lamina; D sori showing minute, fugacious indusia; E adaxial surface of lamina; F adaxial surface of costule; G – HM. costipubens (Araya & Rojas 919, MO); G abaxial surface of lamina; H adaxial surface of lamina; J – MM. dentatum (Grayum 7367, MO); J petiole scale; K base of petiole; L adaxial surface of lamina; M abaxial surface of lamina. drawn by haruto fukuda.

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Fig. 7

Lamina division of three species of Megalastrum. AM. galeottii, basal pinna (Mickel & Leonard 5230, MICH); BM. costipubens, basal and suprabasal pinna (Scamman & Holdridge 7962, GH); CM. longipilosum, basal pinna (Burger & Stolze 5054, GH); DM. longipilosum, basal pinna (Burger & Stolze 5054, F).

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Fig. 8

Lamina division of three species of Megalastrum. AM. macrotheca, leaf (Stolze 1495, US); BM. macrotheca, basal and suprabasal pinnae (Gómez 19290, MO); CM. macrotheca, basal and suprabasal pinnae (Gómez 19291, MO); DM. ctenitoides, medial pinnae (Herrera & Chacón 2794, CR); EM. reductum, basal and suprabasal pinnae (Moran 5060, MO).

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Fig. 9

Indument in three Central American species of Megalastrum. A – GM. ctenitoides.A abaxial surface of rachis, costa, and pinnules; B abaxial surface of rachis; C base of costule abaxially; D scale from rachis; E hairs and scales of costule abaxially; F adaxial surface of segment showing glabrous tissue and hydathodes; G adaxial surface of rachis; H – LM. reductumH abaxial surface of rachis, costa, and pinnules; J filiform scale from rachis; K abaxial surface of costa; L adaxial surface of rachis, costa, and pinnule. M – TM. macrothecaM adaxial surface of rachis, costa, and pinnules; N hair from adaxial surface of rachis; P rachis scale; Q adaxial surface of segment apex; R adaxial surface of costule; S abaxial surface of rachis, costa, and pinnules; T adaxial surface of lobe. A – G from Colombia, MacDougal et al. 3886 (MO), H – L from Moran 5060 (AAU), M – T from Haiti, Ekman 5315 (S). drawn by haruto fukuda.

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Fig. 10

Lamina dissection in two species of Megalastrum. AM. subincisum (Fink 59, NY); BM. gompholepis (Williams et al. 25586, F).

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Fig. 11

Lamina division of three species of Megalastrum. AM. sparsipilosum, medial pinna (Breedlove & Smith 22622, F); BM. heydei, medial pinna (Heyde 310, US); CM. pulverulentum, basal basiscopic pinnule of basal pinna (Ekman 5747, NY).

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Fig. 12

Indument in three Central American species of Megalastrum. A – HM. heydei.A abaxial surface of rachis, costa, pinnules; B hairs of rachis; C base of costa; D strongly denticulate scale from costa abaxially; E abaxial surface of segment; F puberulent adaxial surface of costa; G adaxial surface of costule; H adaxial surface of segment midrib. J – NM. sparsipilosum. J abaxial surface of rachis, costa, pinnules; K strongly denticulate scale from costa abaxially; L detail of abaxial surface of pinnule; M adaxial surface of rachis, costa, and pinnule; N adaxial surface of costa. P – WM. pulverulentum.P adaxial surface of rachis, costa, and pinnule; Q hairs from adaxial surface of costule; R adaxial surface of segment apex; S adaxial surface of rachis; T abaxial surface of rachis, costa, and pinnule; U abaxial surface of rachis, note sparse sessile glands; V gland-tipped hairs from abaxial surface of rachis; W abaxial surface of costule and lamina tissue showing glandular and non-glandular hairs. A – H from Heyde & Lux 3249 (GH), J – N from Breedlove & Smith 32364 (F), P – W from Dominican Republic, Zanoni et al. 18641 (NY). drawn by haruto fukuda.

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Fig. 13

Indument detail of three species of Megalastrum. A – GM. lunense (Herrera Ch. et al. 591, AAU). A basal pinna; B, C petiole and scales; D, E abaxial surface of lamina; F hairs from abaxial surface of lamina between the veins; G adaxial surface of lamina; H – KM. palmense (Mickel 3574, MO). H adaxial surface of the leaf; J, K abaxial surface of the leaf; note short strigose hairs on axes; L, MM. intermedium (Navarro & Rojas 916, K). L abaxial surface of leaf; note capitate-glandular hairs; M adaxial surface of leaf. drawn by haruto fukuda.

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Fig. 14

Silhouettes of two species of Megalastrum. AM. intermedium, medial pinna, acroscopic side on left (Killip 5136, MICH); BM. lunense, leaf (Moran 3186, NY).

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Fig. 15

Indument in three Central American species of Megalastrum. A – EM. longipilosum.A abaxial surface of rachis, costa, pinnules; B rachis scale; C hairs of the costa; D adaxial surface of rachis, costa, pinnules; E hair from adaxial surface of rachis. F – MM. longiglandulosum.F abaxial surface of rachis, costa, pinnules; G hairs on adaxial surface of costules; H adaxial surface of costa; J abaxial surface of rachis, costa, pinnules; K abaxial surface of pinnule showing stalked glandular hairs; L scale from costule; M hairs, gland-tipped and non-glandular, from abaxial surface of lamina. N – VM. subincisum. N adaxial surface of pinnule; P rachis scale; Q hairs on adaxial surface of costule; R segment apex; S adaxial surface of costule; T abaxial surface of pinnule; U abaxial surface of costa; V abaxial surface showing appressed proscales. A – E from Moran 3162 (MO), F – M from Burger & Stolze 5421 (MO), N – V from Jamaica, Clute 132 (US). drawn by haruto fukuda.

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Fig. 16

Lamina dissection in Megalastrum palmense. A proximal pinnae (Maxon 303, NY); B proximal portion of basal pinna (Sundue et al. 1783, NY); C medial pinnules from medial pinna (Lellinger 1379, F).

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Map 1

Distribution of Megalastrum in Central America and the distributions of seven of its species.

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Map 2

Distribution of eight species of Megalastrum in Central America.

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Map 3

Distribution of six species of Megalastrum in Central America.

Key to the Species of Megalastrum in Central America

  • 1. Indusia present and conspicuous, circular, 0.5 – 0.7 mm diam.; sori restricted to the apical portion of the segments; Costa Rica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. M. apicale

  • 1. Indusia absent or inconspicuous, hair-like, minute; sori throughout the length of the segments; Costa Rica and elsewhere

  • 2. Laminae at base 1-pinnate-pinnatifid to 2-pinnate-pinnatisect, medially 1-pinnate-pinnatifid to 2-pinnate

  • 3. Pinna rachises densely pubescent adaxially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14. M. macrotheca

  • 3. Pinna rachises glabrous to subglabrous adaxially, the hairs when present only at base of pinnae rachis

  • 4. Hairs on both surfaces of the leaf rachises 0.1 – 0.2 mm long, 1- or 2-celled, acicular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18. M. reductum

  • 4. Hairs on both surfaces of the leaf rachises c. 1 mm long, 6 – 8-celled, strongly jointed (twisted at the septae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4. M. ctenitoides

  • 2. Laminae at base 3-pinnate-pinnatifid to 4-pinnate-pinnatisect, medially 2- to 3-pinnate-pinnatisect

  • 5. Hairs on the pinna rachises adaxially 0.1 – 0.2 mm long, 1- or 2 (– 4)-celled

  • 6. Laminae between the veins on both surfaces puberulent, the hairs c. 0.1 mm long, acicular, erect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. M. intermedium

  • 6. Laminae between the veins on both surfaces glabrous

  • 7. Leaves up to 1.5 m long; laminae 3-pinnate-pinnatisect at base; scales on the abaxial surface of the pinna rachises inconspicuous, entire, flaccid, appressed; adaxial surface of the veins non-glandular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16. M. palmense

  • 7. Leaves up to 4 m long; laminae 4-pinnate-pinnatisect at base; scales on the abaxial surface of the pinna rachises conspicuous, strongly denticulate, firm, spreading; adaxial surface of the veins glandular, the glands sessile, spherical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. M. heydei

  • 5. Hairs on the pinna rachises adaxially 0.3 – 2.0 mm long, 4 – 12-celled

  • 8. Stalked glands present on the lamina abaxially (these glands c. 0.1 – 0.3 mm long, hairlike with apical cell shiny, enlarged, spherical or clavate)

  • 9. Leaves up to 4 m long; laminae 4-pinnate-pinnatisect at base; basal pinnae up to 1 m long; scales of the pinna rachises abaxially strongly and conspicuously denticulate . . . . . . . . . . . . . . . 17. M. pulverulentum

  • 9. Leaves up to 1.5 m long; laminae 3-pinnate-pinnatisect at base; basal pinnae up to 0.15 – 0.4 m long; scales of the pinna rachises abaxially entire to denticulate

  • 10. Lamina tissue between the veins adaxially pubescent; scales of the pinna rachises c. 1.5 mm wide, ovate to lanceolate, flaccid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. M. lunense

  • 10. Lamina tissue between the veins adaxially not pubescent; scales of the pinna rachises c. 0.2 – 0.4 mm wide, filiform to lanceolate, linear or oblong, firm

  • 11. Lamina abaxially between the veins pubescent with long-stalked glandular hairs only (not intermixed with non-glandular ones); pinna rachis scales dark brown, filiform; Costa Rica (endemic to Osa Peninsula) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. M. longiglandulosum

  • 11. Lamina abaxially between the veins pubescent with long-stalked glandular hairs mixed with acicular ones; pinna rachis scales golden, lanceolate, linear or oblong; Costa Rica, Panama . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. M. atrogriseum

  • 8. Stalked glands absent on the lamina abaxially

  • 12. Scales of the pinna rachises abaxially usually filiform, 1 – 2 × 0.05 – 0.3 mm

  • 13. Pinna rachises abaxially pubescent, often conspicuously and evenly so

  • 14. Hairs on the pinna rachises abaxially 0.6 – 1.3 (– 2) mm long, 4 – 7 (– 12)-celled; veins adaxially pubescent, the hairs 0.5 – 1.5 mm long, 3 – 7-celled . . . . . . . 12. M. longipilosum

  • 14. Hairs on the pinna rachises abaxially 0.2 – 0.6 mm long, 2 – 5-celled; veins adaxially glabrous to pubesecent, the hairs 0.2 – 0.3 (– 0.5) mm long, 2 – 4-celled . . . . . . . . 3. M. costipubens

  • 13. Pinna rachises abaxially glabrous to sparsely pilose

  • 15. Pinna rachises abaxially without hairs; laminar tissue between veins abaxially puberulent, the hairs 0.1 – 0.2 mm long, 1 – 3-celled, acicular, erect or (rarely) glabrous . . . . 6. M. galeottii

  • 15. Pinna rachises abaxially sparsely pilose; laminar tissue between veins abaxially glabrous to sparsely pilose, the hairs 0.5 – 0.8 mm long, 2 – 5-celled, spreading

  • 16. Lamina tissue between the veins abaxially glabrous; hairs of the lamina margins 0.2 – 0.3 mm long; scales of the pinna rachises abaxially 0.2 – 0.3 mm wide, golden brown, not tortuous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. M. glabrum

  • 16. Lamina tissue between the veins abaxially sparsely pilose; hairs of the lamina margins 0.4 – 0.8 mm long; scales of the pinna rachises abaxially 0.1 – 0.2 mm wide, brown, tortuous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. M. mexicanum

  • 12. Scales of the pinna rachises abaxially usually lanceolate, (0.2 –) 2 – 6 × (0.2 –) 0.5 – 0.7 mm

  • 17. Lamina tissue between the veins abaxially pubescent

  • 18. Hairs of the veins adaxially typically 0.3 – 0.4 mm long, 3- or 4-celled . . . . . 20. M. squamosum

  • 18. Hairs of the veins adaxially typically 0.7 – 1.0 mm long, 4 – 6-celled

  • 19. Lamina tissue between the veins adaxially pubescent (especially toward segment apices); scales of the pinna rachises abaxially brown, tortuous . . . . . . . . . . . . . . . 5. M. dentatum

  • 19. Lamina tissue between the veins adaxially glabrous; scales of the pinna rachises abaxially golden brown, flat, firm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. M. atrogriseum

  • 17. Lamina tissue between the veins abaxially glabrous or nearly so (sometimes with uniseriate, reddish appressed proscales)

  • 20. Leaves to 4.6 m long; basal pinnae up to 1 m long; scales of the pinna rachises abaxially strongly denticulate; hairs of the costules adaxially 1.5 – 2.0 mm long, 8 – 12-celled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19. M. sparsipilosum

  • 20. Leaves to 2 m long; basal pinnae up to 0.4 – 0.6 m long; scales of the pinna rachises abaxially entire to denticulate; hairs of the costules adaxially 0.4 – 1.0 mm long, 3 – 8-celled

  • 21. Hairs of the pinna rachises and costules abaxially 0.7 – 1.0 mm long, 6 – 8-celled; scales of the pinna rachises abaxially golden brown; leaves up to 1 m long; Costa Rica, Panama . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. M. glabrum

  • 21. Hairs of the pinna rachises and costules abaxially 0.2 – 0.3 (– 0.5) mm long, 6 – 8-celled; scales of the pinna rachises abaxially dark to light brown; leaves up to 2 m long; Mexico, Guatemala, Honduras

  • 22. Scales of the pinna rachises abaxially light brown, appressed to loosely ascending, lanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21. M. subincisum

  • 22. Scales of the pinna rachises abaxially dark brown, patent, abruptly expanded at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. M. gompholepis

1. Megalastrum apicaleR. C. Moran & J. Pradosp. nov. A Megalastro acrosoro rhachidibus costisque squamis omnino fuscis abaxialiter trichomatibus 0.5 – 0.7 mm longis 3- ad 5-cellularibus vestitis differt. Typus: Costa Rica. Puntarenas: Santa Elena Cloudforest Reserve, 10°20'N, 84°47'W, 2 Feb. 2008, M. Sundue et al. 1743 (holotypus CR!; isotypi INB!, NY!, UC!).

http://www.ipni.org/urn:lsid:ipni.org:names:77105941-1

Rhizomes erect to decumbent; leaves 1.5 – 2.5 m long; scales of the petiole bases 0.6 – 10 × c. 1 mm, linear, twisted, spreading to ascending, dark brown, often shiny denticulate on the margins; laminae 1 – 2 m long, 3-pinnate-pinnatifid at base, 2-pinnate-pinnatifid medially; basal pinnae c. 0.8 m long, strongly inequilateral (elongated basiscopically); pinna rachises abaxially very sparsely glandular, moderately scaly, densely puberulent, the glands 0.1 mm diam., globose, sessile, pale yellowish, shiny, the scales 4 – 5 × 0.5 mm long, lanceolate-oblong, mostly flat (not twisted), sparsely denticulate, dark brown, dull or shiny, the hairs on the abaxial surfaces 0.5 – 0.7 mm long, 1 – 3-celled, the hairs on the adaxial surfaces c. 0.7 – 1.0 mm long, 4 – 6-celled; costules abaxially densely puberulent, moderately scaly, very sparsely glandular, the hairs 0.4 – 0.6 mm long, 1 – 3-celled, spreading to erect, the scales 1.0 – 1.2 mm long, lanceolate (sometimes with an acuminate apex), mostly spreading, brown throughout (not just at point of attachment), entire to sparsely denticulate, adaxially pubescent, the hairs 0.7 – 0.9 mm long, 4 – 6-celled; laminar tissue between the veins abaxially pubescent, the hairs 0.2 – 0.3 mm long, erect, acicular, 1 – 3-celled, glands absent to sparse, yellow to orange, globose, sessile, proscales c. 0.2 mm long, appressed, sparse, adaxially sparsely puberulent, the hairs c. 0.1 – 0.2 mm long, 1- or 2-celled, acicular, erect, glands absent or sparse, globose, sessile, pale yellow; veins obscure on both surfaces, with indumenta on both surfaces like that of the costules; hydathodes evident; lamina margins ciliate, the hairs 0.2 – 0.3 mm long, 3- or 4-celled, non-glandular; indusia present, 0.5 – 0.7 mm wide, circular, dark brown, sparsely pubescent and very sparsely glandular, the hairs 0.2 – 0.3 mm long, 1- or 2-celled, acicular, the glands globose, orangish. Figs 1K – Q and 2B; Map 1.

Distribution. Costa Rica.

Additional Specimens Examined. Costa Rica. Alajuela: Reserva Forestal San Ramón, N of the station, c. 2 km in the mountains, 10°12′40″N, 84°36′20″W, 1000 – 1200 m, 17 April 1991, Bittner 974 (AAU, NY); Reserva Forestal San Ramón, c. 3 km N of station, 10°12′40″N, 84°36′20″W, 1000 – 1200 m, 1 June 1991, Bittner 1058 (CR); La Palma de San Ramón, [10°08′N, 84°33′W], 26 Oct. 1927, Brenes 5766 (F, GH, NY); El Silencio (Los Angeles) de San Ramón, [10°10′N, 84°28′W], 20 Feb. 1933, Brenes 17095 (NY, US); El Silencio (Los Angeles) de San Ramón, [10°10′N, 84°28′W], 20 Feb. 1933, Brenes 17097 (F); La Palma de San Ramón, [10°08′N, 84°33′W], 26 Oct. 1927, Brenes 5766A (NY); near the Continental Divide about 2 to 5 km E and SE of Monteverde, 10°18′N, 84°46′W, 1580 – 1700 m, 17 – 20 March 1973, Burger & Gentry 8696 (CR, F, MO, NY); Along road betwen San Ramón and Bajo Rodríguez, Vic. of Km markers 10 – 11 NW of San Ramón, 10°10′40″N, 84°34′10″W, 1100 m, 3 Sept. 1996, Croat 78854 (MO); 7 miles N of San Ramón, [10°05′N, 84°28′W], 1000 m, 27 July 1967, Evans & Bowers 2943 (MO); Reserva forestal de San Ramón, Río San Lorencito, 10°12′53″N, 84°36′28″W, 800 – 1000 m, 5 Dec. 1986, Herrera et al. 347 (AAU, MO, NY); 11 km N of San Ramón, [10°05′N, 84°28′W], 1000 m, 28 July 1967, Lellinger 750 (CR, MO, US); Cantón de San Ramón. R. B. Monteverde, estación La Casona, sendero Orquídeas, 10°20'N, 84°44'W, 1000 m, 3 May 1995, Martínez 452 (CR, INB, MO); 11 km N of San Ramón, [10°05′N, 84°28′W], 1000 m, 28 July 1967, Mickel 2975 (AAU, MICH, NY, US); Cantón de San Ramón, Cuenca del San Carlos, Los Angeles, 2.1 km de la Escuela La Balsa, naciente de la Quebrada Azul, 10°11′15″N, 84°30′30″W, 1070 m, 1 Dec. 1997, Rojas et al. 4076 (CR, INB); c. 1 km SE of La Balsa de San Ramón (c. 16 km NW of San Ramón), 10°10′N, 83°29.5′W, 1140 – 1190 m, 3 Feb. 1986, Smith et al. 2290 (CR, MICH, MO, NY, US). Cartago: About 10 km S of Tapantí along the new road on the E slope above the Rio Grande de Orosi, 9°42′N, 83°47′W, 1400 – 1600 m, 10 – 24 June 1968, Burger & Stolze 5684 (CR, F, GH, US); Platanillo, [9°49′N, 83°24′W], 650 – 900 m, 24 June 1967, de la Sota 5262 (US); SE of Orosi, c. 2.2 km SSE of Purisil, above Finca La Concordia, in the gorge next to the house on the upper most portion of the upper finca, 1800 m, 8 – 10 Aug. 1970, Lellinger & White 1480 (CR, F, MICH, MO, P, US); c. 22 km E of Turrialba, high ridge above Platanillo, [9°49′N, 83°24′W], 1200 – 1450 m, 22 Aug. 1967, Mickel 3431 (NY); near the entrance to Parque Nacional Tapanti, [9°47′N, 83°48′W], 1270 m, 3 Aug. 1983, Moran 3354 (CR, F, MO, NY); Cantón de Paraiso, Parque Nacional Tapantí, Cuenca del Reventazón, camino por las Torres, entre Rancho Negro y la casa del I.C.E., 9°41′29″N, 83°47′08″W, 1800 m, 24 May 1997, Rojas et al. 3515 (INB); Cantón de Paraíso, Cuenca del Reventazón, Orosí, Río Macho, Est. Biol. Río Macho y alrededores, 9°45′56″N, 83°51′48″W, 1650 – 1800 m, 3 Dec. 1997, Rojas et al. 4162 (CR, INB); La Estrella, [9°47′N, 83°58′W], 26, 27 March 1924, Standley 39206 (US); El Muñeco, [9°48′N, 83°55′W], 1500 m, 19 June 1928, Stork 2671 (MICH, US); El Muñeco, valley S of Navarro Valley, near boundary of Cartago and San José provinces, [9°48′N, 83°55′W], 1400 m, 19 June 1928, Stork 2716 (GH, MICH, US); Parque Nacional Tapantí, Carretera ICE, sendero Arboles Caidos, 9°44′58″N, 83°46′53″W, 1296 m, 4 Feb. 2008, Sundue et al. 1750 (CR, INB, NY, USJ). Guanacaste: Reserva Santa Elena, Santa Elena de Monteverde, 9°44′N, 82°55′W, 1600 – 1700 m, 23 Nov. 1994, Bittner 2300 (AAU, CR). Heredia: Between Río Peje and Río Sardinal, Atlantic slope of Volcán Barva, 10°15.5′N, 84°05′W, 1300 – 1500 m, 10 Nov. 1986, Grayum et al. 7765 (MO). Limón: Limón, Almirante, Cuenca superior del Río Xichiari, 9°45′50″N, 83°19′45″W, 1300 m, 14 Aug. 1995, Herrera 8461 (CR); El Progresso, cabeceras de Río Cariei, Fila Matama, Valle de La Estrella, 9°47′20″N, 83°08′18″W, 1400 m, 26 April 1989, Herrera & Chacón 2791 (CR); Cantón Limón, R.I. Chrripó, a orillas de una de las quebradas del Río Bolley, 9°45′15″N, 83°18′50″W, 1300 – 1400 m, 16 Aug. 1995, Rojas 2318 (CR, INB, MO); Cantón de Limón, Zona Protectora Río Banano, Cuenca del Banano, Valle de la Estrella, Fila Matma, c. 11 km SW del pueblo de Aguas Zarcas, [10°23′N 84°20′W], 1200 – 1300 m, 22 Oct. 2007, Santamaría 6553 (INB); Cantón de Limón, Zona Protectora Río Banano, Cuenca del Banano, Valle de la Estrella, Fila Matma, c. 11 km SW del pueblo de Aguas Zarcas, punto 20A, [10°23′N 84°20′W], 1300 – 1400 m, 24 Oct. 2007, Santamaría 11465 (INB). Puntarenas: Reserva Biológica Monteverde, Brillante trail, [10°18′N, 84°48′W], 25 April 1988, Bigelow & Kukle 50 (CR); Reserva Monteverde, vertiente Pacífico, creca de División Continental (Ventana), [10°18′N, 84°48′W], 1560 – 1600 m, 28 June 1976, Dryer 414 (F); Puntarenas, La Pitahaya, siguiendo la fila entre Río Aranjuez y Quebrada Vueltas, 10°15′30″N, 84°41′00″W, 1400 m, 30 May 1996, Herrera et al. 9094 (CR, K). San José: La Palma de San Ramón, [10°03′N, 83°59′W], 1400 m, 22 June 1910, Brade 156 (BM, GH, MICH, NY, P, S, US); La Palma, [10°03′N, 83°59′W], 1400 m, 18 Aug. 1909, Brade & Brade 349 (BM, GH, S); La Palma, along the Rio Claro (upper Rio La Hondura) along the trail to Guapiles, 10°03′N, 83°58′W, 1000 m, 1 Jan. 1967, Burger 4141 (CR, F, GH, NY, US); La Palma, NE of San Jerónimo, above La Hondura valley, 10°02′N, 84°00′W, 1500 m, 27 May – 1 June 1968, Burger & Stolze 5326 (CR, F, GH, NY, US); La Hondura, la Palma, [10°04′N, 83°58′W], 2500 m, May 1912, Jiménez 603 (P, US); Vic. of La Hondura, 4 km S to 1 km N on route 220, between Volcán Irazú and Volcán Barba, [10°04′N, 83°59′W], 1250 – 1500 m, 25 March 1967, Mickel 2240 (NY); Vic. of La Hondura, 4 km S to 1 km N on route 220, between Volcán Irazú and Volcán Barba, [10°04′N, 83°59′W], 1250 – 1500 m, 25 March 1967, Mickel 2250 (NY); Along the road to La Hondura, [10°04′N, 83°58′W], 1450 m, 8 April 1956, Scamman & Holdridge 7956 (GH, US); La Hondura, [10°04′N, 83°59′W], 1300 m, 15 Aug. 1933, Valerio 1806 (CR); La Hondura, [10°04′N, 83°58′W], 1100 m, 1907, Wercklé s.n. (US).

Habitat. Not known; 650 – 1800 m.

Conservation Status. Least Concern (LC).

Notes. Megalastrum apicale has long been called M. acrosorum (Hieron.) A. R. Sm. & R. C. Moran (e.g., Smith & Moran 1995), a species that occurs only in Colombia, Ecuador, and Peru (both sides of the Andes). Both species are alike in several respects. Both may have sori restricted towards segment apices (Fig. 1N). This character, to which the specific epithet apicale refers, is constant for M. apicale but variable for M. acrosorum (pers. obs.). All other species of Megalastrum have sori distributed throughout the pinnules, not restricted to the apices. Both M. apicale and M. acrosorum also have firm, circular indusia that are minutely pubescent on the surfaces and margins. Their lamina cutting is nearly identical. M. acrosorum, however, differs by whitish or pale brown laminar scales that are dark only at the point of attachment (vs. brown throughout) and shorter hairs on the abaxial surfaces of the rachises of the laminae and pinnae (0.2 – 0.3 vs 0.5 – 0.7 mm long).

2. Megalastrum atrogriseum (C. Chr.) A. R. Sm. & R. C. Moran (Smith & Moran 1987: 77).

Dryopteris atrogrisea C. Chr. (Christensen 1920: 70, f. 15).

Ctenitis atrogrisea (C. Chr.) Ching (1940: 250). Type: Costa Rica, San José, Tablazo, [9°50′N, 84°03′W], in 1906, P. Biolley 70 (holotype C; isotypes BM!, MO!, RB!, US!).

Aspidium karstenianum var. navarrense H. Christ (1906: 56). Type: Costa Rica, Cartago, Navarro, [9°48′N, 83°53′W], C. Wercklé s.n. (lectotype P!, selected here; duplicates BM!, C, CR!, US!).

Rhizomes erect to decumbent; leaves 0.4 – 1.3 m long; scales of the petiole bases 15 – 20 × c. 1 mm, linear, slightly twisted toward the apex, light brown to golden, often shiny, densely denticulate on margins and often on the surfaces; laminae up to 0.8 m long, 3-pinnate-pinnatifid at base, 2-pinnate-pinnatifid medially; basal pinnae 0.2 – 0.3 m long, strongly inequilateral (elongated basiscopically); pinna rachises abaxially with sparse glandular hairs, moderately scaly, densely pubescent, the glands stalked (representing modified smaller hairs with the terminal cell swollen), the scales to 3 – 5 × 0.3 – 0.4 mm long, lanceolate-oblong, flat (not twisted), denticulate, golden brown, shiny, the hairs on the abaxial surfaces of mixed length, 0.2 – 0.8 (– 1.2) mm long, 2 – 5-celled (6- or 7-celled), the hairs on the adaxial surfaces c. 0.4 – 0.7 mm long, 3 – 5-celled, dense, spreading to erect; costules on both surfaces with indument like that of the pinna rachises; laminar tissue between the veins abaxially pubescent, the hairs 0.2 – 0.3 mm long, 2- or 3-celled, erect, acicular or (especially in shorter hairs) gland-tipped with a swollen brownish terminal cell globose sessile glands absent, proscales c. 0.2 mm long, appressed, sparse, adaxially glabrous, sessile or stalked glands absent; veins evident on both surfaces, pubescent abaxially with hairs like those of the laminar tissue, adaxially very sparsely pubescent, the hairs c. 1 mm long, 4- or 5-celled, spreading; hydathodes evident; lamina margins ciliate, the hairs 0.2 – 0.3 mm long, 1- or 2-celled, acicular, mixed with gland-tipped hairs; indusia minute, fugacious, apparently consisting of a cluster of reddish proscales, c. 0.1 mm long, protruding from the centre of the sorus. Figs 3A – F, 4B, and 5A – F; Map 1.

Distribution. Guatemala, Costa Rica, Panama.

Additional Specimens Examined. Guatemala. Cobán, [14°48′N 89°07′W], 2 Aug. 1886, von Türckheim 796 (P). Costa Rica. Alajuela: Berlin, on Cerro Berlin, 10°03'N, 84°27′W, 1300 – 1400 m, 3 April 1982, Barringer et al. 2310 (CR); Cataratas de San Ramón, [10°13′N, 84°37′W], 3 May 1931, Brenes 13665 (F, NY); upper drainage of Río Peñas Blancas below the Monteverde Cloud Forest Reserve, 9°17′N, 84°16′W, 1250 – 1350 m, 25 – 26 Feb. 1977, Burger et al. 10774 (NY); Monteverde, Sendero Chomogo, 10°15′N 84°50′W, 1575 m, 31 Oct. 1986, Hennipman et al. 6589 (U); Monteverde Cloud Forest Reserve, La Ventana, 10°15′N, 84°50′W, 1600 m, 1 Nov. 1986, Hennipman et al. 6613 (CR, U); Monteverde, 10°15′N 84°50′W, 1400 m, 2 Nov. 1986, Hennipman et al. 6652 (U); Reserva forestal Grecia, bosque de el Niño, 10°08′20″N, 84°15′00″W, 1000 m, 16 Jan. 1987, Herrera et al. 402 (AAU, MO); above road to Peñas Blancas, Monteverde, [10°18′N, 84°46′W], 1450 m, 25 Jan. 1973, James s.n. (MO); S of San Ramón, c. 3 km above San Rafael, [10°04′N 84°28′W], 1200 m, 26 July 1970, Lellinger & White 1336 (CR, F); Parque Nacional Juan Castro Blanco, Cuenca del San Carlos, Cuenca del Río Peje, entrando por San Vicente, 10°32′N, 84°29′W, 1700 m, 18 July 2000, Rodríguez et al. 5931 (CR); Region of Zarcero, [10°11′N, 84°24′W], 1200 m, [no date], Smith 189 (US); Region of Zarcero, Atlantic watershed, [10°11′N, 84°24′W], 1370 m, 20 Jan. 1938, Smith H189 (F); S. Luis de Zarcero, [10°11′N, 84°24′W], 1300 m, 15 Feb. 1938, Smith PC107 (F); Region of Zarcero, [10°11′N, 84°24′W], [no date], Smith PC182 (F). Cartago: Carpintera, [9°53′N, 83°49′W], 1850 m, 25 April 1908, Brade 122 (NY, S); La Carpintera, [9°53′N, 83°49′W], 1700 m, 4 April 1909, Brade & Brade 568 (S); tributary of Quebrada Casa Blanca, Tapantí, 9°47′N, 83°48′W, 1350 m, 6 Aug. 1984, Grayum & Sleeper 3713 (CR, MO); Pacayas, [9°55′N, 83°48′W], 1875 m, 1923, Lankester 652 (BM, US); Conventillos, 12 July 1924, Lankester 886 (BM, US); Tapantí, c. 15 km S of Paraíso, [9°47′N, 83°48′W], 1150 m, 20 March 1967, Mickel 1855 (NY); Tapantí, c. 15 km S of Paraíso, [9°47′N, 83°48′W], 1150 m, 3 July 1967, Mickel 2332 (NY); 1 km SW of Cervantes, [9°53′N, 83°48′W], 1400 m, 11 July 1967, Mickel 2640 (NY); Forests near the entrance to Parque Nacional Tapantí, [9°47′N, 83°48′W], 1270 m, 3 Aug. 1983, Moran 3355 (CR, MO, NY); Tapantí, [9°47′N 83°48′W], 1200 m, 22 Sept. 1964, Nisman 18 (GH); Tapantí, [9°47′N, 83°48′W], 1200 m, 22 Sept. 1964, Nisman 21 (GH); Cantón de Paraiso, Parque Nacional Tapantí, Cuenca del Reventazón, Estación de Biología Tropical Río Macho y alrededores, 9°46′00″N, 83°52′00″W, 1580 – 1630 m, 4 March 1993, Rojas 105 (INB); Cantón de Paraiso, Parque Nacional Tapantí, Cuenca del Reventazón, Estación de Biología Tropical Río Macho y alrededores, 9°46′00″N, 83°52′00″W, 1580 – 1630 m, 4 March 1993, Rojas 149 (INB); Cantón de Paraiso, Parque Nacional Tapantí, Cuenca del Reventazón, Estación de Biología Tropical Río Macho y alrededores, 9°46′00″N, 83°52′00″W, 1250 – 1710 m, 31 March 1993, Rojas 232 (INB); Cantón de Paraiso, Parque Nacional Tapantí, Cuenca del Reventazón, Estación de Biología Tropical Río Macho y alrededores, 9°46′00″N, 83°52′00″W, 1590 – 1800 m, 4 April 1993, Rojas 283 (INB); Cantón de Paraiso, Parque Nacional Tapantí, Cuenca del Reventazón, Estación de Biología Tropical Río Macho y alrededores, 9°46′00″N, 83°52′00″W, 1150 – 1750 m, 19 Aug. 1993, Rojas 378 (INB); Cantón de Paraiso, Parque Nacional Tapantí, Cuenca del Reventazón, Estación de Biología Tropical Río Macho y alrededores, 9°46′00″N, 83°52′00″W, 1150 – 1750 m, 19 Aug. 1993, Rojas 378 (INB); Cantón de Paraiso, Parque Nacional Tapantí, Cuenca del Reventazón, Estación de Biología Tropical Río Macho y alrededores, 9°46′00″N, 83°52′00″W, 1590 – 1760 m, 19 Aug. 1993, Rojas 409 (INB); Cantón de Paraiso, Parque Nacional Tapantí, Cuenca del Reventazón, Estación de Biología Tropical Río Macho y alrededores, 9°46′00″N, 83°52′00″W, 1590 – 1760 m, 2 Sept. 1993, Rojas 439 (INB); Cantón de Paraíso, Cordillera de Talamanca, Estación de Biología Tropical Río Macho, 9°45′11″N, 83°51′48″W, 1700 m, 30 May 1994, Rojas 1060 (CR, INB, MO); Pacayas, at the foot of Volcán Turrialba, [9°55′N, 83°48′W], 1400 m, 16 April 1953, Scamman 7093 (GH); Turrialba, near Interamerican Institute, [9°54′N, 83°42′W], 600 m, 9 – 10 March 1953, Scamman 7096 (GH, US); Tapantí, in valley of Río Reventazón, [9°47′N, 83°48′W], 1130 m, 18 March 1956, Scamman & Holdridge 7959 (GH, US); vic. of Orosí, [9°48′N, 83°52′W], 30 March 1924, Standley 39870 (US); El Muñeco, on the Río Navarro, [9°48′N, 83°55′W], 1400 – 1500 m, 6 – 7 March 1926, Standley & Torres 51024 (US); Carpintera #2, [9°53′N, 83°49′W], 2800 m, 3 July 1925, Stork 1184 (MICH, US); Orosi, Monte Sky Forest Reserve, 9°45′58.6″N, 83°51′40.6″W, 1575 m, 11 Feb. 2008, Sundue et al. 1800 (CR, INB, NY, USJ); Tapanti, [9°47′N, 83°48′W], 1200 m, 6 July 1936, Valerio 2248 (US); Guanacaste: Cantón de Liberia, Parque Nacional Guanacaste, Cacão, Estación Cacão, Cerro Cacão, 10°55′43″N, 85°28′10″W, 1100 m, 21 Aug. 1992, Chavarría 484 (INB); Cantón de Liberia, Parque Nacional Guanacaste, Cacão, Estación Cacão, Cerro Cacão, 10°55′43″N, 85°28′10″W, 1100 m, 21 Aug. 1992, Chavarría 493 (INB); Cantón de La Cruz, Parque Nacional Guanacaste, Estación Pitilla, 9 km S de Santa Maria, sendero Fila Orosilito, 10°59′26″N, 85°25′40″W, 700 – 1000 m, 20 May 1992, Moraga 462 (INB); 1 km N of Las Nubes village, 8 km NW of Monteverde, 10°22′N, 84°51′W, 1200 m, 31 Aug. 1989, Zuchowski 9499 (INB); Heredia: Vara Blanca, between Volcán Poás and Barba, N slope of the Central Cordillera, [10°11′N, 84°10′W], 1850 m, 13 July 1940, Chrysler & Roever 5102 (GH); NE of San Isidro de Heredia, [10°01′N, 84°03′W], 1450 m, 25 July 1940, Chrysler & Roever 5576 (US); Braulio Carrillo Park, Zurquí, [10°03′N, 84°02′W], 1700 – 2000 m, March 1983, Gómez 20184 (MO); Vara Blanca, between Poás and Barba volcanoes, [10°11′N, 84°10′W], 1600 – 1700 m, 22 July 1923, Maxon & Harvey 8321 (US); Vara Blanca, between Poás and Barba volcanoes, [10°11′N 84°10′W], 1600 – 1701 m, 23 July 1923, Maxon & Harvey 8340 (US); Virgen del Socorro, 10°17′N, 84°10′W, 1000 m, 11 July 1983, Moran 3172 (CR); Zurquí, [10°03′N, 84°02′W], 1800 m, 24 Feb. 1983, Riba 290 (CR, MO); Vara Blanca de Sarapiquí, N slope of Central Cordillera, [10°11′N, 84°10′W], 1500 – 1750 m, July – Sept. 1937, Skutch 3274 (F, GH, MO, NY, S); Vara Blanca de Sarapiquí, N slope of Central Cordillera, between Poás and Barba volcanoes, [10°11′N, 84°10′W], 1600 m, Feb. 1938, Skutch 3601 (GH, MO, NY, S, US); Yerba Buena, NE of San Isidro, [10°03′N, 84°02′W], 2000 m, 22, 28 Feb. 1926, Standley & Valerio 49230 (US); Yerba Buena, NE of San Isidro, [10°03′N, 84°02′W], 2000 m, 22, 28 Feb. 1926, Standley & Valerio 49421 (US); Yerba Buena, NE of San Isidro, [10°03′N, 84°02′W], 2000 m, 22, 28 Feb. 1926, Standley & Valerio 49692 (US); Yerba Buena, NE of San Isidro, [10°03′N, 84°02′W], 2000 m, 22, 28 Feb. 1926, Standley & Valerio 49731 (US); S slopes of Cerro Zurquí, 5 km N of S. Luis Norte, [10°03′N 84°02′W], 1800 m, 28 March 1973, Stolze 1567 (CR); S slopes of Cerro Zurquí, 5 km N of S. Luis Norte, 10°03′N, 84°02′W, 1800 m, 28 March – 4 April 1973, Stolze 1568 (F, US); Limón: Cantón de Limón, Parque Nacional de Talamanca, antes de la unión de Ríos Lori y Coén, entre Ujarrás y San José de Cabér, 9°24′20″N, 83°13′30″W, 1500 – 1600 m, 3 April 1993, Fernández 933 (INB); Cantón de Limón, N flank of fila de Matama in headwaters of Río Boyei, 9°45′N, 83°19′W, 1200 – 1300 m, 18 Aug. 1995, Grayum 11088 (MO); Cantón de Talamanca, Bratsi, Amubri, Alto Lari, Kivut, quebrada innominada, margen derecha del Río Dapari, 9°24′20″N, 83°05′35″W, 1000 m, 11 March 1992, Herrera 5296 (CR, INB, MO); Cantón de Talamanca, Bratsi, Amubri, Alto Lari, Kivut, afluente innominado del Río Lari, 9°22′50″N, 83°05′10″W, 1500 m, 21 March 1992, Herrera 5410 (INB, MO); Cantón de Talamanca, Cordillera de Talamanca, frente unión Quebrada Kirigú con Río Coén, entre Ujarrás y San José Cabécar, 9°23′25″N, 83°12′55″W, 1550 m, 28 March 1993, Herrera 6064 (CR, INB, MO); Almirante, Cerro entre la cuenca superior del Río Xichiary y la del Río Boyei, 9°45′50″N, 83°20′00″W, 1200 m, 11 Aug. 1995, Herrera 8379 (CR, K); Area no protegida. Almirante, cuenca superior del Río Boyei, rumbo a Fila Matama, 9°44′20″N, 83°17′40″W, 1300 m, 16 Aug. 1995, Herrera 8509 (K, MO); Puntarenas: Reserva Biológica Monteverde, c. 500 m SW of Cloud Forest Reserve headquarters, trail to Campbell’s Meadow, [10°18′N 84°48′W], 13 April 1988, Bigelow & Kukle 21 (CR); vic. of Monteverde Nature Reserve, 10°18′N, 84°47′W, 31 Oct. & 2 Nov. 1975, Burger & Baker 9689 (NY); Just E of Monteverde on the Pacific watershed, 10°18′N, 84°48′W, 1300 – 1450 m, 29 Oct. – 2 Nov. 1975, Burger & Baker 9795 (CR, F, US); Cantón de Golfito, Jiménez, Dos Brazos de Río Tigre, Cerro Rincón, 8°30′30″N, 83°28′00″W, 745 m, 27 Aug. 1990, Herrera 4159 (INB); Pitahaya, area no protegida, Rincón, Los Planes de Monestel, 10°15′20″N, 84°41′50″W, 1400 m, 31 May 1996, Herrera et al. 9103 (K); Cultivated in Las Cruces Tropical Botanical Garden, originally collected on Cerro Zurquí, Prov. of San José, 8°48′N, 82°58′W, 1100 m, Jan. 1974, McAlpin 73-172 (F); Coto Brus, Parque Nacional La Amistad, Cuenca Térraba-Sierpe, sendero sobre el Río Canasta, [9°02′N 82°59′W], 1700 m, 1 Aug. 2000, Ramírez 2400 (INB); Cantón de Golfito, Parque Nacional Corcovado, Península de Osa, Cerro Rincón, nacientes del Río Tigre, 8°31′00″N, 83°28′00″W, 700 – 745 m, 28 Jan. 1998, Rojas et al. 4186 (INB); San José: Tablazo, [9°50′N, 84°03′W], 1 July 1908, Brade 27 (BM, NY); Tablazo, [9°50′N, 84°03′W], 2 June 1910, Brade & Brade 567 (S); La Palma area, northeast of San Jerónimo, above the La Hondura valley, 10°02′N, 84°00′W, 1500 m, 27 May – 1 June 1968, Burger & Stolze 5324 (CR, F, GH, NY, US); Zurquí, [10°03′N, 84°01′W], 2000 m, April 1973, Gómez 3611 (CR); Parque Nacional Braulio Carillo, Estación La Montura, 10°07′N, 83°59′W, 1050 m, 22 Jan. 1984, Gómez et al. 20828 (MO); Parque Nacional Braulio Carrillo, from La Montura to Los Chorritos, 10°07′N 83°59′W, 1200 m, 28 Jan. 1984, Gómez et al. 20928 (CR); Old cart road to Limón, 5.6 km N of San Jeronimo, near La Palma, 10°10′N, 84°00′W, 1400 – 1500 m, 13 Jan. 1987, Hill et al. 17785 (NY); La Palma, on the rd to La Hondura, [9°56′N, 84°05′W], 1500 – 1700 m, 17 – 18 July 1923, Maxon & Harvey 8033 (US); vic. of La Hondura, 4 km S to 1 km N on Rte 220, between Volcán Irazu and Volcán Barba, [10°04′N, 83°59′W], 1250 – 1500 m, 25 March 1967, Mickel 2257 (NY); Between La Palma and La Hondura, 1 – 4 km S of La Hondura, on route 220 between Volcán Irazú and Volcán Barba, [10°04′N, 83°59′W], 1500 m, 9 July 1967, Mickel 2562 (NY); Between La Palma and La Hondura, 1 – 4 km S of La Hondura, on route 220 betwee Volcán Irazú and Volcán Barba, [10°04′N, 83°59′W], 1500 m, 9 July 1967, Mickel 2563 (NY); 1 km N of La Hondura on route 220 between Volcán Irazú and Volcán Barba, [10°04′N, 83°59′W], 1250 m, 9 July 1967, Mickel 2578 (NY, US); Alto La Palma, NW of San José, [10°03′N, 83°59′W], 1450 m, 29 Aug. 1982, Moran 2332 (MO); Parque Nacional Braulio-Carrillo, c. 1 km along road from entrance, 1500 m, 19 July 1983, Moran 3276 (CR, F, MO, NY, US); Dota, Copey, Zapotal de Providencia, Reserva Montaña Fria, 9°31′53″N, 83°49′52″W, 1820 m, 24 Aug. 2004, Rojas 5862 (CR); Cantón de Moravia, P.N. Braulio Carrillo, cuenca del Sarapiquí, túnel Zurquí hasta 5 km, camino a Guapiles, [10°03′30″N, 84°01′W], 1650 m, 12 June 1997, Rojas & Coto 3570 (INB, MO); Cantón de Vázquez de Coronado, Cuenca del Sarpiquí, Bajo La Hondura, 10°03′15″N, 83°59′10″W, 1500 m, 23 April 1999, Rojas & Pacheco 5100 (CR, INB, NY); Cantón de Tarrazu, Valle Central, San Lorenzo, Cerro Pito, 2.5 km SE del Basurero, 9°34′30″N, 84°04′30″W, 1500 m, 18 April 1995, Rojas et al. 1768 (INB); La Palma on the road to La Hondura, [9°21′N, 83°44′W], 1400 – 1500 m, 5 – 6 March 1955, Scamman 7652 (F, GH, NY, US); road to La Hondura, [10°04′N, 83°59′W], 1200 m, 8 April 1956, Scamman & Holdridge 7960 (GH, US); Along unnamed N fork of Río Zurquí (upstream from Hwy. N of tunnel), Cordillera Central, 10°04′N, 84°01′W, 1500 – 1600 m, 18 Jan. 1986, Smith et al. 1685 (AAU, CR, MICH, MO, NY); La Palma, [9°56′N, 84°05′W], 1600 m, 17 March 1924, Standley 38185 (US); 5 miles S of Sta. María, [9°39′N, 83°58′W], 2100 m, 5 Feb. 1928, Stork 1777 (MICH); Sta. María Dota, [9°39′N, 83°58′W], 4 May 1928, Stork 2844 (MICH, US); Parque Nacional Braulio Barrillo, Estacion Transito Zurquí, trail opposite the hwy from the station, c. 500 km from the tunnel, 10°03.595′N 84°00.438′W, 1950 m, 12 Feb. 2008, Sundue & Nitta 1816 (CR, INB, NY, USJ); Old road to La Palma, [10°03′N 83°59′W], 1494 m, 5 Feb. 2008, Sundue et al. 1779 (CR, INB, NY, USJ); Old road to La Hondura, [10°04′N 83°59′W], 1491 m, 5 Feb. 2008, Sundue et al. 1782 (CR, INB, NY, USJ); Tablazo, [9°50′N, 84°03′W], 1877 m, 13 June 1929, Valerio 82 (US); Moravia, San Gerónimo, Bajo La Hondura, 10°05′50″N, 83°58′00″W, 1000 – 1200 m, 13 May 1998, Valverde 935 (CR); Unknown: Monteverde, [10°18′N, 84°48′W], Aug. 1961, James 61 (MO); Pacayas, [9°55′N, 83°48′W], 6 Sept. 1923, Lankester 650 (BM, US); vic. of La Palma, on the rd to La Hondura, [10°04′N, 83°59′W], 1500 – 1700 m, 17 – 18 July 1923, Maxon & Harvey 8016 (US); 1901 – 1905, Wercklé s.n. (MICH). Panama. Bocas del Toro: La Fortuna area, Gualaca to Chiriquí Grande; along oil pipeline road; along continental divide W of road, 8°45′N, 82°17′W, 1300 m, 6 March 1986, Hammel et al. 14657 (MO); vic. of Fortuna Dam, on trail along continental divide, 8°45′N, 82°15′W, 1250 m, 10 March 1988, McPherson 12277B (MO); Chiriquí: Vic. of Gualaca c. 8.5 miles from Planes de Hornito, La Fortuna on road to damsite, [8°53′N, 82°29′W], 1350 m, 10 July 1980, Antonio 5090 (CR, MO); NW del campmento de Fortuna (Hornito), sitio de presa, 1000 – 1200 m, 12 Aug. 1976, Correa et al. 2232 (CR); Monte Rey above Boquete, [8°46′N, 82°25′W], 21 July 1971, Croat 15683 (MO); vic. of branch in road to Cerro Colorado and Escopeta, above Río San Felix near town of San Felix, c. 13 miles N of Río San Felix bridge, [8°16′N, 81°52′W], 800 – 1200 m, 15 March 1976, Croat 33451 (MO, US); vic. of branch in road to Cerro Colorado and Escopeta, above Río San Felix near town of San Felix, c. 13 miles N of Río San Felix bridge, [8°16′N, 81°52′W], 800 – 1200 m, 15 March 1976, Croat 33457 (MO); Cerro Colorado, along rd above San Felix, 29 km above bridge over Río San Felix, 7.0 km above turn off to Escopeta, [8°16′N, 81°52′W], 1500 m, 14 July 1976, Croat 37084 (MO, U); Along road between Gualaca and Fortuna dam site, 10 min NW of Los Planes de Hornito, 8°45′N, 82°17′W, 1260 m, 10 April 1980, Croat 50067 (CR, MO); Along road between Gualaca and Fortuna dam site, 10 min NW of Los Planes de Hornito, 8°45′N, 82°17′W, 1260 m, 10 April 1980, Croat 50082 (MO); Along road between Fortuna lake and Chiriquí Grande, 4 – 4.5 km N of dam over Fortuna lake, 8°43′N, 82°17′W, 1100 – 1135 m, 8 March 1985, Croat 59973 (MO); above Boquete, [8°46′N, 82°25′W], 1100 m, 12 May 1971, D’Arcy 5465 (MO); La Fortuna dam area, N of dam, along Quebrada Arena down stream from road crossing, 8°46′N, 82°14′W, 1000 m, 10 Feb. 1986, Hammel 14445 (MO); trail W from Fortuna Dam Camp to La Fortuna, 8°43′N, 82°14′W, 1300 m, 13 March 1985, Hampshire & Whitefoord 548 (CR); Valley of the Río Piarnasta, about 5 miles E of El Boquete, [8°46′N, 82°25′W], 1525 – 1600 m, 9 – 22 Feb. 1918, Killip 5144 (GH); Valley of Río Piarnasta, above El Boquete, [8°46′N, 82°25′W], 1550 m, 9 Feb. 1918, Killip 5144 (US); Valley of Río Piarnasta, above El Boquete, [8°46′N, 82°25′W], 1550 m, 9 Feb. 1918, Killip 5151 (US); Valley of Río Piarnasta, above El Boquete, [8°46′N, 82°25′W], 1551 m, 22 Feb. 1918, Killip 5377 (US); Vic. of El Boquete, [8°46′N, 82°25′W], 1380 m, 3 Feb. – 15 March 1938, Maurice 670 (GH); c. 5 km NE of Boquete, [9°10′N, 82°16′W], 1700 – 1800 m, 20 March 1977, Skog et al. 4055 (MO, US); Distrito Boquete, Fortuna dam site; along trail following continental divide, [8°46′N, 82°25′W], 1100 m, 8 Feb. 1985, van der Werff & van Hardeveld 6754 (MO, NY).

Habitat. Wet forests; 800 – 2100 (– 2800) m.

Conservation Status. Least Concern (LC).

Notes. The scales on the rachises and costae of Megalastrum atrogriseum are distinctive. They are golden brown, firm, spreading, and mostly flat (i.e., not greatly twisted or tortuous at the apex). Once this scale type is learned, the species is easy to identify. M. squamosum has similar scales that are far more numerous. In Costa Rica and Panama, M. glabrum resembles M. atrogriseum except for being glabrous between the veins on the lamina tissue abaxially, and sparser pubescence on the leaf axes abaxially.

Megalastrum atrogriseum is highly variable in pubescence of the lamina abaxially. A long-pubescent form has hairs 1.0 – 1.2 mm long, 6- or 7-celled, and spreading, whereas the type has hairs 0. 2 – 0.5 mm long, 2 – 4-celled, mostly erect. These differences do not appear to correlate with other morphological characters; thus, it is interpreted as variation within the same species. Also, some forms (such as the type) have mixed stalked-glandular hairs and acicular ones on the abaxial surface, whereas others have mostly acicular hairs.

Megalastrum atrogriseum has at least some stalked glands adaxially. These are modified hairs where the apical cell becomes clavate or spherical. Often the glandular hairs are shorter than the non-glandular ones. M. gilbertii (Clute) R. C. Moran, J. Prado & Labiak, a West Indian species (Moran et al.2009b), also has glandular hairs, but these are shorter (c. 0.1 mm long) and laxer. M. gilbertii further differs by glabrous pinna rachises abaxially and narrower, more flexuose scales on the rachises and pinna rachises.

3. Megalastrum costipubensR. C. Moran & J. Pradosp. nov. A Megalastro galeottii rachidibus pubescentibus abaxialiter differt. Typus: Costa Rica, Limón, Cantón de Pococi, R.N.F.S. Barra del Colorado, Llanura de Tortuguero, Sardinas, 10°38′38″N, 83°44′10″W, 15 – 20 m, 6 June 1996, F. Araya & A. Rojas 919 (holotypus INB!; isotypi MO! 2 sheets, NY! 2 sheets).

http://www.ipni.org/urn:lsid:ipni.org:names:77105942-1

Rhizomes erect to decumbent; leaves up to 1.5 m long; scales of the petiole bases 10 – 20 × 0.4 – 0.5 mm, linear, spreading to ascending, twisted, the apices filiform and uniseriate for a short distance light brown, shiny, conspicuously denticulate on the margins; laminae 1.0 – 1.2 m long, up to 3-pinnate-pinnatisect at the base, 2-pinnate-pinnatisect medially; basal pinnae up to 0.5 m long, strongly inequilateral; pinna rachises abaxially non-glandular (lacking both stipitate or sessile glands), pubescent, sparsely scaly, the scales 1.0 – 1.5 × c. 0.2 mm, linear, the apices filiform, firm, spreading-ascending, brown, shiny, denticulate, the hairs 0.2 – 0.6 mm long, 2 – 5-celled, spreading, adaxially densely pubescent, non-glandular, sparsely scaly, the hairs 0.3 – 0.5 mm long, 3 – 5-celled, spreading, the scales like those of the abaxial surfaces; costules abaxially sparsely pubescent and scaly, non-glandular, subglabrous to sparsely puberulent, sparsely scaly at the base, the hairs 0.2 – 0.3 mm long, 2- or 3-celled, acicular, erect, the scales 0.3 – 0.5 × c. 0.1 mm, brown, twisted, adaxially non-glandular, densely pubescent, hairs c. 0.5 mm long, 3 – 5-celled, strigose, sparsely scaly; laminar tissue between veins abaxially glabrous to subglabrous, the hairs 0.1 – 0.2 mm long, 1 – 3-celled, erect, acicular, adaxially glabrous; veins visible on both surfaces, non-glandular, abaxially sparsely pubescent, the hairs like those of the costules, adaxially glabrous to sparsely pubescent, the hairs 0.2 – 0.3 (– 0.5) mm long, 2 – 4-celled; hydathodes evident; lamina margins ciliate, the hairs 0.1 – 0.3 mm long, 1- or 2-celled, ascending, acicular, non-glandular; indusia minute, fugacious, apparently consisting of a cluster of reddish proscales, c. 0.1 mm long, protruding from the centre of the sorus. Figs 6G – H and 7B; Map 1.

Distribution. Nicaragua, Costa Rica, Panama.

Additional Specimens Examined. Nicaragua. Chontales: [11°50′N, 85°20′W], 1867 – 68, Tate 35 (BM, K); Costa Rica. Alajuela: rd between Cañas and Upala, 4 km NNE of Bijagua on slope leading into Río Zapote, [10°44′N, 85°03′W], 400 m, 24 June 1976, Croat 36281 (CR, MO); Cartago: Turrialba, [9°54′N, 83°42′W], 600 m, 12 June 1910, Brade & Brade 571 (GH, NY); Irazu, [9°59′N, 83°51′W], 2000 m, 10 Aug. 1891, Pittier 307 (B, US); Irazu, [9°59′N, 83°51′W], 2000 m, 10 Aug. 1891, Pittier 312 (US); mts NE of Turrialba, [10°02′N, 83°46′W], 1000 m, 6 Oct. 1928, Stork 2646 (MICH, US); Tuis, [9°51′N, 83°35′W], 720 m, [no date], Tonduz 11334 (GH); Guanacaste: La Tejona, N of Tilarán, [10°31′N 84°58′W], 600 – 700 m, 25 Jan. 1926, Standley & Valerio 45772 (US); Heredia: Finca La Selva, OTS field station, [10°26′N, 84°02′W], 100 m, 2 June 1980, Grayum 2887 (F); upstream from Puerto Vijo c. 4 km at Finca La Selva, [10°26′N, 84°02′W], 125 m, 16 Aug. 1967, Mickel 3553 (NY); Chilamate, [10°27′N 84°04′W], Feb. 1893, Pittier 7480 (US); La Selva, finca of Dr L. R. Holdridge, on the Río Puerto Viejo near the junction with the Sarapiquí, [10°26′N, 84°02′W], 100 m, 28 March – 1 April 1956, Scamman & Holdridge 7962 (CR, GH); Finca La Selva, Sarapiquí region, [10°26′N, 84°02′W], 150 m, 29 Aug. 1961, Webster 6106 (GH, US); Limón: Santa Clara, Las Delicias, [10°12′N 83°39′W], 500 m, Feb. 1897, Biolley 10687 (NY, P, US); Santa Clara, Las Delicias, [10°12′N 83°39′W], 500 m, Feb. 1897, Biolley 10686 (P); Finca Hundrisser, [10°14′N, 83°46′W], Aug. 1909, Brade 404 (NY, S); Suerre, Llanuras de Santa Clara, [10°12′N, 83°45′W], 300 m, April 1896, Donnell-Smith 6895 (NY); Río Toro Amarillo, c. 6 – 7 km S of Guápiles, S of the suspension bridge, [10°12′55″N, 83°47′33″W], 460 – 500 m, 22 June 1975, Lellinger et al. 1866 (US); near Guapiles, at bridge over Río Guácimo, [10°12′55″N, 83°47′33″W], 50 m, 7 Aug. 1982, Moran 2190 (MO); Los Diamantes, USDA Rubber Plant Station, [10°12′49″N, 83°46′33″W], 300 m, 29 – 30 April 1951, Scamman 5888 (GH); Los Diamantes, USDA Rubber Plant Station, [10°12′49″N, 83°46′33″W], 300 m, 9 – 10 April 1953, Scamman 7094 (GH, US); vic. of Guápiles, [10°12′55″N, 83°47′33″W], 300 – 500 m, 12 – 13 March 1924, Standley 37062 (US); Hacienda de Guácimo, [10°13′17″N 83°41′17″W], 120 m, Aug. 1901, Tonduz 14583 (BM, P); Río Toro Amarillo, 2 km S of Guápiles, [10°12′55″N, 83°47′33″W], 17 July 1964, Woodruff s.n. (US); Puntarenas: Osa Peninsula, c. 5 km W of Rincón de Osa, NW of airfield, [8°42′N, 83°31′W], 50 – 200 m, 24 – 30 March 1973, Burger & Gentry 8853 (CR, F, US); Forests of Golfito, Golfo Dulce, [8°38′N, 83°10′W], March 1896, Pittier 9909 (US); San José: Plateau central de San José, [09°56′N 84°05′W], 1100 – 1500 m, 1904, Alfaro 16894 (BM, P). Panama. Chiriquí: roadside from Paso Canoas to Cañas Gordas, 18 miles from Paso Canoas, Quebrada de “Vuelta”, [8°43′60″N, 82°54′00″W], 25 Feb. 1973, Croat 0 (MO); Fortuna, camino de Quebrada Bonita llegando por el Embalse hacia el N, [8°43′N, 82°16′W], 1130 – 1150 m, 8 April 1987, Valdespino et al. 599 (MO).

Habitat. Wet forests; 15 – 1150 m.

Conservation Status. Least Concern (LC).

Notes.Megalastrum costipubens resembles M. galeottii and differs only by the pubescent pinna rachises abaxially. Further study might conclude that these species are the same, but the difference is so striking that we feel it best to name this species to call attention to the variation. Also similar is M. longipilosum (see below), which can be distinguished by the characteristics given in the key.

Megalstrum costipubens is nearly endemic to Costa Rica and Panama, occurring at only one known locality outside this region, in the nearby Nicaraguan department of Chontales. For the purposes of this paper, we have considered this species as an endemic to the mountains of Costa Rica and Panama.

4. Megalastrum ctenitoidesA. Rojas (2001: 470). Type: Costa Rica: Limón: Limón, El Progreso, Fila Matama, Valle de La Estrella, cabeceras del Río Cariei, 9°47′20″ N, 83°08′18″ W, 1400 m, 26 April 1989, G. Herrera & A. Chacón 2794 (holotype INB!; isotypes CR!, MO!, K).

Rhizomes erect to decumbent; leaves 0.8 – 1.0 m long; scales of the petiole bases 3 – 6 × c. 0.5 mm, lanceolate to filiform, firm, dull brown, not twisted, denticulate on the margins and sometimes the surfaces, the teeth occasionally bifid and or black; laminae 0.5 – 0.6 m long, 1-pinnate-pinnatifid at base and middle; rachises abaxially non-glandular (i.e., lacking stipitate glands and sessile spherical ones), scaly, pubescent, the scales 2 – 4 × 0.3 – 0.4 mm, lanceolate, firm, brown, dull, spreading, denticulate to entire, non-bullate, the hairs c. 0.5 mm long, 4- or 5-celled, adaxially non-glandular, pubescent, the hairs c. 1 mm long, 6 – 8-celled, strongly jointed (twisted at the septae); basal pinnae 0.1 – 0.15 m long, equilateral, cut c. half way to the costae; pinna rachises abaxially non-glandular (i.e., lacking stipitate glands and sessile spherical ones), densely pubescent, scaly, the hairs 0.3 – 0.5 mm long, 2 – 4-celled, ascending-appressed, scales like those of the rachises, adaxially glabrous (sometimes a few hairs at the base), the hairs like those abaxially, non-glandular; laminar tissue between the veins on both surfaces, glabrous, non-glandular; veins abaxially obscure, sparsely pubescent, hairs 0.2 – 0.3 mm long, 1 – 3-celled, ascending, adaxially not visible, glabrous; hydathodes evident; lamina margins sparsely ciliate, the hairs 0.1 – 0.3 mm long, 2- or 3-celled, ascending, non-glandular; indusia absent. Figs 8D and 9A – G; Map. 1.

Distribution. Costa Rica, W. Colombia. In Central America known only from the type.

Specimen Examined. Costa Rica. Limón: Limón, El Progreso, Fila Matama, Valle de La Estrella, cabeceras del Río Cariei, 9°47′20″N, 83°08′18″W, 1400 m, 26 April 1989, G. Herrera & A. Chacón 2794 (holotype INB!; isotypes CR!, MO!, K).

Habitat. Wet forests; 1300 – 1400 m.

Conservation Status. Data Deficient (DD); known only from the type and two specimens in Colombia.

Notes.Megalastrum ctenitoides is the only species in Central America with 1-pinnate-pinnatifid leaves. Also distinctive are the adaxially glabrous pinna rachises (sometimes a few hairs occur basally), the basal basiscopic lobes overlapping the leaf rachis, and hairs on the adaxial surface of the leaf rachises strongly jointed (this results from the hairs drying the individual cells collapsing and twisting at right angles to each other). Only one other species in Central America has adaxially glabrous pinna rachises: M. reductum (Fig. 9L). It differs from M. ctenitoides by the characters given in the key.

The protologue cites two specimens from the Department of Nariño, Colombia, that we have been unable to obtain on loan: Betancur et al. 4806 (COL) and Franco 5095 (COL).

5. Megalastrum dentatumA. Rojas (2007: 68, Fig. 1). Type: Costa Rica. Heredia: NW slope of Volcán Barva, c. 2 km by road NE of Los Cartagos, 10°09′N, 84°09′W, 2075 m, 16 March 1986, M. Grayum & G. Yatskievych 6642 (holotype CR; isotype MO!).

Rhizomes erect to decumbent; leaves 1.5 – 2.0 m long; scales of the petiole bases 10 – 20 × c. 0.5 mm, linear-lanceolate to filiform, twisted, brown, often shiny, prominently denticulate on the margins, the teeth usually antrorse; laminae 0.7 – 1.3 m long, 4-pinnate-pinnatisect at base (sometimes 4-pinnate), 3-pinnate-pinnatisect medially; basal pinnae up to 0.5 m long, strongly inequilateral (elongated basiscopically); pinna rachises on both surfaces non-glandular (i.e., lacking stipitate glands and sessile spherical ones), scaly, sparsely puberulent, the scales 4 – 5 × 0.3 – 0.4 mm, often tortuous, linear-lanceolate, often with flared base, non-bullate, spreading, dark brown, dull or shiny, the hairs on the abaxial surfaces 0.1 – 0.2 mm long, 1- or 2-celled; costules abaxially puberulent, scaly, non-glandular, the hairs 0.2 – 0.3 mm long, 1 – 3-celled, erect, acicular, brownish, the scales 1.0 – 1.5 mm long, lanceolate (sometimes with an acuminate apex), brown, denticulate, adaxially pubescent, the hairs 0.7 – 1.0 mm long, 4 – 6-celled; laminar tissue between the veins abaxially non-glandular, puberulent, the hairs 0.1 – 0.2 mm long, 1 – 3-celled, erect, acicular, proscales c. 0.2 mm long, appressed, adaxially pubescent, especially towards the apex of the segments, the hairs 0.2 – 0.3 mm long, 2- or 3-celled; veins evident to obscure on both surfaces, hairs and proscales on both surfaces like that of the costules; hydathodes evident; lamina margins ciliate, the hairs 0.1 – 0.3 mm long, 1 – 3-celled, non-glandular; indusia minute, fugacious, apparently consisting of a cluster of reddish proscales, c. 0.1 mm long, protruding from the centre of the sorus. Figs 2A and 6J – M; Map 1.

Distribution. Costa Rica, Panama.

Additional Specimens Examined. Costa Rica. Cartago: Turrialba, Tayutic, Jicotea, 9°47′05″N, 83°32′40″W, 1200 m, 16 June 1995, Herrera & Cedeño 7929 (CR, K); near el Alto, vic. of Cartago, [9°54′N, 83°57′W], 15 March 1953, Scamman 7013 (GH); Heredia: Río Vueltas, 10°05′N, 84°05′W, 2100 m, 23 May 1969, Gómez 2266 (BM, GH, MO, NY); along headwaters of Río Santo Domingo, c. 3 km E of San Rafael de Vara Blanca, N slope of Volcán Barva, 10°11′N, 83°07′W, 2060 m, 14 April 1986, Grayum 7093 (MO, US); between headwaters of Río San Fernando and Río Sardinal, Atlantic slope of Volcán Barva, 10°12′N, 83°06′W, 1850 – 1880 m, 22 April 1986, Grayum 7367 (MO); Braulio Carillo National Park, 10°15′N 84°10′W, 1830 m, 7 Nov. 1986, Hennipman et al. 6740 (U); Braulio Carillo National Park, 10°15′N 84°10′W, 1865 m, 9 Nov. 1986, Hennipman et al. 6818 (U); Cerro Chompipe, 5 km SE of Volcán Barba, [10°05′N, 84°04′W], 2000 m, 15 April 1971, Lellinger 1702 (F, US); N slope of Cerro Chompipe, c. 10 km NNE of Heredia, [10°05′N, 84°04′W], 2200 m, 8 July 1970, Lellinger & White 1021 (US); end of rt. 113 where rd crosses Río Patria, about 100 – 200 m upstream, 10°08′N, 83°56′W, 2000 m, 27 June 1983, Moran 3050 (B, BM, GH, MO, NY, US); San José de la Montaña, [10°03′N, 84°07′W], 2200 – 2300 m, 25 March 1965, Nisman 163 (GH); Cantón de San Rafael, Reserva Forestal Cordillera Volcánica Central, Cuenca del Río Sarapiquí, costado N de Cerro Chompipe, 10°05′15″N, 84°04′20″W, 2000 – 2100 m, 28 Nov. 1997, Rojas et al. 4059 (INB, NY); Cerro Chompipe, 2130 m, 5 Feb. 2008, Sundue et al. 1789 (CR, INB, NY, USJ); Limón: Cantón de Talamanca, Bratsi, Amubri, Alto Lari, Kivut, afluente innominado del Río Lari, 9°22′45″N, 83°06′15″W, 1900 m, 25 March 1992, Herrera 5488 (MO); Puntarenas: Forests E of hairpin c. 1 km SE of Las Alturas de Coto Brus, on road from San Vito, 8°56′N, 82°50′W, 1390 – 1440 m, 13 July 1985, Grayum & Hammel 5674 (CR, MO); Reserva Indígena Ujarrás, Buenos Aires, 9°17′50″N, 83°15′30″W, 1520 m, 11 March 1993, Herrera 5867 (INB); Cantón de Coto Brus, Zona Protectora Las Tablas, Cuenca Terraba-Sierpe, Sendero Sura, 8°57′21″N, 82°44′30″W, 2050 m, 17 Feb. 1998, Navarro V. & Rojas 860 (INB, MO, US); Cantón de Coto Brus, Zona Protectora Las Tablas, Cuenca Terraba-Sierpe, Sendero Sura, 8°57′21″N, 82°44′30″W, 2050 m, 17 Feb. 1998, Navarro V. & Rojas 862 (INB, MO); Cantón de Coto Brus, Zona Protectora Las Tablas, Sabalito, Las Alturas de Cotón, Est. Biol. Las Alturas, 8°57′15″N, 82°50′10″W, 1580 m, 23 Dec. 1993, Rojas et al. 754 (CR, INB). San José: Grenadilla, [9°56′N, 84°02′W], 1200 m, 26 Feb. 1910, Brade & Brade 569 (GH, NY, S); 10 km N of San Rafael de Heredia on Volcán Barva, 1950 m, 30 July 1967, Lellinger 793 (US); 10 km N of San Rafael de Heredia on Volcán Barva, [10°08′N, 84°06′W], 1950 m, 30 July 1967, Mickel 3009 (NY); La Hondura, 10°04′N, 83°59′W, 1300 m, 11 Nov. 1964, Nisman 97 (CR, GH). Panama. Chiriquí: Distr. Bugaba, Cerro Punta, from STRI house to nearby ridge, 8°52′N, 82°33′W, 2200 m, 25 Jan. 1985, van der Werff & Herrera 6376 (AAU, MO, NY).

Habitat. Not known; 1200 – 2300 m.

Conservation Status. Least Concern (LC).

Notes.Megalastrum dentatum is characterised by densely and conspicuously scaly petioles and rachises, lanceolate scales on the pinna rachises abaxially, and minute (0.1 – 0.2 mm long) acicular, erect hairs between the veins abaxially. The specific epithet dentatum refers to the long teeth on the scales of the lamina and pinna rachises (Rojas 2007). M. dentatum resembles M. galeottii but differs by leaf axes conspicuously scaly (vs. inconspicuously scaly), and lanceolate scales on the pinna rachises abaxially (versus linear to filiform). When large laminae of both species are compared, M. dentatum can be seen to be more finely cut, being 3-pinnate-pinnatisect medially, whereas M. galeottii is 2-pinnate-pinnatisect medially.

Megalastrum dentatum also resembles M. squamosum (treated below) by having densely scaly leaf axes. The scales of M. squamosum, however, differ by being wider (0.2 – 0.3 mm), flat (not twisted), straight (not tortuous), and yellow brown. In these respects, the scales of M. squamosum resemble those of M. atrogriseum.

Moran 3050 is atypical by lamina tissue between the veins adaxially glabrous or very sparsely pubescent. Usually, the species is pubescent between the veins adaxially, especially toward the segment apices.

6. Megalastrum galeottii (M. Martens) R. C. Moran & J. Prado, comb. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77105949-1

Polypodium galeottii M. Martens (1842: 43, pl. 7, f. 3.); Phegopteris galeottii (M. Martens) Fée (1852: 243); Dryopteris galeottii (M. Martens) C. Chr. (Christensen 1905: 267). Type: Mexico, Veracruz, Mirador, [18°47′N, 96°36′W], 1000 m, June – Oct. 1840, H. Galeotti 6321 (lectotype BR-627987!, selected by Smith (1981); duplicate BR-627822!, photo US!).

Phegopteris stenolepis Fée (1857: 89). Type: Mexico, Veracruz, near Huatusco, J. G. Schaffner 239 (lectotype K!, selected by Smith (1981); duplicate P!).

Megalastrum longipilosum A. Rojas var. glabrescens A. Rojas (2007: 19, f. 3). Type: Costa Rica. Alajuela: Cantón de Alajuela, Cuenca del Sarapiquí, cerca de la unión de la Quebrada Ten Fé y Río Cariblanco, 10°15′45″N, 84°11′35″W, 840 – 950 m, 2 Dec. 1997, A. Rojas et al. 4132 (holotype CR; isotype INB!).

Rhizomes erect to decumbent; leaves up to 1.5 m long; scales of the petiole bases 10 – 20 × 0.4 – 0.5 mm, linear, light brown, shiny, conspicuously denticulate on the margins, twisted, the teeth often bifid, the apices filiform and 1-celled for a short distance; laminae 1.0 – 1.2 m long, up to 3-pinnate-pinnatisect at the base, 2-pinnate-pinnatisect medially; basal pinnae up to 0.5 m long, strongly inequilateral (elongated basiscopically); pinna rachises abaxially non-glandular (lacking both stipitate or sessile glands), sparsely scaly, hairs absent, the scales 1.0 – 1.5 × 0.05 – 0.2 mm, linear to filiform, spreading-ascending, brown, shiny, denticulate, adaxially pubescent, sparsely scaly, the hairs 0.3 – 0.5 mm long, 3- or 4-celled, spreading, the scales like those of the abaxial surfaces; costules non-glandular, subglabrous to sparsely puberulent, sparsely scaly at the base, the hairs 0.1 – 0.2 mm long, 1 – 3-celled, acicular, erect, adaxially non-glandular, densely pubescent, hairs c. 0.5 mm long, 3 – 5-celled, strigose to spreading; laminar tissue between veins abaxially glabrous to (more commonly) densely puberulent, the hairs 0.1 – 0.2 mm long, 1 – 3-celled, erect, acicular, adaxially glabrous; veins visible on both surfaces, abaxially sparsely pubescent to glabrous, the hairs like those of the costules, adaxially glabrous to sparsely pubescent, the hairs 0.3 – 0.4 mm long, 1 – 3-celled, strigose to spreading; hydathodes evident; lamina margins ciliate, the hairs 0.2 – 0.3 mm long, 1- or 2-celled, acicular, ascending, non-glandular; indusia minute, fugacious, apparently consisting of a cluster of reddish proscales, c. 0.1 mm long, protruding from the centre of the sorus. Figs 6A – F and 7A; Map 1.

Distribution. Mexico, Belize, Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama.

Additional Specimens Examined. Mexico. Chiapas: Mun. Cintalapa, ridge 3 km E of Francisco Madero, NE of Cintalapa, [16°42′N, 93°55′W], 1250 m, 4 Oct. 1974, Breedlove 38082 (MO); Mun. La Independencia, 12 km from Laguna Tsiskaw on road to Ixcán, 1250 m, 19 Oct. 1974, Breedlove 38931 (MO); Mun. Berriozábal, 13 km N of Berriozábal, [16°47′N, 93°19′W], 1000 m, 2 Nov. 1971, Breedlove & Smith 21602 (MICH, MO, NY); Mun. Ocosingo, Laguna Ocotal Grande, [17°06′N, 91°31′W], 800 m, 6 Feb. 1973, Breedlove & Smith 33078 (MICH); between El Carmen and Solosuchiapa, [16°39′N, 92°54′W], 1907, Collins & Doyle 234 (US); Ejida Las Golandrias, lower slopes of Cerro Ovando, along road between Golandrinas and Los Cacaos, 15°27′N, 92°38′W, 800 – 900 m, 22 Aug. 1996, Croat 78537 (MO); no locality given, 1864 – 70, Ghiesbreght 365 (GH, K, NY); Volcán Tacaná, [15°07′48″N, 92°06′43″W], 30 March 1939, Matuda 2915 (MICH); Finca Irlandia, May 1914, Purpus 7230 (F, GH, MO, NY, US); 1914, Purpus 7255 (US); Phoenia, 1927, Purpus 11086 (MICH); Guerrero: Montes de Oca, San Antonio – Buenos Aires, [18°10′N, 101°47′W], 28 April 1938, Hinton 14060 (GH, MO, NY); Oaxaca: Road from Oaxaca to Tuxtepec Km 149 –153, [17°55′N, 95°59′W], 3 Aug. 1967, Hellwig 454 (NY); Comaltepec, Distr. Ixtlán, Puerto Eligio, 17°45′N, 96°30′W, 700 m, 5 Aug. 1987, López-G 1 (MO, NY); Distr. of Villa Alta, valley of the Yelagago R., c. 20 miles NE of Villa Alta, 17°25′N, 96°05′W, 1000 – 1200 m, 30 July 1962, Mickel 1077 (MICH); Distr. of Villa Alta, valley of the Yelagago R., c. 20 miles NE of Villa Alta, 17°25′N, 96°05′W, 1000 – 1200 m, 30 July 1962, Mickel 1082 (MICH, NY, US); Road from Ixtlan to Tuxtepec, 24 km S of Valle Nacional Km 85, 800 m, 26 July 1964, Mickel 1455 (MICH, NY, US); Distr. Tuxtepec, 4 – 9 km S of Valle Nacional on Route 175, 800 – 2200 m, 31 July 1971, Mickel 5892 (NY); Distr. Tuxtepec, 4 – 9 km S of Valle Nacional on Route 175, [17°55′N, 95°59′W], 800 – 2200 m, 31 July 1971, Mickel 5923 (MICH, NY, US); Juquila, 28 – 29 km N of San Gabriel, [16°04′N, 97°05′W], 1500 – 1700 m, 10 Aug. 1971, Mickel 6191 (AAU, NY, US); Distr. Ixtlán, 29 km S of Valle Nacional, 80 km N of Ixtlán de Juárez, trail E of Route 175 at Campamento Vista Hermosa toward Ladú, 1 hr hike down to Río de la Trucha, 1500 – 2000 m, 13 Aug. 1971, Mickel 6376 (NY); Distr. Ixtlán, 29 km S of Valle Nacional, 80 km N of Ixtlán de Juárez, trail E of Route 175 at Campamento Vista Hermosa toward Ladú, 1 hr hike down to Río de la Trucha, 1500 – 2000 m, 13 Aug. 1971, Mickel 6396 (NY); Distrito Ixtlán, 29 km S of Valle Nacional, 80 km N of Ixtlán de Juárez, [17°22′N, 96°29′W], 500 – 650 m, 13 Aug. 1971, Mickel 6401 (NY); Cuicatlán, vic. of Teutila, [17°58′N, 96°43′W], 1000 m, 25 Sept. 1973, Mickel 7268 (AAU, NY); Distr. Putla, 15 km S of Putla, [17°01′N, 97°55′W], 2800 m, 10 Oct. 1969, Mickel & Hellwig 3949 (NY); Dtto. Pochutla, 45 km N of Pochutla, 13 km N of Candelaria, [18°19′N, 101°40′W], 1100 m, 1 Oct. 1970, Mickel & Leonard 5230 (MICH, NY); vic. of Cafetal Concordia, 400 – 650 m, 1 – 15 April 1933, Morton & Makrinius 2397 (US); Veracruz: country of Cordova, [15°52′N, 96°55′W], 1889 – 91, Fink 59 (NY); Valle de Cordova, [18°51′N, 96°55′W], 1 March 1866, Bourgeau 2016 (MO, P, S); Valle de Cordova, La Trinidad, [18°51′N, 96°55′W], 16 April 1866, Bourgeau 2277 (GH, MO, NY, P); Cantón de Cordoba, [18°59′N, 96°54′W], 1200 m, 20 Dec. 1897, Conzatti & González 594 (GH, P); Cantón de Cordoba, [18°59′N, 96°54′W], 1200 m, 25 Dec. 1897, Conzatti & González 615 (GH, MICH, P); Cordova, [18°51′N, 96°55′W], 1889 – 91, Fink 55 (NY); 2000 m, May 1842, Liebmann 649 (BM); Zacualpan, [15°10′N, 92°37′W], Oct. 1906, Purpus 1980 (F, GH, MO, NY); Zacualpan, [15°10′N, 92°37′W], Nov. 1908, Purpus 4254 (MICH, P); [locality illegible], May 1930, Purpus 13060 (F, S); Zacualpan, [15°10′N, 92°37′W], June 1929, Purpus 13060 (GH, MICH, MO); Zacualpan, [15°10′N, 92°37′W], Nov. 1931, Purpus 15735 (MICH); Barranca de Tenampa, [19°15′N, 96°53′W], Sept. 1934, Purpus 16624 (K); near Cordova, [18°51′N, 96°55′W], 1908, Spence 75 (GH); Ixhuatlan, El Presidio, [18°40′N, 96°45′W], 950 m, 27 June 1979, Ventura-A 16270 (NY). Belize. Camp 34, [16°35′N, 87°58′W], 850 m, 11 May 1934, Schipp 762 (GH). Guatemala. Alta Verapaz: vic. of Sepacuité, [15°25′60″N, 89°45′00"W], 1000 m, June 1904, Cook & Doyle 329 (US); Cobán, [15°28′N, 90°22′W], [no date], Salvin s.n. (GH); Cubilquitz, [15°40′N, 90°25′W], 350 m, April 1901, von Türckheim 1052 (NY); Pansamalá, [15°27′N, 90°01′W], 1200 m, Sept. 1896, von Türckheim 1056 (B, GH, K, NY, US); Cubilquitz, [15°40′N, 90°25′W], 350 m, Oct. 1904, von Türckheim 1069 (P); Cubilquitz, [15°40′N, 90°25′W], 350 m, April 1901, von Türckheim 8052 (B, K, NY, US); Cubilquitz, [15°40′N, 90°25′W], 350 m, Oct. 1904, von Türckheim 8811 (US); Jalapa: Los Amates, Finca Alsacia, [14°40′N, 89°42′W], 165 Aug. 1936, Hatch & Wilson 25 (F, US); Unknown: Choctum, 1862, Salvin s.n. (GH, K). Honduras. Atlántida/Yoro: Cordillera Nombre de Dios, [15°40′N, 86°40′W], 700 – 1000 m, April 1979, Gómez 7048 (CR); Comayagua: Cordillera de Comayagua, near Coyocutena, [14°37′N, 87°30′W], 1500 m, 14 April 1957, Molina R. 8084 (F, US); near El Achote, hills above plain of Siguatepeque, [14°30′N, 87°50′W], 1350 m, 26 July 1936, Yuncker et al. 6098 (GH, MO, NY, US); Ocotepeque: Cordillera Merendón, 10 km from Nueva Ocotepeque, rd to El Portillo, [14°25′N, 85°10′W], 1500 m, 26 Aug. 1968, Molina R. 22217 (F, MO, NY); Olancho: Río del Real, 8 km W de Catacamas, [14°48′N, 85°54′W], 800 m, 3 May 1987, Ortega-U 282 (MO); Santa Bárbara: Lake Yojoa, [14°59′N, 87°59′W], 23 Aug. 1951, Ray 2141 (MO); El Sauce, above Lake Yojoa, [14°24′N, 87°46′W], 1000 m, 9 April 1951, Williams & Molina 17686 (US); Yoro: Cordillera Nombre de Dios, hills S of San José de Texíguat, 15°29′N, 87°26′W, 300 – 400 m, 17 May 1991, Davidse et al. 34476 (MO). El Salvador. Santa Ana: Parque Nacional Los Volcanes, Sector el Paraiso, La Joya, 13°52′N, 89°38′30″W, 1800 m, 23 June 2005, Monterrosa et al. 961 (NY). Nicaragua. Chontales: 4 km NNW of Cuapa, ridgetops and summits of Cerro Oluma, 12°18′N, 85°23′30″W, 700 – 775 m, 23 Sept. 1983, Nee 28419 (CR); Granada: summit of Volcán Mombacho, [11°46′60″N, 85°54′00″W], 1600 m, 18 Dec. 1940 – 9 Feb. 1941, Grant 827 (GH); Volcán Mombacho, [11°46′60″N, 85°54′00″W], 1300 m, 5 Jan. 1967, Molina R. 20024 (F, NY, US); Volcán Mombacho, Plan de Las Flores, [11°49′N, 85°58′W], 1300 m, 17 Jan. 1980, Moreno 624 (CR); Jinotega: road to Hacienda Fundador which turns off Highway 3 c. halfway between Jinotega and Matagalpa, [14°43′N, 84°58′W], 1400 m, 8 Aug. 1977, Croat 43092 (CR); Laguna Miraflores, c. 26.1 km by rd NE of Hwy 1 at Estelí, 13°15′N, 86°15′W, 1250 – 1300 m, 10 – 11 June 1981, Henrich & Stevens 357 (CR, NY); Laguna Miraflores, c. 26.1 km by rd NE of Hwy 1 at Estelí, 13°15′N, 86°15′W, 1250 – 1300 m, 10 – 11 June 1981, Stevens 254 (CR); Matagalpa: La Carlota, carretera Matagalpa-Tuma, [13°03′N, 84°43′W], 1400 – 1500 m, Oct. 1975, Gómez et al. 6291 (CR); Sta. María de Ostuma, carretera Matagalpa-Jinotega, [12°55′N, 85°55′W], 1500 m, Nov. 1975, Gómez et al. 6383 (CR); El Arenal between Aranjuez and Sta. Martha, [13°01′N, 85°55′W], 1400 m, 7 March 1967, Molina R. 20337 (F, GH, NY, US); between El Triunfo and Fuente Pura, NE above Sta. María de Ostuma, [12°57′N, 85°58′W], 1800 m, 8 March 1967, Molina R. 20397 (F, NY); Cañada Yasira, [12°55′N, 85°55′W], 1000 m, Aug. 1893, Rothschub 213 (BR); Cordillera Central, Finca Santa María de Ostuma, [12°57′N, 85°58′W], 1300 – 1500 m, 30 Nov. – 4 Dec. 1973, Williams & Molina 42616 (F, GH, US); Cordillera Central, Finca Santa María de Ostuma, between Matagalpa and Jinotega, [12°57′N, 85°58′W], 1300 – 1500 m, 8 – 15 Jan. 1963, Williams et al. 23646 (F); Cordillera Central, Finca Santa María de Ostuma, between Matagalpa and Jinotega, [12°57′N, 85°58′W], 1300 – 1500 m, 8 – 15 Jan. 1963, Williams et al. 23957 (F, US); Disparate de Potter, St. María de Ostuma, Cordillera Central, [12°57′N, 85°58′W], 1600 m, 15 Jan. 1965, Williams et al. 27688 (F, NY); Nueva Segovia: Río Solonlí, 5 km N de Jalapa, [13°56′N, 86°12′W], 700 – 950 m, 5 April 1977, Neill 1619 (CR). Costa Rica. Alajuela: Alajuelita, [9°54′N, 84°06′W], 1300 m, March 1902, Alfaro 8076 (B, NY, US); Buena Vista, rd to San Carlos Valley, [10°17′N, 84°28′W], 600 m, 16 April 1903, Cook & Doyle 157 (US); Reserva forestal de San Ramón, 10°12′53"N, 84°36′28″W, 800 – 1200 m, 3 Dec. 1986, Herrera et al. 315 (AAU, MO); Aguacate, [10°02′N 84°28′W], Aug. 1857, Hoffmann 774 (BR); Hacienda La Argentina, Grecia, [10°04′N, 84°18′W], Dec. 1917, Jiménez 1140 (CR, NY); Finca La Constancia, Buena Vista, San Carlos, 850 m, 3 March 1963, Jiménez 446 (CR, F); Finca Los Ensayos, Buena vista de Zarcero, 10°16′N, 84°27′W, 900 – 1000 m, 26 Feb. 1983, Judziewicz 4411 (CR, MO); 12 km N of San Ramón, [10°13′N, 84°37′W], 900 m, 27 July 1967, Lellinger 735 (US); Cantón de San Ramón, Reserva Forestal San Ramón, Reserva Biológica Alberto Brenes, 10°13′10"N, 84°36′00″W, 800 – 1000 m, 6 May 1966, Moraga 351 (INB, MO); c. 20 km N of San Ramón, Univ. of San Ramón’s Biological Station, 10°13′N, 84°37′W, 1000 m, 17 July 1983, Moran 3186A (F); Cantón de Alajuela, Virgen del Socorro, 10°15′25″N, 84°10′20″W, 720 – 870 m, 23 Oct. 1993, Rojas 566 (INB); Cantón de Upala, Finca Zapote, Bijagua, Upala, 10°45′05″N, 85°04′35″W, 500 – 530 m, 9 July 1994, Rojas 1296 (INB); Cantón de San Ramón, Cuenca de San Carlos, Los Angeles a 4.5 km de la Escuela la Balsa, naciente del Río San Luis, 10°12′15′N, 84°32′15′W, 900 – 1250 m, 1 Dec. 1997, Rojas et al. 4101 (CR, INB); USDA Experimental Station, Los Diamantes, on Río Sta. Clara, 1.6 km E of Guapiles, [10°12′49″N, 83°46′33″W], 200 m, 29 – 30 April 1951, Scamman 5936 (GH, MICH); near Zapote on the road to Vila Quesada, [10°13′N, 83°25′W], 1200 m, 24 – 27 March 1955, Scamman 7595 (GH, US); Canyon of Río Cariblanco and W slope and summit of ridge between Río Cariblanco and Quebrada Quicuyal, SW of Cariblanco, 10°16′N, 84°12′W, 840 – 950 m, 22 Jan. 1986, Smith et al. 1863 (CR); Cartago: Río Turrialba, [9°54′N 83°39′W], 500 m, March 1894, Donnell-Smith 5086 (US); Santa Barbara, Cerro Peña Blanca y Palomo, [9°47′N, 83°51′W], 1100 m, Feb. 1979, Gómez 6863 (CR); lowermost slope of Alto Perlas at N end, Tapantí, 9°46′30"N, 83°51′00"W, 1300 – 1400 m, 12 Aug. 1984, Grayum & Jacobs 3806 (MO); lowermost slope of Alto Perlas at N end, Tapantí, 9°46′30″N, 83°48′00″W, 1300 – 1400 m, 12 Aug. 1984, Grayum & Jacobs 3811 (CR, MO); Turrialba, CATIE, 9°52′20″N, 83°38′30″W, 580 m, 15 June 1994, Herrera 7155 (CR); Cantón de Paraíso, Cuenca del Reventazón, Orosí, Río Macho, Est. Biol. Río Macho y alrededores, 09°46′N, 83°52′W, 1100 – 1850 m, 4 Oct. 1994, Mejía 1586 (INB); Tapantí, c. 15 km S of Paraíso, [9°43′N, 83°47′W], 1150 m, 20 March 1967, Mickel 1845 (NY); Tapantí, c. 15 km S of Paraíso, [9°43′N, 83°47′W], 1150 m, 20 March 1967, Mickel 1853 (NY); 1 km SW of Cervantes, [9°53′N, 83°48′W], 1400 m, 11 July 1967, Mickel 2633 (NY); c. 22 km E of Turrialba, high ridge above Platanillo, [9°49′N, 83°24′W], 1200 – 1450 m, 22 Aug. 1967, Mickel 3433 (NY, US); Cantón de Turrialba, Tayutic, Jicotea, 9°47′15″N, 83°32′50″W, 100 – 1600 m, 22 June 1995, Rojas et al. 2060 (INB); near el Alto, vic. of Cartago, [9°54′N, 83°57′W], 30 March 1951, Scamman 5883 (GH); Forests of Juan Viñas, Atlantic slope, [9°54′N, 83°45′W], 1135 m, 25 Jan. 1890, Tonduz 1838 (US); Tapantí, [9°43′N, 83°47′W], 1200 m, 6 July 1936, Valerio 2250 (US); Navarro, [9°48′N, 83°53′W], 1400 m, 1905, Wercklé s.n. (US); Valle de Navarro, [9°48′N 83°53′W], 1400 m, 1905, Wercklé s.n. (BM); Guanacaste: W slope of Cerro Nubes, c. 2 km E of Silencio de Tilarán, 10°28′N, 84°53′W, 900 m, 26 Jan. 1985, Grayum et al. 5008 (CR, MO); Las Nubes de Tilerán, Quebradas Valdivia and Azul, 10°23′N, 84°52′W, 1000 – 1150 m, 10 Nov. 1990, Ivey & Haber 141 (CR, MO); Heredia: Tirimbina, 10°24′N, 84°7′W, 150 – 250 m, 12 – 15 Aug. 1971, Burger & Burger 8024 (F); Virgen del Socorro, 10°17′N, 84°10′W, 600 – 800 m, 31 Aug. 1983, Chacón & Herrera 1160 (CR, MO); Plaines de Sta. Clara, 100 m, Sept. 1896, Cooper 10271 (US); Forests of Río Vueltas, [9°50′N, 83°42′W], 2100 m, 23 May 1969, Gómez 2272 (F, GH, NY); Finca La Selva, OTS field station, [10°26′N, 84°02′W], 100 m, 13 Aug. 1979, Grayum 2331 (CR); Hills to the S of the Río Sarapiquí at Chilamate de Sarapiquí, 10°27′N, 84°04′W, 60 – 100 m, 4 July 1985, Grayum et al. 5554 (CR, MO); Cantón Sarapiquí, Puerto Viejo, La Selva Biological Station, 10°26′N, 84°01′W, 50 m, 20 July 2001, Jones et al. 139 (CR); La Selva, Puerto Viejo de Sarapiquí, [10°26′N, 84°02′W], 100 m, 31 Oct. 1964, Nisman 89 (CR, GH); Cantón Sarapiquí, Puerto Viejo, La Selva Biological Station, 10°26′N, 84°01′W, 50 m, 19 Oct. 2001, Olivas 301 (CR); Forets de Tsaki, [9°31′N, 82°51′W], 200 m, April 1895, Pittier 9472 (US); Forets de Tsaki, [10°12′N, 83°39′W], 500 m, Feb. 1897, Pittier 10684 (US); Forets de Tsaki, [09°31′N, 82°51′W], 200 m, April 1895, Tonduz 9481 (U); Limón: La Emilia, Llanuras de Santa Clara, [10°13′N, 83°47′W], 301 m, April 1897, Alfaro 6893 (GH, US); Suerre, Llanuras de Santa Clara, [10°12′N, 83°45′W], 300 m, April 1896, Alfaro 6895 (B, NY, US); Santa Clara, Las Delicias, [10°12′N 83°39′W], 500 m, Feb. 1897, Biolley 10686 (P); Suerre, Llanuras de Santa Clara, [10°12′N, 83°45′W], 300 m, April 1896, Donnell-Smith 6895 (GH, NY, US); Cantón de Talamanca, Valle de Talamanca, Amubri, Katsi, [9°30′40″N, 82°56′20″W], 100 m, 16 June 1994, Gallardo 211 (INB, MO); USDA Experimental Station, Los Diamantes, on Río Sta. Clara, 1.6 km E of Guapiles, [10°12′49″N, 83°46′33″W], 200 m, July 1949, Holm & Iltis 339 (BM, F, MO, NY, US); USDA Experimental Station, Los Diamantes, on Río Sta. Clara, 1.6 km E of Guapiles, [10°12′49"N, 83°46′33′W], 200 m, 12 July 1949, Holm & Iltis 411 (BM, F, MO, NY, US); 3 km SW of Suretka, RECOPE test drill site, [9°35′N, 82°57′W], 70 m, 6 July 1983, Moran 3111 (CR, F, MO, NY); Puntarenas: S side of Quebrada Bonita, to c. 1 km E of Costanera hwy, Carara Reserve, 9°47′N, 84°37′W, 30 – 40 m, 11 Jan. 1985, Grayum 4769 (CR, MO); San Vito de Coto Brus to Ciudad Neily, NE slopes of Fila de Cal, 8°41′N, 82°56′W, 500 – 620 m, 12 July 1985, Hammel & Grayum 14186 (CR, MO); Cantón de Buenos Aires Olán, siguiendo filas en cuenca superior de Río Cabagra, 9°17′40"N, 83°11′50"W, 1700 m, 24 Sept. 1989, Herrera 3538 (CR, INB, MO); Cantón de Golfito, Jiménez, Alto Carbonera, Cerro Osa, cabeceras de Quebrada Sombrero, 8°25′20″N, 83°18′20″W, 250 m, 20 Sept. 1990, Herrera 4338 (INB); Monteverde, SE of Welfords Tract, 1400 m, 8 Jan. 1972, James s.n. (MO); Cultivated in Las Cruces Tropical Botanical Garden, originally collected on Cerro Zurqui, Prov. Of San Jose, [8°48′N, 82°58′W], 1100 m, Jan. 1974, McAlpin 70-130 (F); vic. of biological field station at Finca Wilson, 5 km S of San Vito de Java, [8°47′N, 82°57′W], 1100 – 1200 m, 2 – 6 Aug. 1967, Mickel 3171 (NY); Cantón de Coto Brus, Zona Protectora Las Tablas, Cuenca Terraba-Sierpe, 8°55′58″N, 82°45′33″W, 1800 m, 13 Feb. 1999, Mora-C. 63 (INB, NY); Cantón de Coto Brus, Zona Protectora Las Tablas, Cuenca Terraba-Sierpe, sitio Coto Brus, orillas de la Quebrada Surú, 8°57′20″N, 82°51′01″W, 1600 m, 9 Feb. 1999, Mora-C. et al. 6 (INB); Cantón Coto Brus, near San Vito, Robert & Catharine Wilson Botanical Garden, near greenhouse, 8°47′03″N, 82°57′38″W, 1180 m, 19 June 2007, Moran 8176 (CR, INB, NY, USJ); Cantón Coto Brus, near San Vito, Robert & Catharine Wilson Botanical Garden, near greenhouse, 8°47′03″N, 82°57′38″W, 1180 m, 19 June 2007, Moran & Loriga 8133 (CR, INB, NY, USJ); drainage of Coto Brus R., 1470 m, 22 Feb. 1965, Nisman 149 (CR, GH); Cantón de Golfito, Valle de Coto Colorado, Golfito, Cacão, 8°37′35″N, 83°11′05″W, 50 – 130 m, 12 June 1994, Rojas 1177 (INB); Cocos Island, 5°32′21″N, 87°03′26″W, 50 – 300 m, 22 June 1997, Rojas 3649 (CR, INB, MO, NY); Cocos Island, 5°32′N, 87°05′W , 30 July 2001, Trusty 146 (US); San José: Cerro Turubares, [9°48′N, 84°28′W], 500 m, 12 Feb. 1910, Brade 570 (S); Hacienda El Rodeo, 9°55′N, 84°15′W, 700 m, 22 Feb. 1982, Burger et al. 11927 (CR, F); Escazú, San Antonio, 9°52′40″N, 84°08′13″W, 1850 m, 9 Sept. 2004, Chacón 6024 (CR); hills at SW part of Montañas Jamaica, c. 2.5 km NE of Bijagual de Turrubares, Carara Reserve, 9°45′N, 84°33′W, 460 – 520 m, 26 June 1985, Grayum et al. 5487 (CR, MO); Zona Protectora La Cangrega, along Río Negro, c. 1.5 km E of Santa Rosa de Puriscal, 9°42′N, 84°23′W, 320 m, 14 May 1987, Grayum et al. 8326 (CR, MO); Los Cuadros, Guadalupe, [9°57′N 84°03′W], 1300 m, Dec. 1912, Jiménez 808 (P, US); San Juan, [09°58′N 84°05′W], May 1913, Jiménez 891 (P); Bosques de Carara, Puriscal, [9°51′N 84°19′W], 400 m, 12 Feb. 1913, Jiménez 897 (P); WSW of San José, SW of Santiago de Puriscal, between Cerbantana and Crecedes Sur, [9°49′N, 84°20′W], 1100 m, 13 Aug. 1970, Lellinger & White 1576 (MICH, MO, US); Cantón de Puriscal, Zona Protectora La Cangreja, Cuenca del Río Negro, San Martín de Puriscal, E de La Fila Vara Blanca, 9°44′12″N, 84°23′28″W, 800 m, 20 Nov. 1993, Morales 2036 (CR, INB, MO); Acosta, Palmichal, Zona Protectora Cerros de Escazú, 9°50′32″N, 84°10′21″W, 1450 m, 23 June 2004, Quesada 5770 (CR); Finca Ortuna, Desamparados, [9°54′N 84°04′W], 1200 m, 15 March 1956, Scamman & Holdridge 7961 (GH, US); San Francisco de Guadalupe, [9°57′N, 84°04′W], 1170 m, Jan. 1896, Tonduz 9868 (B, US); Unknown: 1901 – 1905, Wercklé s.n. (MICH, NY); 1901 – 1905, Wercklé s.n. (NY, US); Panama. Bocas del Toro: A orillas del Changuinola y bosque alrededor de Corriente Grande, [8°56′N, 82°25′W], 25 Feb. 1980, Correa et al. 3978 (US); between Fortuna and Chiriquí Grande, 2.2 miles N of Continental Divide, 6.3 miles N of bridge over Fortuna Lake, 8°45′N, 82°16′W, 820 m, 12 March 1985, Croat & Grayum 60386 (AAU, MO, NY); between Fortuna and Chiriquí Grande, 1.2 miles N of Continental Divide, 5.3 miles N of bridge over Fortuna Lake, 8°45′N, 82°16′W, 910 m, 12 March 1985, Croat & Grayum 60435 (MO); Chiriquí: vic. of El Boquete, [9°10′N, 82°16′W], 1000 – 1500 m, 16 Feb. 1918, Cornman 973 (US); vic. of Boquete, [8°46′N, 82°25′W], 1000 – 1500 m, 18 Feb. 1918, Cornman 1003 (F, US); Palo Sante, 3 miles N of Volcán, [8°46′N, 82°40′W], 19 Feb. 1971, Croat 13570 (MO); vic. of Fortuna Dam in Valley of Río Chiriquí, along aqueduct to water source for IRHE facilites at dam, 8°45′N, 82°18′W, 1200 – 1300 m, 20 June 1987, Croat 66527 (MO); vic. of Fortuna Dam, valley of Río Chiriquí, along aqueduct trail for water supply for IRHE facilities, 8°45′N, 82°18′W, 1100 – 1200 m, 21 June 1987, Croat 66561 (MO); vic. of el Boquete, [9°10′N, 82°16′W], 1300 m, 14 Feb. 1918, Killip 5211 (GH, US); vic. of el Boquete, [9°10′N, 82°16′W], 1000 – 1300 m, 2 – 8 March 1911, Maxon 4952 (NY, US); Coclé: vic. of El Valle, 8°35′N, 80°07′W, 800 – 1000 m, 5 Sept. 1938, Allen 788 (GH, MICH, MO, NY, US).

Habitat. Not known; 50 – 2800 m.

Conservation Status. Least Concern (LC).

Notes.Megalastrum galeottii is characterised by pinnae rachises abaxially with filiform to linear scales and without hairs. Furthermore, most specimens are puberulent between the veins abaxially with hairs 0.1 – 0.2 mm long, erect, and acicular (specimens glabrous between the veins abaxially are Grayum et al. 5008, Correa et al. 3978, Jiménez 446, Cook & Doyle 157, and Wercklé s.n.). We consider this absence of hairs between the veins abaxially as part of the variation within the species because no other character appears to correlate with it. Also variable in M. galeottii is the length of the hairs on the adaxial surface of the laminae. Some specimens (e.g., Mickel 6396, Molina R. 8084) have longer hairs (0.8 – 1.2 mm, 4 – 6-celled) than usual (0.3 – 0.4 mm long, 1 – 3-celled).

Megalastrum costipubens is similar but differs by dense puberulence on the pinna rachises and costae abaxially. M. longipilosum also differs by pubescent pinna rachises abaxially and further differs by longer (0.6 – 1.3 (– 2) mm, 4 – 7 (– 12)-celled) hairs on the laminae adaxially.

Megalastrum galeottii can be confused with M. dentatum. The best way to distinguish them is by the scales on the abaxial surface of the pinna rachises and costae. In M. galeottii these scales are inconspicuous because they are generally sparse and ascending to appressed. In M. dentatum, however, the scales are widely spreading and immediately conspicuous to the naked eye (also along the petiole and rachises). With magnification, the scales of M. galeottii appear linear to filiform, c. 0.2 mm wide, with the cells obscure, whereas those of M. dentatum appear decidedly lanceolate, 0.3 – 0.4 mm wide, with more evident cells (nearly subclathrate). If large laminae of both species are compared, M. galeottii can be seen to be less finely divided, being only 2-pinnate-pinnatisect medially in contrast to M. dentatum which is 3-pinnate-pinnatisect medially.

Croat 27413 might represent a hybrid between Megalastrum galeottii and M. atrogriseum. It is minutely glandular on the laminae abaxially and has golden brown, firm scales like M. atrogriseum, but it resembles M. galeottii by the minute acicular hairs on the lamina abaxially and glabrous pinna rachises abaxially. Its spores appear misshapen and irregular. The indument of the abaxial surface between the veins only occurs in the soriferous parts of the leaf, not in the non-soriferous portions.

In a few specimens, stipitate-glandular hairs can be found between the veins abaxially (e.g., Conzatti 615, P). Such hairs are rare and occur among the typical acicular, erect hairs, from which they seem modified and differ only by their glandular apices.

Mickel 1077 is a typical by veins pilose adaxially (the c. 0.5 – 1.5 mm long). This suggests Megalastrum longipilosum from Costa Rica and Panama, but the specimen is otherwise typical of M. galeottii.

7. Megalastrum glabrumR. C. Moran & J. Pradosp. nov. A Megalastro atrogriseo lamina abaxialiter inter venas glabra et costulis venisque abaxialiter sparsim pubescentibus vel subglabris distinguitur. Typus: Costa Rica, Puntarenas, Monte Verde Cloudforest Reserve, 10°20′19.7″N, 84°47′34.1″W, 1709 m, 1 Feb. 2008, M. Sundue et al. 1737 (holotypus INB!; isotypi CR!, NY!, SP!, UC!).

http://www.ipni.org/urn:lsid:ipni.org:names:77105943-1

Rhizomes erect to decumbent, the scales golden, denticulate on both the margin and to a lesser extent on the surfaces (this sometimes hard to see); leaves up to 1.0 m long (estimate); scales of the petiole bases 0.6 – 10 × c. 1 mm, linear, twisted, spreading to ascending, yellow or golden, often shiny, denticulate; laminae up to 0.4 – 0.8 m long, 4-pinnate to 3-pinnate-pinnatisect at base, 2-pinnate-pinnatisect medially; basal pinnae c. 0.4 m long, strongly inequilateral (elongated basiscopically); pinna rachises abaxially glabous or nearly so, non-glandular, sparsely scaly, the scales to c. 1 – 2 × 0.2 – 0.3 mm long, linear, flat (not twisted), not tortuous, firm, spreading, golden brown, shiny, entire or sparsely denticulate; costules abaxially very sparsely pubescent, non-glandular, without scales, the hairs 0.4 – 0.6 mm long, 4- or 5-celled, spreading, adaxially densely pubescent, the hairs 0.7 – 1.0 mm long, 6 – 8-celled, often substrigose; laminar tissue between the veins glabrous on both surfaces, non-glandular, a few proscales present abaxially; veins visible on both surfaces, sparsely pubescent, the hairs 0.4 – 0.6 mm long, 4- or 5-celled; lamina margins ciliate, the hairs 0.2 – 0.3 mm long, 3- or 4-celled, non-glandular; indusia minute, fugacious, apparently consisting of a cluster of reddish proscales, c. 0.1 mm long, protruding from the centre of the sorus. Figs 3G – J and 4A; Map 1.

Distribution. Costa Rica, Panama.

Additional Specimens Examined. Costa Rica. Guanacaste: Las Nubes, 1 km N of Las Nubes Village, 8 km NW of Monteverde, 10°22′N, 84°51′W, 1200 m, 31 Aug. 1989, Haber & Zuchowski 9499 (MO); Puntarenas: c. 2 km SE of Monteverde, Pacific side, 10°18′N, 84°48′W, 1500 – 1550 m, 18 – 21 March 1973, Burger & Gentry 8566 (F, U); Monteverde, [10°18′N, 84°48′W], 20 June 1967, de la Sota 5026 (US); Monte Verde Cloudforest Reserve, above Quebrada Cuecha, [10°20′N, 84°47′W], 1540 – 1600 m, 6 Jan. 1976, Dryer 110 (F); rd from Pensiíon Flor-Mar to Monteverde Cloud Forest Reserve, 10°18′N, 84°48′W, 1400 – 1500 m, 8 Jan. 1987, Hill et al. 17693 (NY); Monteverde, [10°18′N, 84°48′W], 9 Jan. 1961, James 63 (MO); Monteverde, [10°18′N, 84°48′W], 1275 m, 16 Oct. 1963, Jiménez 1182 (F); Monteverde, [10°20′N, 84°47′W], 1600 m, March 1959, Palmer 112 (NY); Monteverde, [10°18′N, 84°48′W], 1500 m, 15 March 1960, Palmer 125 (CR, NY); Monteverde, [10°20′N, 84°47′W], 1500 m, 15 March 1960, Palmer 148 (NY); Monteverde Cloudforest Reserve, 10°20′19.7"N, 84°47′34.1"W, 1709 m, 1 Feb. 2008, Sundue et al. 1724 (CR, INB, NY, USJ); Monteverde Cloudforest Reserve, 10°20′19.7″N, 84°47′34.1″W, 1709 m, 1 Feb. 2008, Sundue et al. 1733 (CR, INB, NY, USJ). Panama. Chiriquí: vic. of El Boquete, [9°10′N, 82°16′W], 1000 – 1500 m, 7 Feb. 1918, Cornman 862 (US).

Habitat. Wet forests; 1000 – 1700 m.

Conservation Status. Least Concern (LC).

Notes.Megalastrum glabrum resembles M. atrogriseum by the flat (not twisted), yellowish, wide scales on pinna rachises and costules abaxially; however, it differs by the laminar tissue between the veins glabrous abaxially and the sparse pubescence on the costules and veins abaxially (versus dense pubescence). The first characteristic is referred to by the specific epithet glabrum. With the exception of the single collection from Panama, all specimens of this species have been collected in Costa Rica on the drier Pacific side of the Monteverde Cloudforest Reserve.

8. Megalastrum gompholepisR. C. Moran & J. Pradosp. nov.Megalastro subinciso rhachidibus pinnarum abaxialiter glabris et divisionibus laminarum similis, sed ab eo rhachidibus et costis abaxialiter squamis atrofuscis patentibus basibus abrupte dilatatis indutis differt. Typus: Guatemala, San Marcos, near Aldea Fraternidad, between San Rafael Pie de la Cuesta and Palo Gordo, W facing slope of the Sierra Madre Mts, [14°56′N, 91°49′W], 1800 – 2400 m, 10 – 18 Oct. 1963, L. O. Williams et al. 25752 (holotypus NY!; isotypi F!, GH!, MO!, US!).

http://www.ipni.org/urn:lsid:ipni.org:names:77105944-1

Leaves up to 2 m long; scales of the petiole base 10 – 30 × 0.1 – 0.3 mm, linear, brown, shiny, sparsely and minutely denticulate on the margins, the teeth often bifid; laminae 1 – 1.5 m long, up to 3-pinnate-pinnatisect at the base, 2-pinnate-pinnatisect medially; basal pinnae c. 0.5 m long, strongly inequilateral (elongated basiscopically); pinna rachises abaxially non-glandular (lacking both stipitate or sessile glands), subglabrous, scaly, the scales 0.2 – 5 × c. 0.5 mm long, the base usually expanded to slightly bullate, the apices filiform, patent, dark brown, shiny, denticulate, the hairs c. 0.2 – 0.3 mm long, 1- or 2-celled, erect, acicular, adaxially densely pubescent, the hairs 0.4 – 1 mm long, 3 – 5-celled, erect to slightly strigose, acicular; costules on both surfaces with indumenta like that of the pinna rachises; laminar tissue between veins on both surfaces non-glandular, glabrous, proscales present, appressed, reddish; veins on both surfaces obscure, sparsely pubescent to glabrous, the hairs c. 0.2 mm long, erect, acicular, adaxially sparsely pubescent to glabrous, hairs 0.5 – 0.7 mm long, 2- or 3-celled; hydathodes evident; lamina margins ciliate, the hairs 0.3 – 0.5 mm long, 2- or 3-celled, acicular, substrigose, non-glandular; indusia absent. Figs 1A – E and 10B; Map 2.

Distribution. Southern Mexico, Guatemala.

Additional Specimens Examined. Mexico. Chiapas: Motozintla de Mendoza, 45 – 50 km NE of Huixtla along rd to Motozintla, [15°22′N, 92°13′W], 1900 m, 17 Nov. 1971, Breedlove & Smith 22633 (F, MICH, MO, NY); Siltepec, ridge above Siltepec on the rd to Huixtla, [15°33′N, 92°19′W], 2000 – 2400 m, 18 Jan. 1973, Breedlove & Smith 31884 (F, NY); Unión Juárez, Volcán Tacaná por el camino de Talquián, por la linea divisoria con Guatemala, [15°40′N, 92°04′W], 2200 – 2400 m, 6 Feb. 1987, Martínez-S et al. 19692 (NY); Cerro del Boqueron, [15°10′N, 92°16′W], Aug. 1913, Purpus 6768 (NY). Guatemala. Quezaltenango: El Pocito, S of San Martín Chile Verde, on rd to Colombia, [14°49′N, 91°39′W], 2200 m, 27 Jan. 1941, Standley 85060 (F, NY); above Mujuliá, between San Martín Chile Verde and Colomba, [14°43′N, 91°46′W], 1800 m, 1 Feb. 1941, Standley 85477 (F, US); above Mujuliá, between San Martín Chile Verde and Colomba, [14°43′N, 91°46′W], 1800 m, 1 Feb. 1941, Standley 85554 (F); San Marcos: along Hwy 1, c. 9.2 miles W of San Pedro Sac towards San Rafael Pie de la Cuesta, [14°56′N, 91°55′W], 2300 m, 1 Jan. 1973, Luteyn & Almeda 3471 (F); along rd above Barranco Eminencia, [14°56′N, 91°55′W], 2700 m, 14 March 1939, Standley 68520 (F, US); Barranco Eminencia, rd between San Marcos and San Rafael Pie de la Cuesta, in upper part of the barranco between Finca La Lucha and Buena Vista, [14°56′N, 91°55′W], 2500 – 2700 m, 6 Feb. 1941, Standley 86322 (F); slopes of barrancos tributary to and bordering Río Vega, between San Rafael at NE portion of Volcán Tacaná and Guatemala-Mexico line, [15°14′N, 92°05′W], 2500 – 3000 m, 21 Feb. 1949, Steyermark 36301 (F, US); between Canjulá and La Unión Juárez, near SE portion of Volcán Tacaná, [15°40′N, 92°04′W], 2000 – 3000 m, 22 Feb. 1940, Steyermark 36422 (F); near Aldea Fraternidad, between San Rafael Pie de la Cuesta and Palo Gordo, W facing slope of the Sierra Madre Mts, [14°56′N, 91°49′W], 1800 – 2400 m, 10 – 18 Oct. 1963, Williams et al. 25586 (F); near Aldea Fraternidad, between San Rafael Pie de la Cuesta and Palo Gordo, W facing slope of the Sierra Madre Mts, [14°56′N, 91°49′W], 1800 – 2400 m, 10 – 18 Oct. 1963, Williams et al. 26219 (F, GH); outer slopes of Tajumulco Volcano, Sierra Madre Mts, c. 8 – 10 km W of San Marcos, [15°03′N, 91°54′W], 2300 m, 31 Dec. – 1 Jan. 1965, Williams et al. 26975 (F, GH); outer slopes of Tajumulco Volcano, Sierra Madre Mts, c. 10 km W of San Marcos, [15°03′N, 91°54′W], 2400 – 2700 m, 3 Jan. 1965, Williams et al. 27211 (F, US); outer slopes of Tajumulco Volcano, Sierra Madre Mts, c. 10 km W of San Marcos, [15°03′N, 91°54′W], 2400 – 2700 m, 3 Jan. 1965, Williams et al. 27216 (F, US).

Habitat. Wet forests, montane rain forests, cloud forests; 1800 – 3000 m.

Conservation Status. Least Concern (LC).

Notes. The specific epithet is derived from the Greek, gompho-, nail, and lepis, scale. It refers to the scales on the rachises, costae, and costules abaxially that are often gomphoid, or nail-shaped. Such scales have an abruptly widened base resembling the head of the nail and an elongated apical portion resembling the shaft of the nail. These scales are typically patent, conspicuous, and slightly bullate at base. Also distinctive are the nearly hairless rachises, costae, and costules abaxially; in contrast, most species of Megalastrum are pubescent on the abaxial surfaces of these axes.

Specimens of Megalastrum gompholepis have previously been identified as M. subincisum (Willd.) A. R. Sm. & R. C. Moran (Mickel & Smith 2004; Stolze 1981; Smith 1981); however, we treat that species in a narrow sense (type from Caracas, Venezuela), occurring in Mexico, Honduras, the Antilles, and northern Venezuela and Colombia (Moran et al.2009b). M. subincisum also has axes glabrous abaxially, but its scales on these axes differ by being lanceolate, flat (never slightly bullate at base), light brown, and appressed to slightly spreading.

9. Megalastrum heydei (C. Chr.) R. C. Moran & J. Prado, comb. et stat. nov.

http://www.ipni.org/urn:lsid:ipni.org:names:77105950-1

Dryopteris karsteniana var. heydei C. Chr. (Christensen 1920: 77). Ctenitis pulverulenta (Poir.) Copel. var. heydei (C. Chr.) Stolze (1977: 43). Megalastrum pulverulentum (Poir.) A. R. Sm. & R. C. Moran var. heydei (C. Chr.) A. R. Sm. & R. C. Moran (Smith & Moran 1987: 77). Type: Guatemala, Santa Rosa, Río de los Esclavos, E. T. Heyde & Lux s.n. [Donn. Sm. 3249, pro parte] (lectotype US-258590!, selected here; duplicates GH!, MO!, US-246305!).

Rhizomes not seen; leaves up to 4 m long (estimate); scales of the petiole base not seen; laminae up to 2.5 m long (estimate), 4-pinnate-pinnatifid at base, 3-pinnate-pinnatisect medially; basal pinnae c. 0.7 m long, strongly inequilateral (elongated basiscopically); pinna rachises glabrescent but when young with sparse glands, hairs, and scales, the glands spherical, c. 0.1 mm wide, the hairs c. 0.2 mm long, 2- or 3-celled, spreading, the scales to 1.0 – 1.8 mm long, lanceolate to linear, loosely appressed to spreading, brown, distinctly toothed, the margins sometimes slightly darker than the scale body, adaxially sparsely glandular, densely puberulent, the glands spherical, sessile, the hairs 0.1 – 0.2 mm long, 1 – 3-celled, dense, spreading, brown to reddish brown; costules with indument like that of the pinna rachises; laminar tissue between the veins non-glandular, glabrous on both surfaces, proscales present abaxially, sparse; veins abaxially evident, sparsely puberulent, the hairs 0.2 – 0.3 mm long, 2- or 3-celled, adaxially obscure, sparsely glandular, subglabrous, the glands c. 0.1 mm long, spherical, sessile; hydathodes evident; lamina margins sparsely ciliate to subglabrous, the hairs 0.1 – 0.2 mm long, 1- or 2-celled, non-glandular; indusia minute, fugacious, apparently consisting of a cluster of reddish proscales, c. 0.1 mm long, protruding from the centre of the sorus. Figs 11B and 12A – H; Map 2.

Distribution. Guatemala; known only from two collections.

Additional Specimen Examined. Guatemala. Without locality, 1892, Heyde 310 (US).

Habitat. Presumably in wet forests; 800 – 2200 m.

Conservation Status. Data Deficient (DD).

Notes. Because of its large, highly divided leaves (up to 4 m long) and distinctly dentate laminar scales, Megalastrum heydei resembles M. pulverulentum and M. semipubescens. It differs from these two species by minute (c. 0.1 mm long), often strigose hairs on both surfaces of the pinna rachises and costules. In contrast, these hairs in M. pulverulentum and M. semipubescens are 1 – 2 mm long. The short hairs on the axes of M. heydei resemble those of M. palmense, a species that differs by broad, flaccid, dull brown, entire laminar scales and less divided laminae (2-pinnate-pinnatisect medially). M. heydei is endemic to Guatemala, whereas M. palmense occurs only in the mountains of Costa Rica and Panama. In the two specimens available, the lamina has dried nearly black on the adaxial surface.

As pointed out by Stolze (1981), the type collection is mixed. Some Heyde and Lux specimens bearing Donnell-Smith’s distribution number 3249 represent Ctenitis excelsa (for instance, US sheet numbers 258596 and 830988).

Martínez 12943 (MO) was cited by Smith & Moran (1995) as this taxon from Honduras. It actually represents our new species, Megalastrum sparsipilosum.

10. Megalastrum intermediumR. C. Moran & J. Pradosp. nov.Megalastro lunensi et M. palmensi similis, sed a M. lunensi trichomatibus rhachidum pinnarum costularumque c. 0.1 mm longis saepe dense strigosis, a M. palmensi glandulis sphaericis sessilibus stipitatisque et rhachidibus pinnarum abaxialiter squamis 0.2 – 0.3 mm latis lanceolatis indutis differt. Typus: Costa Rica, Puntarenas, Cantón de Coto Brus, Cuenca Terraba–Sierpe, Mellizas, Finca Cafrosa, Sendero Ripario, 8°53′15″N, 82°46′15″W, 1300 – 1350 m, 19 Feb. 1998, E. Navarro V. & A. Rojas 916 (holotypus INB!; isotypus MO 3 sheets!).

http://www.ipni.org/urn:lsid:ipni.org:names:77105945-1

Rhizomes erect; leaves up to 1.5 m long (estimate); scales of the petiole bases 5 – 10 × 0.5 – 0.7 mm, lanceolate, twisted, spreading to ascending, brown, shiny, sparsely denticulate on the margins; laminae up to 0.4 – 0.8 m long, 3-pinnate-pinnatisect to 4-pinnate at base, 2-pinnate-pinnatisect to 3-pinnate medially; basal pinnae 0.3 – 0.4 m long, strongly inequilateral (elongated basiscopically); pinna rachises on both surfaces densely puberulent, without capitate-glandular hairs but with brown spherical sessile glands, sparsely scaly, the scales to 2 – 3 × 0.2 – 0.3 mm long, lanceolate, slightly twisted, flaccid, spreading, entire or nearly so, brown, shiny, the hairs 0.1 – 0.2 mm long, 1- or 2-celled, mostly substrigose; costules abaxially densely puberulent, scaly, glandular, the hairs 0.1 – 0.2 mm long, 1- or 2-celled, erect, acicular, the scales c. 1 × 0.2 – 0.3 mm, brown shiny lanceolate, subentire; the glands stalked and sessile-spherical, adaxially densely puberulent, the hairs 0.1 – 0.3 mm long, 1 – 3-celled mostly substrigose, the glands spherical, sessile brown, the scales c. 0.2 mm long, brownish, filiform; laminar tissue between the veins abaxially sparsely puberulent, glandular, the hairs 1 or 2-celled, erect, acicular, the glands both stalked and sessile, yellowish, a few proscales present abaxially, adaxially similar in indument but more sparsely so; veins visible on both surfaces, sparsely pubescent like the lamina tissue; hydathodes conspicuous; lamina margins densely ciliate, the hairs 0.2 – 0.3 mm long, 1 – 3-celled, acicular, non-glandular; indusia minute, fugacious, apparently consisting of a cluster of reddish proscales, c. 0.1 mm long, protruding from the centre of the sorus. Figs 13L, M and 14A; Map 2.

Distribution. Costa Rica, Panama.

Additional Specimens Examined. Costa Rica. Puntarenas: above San Vito at Finca Wilson, [8°49′N, 82°58′W], 2 & 4 April 1967, Evans & Bowers 3050 (U); Upper Río Burú, [9°01′N 82°51′W], 2010 m, 19 Aug. 1983, Gómez et al. 21672 (CR, MO); vic. of biological station at Finca Wilson, 5 km S of San Vito de Java, [8°49′N, 82°58′W], 1100 – 1200 m, 2 – 6 Aug. 1967, Lellinger 814 (US); Cantón de Coto Brus, Zona Protectora Las Tablas, Cuenca terraba-Sierpe, 8°58′30"N, 82°46′15"W, 1960 m, 18 Feb. 1998, Navarro V. 906 (INB, MO); Cañas Gordas, [8°45′N, 82°55′W], 1100 m, Nov. 1897, Pittier 10993 (US); Cantón de Coto Brus Z.P. Las Tablas, Cuenca Térraba-Sierpe, 1900 m, 13 Aug. 1997, Quesada 2017 (INB, MO); Zona Protectora Las Tablas, Sabalito, Las Alturas de Cotó, Estación Biológica Las Alturas, sendero a Cerro Echandi, postes 40 – 57, 8°59′N, 82°43′W, 2070 – 2230 m, 28 Dec. 1993, Rojas 853 (INB, MO). Panama. Bocas del Toro: La Pata del Cedro, 9°58′N, 82°56′W, 1500 m, 7 March 2004, Monro & Alfaro 4230 (MO); Chiriquí: El Boquete, trail back of O’Donnell’s, 9°10′N, 82°16′W, 14 March 1918, Cornman 1138 (MO, S); Ojo de Agua, property of Ratidon Hartmann, vic. of Sta. Clara, between Volcán and Río Sereno, 8°51′N, 82°45′W, 1520 m, 17 June 1987, Croat 66317 (AAU, MO); Río Quebrada, [8°26′N, 82°26′W], 1650 m, 2 Aug. 1918, Killip 5136 (GH, MICH); Río Quebrada, [8°26′N, 82°26′W], 1650 m, 2 Aug. 1918, Killip 5502 (US).

Habitat. Wet forests; 1100 – 2200 m.

Conservation Status. Least Concern (LC).

Notes.Megalastrum intermedium appears intermediate between M. lunense and M. palmense (thus the specific epithet intermedium). It resembles M. lunense by the pinna rachises and costules with scales spreading, shiny brown, and denticulate, and laminae abaxially between the veins with a mixture of sessile and stalked glands. It resembles M. palmense by short (c. 0. 1 mm long), dense, often strigose hairs on both surfaces of the pinna rachises.

11. Megalastrum longiglandulosumR. C. Moran & J. Pradosp. nov. A Megalastro gilbertii rhachidibus pinnarum abaxialiter dense pubescentibus ac squamis filiformibus indutis et pinnulis inter venas abaxialiter trichomatibus usque ad 3 mm longis vestitis differt. Typus: Costa Rica. Puntarenas: Osa Peninsula, c. 4 miles W of Rincón de Osa, near airfield, 8°42′N, 83°31′W, 30 m, 4 – 7 June 1968, W. C. Burger & R. G. Stolze 5421 (holotypus F-2 sheets!; isotypi CR!, GH-2 sheets!, US!).

http://www.ipni.org/urn:lsid:ipni.org:names:77105946-1

Megalastrum longipilosum A. Rojas var. glandulosum A. Rojas (2007: 20, f. 3). Type: Costa Rica, Puntarenas, Cantón de Osa, Sierpe, San Juan, cuenca media del Río San Juan, siguiendo su curso aguas arriba, 8°43′20"N, 83°32′20"W, 100 – 400 m, 5 Nov. 1990, G. Herrera 4580 (holotype CR!; isotype MO!).

Rhizomes compact, ascending (from original description); leaves up to 1.5 m long; scales of the petiole bases 15 – 25 × 0.3 – 0.6 mm, linear, spreading-ascending, brown, shiny, denticulate on the margins; laminae up to 0.9 m long, 3-pinnate-pinnatisect at base, 2-pinnate-pinnatisect medially; basal pinnae 15 – 40 cm long, strongly inequilateral (elongated basiscopically); pinna rachises abaxially glandular (both stipitate and sessile glands present), densely pubescent, scaly, the scales 0.8 – 1.0 × c. 0.2 mm, linear-lanceolate, non-bullate, mostly appressed, shiny, brown, sparsely denticulate, adaxially densely pilose, sparsely glandular, sparsely scaly, hairs c. 1.0 mm long, 6 – 8-celled, spreading, glands spherical and sessile and stalked, c. 0.2 mm long; costules sparsely pilose, glandular, hairs 0.3 – 0.5 mm long, 3 – 5-celled, erect, the glands both sessile and spherical, and stalked-glandular, adaxially like the pinna rachises adaxially; laminar tissue between the veins abaxially densely glandular, the glands 0.2 – 0.3 mm long, 2 or 3-celled, erect, capitate, the terminal cell yellow, shiny, these mixed with hairs 0.2 – 0.3 mm long, acicular, 1 or 2-celled, adaxially without hairs, non-glandular or with a few sparse sessile spherical glands; veins visible on both surfaces, abaxially with indumenta like the laminar tissue, adaxially sparsely pilose, glandular, the hairs 0.4 – 0.8 mm long, 4 – 6-celled, glands sessile, spherical; hydathodes evident; lamina margins pubescent, hairs 0.3 – 0.5 mm long, 3 or 4-celled, ascending, acicular, non-glandular; indusia absent. Fig. 15F – M; Map 2.

Distribution. Costa Rica, endemic to the Osa Peninsula.

Additional Specimens Examined. Costa Rica. Puntarenas: Osa Peninsula, on ridge 9.5 km W of Rincón de Osa, [8°42′N, 83°31′W], 600 m, 17 July 1967, Mickel 2756 (NY, US).

Habitat. Wet forests; 30 – 600 m.

Conservation Status. Least Concern (LC).

Notes.Megalastrum longiglandulosum bears dense, erect, long-stalked capitate-glandular hairs between the veins abaxially (the specific epithet refers to this character). In this it resembles M. gilbertii (Clute) R. C. Moran, J. Prado & Labiak, a West Indian species, but it differs by filiform scales and densely pubescent pinna rachises abaxially (M. gilbertii has wider scales and hairless pinna rachises abaxially). The size of the glandular hairs on the laminar tissue between the veins abaxially is also different: those on M. longiglandulosum are up to 0.3 mm long, whereas those on M. gilbertii are c. 0.1 mm long.

Two Central American species, Megalastrum lunense and M. atrogriseum, also have long-stalked capitate-glandular hairs on the laminae abaxially, but their hairs are usually mixed with many non-glandular, acicular hairs of variable length. These two species also differ from M. longiglandulosum by the scales on the abaxial surfaces of the pinna rachises: the scales of M. lunense are ovate-lanceolate and flaccid, whereas those of M. atrogriseum are linear, flat (not twisted or tortuous), and golden brown.

12. Megalastrum longipilosumA. Rojas (2007: 18, f. 2). Type: Costa Rica. Puntarenas: Cantón de Coto Brus, Cuenca Térraba-Sierpe, Estación Pittier, Sendero Río Canasta, 9°01′32″N, 82°57′45″W, 1670 m, 26 Nov. 1997, A. Rodríguez & L. Vargas 2807 (holotype CR!; isotypes INB!, NY!).

Rhizomes erect; leaves up to 1.2 m long; scales of the petiole bases 10 – 20 × c. 1.0 mm, linear, spreading, brown, shiny, sparsely denticulate on the margins; laminae up to 0.8 m long, 3-pinnate-pinnatifid at base (2-pinnate-pinnatifid in Mexican plants), 2-pinnate-pinnatifid medially; basal pinnae 15 – 20 cm long, strongly inequilateral (elongated basiscopically); pinna rachises abaxially non-glandular (both stipitate and sessile glands lacking), densely pilose, scaly, the scales 1.0 – 1.5 × c. 0.2 mm, linear-lanceolate, non-bullate, spreading, shiny brown, entire, adaxially densely pilose, 1.0 – 1.5 mm long, 4 – 7-celled, more sparsely scaly; costules abaxially densely to sparsely pilose, hairs 0.3 – 0.7 mm long, 3 – 5-celled, spreading to slightly strigose, adaxially scales absent, the hairs 0.8 – 1.5 mm long, 3 – 7-celled; laminar tissue between the veins on both surfaces non-glandular, glabrous; veins visible on both surfaces, non-glandular, sparsely pilose, the hairs like those of the costules; hydathodes evident; lamina margins pilose, the hairs 0.3 – 0.4 (– 0.8) mm long, 1 – 3 (– 6)-celled, spreading to ascending, non-glandular; indusia minute, fugacious, apparently consisting of a cluster of reddish proscales, c. 0.1 mm long, protruding from the centre of the sorus. Figs 7C, D and 15A – E; Map 2.

Distribution. Costa Rica, Panama; W. Ecuador, Bolivia.

Additional Specimens Examined. Costa Rica. Alajuela: Reserva Forestal San Ramón, N of the station, c. 2 km in the mts, 10°12′40″N, 84°36′20″W, 1000 m, 18 April 1991, Bittner 985 (AAU); E of the Río San Rafael and S of the hot springs, W of La Marina, 10°23′N, 84°23′W, 500 m, 19 May 1968, Burger & Stolze 5054 (CR, F, GH, US); Guatuso, Parque Nacional Volcán Tenorio, Cuenca del Río Frío, El Pilón, sector La Catarata, 800 m, 11 May 2000, Chaves et al. 466 (INB, NY); above Bijagua, slopes of Miravalles, [10°44′N, 85°03′W], 1500 m, Nov. 1982, Gómez 19056 (F, MO); Reserva Monteverde, Poco Sol, 13 km S of Fortuna, 10°21′N, 84°41′W, 700 – 900 m, 20 Aug. 1989, Haber & Zuchowski 9354 (INB, MO); c. 20 km N of San Ramón, Reserva Biológica Alberto Brenes, [10°13′N, 84°37′W], 1100 m, 17 July 1983, Moran 3258 (MO); Cantón de San Ramón, Cuenca del San Carlos, cerca de la Reserva Forestal San Ramón, 10°14′35″N, 84°33′40″W, 760 – 930 m, 11 Nov. 1997, Rojas et al. 3895 (INB, MO); Cantón de Guatuso, Parque Nacional Tenorio, Cuenca del Río Frio, Estación Quebradón, camino a Lago Cote, Fila Vieja Dormida, 10°36′35″N, 84°55′35″W, 700 – 1000 m, 9 June 1998, Rojas et al. 4630 (INB); Cartago: Capelladas, [09°55′N 83°47′W], 1100 m, April 1901, Alfaro 16552 (P); Guanacaste: Parque Nacional Guanacaste La Cruz, 9 km S de Santa Cecília, Estación Pitilla, 10°59′26″N, 85°25′40″W, 700 m, 28 Sept. 1990, Cháves 159 (CR, F); slopes of Miravalles, above Bijagua, [10°44′N, 85°03′W], 1500 m, Nov. 1982, Gómez et al. 19073 (AAU, MO); Liberia, Parque Nacional Guanacaste, Estación Cacão, 10°55′N, 85°28′W, 1100 – 1500 m, 5 Sept. 1996, Rojas & Mata 3075 (INB, MO, NY); Heredia: Virgen del Socorro, [10°17′N, 84°10′W], 1000 m, 11 July 1983, Moran 3162 (F, MO); Limón: SW of Siquirres, on rd to Turrialba between Moravia and Guayacán, [10°02′31″N, 83°32′57″W], 700 m, 4 Aug. 1970, Lellinger & White 1442 (US); Los Diamantes, USDA Rubber Plant Station, [10°12′49″N, 83°46′33″W], 300 m, 29 – 30 April 1951, Scamman 5938 (GH); Puntarenas: San Vito, [8°49′N, 82°58′W], 1500 m, 28 June 1967, de la Sota 5210 (US); near San Vito, along road and in cloud forest above and W of Finca Wilson, [8°48′N, 82°58′W], 1200 m, 3 – 5 Aug. 1967, Evans 3082 (MO, U); Cantón de Osa, Fila Costeña, cabeceras del Río Piedras Blancas, Cerro Anguciana, faldas al Oeste, 8°48′56″N, 83°10′37″W, 1400 – 1600 m, 10 Dec. 1983, Hammel 19276 (INB, MO); 3 – 5 km NW of the biological station at Finca Wilson, 5 km S of San Vito de Java, [8°47′N, 82°57′W], 1300 – 1400 m, 3 – 5 Aug. 1967, Lellinger 833 (CR, MO); vic. of biological station at Finca Wilson, 5 km S of San Vito de Java, 1 – 4 km SW of station, [8°48′N, 82°58′W], 1200 – 1400 m, 4 Aug. 1967, Mickel 3116 (NY, US); 5 km S of San Vito de Java, vic. of biological station at Wilson finca, [8°48′N, 82°58′W], 1100 – 1400 m, 22 March 1967, Mickel 2049 (NY); Coto Brus, Parque Nacional La Amistad, Valle del Silencio, sector de acampar a los jardines, 9°07′15″N, 82°57′55″W, 2500 m, 14 April 1996, Quesada et al. 1474 (INB, MO, NY); San José: Parque Nacional Braulio Carrillo, at the intersection of Hwy 32 and Río Sucio, 12 Feb. 2008, Sundue & Nitta 1818 (CR, INB, NY, USJ); Unknown: [no date], James 73 (MO). Panama. Chiriquí: vic. of Fortuna Dam, in valley of Río Chiriquí, along aquaduct trail for water supply to IRHE facilities, 8°45′N, 82°18′W, 1100 – 1200 m, 21 June 1987, Croat 66574 (MO); Veraguas: 5 miles W of Santa Fe on road past Escuela Agrícola Alto Piedra on Pacific side of divide, [8°29′N, 81°06′W], 800 – 1200 m, 18 March 1973, Croat 23007 (MO, US).

Habitat. Cloud forests; 300 – 2500 m.

Conservation Status. Least Concern (LC).

Notes.Megalastrum longipilosum is characterised by pinna rachises abaxially sparsely pilose and costules and veins adaxially pilose, the hairs typically 0.8 – 1.5 mm long, 3 – 7-celled. The scales on the pinna rachises abaxially are filiform, and the laminar tissue between the veins abaxially is glabrous. The pinnules are obtuse and often close-set compared to those of other species in the genus, most of which have acute or acuminate pinnules.

Megalastrum galeottii is similar in size and lamina cutting but differs by pinna rachises glabrous abaxially, shorter hairs adaxially on the laminae, and (in most specimens) the presence between the veins abaxially of minute (c. 0.1 – 0.2 mm long) acicular, erect hairs. Also similar to M. longipilosum is M. costipubens, which can be distinguished by the pubescence characters given in the key.

Several specimens of Megalastrum longipilosum from the Fila Costeña of Costa Rica (e.g., Lellinger 833, Evans & Bowers 3082) are atypical by laminae densely pilose along the veins on both surfaces. The hairs are extremely long (1 – 3 mm) and many-celled (7 – 12-celled). We interpret this as part of the variation within the species because no other characteristic appears to correlate.

13. Megalastrum lunense (H. Christ) A. R. Sm. & R. C. Moran (Smith & Moran 1987: 128).

Aspidium lunense H. Christ (1906: 55); Dryopteris lunensis (H. Christ) C. Chr. (Christensen 1913: 35); Ctenitis lunensis (H. Christ) Lellinger (1977: 709). Type: Costa Rica. Cartago: Luna, [9°48′N 83°53′W], in 1905, C. Wercklé s.n. (lectotype 2 sheets: P-00600390!, P-00600389!, selected here; duplicates P-00600388!, fragm. BM! and U! ex P!).

Dryopteris pansamalensis C. Chr. (Christensen 1920: 72); Ctenitis pansamalensis (C. Chr.) Ching (1940: 250); Megalastrum pansamalense (C. Chr.) A. R. Sm. & R. C. Moran (Smith & Moran 1987: 128). Type: Guatemala. Alta Verapaz: Pansamalá, von Türkheim s.n. [Donn. Sm. 1005] (lectotype US-831791!, selected by Mickel & Smith (2004); duplicates UC!, US-831790!, fragm. MO!).

Ctenitis skutchii Lellinger (1985: 375, f. 6); Megalastrum skutchii (Lellinger) A. R. Sm. & R. C. Moran (Smith & Moran 1987: 129). Type: Costa Rica. San José: Vic. of El General, [9°30′N, 83°40′W], 1190 m, Dec. 1935, A. F. Skutch 2337 (holotype US!; isotypes F!, GH!, K, MICH!, MO!, NY!, S!).

Rhizomes erect; leaves up to 1.5 m long (estimate); scales of the petiole bases 10 – 15 × c. 2.5 mm, ovate to lanceolate, slightly twisted, spreading to ascending, brown, shiny, denticulate on the margins; laminae up to 0.3 – 0.8 m long, 3-pinnate-pinnatisect, 2-pinnate-pinnatisect medially; basal pinnae c. 0.2 – 0.4 m long, strongly inequilateral (elongated basiscopically); pinna rachises abaxially densely pilose, non-glandular (i.e., without capitate-glandular hairs or sessile spherical glands), scaly, the scales to c. 2 – 3 × c. 1.5 mm, ovate to lanceolate, flat (not twisted), flaccid, spreading, brown, shiny, entire or nearly so, brown, shiny, adaxially non-glandular, densely pilose, the hairs 1 – 1.3 mm long, 6 – 9-celled, spreading; costules on both surfaces with indument like the pinna rachises but with a few sessile and stalked glands; laminar tissue between the veins abaxially puberulent, the hairs 0.1 – 0.3 mm long, 3-celled, erect, acicular, the glands stalked, c. 0.1 – 0.2 mm long, capitate, these mixed with sparse sessile spherical glands, a few proscales present abaxially, adaxially with similar indument but sparser; veins visible on both surfaces, sparsely pubescent like the lamina tissue; hydathodes conspicuous; lamina margins ciliate, the hairs 0.2 – 0.3 mm long, 1 – 3-celled, erect, acicular, non-glandular; indusia absent. Figs 13A – G and 14B; Map 2.

Distribution. Mexico, Belize, Guatemala, Nicaragua, Costa Rica, Panama.

Additional Specimens Examined. Belize. Toledo: Columbia R. Forest Reserve, Little Quartz Ridge, middle slopes on S flank, N of Camp 2, 16°23′50″N, 89°05′36″W, 850 m, 18 Feb. 1997, Holst 5820 (MO). Costa Rica. Alajuela: San Ramón, Angeles, Reserva Biológica Alberto Brenes, 10°12′40″N, 84°36′20″W, 1100 m, 16 April 1991, Bittner 970 (CR); Reserva Forestal San Ramón, Quebrada Cacical, 10°14′13″N, 84°36′22″W, 2 May 1987, Herrera et al. 591 (AAU, MO); N of San Ramón, c. 4 km N of Balsa along rd to Colonia Palmareña, 10°11′N, 84°30′W, 1300 m, 23 July 1970, Lellinger 1233 (CR, MO, US); c. 20 km N of San Ramón, Univ. of San Ramón’s Biological Station, 10°13′N, 84°37′W, 1000 m, 17 July 1983, Moran 3186, pro parte (CR, GH, MO, NY, US); c. 20 km N of San Ramón, Univ. of San Ramón’s Biological Station, 10°15′N, 84°30′W, 800 – 900 m, 27 Feb. 1988, Moran 4122 (CR, MO); Cantón de San Ramón, Reserva Biológica Monteverde, Estación Aleman’s, 10°22′00"N, 84°48′W, 1200 – 1500 m, 20 May 1995, Rojas 1871 (INB); Cantón de San Ramón, Reserva Biológica Monteverde, Estación Aleman’s, 10°22′00″N, 84°48′W, 1200 – 1500 m, 20 May 1995, Rojas 1898 (F, INB); Cantón de Guatuso, Parque Nacional Tenorio, Cuenca del Río Frio, Estación Quebradón, camino a Lago Cote, Fila Vieja Dormida, 10°36′35″N, 84°55′35″W, 700 – 1000 m, 9 June 1998, Rojas et al. 4631 (CR, INB, NY); Cantón de Upala, Cuenca del Pizote, Estación San Cristóbal, Quebrada Cucaracho, 10°52′58″N, 85°24′05″W, 600 – 625 m, 11 Aug. 1998, Rojas et al. 4736 (CR, INB); Cantón de Alajuela, Cuenca del Sarapiqui, cerca de la union de la Quebrada Ten Fe y Río Cariblanco, 10°15′45″N, 84°11′35″W, 840 – 950 m, 2 Dec. 1987, Rojas et al. 4129 (INB); N slope of ridge along quebrada draining E to Río Cataratitas, c. 20 km NW of San Ramvalley of Santa Elena along rd to Ixcan, 30 km E of paved rd at San Ramón, 10°13′N, 84°32′W, 850 m, 3 Feb. 1986, Smith 2256 (CR, MO); Cartago: San Juan del Norte, 9°48′N, 84°04′W, 1100 m, 15 March 1955, Scamman 7654 (GH); Valle de Navarro, [9°48′N 83°53′W], 1400 m, 1905, Wercklé s.n. [Inst. Physico-geogr. Nat. costarricense 16788] (BM); Guanacaste: Parque Nacional Guanacaste, La Cruz, Sta. Cecilia, Estación Pitilla, cunca superior de Quebrada Nacho, 10°59′10″N, 85°26′10″W, 800 m, 24 May 1990, Herrera 3896 (INB, MO); Limón: Cantón de Limón, along divide between Río Xikiari and Río Boyei, above Cabécar village of Almirante, 9°46′30″N, 83°20′00″W, 1100 – 1150 m, 13 Aug. 1995, Grayum 10914 (INB); c. 3 km SW of Suretka, RECOPE test drill site, near entrance, [9°35′N, 82°57′W], 70 m, 6 July 1983, Moran 3117 (CR, F); Cantón de Matina, Zona Protectora Pacuare, Cuenca del Matina, bosques aledaños a la comunidad Las Brisas, 9°59′39"N, 83°27′11"W, 500 m, 16 July 1998, Vargas 3809 (INB); Puntarenas: Fila el Tigre, SE of Las Alturas, 8°56′N, 82°51′W, 1350 – 1450 m, 29 Aug. 1983, Davidse 24225 (CR, MO); Cantón de Coto Brus, Zona Protectora Las Tablas, Las Alturas de Cotó, Sabalito, Estación Biológica Las Alturas, sendero a Cerro Echandi, postes 40 – 58, 8°57′15″N, 82°50′10″W, 1580 – 1620 m, 19 Dec. 1993, Rojas 699 (INB); San José: vic. of El General, [9°30′N, 83°40′W], 950 m, Jan. 1937, Skutch 3113 (NY, US); vic. of El General, [9°30′N, 83°40′W], 1190 m, Dec. 1935, Skutch 2337, pro parte (K, MICH); Unknown: along Interamerican Hwy, 14 April 1951, Scamman 5937 (US). Guatemala. Alta Verapaz: Chaciraciha, Finca Seamay, Senahu, [15°31′60″N, 89°46′60″W], 30 July 1936, Hatch & Wilson 210 (US); trail between Sepacuité and Secanquim, [15°31′60"N, 89°46′W], 550 – 900 m, 12 Jan. 1905, Maxon & Hay 3268A (BM, US); near Chirriacté, on the Petén Hwy, [15°40′60″N, 89°58′60″W], 900 m, 9 April 1941, Standley 91653 (F); near Finca Pansamala, [15°27′N, 90°01′W], 1300 m, Sept. 1886, von Türckheim 1055 (MO); Finca Volcán, 1000 m, 21 Feb. 1934, Wilson 234 (F). Mexico. Chiapas: La Trinitaria, 15 km ENE of Dos Lagos above Sta. Elena, [16°8′0″N, 91°47′0″W], 1000 m, 29 Dec. 1981, Breedlove 56669 (NY); La Independencia, valley of Santa Elena along rd to Ixcan, 30 km E of paved rd at Montebello, 800 m, 29 Nov. 1976, Breedlove 42014 (MO). Nicaragua. Zelaya: Costado suroeste de Cerro El Hormiguero, 13°44′10″N, 84°59′50″W, 900 – 1000 m, 18 April 1979, Grijalva 468 (CR); along trail from Cerro El Inocente toward Cerro Saslaya, near source of Caño Majagua, 13°46′N, 85°01′W, 1050 – 1150 m, 8 March 1978, Stevens et al. 6666 (CR). Panama. Coclé: Valle de Antón, [8°37′N, 80°07′W], 1000 m, 5 June 1939, Alston 8743 (CR); Cerro Pilón, [8°33′0″N, 80°42′0″W], 900 – 1173 m, 16 March 1973, Liesner 780 (MO); La Mesa, 2 km W of Cerro Pilón, 8°33′0″N, 80°42′0″W, 900 m, 22 July 1976, Sullivan 524 (MO); Veraguas: Cerro Tute, c. 2 km N of Santa Fé, along ridge trail to summit, N of escuela primaria Alto de Piedra, 8°31′N, 84°04′W, 900 – 1000 m, 5 Feb. 1988, Moran 4019 (MO); c. 4 miles N of Santa Fé, where road crosses the “primer brazo del Río Calovébora”, 8°34′N, 81°07′W, 600 m, 7 Feb. 1988, Moran 4075 (MO).

Habitat. Wet forests; 70 – 1600 m.

Conservation Status. Least Concern (LC).

Notes.Megalastrum lunense is characterised by petioles and pinna rachises with wide (up to 2.5 mm) ovate to lanceolate scales, pinna rachises and costules abaxially with hairs up to 1 mm long and 4 – 7-celled, abaxial surfaces of the laminae with glands varying from stalked to sessile. Similar is M. palmense, which differs by scales flaccid and appressed to the petioles and leaf axes, lack of glands (both stipitate and sessile), and minute (c. 0.1 mm long) substrigose pubescence on the leaf axes abaxially.

Megalastrum atrogriseum also has long hairs abaxially, and these can be mixed with sessile and stalked glands. It differs by its pinna rachis scales that are narrower (0.3 – 0.4 mm), flat (not twisted or tortuous), yellowish brown, and denticulate.

14. Megalastrum macrotheca (Fée) A. R. Sm. & R. C. Moran (Smith & Moran 1987: 128).

Phegopteris macrotheca Fée (1866: 56); Dryopteris macrotheca (Fée) C. Chr. (Christensen 1913: 35); Ctenitis macrotheca (Fée) Ching (1940: 250). Type: Guadeloupe, 1864, F. L’Herminier s.n. (lectotype P-00600368!, selected here; duplicates P-00610561!, RB!, fragm. BM! ex P!).

Polypodium biseriale Baker (in Hooker & Baker 1867: 309); Nephrodium biseriale (Baker) Diels (1899: 170); Dryopteris biserialis (Baker) C. Chr. (Christensen 1905: 254); Ctenitis biserialis (Baker) Lellinger (1975: 108); Megalastrum biseriale (Baker) A. R. Sm. & R. C. Moran (Smith & Moran 1987: 127). Type: Ecuador, R. Spruce 4656 (lectotype K-000200146!, selected here; duplicates K-000227603!, fragm. BM!).

Polypodium mazei E. Fourn. ex Baker (1891: 457); Polypodium connexum var. mazei (Fourn. ex Baker) E. Fourn. ex Duss (1903: 56); Dryopteris mazei (Baker) C. Chr. (Christensen 1905: 277). Type: Guadeloupe, 1050 – 1100 m, 1883 – 1884, H. P. Mazé ‘643, 786’ (syntypes K!; isosyntypes NY!, P!, S!).

Dryopteris subincisa (Willd.) Urb. var. haitiensis Brause (in Urban 1922: 67); Dryopteris haitiensis (Brause) Urb. & Maxon (Urban & Maxon 1924: 91); Ctenitis haitiensis (Brause) Lellinger (1984: 56); Megalastrum haitiense (Brause) A. R. Sm. & R. C. Moran (Smith & Moran 1987: 128). Type: Haiti, Mt Blanche, Morne de la Hotte, 1400 m, E. L. Ekman 556 (holotype B; fragment NY!, fragment and photo BM!, US ex B).

Ctenitis bidecorata Lellinger (1985: 373, f. 5); Megalastrum bidecoratum (Lellinger) A. R. Sm. & R. C. Moran (Smith & Moran 1987: 127). Type: Costa Rica, Cartago, Estrella-Sta. María road, [9°47′N 83°58′W], 230 m, 21 April 1928, H. Stork 1518 (holotype US! photo CR! ex US).

Leaves up to 1.0 m long; scales of the petiole bases 0.7 – 1.0 × 0.1 – 0.2 cm, lanceolate, spreading, brown, sparsely denticulate on the margins; laminae up to 0.8 m long, 2-pinnate-pinnatifid at base, 1-pinnate-pinnatisect medially; basal pinnae 10 – 20 cm long, subequilateral to inequilateral; pinnules broadly adnate to the pinna rachis, 3 – 6 cm long; pinna rachises abaxially non-glandular, densely pubescent, scaly, the scales 3 – 5 mm long, lanceolate, non-bullate, brown, entire to sparsely denticulate, non-bullate, rachises adaxially non-glandular, lacking scales, densely pubescent, hairs 1.0 – 1.2 mm long, 5 – 8-celled, ascending, appressed, or spreading brownish; costules on the abaxial surface non-glandular, puberulent and scaly, the hairs 0.2 – 0.4 mm long, 2 or 3-celled, erect to spreading, acicular, the scales like those on the pinna rachises abaxially but smaller, 2 – 2.5 mm long, the adaxial surface sparsely pubescent, the hairs 0.4 – 0.5 mm long, substrigose; laminar tissue between veins glabrous on both surfaces; veins visible on both surfaces, abaxially non-glandular, very sparsely pubescent, non-scaly, the hairs c. 0.2 mm long, 1- or 2-celled, the scales c. 0.3 mm long, uniseriate, appressed, reddish, adaxially non-glandular, glabrous; laminar margins sparsely ciliate, non-glandular, the hairs 0.3 – 0.4 mm long, 1 – 3-celled, spreading to ascending; indusia absent. Figs 8A – C and 9M – T; Map 2.

Distribution. Cuba, Haiti, Guadeloupe, Costa Rica, Panama, Colombia, Ecuador, Peru, Bolivia.

Additional Specimens Examined. Costa Rica. Alajuela: Cantón de San Ramón, Reserva Biológica Monteverde, Estación La Casona, sendero Orquídeas, 10°20′00"N, 84°44′W, 1000 m, 3 May 1995, Martínez 433 (INB); Cartago: Tapantí Reserve, 9°43′N, 83°47′W, 1400 – 1700 m, 7 Dec. 1982, Gómez 19290 (MO); Tapantí Reserve, 9°43′N, 83°47′W, 1400 – 1700 m, 7 Dec. 1982, Gómez 19291 (MO); N and S slopes of ridge on E side of Río Grande de Orosí, opposite mouth of Río Humo, c. 6 km upstream from Tapantí, 9°43′N, 83°47′W, 1500 – 1800 m, 24 Nov. 1984, Grayum et al. 4538 (MO); along trail leading E into mountains from road into Tapantí Reserve, c. 1 km S of jct. of Quebrada Salto and Río Grande de Orosí, 9°43′N, 83°47′W, 1500 – 1800 m, 1 Feb. 1986, Smith et al. 2130 (CR, MO, NY); along road above Río Grande de Orosí, 12 to 16 km S of Tapantí, 9°42′N, 83°46′W, 1500 – 1600 m, 25 March 1973, Stolze 1495 (US); Heredia: Porrosatí, [10°06′N, 84°06′W], 1900 m, April 1973, Gómez 3553 (CR); Porrosatí, [10°06′N, 84°06′W], 1900 m, April 1973, Gómez 3564 (CR); NW slope of Volcán Barva, between Laguna de Barva and base of Cerros La Marías, 10°08′N, 84°07′W, 2450 – 2800 m, 28 April 1986, Grayum 7467 (AAU, CR, MO); Headwaters of Río Santo Domingo, c. 3 km E of San Rafael de Vara Blanca, N slope of Volcán Barva, 10°11′N, 84°07′W, 2060 – 2100 m, 16 April 1986, Grayum 7178 (CR, MO); vic. of water-filled crater NE of summit of Volcán Barva and SE of Laguna Danta, 10°08′N, 84°06′W, 2500 – 2600 m, 26 April 1986, Grayum et al. 7405 (MO); Braulio Carillo National Park, trail between OTS Station La Selva and Volcán Barva, 2500 m, 30 April 2003, Kluge 6357 (GOET); Volcán Barva, [10°08′N, 84°06′W], 2800 m, 29 July 1926, Valerio 28 (US); Limón: Cantón de Talamanca, along Quebrada Kuisa (tributary of Río Lori) near crossing of Ujarrás-San José Cabécar trail, Cordillera de Talamanca, 9°20′30″N, 83°14′00″W, 2170 m, 16 March 1993, Grayum 10315 (CR, INB, MO); Puntarenas: Cantón de Coto Brus, Zona Protectora Las Tablas, Sabalito, Las Alturas de Cotó, Est. Biol. Las Alturas, senderoa Cerro Echandi, postes 40 – 57, 8°59′05″N, 82°43′10″W, 2070 – 2230 m, 28 Dec. 1993, Rojas 842 (CR, INB); Cantón de Coto Brus, Zona Protectora Las Tablas, Cuenca Terraba-Sierpe, sitio Coto Brus, orillas de la Quebrada Surú, 8°58′30″N, 82°46′15″W, 1960 m, 12 Feb. 1999, Rojas et al. 4910 (CR, INB, NY); San José: Zurquí, [10°03′N, 84°01′W], 1900 m, 4 April 1976, Gómez 3525 (CR). Panama. Bocas del Toro: E branch, headwaters of Colubre R., [9°21′30″N, 83°40′00″W], 2300 – 2500 m, 2 March 1984, Gómez et al. 22324 (CR, MO); above Boquete, Holcomb’s Trail, between Camp I and the Divide, [9°10′N, 82°16′W], 1675 – 1800 m, 19 Feb. 1918, Killip 5336 (US).

Habitat. Wet forests, shady ravines; 1400 – 2800 m (in Antilles 900 – 2100 m, Moran et al.2009b).

Conservation Status. Least Concern (LC).

Notes.Megalastrum macrotheca is characterised by laminae 2-pinnate-pinnatisect at the base and pinna rachises abaxially pubescent (hairs 0.4 – 0.6 mm long) and conspicuously scaly. It resembles M. reductum (see below).

The Central American plants are generally smaller and less divided than those in the West Indies. Often a single leaf will fit on a herbarium sheet, even if not folded. Larger leaves have distinctly inequilateral basal pinnae, whereas those of smaller leaves may sometimes be nearly equilateral (Fig. 8A – C).

We select the lectotype above of Megalastrum biseriale because this sheet was annotated by Christensen (1920) as “type,” whereas he did not annotate the second sheet as such.

15. Megalastrum mexicanumR. C. Moran & J. Pradosp. nov. A Megalastro atrogriseo squamis petioli angustioribus et lamina utrinque pilis sparsis 1.0 – 1.5 mm longis vestita differt; etiam praesentia in altitudinibus inferioribus differt. Typus: Mexico. Oaxaca, Mun. Sta. María Cimalapa, c. 13 km S de Sta. María por la vered al Río Negro que sale a media bajada al cañon de la vereda al Paso Napajo-ua, 16°50′N, 94°40′W, 400 m, 30 Aug. 1987, H. Hernández G. 1414 (holotypus NY 2 sheets!; isotypus CHAPA).

http://www.ipni.org/urn:lsid:ipni.org:names:77105947-1

Rhizomes erect to decumbent; leaves up to 1.3 m long; scales of the petiole bases 15 – 20 × c. 0.3 – 0.4 mm, linear-filiform, ascending, twisted toward the apex, light brown, often shiny, the margins sparsely denticulate on the margins, the surfaces smooth; laminae up to 0.4 – 0.88 m long, 3-pinnate-pinnatisect at base, 2-pinnate-pinnatisect medially; basal pinnae 0.15 – 0.2 m long, strongly inequilateral (elongated basiscopically); pinna rachises abaxially non-glandular, sparsely scaly, sparsely pilose, the scales to 1 – 2 × 0.1 – 0.2 mm long, filiform, tortuous, spreading, brown, shiny, sparsely denticulate, the hairs on the abaxial surfaces 2.5 – 3.0 mm long, 6 – 10-celled, spreading, the hairs on the adaxial surfaces c. 2.0 – 2.5 mm long, 6 – 8-celled, dense; costules on both surfaces with indument like that of the pinna rachises; laminar tissue between the veins abaxially non-glandular (both gland-tipped hairs and sessile globose ones absent), sparsely pilose, the hairs 0.5 – 0.8 mm long, 2 – 5-celled, erect, acicular, proscales sparse, c. 0.2 mm long, appressed, adaxially glabrous, sessile or stalked glands absent; veins evident on both surfaces, sparsely pilose abaxially with hairs like those of the laminar tissue, sparsely pilose adaxially, typically only 2 – 4 hairs per vein, the hairs 0.5 – 0.8 mm long, 2 – 5-celled; hydathodes evident; lamina margins pilose, the hairs 0.4 – 0.8 mm long, 3 – 5-celled, acicular, spreading, non-glandular; indusia absent. Figs 3K – N and 4C; Map. 2.

Distribution. Mexico.

Additional Specimens Examined. Mexico. Chiapas: Cerro del Roblar, near Ixtacomitan, [17°25′N, 93°06′W], 360 m, 12 July 1890, Rovirosa 817 (NY); Hidalgo: below Chapulhuacan, [21°10′N, 98°54′W], 0 m, 27 Nov. 1937, Kenoyer s.n. (MO); Oaxaca: Sta. María de Chimalapa, Cabecera del Arroyo Milagrito, c. 13 km SE de Sta. María por la vereda al Río Negro, 16°51′30″N, 94°39′0″W, 400 m, 26 Sept. 1985, Hernández 1616 (NY); Sta. María de Chimalapa, Paso Piedra de Tigre (Cautza), c. 8 km W de Sta. María, 16°53′N, 94°43′W, 220 m, 26 Sept. 1985, Hernández 1962 (MO, NY); Sta. María de Chimalapa, Cabecera del Río Escolapa, c. 15 km S de Sta. María, 16°50′N, 94°41′W, 400 m, 21 Oct. 1985, Hernández & González 1704 (NY); Veracruz: San Andrés Tuxtla, Estación Biol. Tropical Los Tuxtlas, 18°34′N, 95°04′W, 250 m, 8 Jan. 1987, Sinaca C. 1154 (MO).

Habitat. Not known; 220 – 400 m.

Conservation Status. Least Concern (LC).

Notes. Specimens of Megalastrum mexicanum have previously been identified as M. atrogriseum (e.g., Mickel & Smith 2004). M. mexicanum differs from that species by sparser hairs on and between the veins on both surfaces of the laminae and narrower petiole scales (0.2 – 0.5 mm vs. 0.5 – 1.5 mm wide). The two species also differ in geographic distribution and elevation: M. mexicanum is endemic to Mexico (thus the specific epithet) and occurs from 220 – 400 m, whereas M. atrogriseum is endemic to Costa Rica and Panama and occurs from (600 –) 1000 – 2000 m.

Megalastrum galeotti, which is common in Central America, differs by pinna rachises glabrous abaxially, lamina tissue between the veins abaxially glabrous or (more commonly) puberulent by minute (0.1 – 0.2 mm long) hairs, and laminar margins with hairs 0.1 – 0.2 mm long.

16. Megalastrum palmense (Rosenst.) A. R. Sm. & R. C. Moran (Smith & Moran 1987: 128).

Dryopteris subincisa var. palmensis Rosenst. (Rosenstock 1925: 11); Ctenitis palmensis (Rosenst.) Lellinger (1977: 709). Type: Costa Rica, San José, La Palma, [10°03′N 83°59′W], 1400 m, 20 March 1909, A. Brade & C. Brade 215 (lectotype UC!, selected by Smith & Moran (1995); duplicates BM!, NY!, GH!).

Rhizomes erect, the scales 5 – 10 mm long, entire, golden, flaccid; leaves up to 1.5 m long; scales of the petiole base 1 – 7 × 0.2 – 1 mm, generally inconspicuous, linear to lanceolate, flaccid, tightly appressed to the petiole, brown, dull, entire to denticulate or erose on the margins; laminae up to 1.5 m long, 3-pinnate-pinnatifid at base, 2-pinnate-pinnatifid medially; basal pinnae c. 0.4 m long, strongly inequilateral (elongated basiscopically); pinna rachises abaxially non-glandular (lacking both stipate and sessile glands), densely puberulent, sparsely scaly, the scales to 6 mm long, ovate to oblong, flaccid, appressed, dull brown, entire to erose, the hairs c. 0.1 mm long, 1 or 2 (– 4)-celled, spreading; adaxially densely and evenly puberulent, the hairs 0.1 mm long, 1- or 2-celled, spreading to antrorsely strigose; costules abaxially sparsely pubescent and scaly, the hairs and scales like those on the rachis, adaxially pubescent, the hairs 0.1 mm long, 1 – 3-celled, spreading to antrorsely strigose; laminar tissue between the veins on both surfaces non-glandular, glabrous, proscales present, sparse; veins obscure on both surfaces, non-glandular, glabrous; hydathodes evident; lamina margins ciliate, the hairs 0.1 – 0.2 mm long, 1 – 3-celled, non-glandular; indusia absent. Figs 13H – K and 16A – C; Map 3.

Distribution. Costa Rica, Panama.

Additional Specimens Examined. Costa Rica. Alajuela: Cantón de San Ramón, Reserva Forestal San Ramón, Cordillera de Tilarán, Estación Río San Lorenzo, 10°13′00″N, 84°35′20″W, 800 – 1000 m, 22 April 1994, Fuentes 760 (CR, INB, MO); Angel Falls, 5 km N of Varablanca, on rd to Puerto Viejo, 10°11′N, 84°10′W, Aug. 1967, Mickel 3574 (NY, US); Cartago: c. 10 km S of Tapaní, 9°42′N, 83°47′W, 1400 – 1600 m, 10 – 24 June 1968, Burger & Stolze 5693 (F, NY, US); Coliblanco, 9°57′N, 83°48′W, 1950 m, 30 April – 2 May 1906, Maxon 303 (NY, US); Cantón de Paraíso, Cuenca del Reventazón, entrando por la Hacienda Queveri, antes de llegar a Tres Picos, 9°44′20″N, 83°50′30″W, 2000 – 2200 m, 3 April 1997, Rojas et al. 3404 (CR, INB); along trail leading E into mts from rd into Tapantí Reserve, c. 1 km S of jct. of Quebrada Salto and Río Grande de Orosí, ″Quebrada Val Verde″, 9°43′N, 83°47′W, 1500 – 1800 m, 1 Feb. 1986, Smith et al. 2129 (CR, MO, NY); near La Sierra, c. 25 km N of Cartago, Cordillera de Talamanca, 9°45′N, 83°54′W, 2000 m, 23 Jan. 1965, Williams et al. 28017 (F, NY); near La Sierra, c. 25 km N of Cartago, Cordillera de Talamanca, 9°45′N, 83°54′W, 2000 m, 23 Jan. 1965, Williams et al. 28077 (F, NY); Heredia: banks of Sardinal R., 10°16′N, 84°03′W, 1800 m, 1 May 1969, Gómez 2193 (F, GH, NY); Braulio Carillo National Park, 10°15′N, 84°10′W, 1865 m, 9 Nov. 1986, Hennipman et al. 6813 (U); Cinchona, above the Upper Sarapaquí Valley, [10°13′N, 84°10′W], 1100 m, 20 March 1955, Scamman s.n. (GH); Yerba Buena, NE of San Isidro, [10°03′N, 84°02′W], 2000 m, 22, 28 Feb. 1926, Standley & Valerio 49940 (US); Limón: Cantón de Talamanca, along Quebrada Kuisa (tributary of the Río Lori) near crossing of Ujarrás-San José Cabécar trail, 9°20′30″N, 83°14′00″W, 2170 m, 16 March 1993, Grayum 10316 (INB, MO); Cantón de Talamanca, Parque Nacional Talamanca, 500 m abajo unión Quebrada Kirigú en Río Coén, entre Ujarrás y San José Cabécar, 9°23′30″N, 83°12′45″W, 1650 m, 31 March 1993, Herrera 6116 (INB, MO); Puntarenas: Cantón de Coto Brus, Parque Internacional La Amistad, Cordillera de Talamanca, Estación Pittier, sendero a Río Gemelo, 9°01′30″N, 82°57′40″W, 1650 m, 30 Jan. 1995, Fletes 27 (INB, MO); Cantón de Coto Brus, Parque Nacional La Amistad, Cuenca Térraba-Sierpe, Estación Pittier, Cerro Gemelo, 9°02′53″N, 82°55′55″W, 2600 – 2700 m, 23 May 1996, Moraga & Rojas 486 (INB, MO); Cantón de Coto Brus, Zona Protectora Las Tablas, Cuenca Terraba-Sierpe, sitio Coto Brus, 8°58′30″N, 82°46′15″W, 1960 m, 18 Feb. 1998, Navarro V. 4334 (INB); Cantón de Coto Brus, Zona Protectora Las Tablas, Las Alturas de Cotó, Sabalito, Estación Biológica Las Alturas, sendero a Cerro Echandi, postes 40 – 57, 8°59′05″N, 82°43′10″W, 2070 – 2230 m, 28 Dec. 1993, Rojas 840 (INB); Cantón de Coto Brus, Zona Protectora Las Tablas, Sabalito, Las Alturas de Cotó, Est. Biol. Las Alturas, senderoa Cerro Echandi, postes 40 – 57, 8°59′05″N, 82°43′10″W, 1500 – 1600 m, 3 Feb. 1999, Rojas 4879 (INB); San José: valley of the Río Hondura, below La Palma, NE of San Jeronimo, 10°03′N, 83°58′W, 1000 m, 15 May 1968, Burger & Stolze 4871 (CR, F); Dota, Copey, Zapotal de Providencia, Reserva Montaña Fria, 1 – 2 km hacia Providencia, 9°32′05″N, 83°50′19″W, 1720 m, 26 Aug. 2004, Gómez 6006 (CR); c. 15 km N of Tres Ríos, c. 4 km N of Cascajal, near where road fords a stream, [10°00′N, 83°58′W], 1600 m, 2 Aug. 1970, Lellinger 1379 (BM, CR, F, GH, US); vic. of La Palma on the road to La Hondura, [10°04′N, 83°59′W], 1500 – 1700 m, 17 – 18 July 1923, Maxon & Harvey 8062 (US); vic. of La Hondura, 4 km S to 1 km N on Rte 220, between Volcán Irazu and Volcán Barba, [10°04′N, 83°59′W], 1250 – 1500 m, 25 March 1967, Mickel 2244 (NY); vic. of La Hondura, 4 km S to 1 km N on Rte 220, between Volcán Irazu and Volcán Barba, [10°04′N, 83°59′W], 1250 – 1500 m, 25 March 1967, Mickel 2247 (NY); Cantón de Dota, Savegre Lodge, [9°33′04″N, 83°48′29″W], 2200 m, 24 June 2007, Moran & Triana-Moreno 8015 (CR, INB, NY, USJ); Cantón de Moravia, Parque Nacional Braulio Carrillo, 10°03′30″N, 84°01′00″W, 1650 m, 12 June 1997, Rojas & Coto 3571 (INB, MO); La Hondura, [10°04′N, 83°59′W], 1300 – 1700 m, 16 March 1924, Standley 37756 (US); La Palma, c. 9 km NE of San Jerónimo, 10°02′N, 84°00′W, 1500 m, 23 March 1973, Stolze 1433 (F, US); Old road to La Hondura, [10°04′N, 83°59′W], 1491 m, 5 Feb. 2008, Sundue et al. 1783 (CR, INB, NY, USJ). Panama. Chiriquí: Monte Azul, 1.4 miles N of Entre Ríos on E slopes of Cerro Punta, 3 miles by rd from town of Cerro Punta, 8°52′N, 82°33′W, 2250 m, 22 Nov. 1979, Antonio 2722 (MO); Guadalupe, Cerro Punta, Finca Maduro, 8°52′N, 82°33′W, 2000 m, 23 April 1982, Caballero 143 (MO); Guadalupe, Cerro Punta, Finca Maduro, 8°52′N, 82°33′W, 2000 m, 23 April 1982, Caballero 136 (MO); vic. of el Boquete, [9°10′N, 82°16′W], 1000 – 1501 m, 18 Feb. 1918, Cornman 998 (S, US); vic. of Boquete, Cerro Pate de Macho, SW slope, 8°46′N, 82°25′W, 1800 – 1950 m, 19 June 1987, Croat 66424 (MO); vic. of Las Nubes, 2.7 miles NW of Río Chiriquí Viejo, W of Cerro Punta, 8°52′60″N, 82°35′60″W, 2200 m, 27 Feb. 1973, Croat 22407 (MO, US); vic. of Boquete, Cerro Pate de Macho, SW slope, 8°46′N, 82°25′W, 1800 – 1950 m, 19 June 1987, Croat 66425 (MO); Boquete Distr., Bajo Chorro, 8°46′N, 82°26′W, 1800 m, 15 Feb. 1938, Davidson 293 (GH, US); Parque Internacional La Amistad, Las Nubes – Cerro Punta, [8°52′N, 82°34′W], 9 April 1995, González 155 (K); valley of the Río Caldera, from El Boquete to the Cordillera, [8°46′N, 82°25′W], 1400 – 1600 m, 5 – 19 Feb. 1918, Killip 5061 (US); valley of Río Quebrada, above El Boquete, [8°46′N, 82°25′W], 1650 m, 8 Feb. 1918, Killip 5501 (US); valley of Río Caldera, from El Boquete to the Cordillera, [8°46′N, 82°25′W], 1400 – 1600 m, 6 Feb. 1918, Killip 5053 (US); along trail between north fork of Río Palo Alto and Cerro Pate Macho, c. 6 km NE of Boquete, 8°48′N, 82°23.5′W, 1600 – 2000 m, 6 Feb. 1986, Smith et al. 2316 (MO); Distr. Bugaba, Cerro Punta, around STRI house, 8°52′N, 82°33′W, 2200 m, 27 Jan. 1984, van der Werff & Herrera 6514 (MO); Distrito de Bugaba, Santa Clara, Cerro Pando, 8°50′N, 82°44′W, [no date], van der Werff & Herrera 7240 (MO); Unknown: Río Caldera, [8°46′N, 82°25′W], 1650 m, 17 – 19 Feb. 1918, Killip 5124 (US).

Habitat. Wet forests; 1100 – 2700 m.

Conservation Status. Least Concern (LC).

Notes.Megalastrum palmense can be distinguished from nearly all its congeners by the short (c. 0.1 mm long), often antrorsely strigose hairs on both surfaces of the pinna rachises and costules. Having hairs of the same length on both surfaces of the pinna rachises is unusual; nearly all other species of Megalastrum have longer hairs on the adaxial surfaces. The only species of Megalastrum in Central America with similar hairs on the pinna rachises and costae is M. heydei, endemic to Guatemala. M. heydei belongs to the M. pulverulentum group, which differs from M. palmense (and other species of Megalastrum) by large leaves (3 – 4 m long), more highly divided laminae (up to 4-pinnate-pinnatisect), and strongly and conspicuously dentate scales on the pinna rachises and costules.

Also distinctive to Megalastrum palmense are the scales on the petioles, rachises, and costules. These scales are lax, flaccid, dull brown, and entire. Upon drying, they often conform closely to the surface. Often the petiole scales appear to be absent, but in fact they are present and tightly appressed to the petiole base. No other species of Megalastrum has similar scales.

Megalastrum palmense also lacks hairs and glands (both stalked and spherical) on both surfaces of the laminae. These characters serve to distinguish it from many congeners in Costa Rica and Panama, such as M. apicale, M. galeottii, and M. lunense.

An unusual specimen that will key to Megalastrum palmense is Moran & Triana-Moreno 8015, from Costa Rica. It differs from typical M. palmense by its petiole scales that are spreading, linear, firm, and denticulate. Its lamina pubescence resembles that of M. palmense except that the costules abaxially are sparsely and minutely acicular pubescent (vs. glabrous) and the hairs on the adaxial surface of the rachis and costae are longer than the hairs on the abaxial surfaces. The scales on the abaxial surface of the rachis and costae resemble those of M. palmense, but some are longer, denticulate, and ascending-spreading, not appressed. This specimen might represent a hybrid involving M. palmense, but we are not sure what species the other parent is.

The protologue of Dryopteris palmensis lists Brade 215a as one of the syntypes. This specimen is not at S or UC where Rosenstock types are often found. We have examined three specimens collected on the same date, locality, and elevation that are Brade 215, and consider these specimens as syntypes. Both are annotated in Rosenstock’s hand as “palmensis var. nov.” The other syntype cited (Costa Rica, La Palma, 1450 m, 17 March 1910, A. Brade & C. Brade s.n.) was not found among the specimens requested on loan. For these reasons, we select Brade 215 (NY) as the lectotype.

17. Megalastrum pulverulentum (Poir.) A. R. Sm. & R. C. Moran (Smith & Moran 1987: 129).

Polypodium pulverulentum Poir. (Poiret in Lamarck 1804: 555); Aspidium lutescens Willd. (Willdenow 1810: 272), nom. superfl.; Dryopteris pulverulenta (Poir.) C. Chr. (in Urban 1925: 305); Ctenitis pulverulenta (Poir.) Copel. (Copeland 1947: 124). Type: Plumier, Traité Foug. Amér. 27, pl. 34!, 1707, lectotype, selected by Proctor (1985), illustrating a plant from Hispaniola.

Polypodium barbatum Desv. (Desvaux 1827: 242). Type: Hispaniola, without exact locality, s.d., A. N. Desvaux s.n. (holotype P! photo US! ex P).

Polypodium karstenianum Klotzsch (1847: 390); Phegopteris karsteniana (Klotzsch) Mett. (Mettenius 1858: 314); Nephrodium villosum var. karstenianum (Klotzsch) Jenman (1896: 114); Dryopteris subincisa var. karsteniana (Klotzsch) C. Chr. (Christensen 1905: 295); Dryopteris karsteniana (Klotzsch) Hieron. (Hieronymus 1907: 348); Ctenitis karsteniana (Poir.) Vareschi (1969: 405); Megalastrum karstenianum (Klotzsch) A. Rojas (2001: 483). Type: Venezuela, H. Karsten II, 3 (lectotype B!, selected here; duplicates H photos B!, US! ex H).

Phegopteris hirsuta Fée (1852: 248). Type: Venezuela [“Mexico,” in error], Mérida, 2000 m, July 1846, H. Funck & L. J. Schlim 975 (lectotype P-00600642!, selected here; duplicates BM!, P!).

Aspidium erythrostemma H. Christ (1904: 961). Dryopteris erythrostemma (H. Christ) C. Chr. (Christensen 1905: 263). Type: Costa Rica, 1903, C. Wercklé s.n. (lectotype P-00600398!, selected here).

Leaves up to 4 m long; scales of the petiole bases 2 – 4 × c. 0.05 cm, linear-lanceolate, spreading, light brown to golden denticulate; laminae up to 3 m long, 4-pinnate-pinnatisect at base, 3-pinnate-pinnatisect medially; basal pinnae c. 1.4 m long, strongly inequilateral (elongated basiscopically); pinnules short-stalked, 35 – 70 cm long; pinna rachises glandular, pubescent, scaly, the glands c. 0.1 mm long, spherical, sessile, yellowish, the scales 2 – 3 mm long, linear-lanceolate, dark brown, denticulate to ciliate (the cilia often darker than the body of the scale, especially towards the apex), non-bullate, the hairs 1.0 – 1.5 mm long, 5 – 8-celled, rachises adaxially glandular, densely pubescent, the hairs and glands like those abaxially; costules on the abaxial surface glandular, pubescent and scaly, the glands both sessile and stalked, c. 0.1 mm long, spherical, sessile, yellowish, the hairs 1 – 2 mm long, 5 – 8-celled, spreading, the scales like those on the pinna rachises abaxially but smaller, c. 1 mm long, the adaxial surface pubescent, the hairs 0.5 – 1.5 mm long, 3 – 6-celled, spreading; laminar tissue between veins pubescent and glandular on both surfaces, the hairs 1 – 2 mm long, 5 – 8-celled, the glands both sessile and stalked, rounded, yellowish; veins visible on both surfaces, abaxially glandular, sparsely pubescent, non-scaly, the hairs c. 0.5 – 1.0 mm long, 3- or 4-celled, the scales c. 0.3 mm long, uniseriate, appressed, reddish, adaxially non-glandular, sparsely pubescent, the hairs 0.4 – 1.0 mm long, 4 – 6-celled, tortuous; laminar margins ciliate and glandular, the hairs 0.4 – 1.0 mm long, 3- or 4-celled, spreading, the glands few and stalked; indusia absent. Figs 11C and 12P – W; Map 3.

Distribution. Mexico, Guatemala, Honduras (Stolze 1981), Costa Rica, Panama, Venezuela, Colombia, Jamaica, Haiti, Dominican Republic, Guadeloupe, Martinique, St. Vincent, Dominica, Saba.

Additional Specimens Examined. Mexico. Oaxaca: Distr. Teotitlan, Mun. San José de Tenango, San Martín Caballero, 18°06′63″N, 96°39′61″W, 1600 m, 3 Jan. 1995, Ingela 95-M141 (NY). Guatemala. Alta Verapaz: along road between Tactic and the divide on road to Tamahú, [15°19′60″N, 90°17′60″W], 1500 – 1600 m, 1 – 7 April 1941, Standley 91509 (F, U, US). Costa Rica. Cartago: El Muñeco, the valley S of Navarro Valley, near boundary of San José and Cartago provinces, [9°48′N, 83°55′W], 7 April 1928, Stork 2849 (MICH, NY, US); Heredia: near Porrosati on the S slope of Volcán Barba, [10°07′N, 84°07′W], 2200 m, 22 June 1968, Burger & Stolze 6005 (CR, F, GH, US); Escuela de Porrosati, toward Volcán Barba, [10°06′N, 84°07′W], 2000 m, 17 Aug. 1940, Chrysler & Roever 5511 (US); below the crest of Barba Volcano, [10°08′84°06′W], 2000 m, 23 Jan. 1964, Lems 640129 (NY); above San José de la Montaña on S slope of Volcán Barba, 1900 m, 10 Oct. 1964, Nisman 31 (CR, GH); above San José de la Montaña on S slope of Volcán Barba, 1900 m, 14 Oct. 1964, Nisman 47 (CR, GH); Puntarenas: Cantón de Coto Brus, Zona Protectora Las Tablas, Cuenca Térraba-Sierpe, sitio Coto Brus, 1960 m, 12 Feb. 1999, Jiménez et al. 47 (INB); Cantón de Coto Brus, Zona Protectora Las Tablas, Cuenca Térraba-Sierpe, sitio Coto Brus, orillas de la Quebrada Suru, 1960 m, 12 Feb. 1999, Rojas et al. 4911 (INB); Las Alturas de Coton, c. 20 miles N of San Vito de Java, [8°47′N, 82°57′W], 1700 m, Aug. 1975, Unknown s.n. (F, GH); San José: Cuenca del Pirris-Damas, Cerros de Caraigres, Quebrada Concha, camino viejo a Bijagual, 9°42′18″N, 84°07′51″W, 1500 m, 5 Dec. 1996, Hammel et al. 20559 (CR, INB, MO); Finca Río Blanco, along rd 3 km E of Cope de Dota, [9°38′N 83°55′W], 2100 m, 17 June 1975, Lellinger et al. 1787 (US); Cantón de Aserrí, Zona Protectora Cerros de Escazú, Cuenca del Río Grande de Tárcoles, El Cedral, Alto Hierbabuena, 9°50′30″N, 84°06′35″W, 2150 m, 6 Nov. 1993, Morales 1958 (CR, INB, MO); Cantón de Dota, Hotel Savegre, 9°33′45″N, 83°48′29″W, 2200 m, 23 June 2007, Moran & Hernández 8029 (CR, INB, NY); Acosta, Palmichal, Zona Protectora Cerros de Escazú, 9°50′55″N, 84°19′50″W, 1600 m, 24 June 2004, Rojas 5809 (CR); vic. of Sta. María de Dota, [9°39′N, 83°58′W], 1500 – 1800 m, 14 – 26 Dec. 1925, Standley 42113 (US); Unknown: San Jeronimo, 1500 m, April 1910, Wercklé 577 (CR). Panama. Chiriquí: Bajo Grande-Cerro Punta, Finca de Toño Ríos, [8°52′60"N, 82°34′60 W], 2000 m, 7 Sept. 1982, Caballero 191 (MO); El Boquete, trail to Denham’s, [9°10′N, 82°16′W], 10 April 1918, Cornman 1282 (MICH, MO); 2 miles N of El Hato del Volcán, [8°46′N, 82°37′W], 30 May 1970, Croat 10656 (MO).

Habitat. Wet forests, shady ravines; 1500 – 2200 m (in West Indies from 0 – 1600 m; Moran et al.2009b).

Conservation Status. Least Concern (LC).

Notes.Megalastrum pulverulentum is densely pubescent on both surfaces by long (1 – 2 mm) whitish hairs (Fig. 12P – W). The scales on the pinna rachises abaxially are more prominently toothed than any other species in Central America. The teeth are usually longest and darkest toward the apex of the scale. Yellowish glands are present on both surfaces and may be either sessile or stalked (Fig. 12R, U, W). Although difficult to describe, the laminar cutting (Fig. 11C) is distinctive compared to other decompound species in the genus.

Megalastrum pulverulentum is one of the largest species in the genus. Its leaves reach 4 m long, with basal pinna up to 1.4 m long. The two collections from Mexico and Guatemala, however, have leaves about 1 m long but are otherwise typical.

In the past, the name Megalastrum pulverulentum was widely applied to plants from Mexico to northern Argentina. Our unpublished studies suggest that this species in the wide sense is an aggregate of several different ones. In Central America, we distinguish two species, M. heydei and M. sparsipilosa, that were previously identified as M. pulverulentum. In a previous paper (Moran et al. 2009b), we included Ecuador, Peru, Bolivia, and northern Argentina in the range of this species; however, specimens previously identified as M. pulverulentum from these countries represent several new species that we will describe in a treatment of the genus for the Andes.

Although Christ cited Wercklé 118 and 136 in the protologue of Aspidium erythrostemma, the only specimen at P labelled in his hand as “A. erythrostemma, sp. nov.” does not have a number. We choose this specimen as the lectotype.

Monterrosa & Monro (2008) reported this species from El Salvador based on Monterrosa & Martínez 944 (LAGU) and Monterrosa et al. 961 (LAGU, NY). These specimens, however, represent M. galeottii.

18. Megalastrum reductumA. Rojas (2008: 45, f. 6A – C). Type: Ecuador, Guayas/Cañar/Chimborazo/Bolívar, foothills of the W cordillera near the village of Bucay, [2˚40′S, 79˚40′W], 305 – 365 m, 8 – 15 June 1945, W. H. Camp 3786 (holotype US; isotypes F!, K!, NY!, P!).

Megalastrum dorsiglabrum A. Rojas (2008: 45, f. 6A – C). [originally published as “dorsoglabrum”]. Type: Panama, Coclé, El Valle de Antón, [8°37′N, 80°07′W], 1000 m, 4 June 1934, A. H. G. Alston 8730 (holotype CR!; isotype US).

Rhizomes up to 10 cm long, erect, the scales 10 – 20 mm long, linear, firm, densely and finely setulose on margins and surfaces, golden brown; leaves up to 1.2 m long; scales of the petiole base 10 – 20 × 1 – 2 mm, linear, firm, spreading, denticulate, setulose on surfaces, brown, usually shiny; laminae up to 0.8 m long, 2-pinnate-pinnatisect at base and medially; rachises of the laminae pubescent, the hairs 0.1 – 0.2 mm long, 1- or 2-celled, spreading or appressed-ascending; basal pinnae c. 20 cm long, equilateral or nearly so, the segments adnate; pinna rachises abaxially non-glandular, densely puberulent, sparsely scaly, the hairs 0.1 – 0.2 mm long, 1- or 2-celled, spreading or appressed-ascending to strigose, acicular, the scales 3 – 4 × c. 0.1 mm, filiform, loosely spreading, often tortuous, dull brown, entire, adaxially glabrous or pubescent, sparsely scaly or lacking scales, hairs like those abaxially, the scales c. 1 × 0.1 mm, filiform, entire, appressed, dull, brown; costules abaxially eglandular, glabrous to sparsely scaly and puberulent, the scales filiform, dull brown, entire, hairs c. 0.1 mm long, 1 or 2-celled, acicular, ascending to strigose, adaxially glabrous or subglabrous to sparsely pubescent, the hairs like those on the pinnae rachises adaxially; laminar tissue between the veins abaxially non-glandular, glabrous to puberulent, the hairs c. 0.1 mm, 1- or 2-celled, acicular, erect, adaxially non-glandular, glabrous; veins abaxially visible, sparsely puberulent with hairs like those on the costules, adaxially obscure, glabrous to sparsely pubescent, the hairs like those on the pinna rachises; hydathodes evident; lamina margins ciliate, non-glandular, the hairs 0.1 – 0.2 mm long, 1- or 2-celled, erect, acicular, ascending to substrigose; indusia absent. Figs 8E and 9H – L; Map 3.

Distribution. Eastern Panama, western Colombia, western Ecuador, eastern slope of the Andes in central Peru.

Additional Specimens Examined. Panama. Darién: along ridge trail from Cana up the Cerro Pirre Massif, 7°50′N, 77°40′W, 1000 – 1300 m, 3 May 1990, Moran 5060 (AAU, F, MO, NY, US).

Habitat. Wet forests; 1000 – 1300 m in Panama (100 – 800 m in Colombia and Ecuador, 1000 – 1700 in Peru).

Conservation Status. Least Concern (LC).

Notes.Megalastrum reductum is characterised by basal pinnae equilateral or nearly so, the first two proximal pinnules often slightly reduced (thus the name of the specific epithet), and the pinna rachises abaxially evenly puberulent (hairs 0.1 – 0.2 mm long, 1- or 2-celled) and with sparse filiform scales. Acicular hairs c. 0.1 mm long occur between the veins abaxially, especially near the costules. The rhizome scales appear puberulent on both surfaces, this apparently from the upturned and projecting ends of the cells. Some South American specimens have longer and more highly cut leaves than the Panamanian plants (pers. obs.), being 2-pinnate-pinnatisect medially with free pinnules.

Adaxially, Megalastrum reductum varies in the amount of pubescence along the pinna rachises. Some specimens, which have been named M. dorsiglabrum A. Rojas (Rojas 2008), are glabrous adaxially, whereas others are pubescent (e.g., the type of M. reductum). Some specimens are intermediate, with a few hairs at the base of the pinna rachises adaxially (e.g., Moran 5060). No other difference appears to correlate with the amount of pubescence adaxially; therefore, we recognise a single species.

This species occurs on both sides of the Andes, but the populations are slightly different. Those from Panama and the western side of the Andes in Colombia and Ecuador show what is termed (Moran 1995, 1996) a Mesoamerican-Chocó distribution. The type is from this region. Specimens on the eastern side of the Andes in central Peru differ from those of Ecuador, Colombia, and Panama by pinna rachises subglabrous abaxially (very few minute acicular hairs present) and rhizome scales with scattered blackish streaks caused by indurate cells. We feel this is best interpreted as part of the variation in a single species.

19. Megalastrum sparsipilosumR. C. Moran & J. Pradosp. nov.Megalastro pulverulento quoad folia usque ad 4 m longa et 4-pinnato-pinnatisecta simile, sed ab eo pinnarum rhachidibus costulis et venis subglabris vel abaxialiter sparsim pubescentibus glandulsisque differt. Typus: Mexico, Chiapas: Mun. Yajalón, steep slopes of Ahk’ubal Nab above Yajalón, 17°13′N, 92°19′W, 1200 m, 6 April 1973, D. E. Breedlove 34621 (holotypus NY 2 sheets!; isotypi F!, MO!).

http://www.ipni.org/urn:lsid:ipni.org:names:77105948-1

Rhizomes to 0.4 m tall; leaves up to 4.6 m long; scales of the petiole base 20 – 50 × 2 – 3 mm, linear-lanceolate, spreading, golden to yellowish brown, denticulate throughout; laminae up to 3 m long, 4-pinnate-pinnatisect at base, medially 3-pinnate-pinnatisect; basal pinnae c. 1 m long, strongly inequilateral (elongated basiscopically); pinna rachises abaxially sparsely glandular, sparsely scaly, glabrous to sparsely pilose, the glands c. 0.1 mm wide, spherical, brownish, the scales to 2 – 3 × 0.2 – 0.5 mm, linear to lanceolate, appressed to slightly spreading, strongly denticulate, the hairs 1.0 – 1.5 mm long, c. 8-celled, adaxially glandular, densely pilose, the glands c. 0.1 mm wide, spherical, sessile, brownish, the hairs 1 – 2 mm long, 7 – 9-celled, spreading, light brown, the scales like those abaxially; costules abaxially sparsely glandular, sparsely pilose to nearly glabrous, sparsely scaly, the hairs 1.5 – 2.0 mm long, 8 – 12-celled, erect, the scales 0.8 – 1.0 mm long, strongly denticulate, appressed to slightly spreading, reddish brown, slightly shiny, the scales like those on the pinna rachises, proscales numerous, appressed, adaxially with indumenta like that of the pinna rachises; laminar tissue between the veins abaxially non-glandular, glabrous, adaxially sparsely glandular to nearly non-glandular; veins on both surfaces visible, abaxially non-glandular, sparsely pilose, the hairs 0.5 – 0.7 mm long, 3- or 4-celled, adaxially pilose, sparsely glandular, the hairs 1.0 – 2.0 mm long, 3 – 8-celled, spreading, the glands sessile, spherical; hydathodes evident; lamina margins sparsely ciliate to nearly glabrous, the hairs 0.2 – 0.4 mm long, 2 – 4-celled, non-glandular; indusia minute, fugacious, apparently consisting of a cluster of reddish proscales, c. 0.1 mm long, protruding from the centre of the sorus. Figs 11A and 12J – N; Map 3.

Distribution. Mexico, Honduras.

Additional Specimens Examined. Mexico. Chiapas: Rayón, Selva Negra 10 km above Rayón Mezcalapa along road to Jitotol, [17°10′N, 92°10′W], 1700 m, 25 Jan. 1973, Breedlove 32364 (F, NY); La Independencia, third ridge along logging road from Las Margaritas to Campo Alegre, [15°20′N, 92°34′W], 2300 m, 18 Feb. 1973, Breedlove 33687 (F, NY); SE side of Volcán Tacaná above Talquian, 2200 m, 16 Jan. 1973, Breedlove 31677 (F, NY); Motozintla de Mendoza, 45 – 50 km NE of Huixtla along rd to Motozintla, [15°22′N, 92°13′W], 1900 m, 17 Nov. 1971, Breedlove & Smith 22622 (F, MO, NY); Hidalgo: Chapulhuacán, [21°10′N, 98°54′W], 0 m, 27 Nov. 1937, Kenoyer 720 (F); Chapulhuacán, [21°10′N, 98°54′W], 0 m, 23 Nov. 1937, Kenoyer 689 (F); Puebla: Tehuacán area, above Teotitlan del Camino on the road to Huautla, [18°28′N, 97°23′W], 0 m, 3 Aug. 1961, Smith et al. 4139 (F, US); Teziutlán, Agua de Obispo, [19°48′N, 97°21′W], 1400 m, 28 Jan. 1970, Ventura A 399 (MICH, NY); San Marcos: between Finca El Porvenir and Loma Corona, 9 miles W of el Porvenir, SW facing slopes of Volcán Tajumulco, [15°01′N, 91°55′W], 0 m, 14 March 1949, Steyermark 37750 (F, US); vic. of Tajumulco, NW slopes of Volcán Tajumulco, [15°01′N, 91°55′W], 0 m, 28 Feb. 1940, Steyermark 36867 (F, US); Veracruz: Huayacocotla, El Salto, [20°39′N, 98°26′W], 1650 m, 13 March 1980, Ramírez 645 (F); 10 km de la carretera Teziutlán – Nautla, [20°13′N, 96°46′W], 0 m, Dec. 1945, Sánchez 345 (US). Honduras. La Paz: Aldea Las Marías, Cordillera Guajiquiro, Montaña Verde, [14°37′N, 88°20′W], 2100 m, 23 May 1964, Molina R. & Molina 14068 (F, US); Lempira: Celaque National Park, 2040 m, 13 Nov. 1991, Moran 5527 (AAU, MO); Santa Barbara: 7 km N of El Mochito, on the NE slopes of Mt Santa Barbara, 14°55′N, 88°07′W, 1880 m, 22 Nov. 91, Moran 5651 (MO).

Habitat. Cloud forests, wet forests; 1200 – 2800 m.

Conservation Status. Least Concern (LC).

Notes.Megalastrum sparsipilosum is characterised by large leaves (to 4.5 m long), laminae sparsely pilose on both surfaces, and strongly denticulate scales on the axes. It resembles M. heydei and M. pulverulentum in the large size and dissection of the leaves but differs from the former by longer hairs on both surfaces of the pinna rachises (these hairs only 0.1 – 0.2 mm long in M. heydei), and from the latter species by sparsely pilose pinna rachises, costules, and veins on abaxial surface of the laminae (versus densely pubescent in M. pulverulentum). The specific epithet sparsipilosum refers to the sparse pubescence on the abaxial surface of the laminae, especially when compared to M. pulverulentum.

Specimens of Megalastrum sparsipilosum from Chiapas and Guatemala are more sparsely pubescent than those from Honduras. In some, it is difficult to find hairs on the lamina margins — a characteristic unusual in the genus. The variation in pubescence does not appear to correlate with other characters and is therefore interpreted here as variation in a single species.

20. Megalastrum squamosumA. Rojas (1996 42, f. 6). Type: Costa Rica. Limón: Cordillera de Talamanca, Reserva Indígena Hitoy Cerere, siguiendo la fila entre Río Hitoy and Río Cerere, 9°38′35″N, 83°05′30″W, 300 m, 26 Feb. 1989, G. Herrera & Solís 2458 (holotype INB!; isotypes CR!, MO!).

Rhizomes erect; leaves up to 2 m long; scales of the petiole base 10 – 25 × 0.2 – 1 mm, dense, spreading to deflexed, linear, flat (not twisted), yellowish brown, often shiny, denticulate throughout; laminae up to 1.5 m long, 3-pinnate-pinnatifid at base, 2-pinnate-pinnatisect medially; basal pinnae c. 0.4 m long, strongly inequilateral (elongated basiscopically); pinna rachises abaxially non-glandular (neither stalked or sessile glands present), sparsely pubescent to glabrescent, the hairs 0.2 – 0.4 mm long, 3 – 5-celled, moderately to densely scaly, the scales to 2 – 4 × 0.2 – 0.3 mm, linear, flat (not twisted), spreading, yellowish brown, often shiny sparsely denticulate throughout; adaxially densely pubescent, the hairs 0.5 – 1 mm long, 5 – 8-celled, whitish, spreading to antrorsely strigose; costules abaxially sparsely pubescent and scaly, the hairs like those on the rachis, the scales 1.0 – 1.2 × c. 0.1 mm, filiform, brown, shiny, adaxially the hairs like those on the pinna rachises, scales absent; laminar tissue between the veins abaxially non-glandular (neither stipitate nor sessile glands present), abaxially sparsely puberulent, the hairs c. 0.1 – 0.2 mm long, 1-celled, erect, acicular, adaxially glabrous; veins visible on both surfaces, abaxially non-glandular, pubescent, the hairs 0.1 – 0.2 mm long, 2- or 3-celled, adaxially glabrous or with 1 – 3 hairs, the hairs 0.3 – 0.4 mm long, 3- or 4-celled, spreading, proscales present, appressed, reddish; hydathodes evident; lamina margins ciliate, the hairs 0.1 – 0.3 mm long, 1 – 3-celled, non-glandular; indusia minute, fugacious, apparently consisting of a cluster of reddish proscales protruding from the centre of the sorus. Figs 1F – J and 4D; Map 3.

Distribution. Talamanca mountains of Costa Rica and Panama.

Additional Specimens Examined. Costa Rica. Limón: Hitory Cerere Reserve and vic. In Valle La Estrella S of Finca Concepción, from station to top of ridge Miramar or Los Jabillos, 9°42′N, 83°02′W, 140 – 500 m, 1 Aug. 1985, Hammel & Grayum 14332 (CR, MO); Reserva Biol. Hitoy Cerere, siguiendo la fila frente a la confluencia de los dos afluentes principales, Cordillera de Talamanca, 9°38′35"N, 83°05′30"W, 300 m, 25 Feb. 1980, Herrera 2455 (MO); Siquirres, Las Brisas de Pacuarito, sector entre el camino principal y la márgen izquierda de la Quebrada Azul, 10°00′10"N, 83°28′50"W, 400 m, 6 Nov. 1995, Herrera 8717 (CR, K); Cantón de Limón, Reserva Biológica Hitoy Cerere, Cuenca del Estrella, Sendero Espavel, 9°39′40″N, 83°01′45″W, 520 m, 13 May 1998, Rojas et al. 4580 (CR, INB); Cantón de Siquirres, Zona Protectora Pacuare, Cuenca del Matina, Valle Escondido, entrando por las Brisas de Pacuarito, entre la parte alta de la Fila de Río Danta, 9°59′15″N, 83°28′20″W, 400 – 800 m, 14 April 1999, Rojas et al. 5062 (INB). Panama. Veraguas: 0.2 miles beyond fork in road at Escuela Agrícola Alto Piedra on road to Río Calovébora, [8°29′N, 81°06′W], 750 m, 3 April 1976, Croat & Folsom 33928 (MO, US); 0.5 miles beyond fork in road at Escuela Agrícola Alto Piedra on road to Río Calovébora, [8°29′N, 81°06′W], 750 m, 3 April 1976, Croat & Folsom 33913 (MO, NY); 0.6 miles beyond Escuela Agrícola Alto Piedra, [8°29′N, 81°06′W], 730 m, 4 April 1976, Croat & Folsom 34046 (MO, US).

Habitat. Wet forests; 140 – 800 m.

Conservation Status. Least Concern (LC).

Notes.Megalastrum squamosum is distinguished by petioles, rachises, and costae densely and conspicuously scaly. No other Central American species is as densely or conspicuously scaly. Sometimes the scales on the leaf axes are rubbed off and therefore absent, but the typical dense scales can be seen on the petiole bases. The scales are unlike those of many species in the genus by being firm, flat (not twisted), and yellowish brown. They resemble the scales of M. atrogriseum but are much larger. The scales of many other species in Central America differ by being dull, narrower, shorter, filiform, and twisted or tortuous toward the apex.

Another helpful distinguishing character is the minute (0.1 mm long), acicular hairs on the abaxial surface of the laminae between the veins. These hairs are mostly erect and can be easily overlooked when viewed from above. Similar hairs occur in Megalastrum galeottii, a species that differs by shorter and filiform scales that are scattered on the petioles, rachises, and costae.

See Megalastrum dentatum for comparison with that species.

21. Megalastrum subincisum (Willd.) A. R. Sm. & R. C. Moran, (Smith & Moran 1987: 129).

Polypodium subincisum Willd. (Willdenow 1810: 202); Phegopteris subincisa (Willd.) Fée (1852: 243); Nephrodium villosum var. subincisum (Willd.) Hook. ex Sodiro (1883: 54); Nephrodium villosum var. subincisum (Willd.) Jenman (1896: 114), comb. superfl.; Dryopteris subincisa (Willd.) Urb. (Urban 1903: 19); Aspidium subincisum (Willd.) H. Christ (1906: 56); Ctenitis subincisa (Willd.) Ching (1940: 250). Type: Venezuela, Federal Distr., Caracas, [10˚30′N, 66˚55′W], F. Bredemeyer s.n. (holotype B-W 19701!).

Leaves up to 2 m long; scales of the petiole bases 2.0 – 2.5 × 0.1 – 0.2 cm, linear-lanceolate, spreading, brown, denticulate on the margins; laminae up to 1.5 m long, 3-pinnate-pinnatisect at base, 2-pinnate-pinnatisect medially; basal pinnae 0.4 – 0.6 m long, strongly inequilateral (elongated basiscopically); pinnules short-stalked, 4 – 15 cm long; pinna rachises abaxially non-glandular, without hairs, scaly, the scales 5 – 6 × 0.5 – 0.7 mm long, lanceolate, non-bullate, golden brown, appressed to loosely ascending, rachises adaxially non-glandular, densely pubescent, the hairs 0.5 – 0.8 mm long; 4- or 5-celled, the scales like those on the abaxial surface; costules on the abaxial surface non-glandular, sparsely pubescent (especially distally) and scaly, the hairs 0.2 – 0.3 (– 0.5) mm long, 2 – 4-celled, erect to spreading or substrigose, acicular, the scales like those on the pinna rachises abaxially but smaller, 3 – 5 × 0.3 – 0.7 mm, lanceolate, enlarged basally, the adaxial surface pubescent, the hairs 0.5 – 1.0 mm long, 4- or 5-celled, substrigose; laminar tissue between veins glabrous on both surfaces, or near the costules and sinuses very sparsely puberulent abaxially, the hairs c. 0.1 mm long, erect, acicular; veins visible on both surfaces, abaxially non-glandular, sparsely pubescent, non-scaly, the hairs c. 0.5 – 0.8 mm long, 2- or 3-celled, spreading, the scales c. 0.3 mm long, uniseriate, appressed, reddish, adaxially non-glandular, sparsely pubescent, the hairs 0.8 – 1.0 mm long, 2 – 4-celled; laminar margins ciliate, the hairs 0.2 – 0.3 mm long, 1- or 2-celled, spreading to substrigose, non-glandular; indusia absent. Figs 10A and 15N – V; Map 3.

Distribution. Mexico, Honduras, Cuba, Jamaica, Haiti, Dominican Republic, Guadeloupe, Dominica, Venezuela, Colombia.

Additional Specimens Examined. Mexico. Chiapas: La Trinitaria, E of Laguna Tzikaw, Monte Bello Natl Park, [16°07′N, 92°2′W], 1300 m, 23 Jan. 1974, Breedlove & Smith 32211 (NY); Veracruz: country of Cordova, [15°52′N, 96°55′W], 1889 – 91, Fink 59, pro parte (MO, NY). Honduras. Comayagua: San Juanillo, Reserva Biológica Cordillera de Montecillos, 14°30′N, 87°53′W, 2020 m, 2 March 1991, House 868 (MO); Francisco Morazán: SW of San Juancito, Montaña La Tigra, [13°07′N, 86°58′W], 2100 m, 31 May 1962, Molina R. 10659 (F); Montaña La Tigra, [13°07′N, 86°58′W], 2000 m, 13 July 1964, Molina R. 14499 (F, GH, US); Montaña La Tigra, 30 km NE of Tegucigalpa, [13°07′N, 86°58′W], 2000 m, 22 April 1983, Pineda 255 (MO); Montaña La Tigra, 30 km NE of Tegucigalpa, [13°07′N, 86°58′W], 1600 m, 13 May 1984, Soihet 201 (MO); Santa Bárbara: 7 km N of el Mochito, on the NE slopes of Mt Sta. Bárbara, 14°55′N, 88°07′W, 1880 m, 22 Nov. 1991, Moran 5648 (MO).

Habitat. Wet forests, shady ravines; 1300 – 2100 m (in the Antilles, 900 – 1700 m; Moran et al.2009b).

Conservation Status. Least Concern (LC).

Notes.Megalastrum subincisum is characterised by lanceolate, golden brown scales on the pinna rachises and costules abaxially (Fig. 15P, Q), pinnae rachises hairless abaxially, and costules subglabrous abaxially (sparse hairs present distally). The laminar tissue between the veins is glabrous abaxially. In Central America M. subincisum most resembles M. galeottii, a species that can distinguished by filiform, dark brown scales on the pinna rachises abaxially, and in many specimens by minute (0.1 – 0.2 mm long) acicular erect hairs on the lamina tissue between the veins abaxially. When M. galeottii is glabrous between the veins abaxially, it can be distinguished by the narrower, filiform scales of the pinna rachises. Also similar are M. costipubens and M. longipilosum, but they differ by pubescent pinna rachises abaxially and narrower scales on the pinna rachises abaxially.

Christensen (1920) construed Megalastrum subincisum in the wide sense, occurring from Mexico to Bolivia. He admitted much variation in the presence versus absence of hairs and glands and the type of laminar scales. We find, however, that indument characteristics can be used to divide M. subincisum in the broad sense (and other species in the genus) into additional species that often correspond to elevation and geography. In Central America, species that we segregate from M. subincisum sensu lato are M. costipubens, M. galeottii, M. gompholepis, M. longiglandulosum, and M. longipilosum. In our restricted sense, M. subincisum is more narrowly distributed than conceived by Christensen, occurring only in Mexico and Honduras, the West Indies, and northern Venezuela and Colombia.

Acknowledgements

This research was funded by a grant to the senior author from the United States National Science Foundation (DEB 0717056). We thank the curators of AAU, B, BM, BR, F, GH, HB, HBR, HUA, K, MBM, MICH, MO, NY, P, R, RB, S, SP, SPF, U, UC, UPCB, and US for loans. We thank Alejandra Vasco (NY) for help generating the lists of specimens examined and index to collectors’ names and numbers, Roy Gereau (MO) for help with the Latin diagnoses, and Hannah Stevens (GIS lab at the New York Botanical Garden) for help with the distribution maps. The drawings of indument were made by Mr Haruto Fukuda.

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© The Board of Trustees of the Royal Botanic Gardens, Kew 2010