Megalastrum (Dryopteridaceae) of the Circumaustral Region: Chile, Argentina, and Southern Islands of the Atlantic, Pacific, and Indian Oceans

Megalastrum (Dryopteridaceae) of the Circumaustral region: Chile, Argentina, and Southern Islands of the Atlantic, Pacific, and Indian Oceans

Michael Sundue , 1 , 3 Germinal Rouhan ,2 and Robbin Moran1 1 The New York Botanical Garden, Bronx, New York 10456-5126, U. S. A. 2 Muséum national d’Histoire naturelle, UMR CNRS 7205 ‘Origine, Structure et Evolution de la Biodiversité’, Botanique, 16 rue Buffon CP39, F-75005 Paris, France 3 Author for Correspondence ( [email protected] )

Communicating Editor: Kenneth M. Cameron

Abstract—In this treatment we revise the circumaustral fern species of Megalastrum found primarily between 33–45° latitude South. These species are found in the Valdivian temperate rain forests of Chile and Argentina, the Juan Fernández Islands in the South Pacific, the Tristan da Cunha archipelago in the southern Atlantic, and Île Amsterdam and Île Saint-Paul in the southern Indian Ocean. Megalastrum is absent from Australia and New Zealand. Seven species are treated in total. Two species and two varieties of Megalastrum have generally been recognized in Chile and the Juan Fernández Islands. In this treatment, we recognize four species for the region, one of which is elevated to the species rank ( M. glabrius stat. nov.), and one which is new ( M. masafuerae sp. nov.). Within the Tristan da Cunha archipelago, two species occur: M. aquilinum comb. nov., and another here described as new, M. peregrinum sp. nov. Île Amsterdam and Île Saint-Paul harbor one species that is described here as new: M. taafense sp. nov. For each species we provide a key, descriptions, complete synonymy, discussions, specimens examined, and illustrations.

Keywords— Fern , Juan Fernández archipelago , Oceanic islands , pteridophyte , TAAF , Taxonomy , Trade winds , Tristan da Cunha archipelago.

This paper treats the circumaustral species of Megalastrum 2004 ) and some species of Polystichum Roth such as P. punc- Holttum occurring between 33°–45° south latitude. These tiferum C. Chr. (Christensen, 1931 ). species are found in the Valdivian temperate rain forests The above characteristics were first pointed out by of Chile and Argentina, the Juan Fernández Islands in the Christensen (1913 , 1920 ), who treated the species now called south Pacific, the Tristan da Cunha archipelago in the south Megalastrum as the “group of Dryopteris subincisa ” of sub- Atlantic, and Île Amsterdam and Île Saint-Paul in the south gen. Ctenitis. He assigned about 30 species to the group, of Indian Ocean. Megalastrum is absent from Australia and New which two species, each with a second variety, occurred in Zealand. This study is one in a series of papers treating the Chile and the Juan Fernández Islands. These four taxa have genus in large geographic regions. So far we have published a been accepted in recent floristic accounts within the region treatment of the 18 species of Megalastrum in Brazil, Paraguay, ( Marticorena and Rodríguez 1995 ). Since Christensen’s mono- and Uruguay ( Moran et al. 2009a ) and the nine species in the graph, no further comprehensive study has been published West Indies (Moran et al. 2009b). Other papers in this series for the species of Megalastrum occurring in Chile and the Juan will treat separately the remaining species; that is, those in Fernández Islands despite many subsequent collections. Africa-Madagascar-La Réunion, Central America, and the We also examined material from the oceanic islands of Andean region of South America. Between any two of these Tristan da Cunha, Île Amsterdam, and Île Saint-Paul from regions there is little or no overlap of species. where Christensen later reported (1940) a third species that For a review of the taxonomic history and geography of he believed was closely related to the Chilean species. Megalastrum , see Smith and Moran (1987) and Moran et al. The purpose of our study was to review the species of (2009a) . Christensen (1913 , 1920 ) was the first to point out the Megalastrum throughout the circumaustral region, examine distinctiveness of the group. He noted its characteristic lamina relevant type specimens, and provide an up-to-date taxo- cutting and venation; namely, as one goes distally along the nomic treatment. pinna, the basal basiscopic pinnules gradually become decurrent and broadly adnate to the pinna rachis. Correspondingly, Methods the vein that supplies the broadly adnate segment or lobe Herbarium specimens were borrowed from 25 herbaria, 17 of which arises from the pinna rachis, not the costule. Among ferns, had specimens from the region (see Acknowledgments). These specimens this combined cutting and venation pattern is unique. represent 108 separate gatherings. To show the variation in lamina cut- Christensen (1913 , 1920 ) pointed out another helpful char- ting, silhouettes were prepared from herbarium specimens for all spe- acteristic: the form of the hairs on the adaxial surfaces of cies. To do this, digital images were taken of basal pinna, and the images were then adjusted to provide a white background and a black lamina. pinna rachises. Generally, these are coarse, whitish, septate, Geographic coordinates were estimated for most of the specimens because sharp-tipped, and antrorsely strigose or spreading (Smith and this information was often not provided on the labels. Our estimates of Moran 1987 ). geographic coordinates are given in brackets for the herbarium specimens Another distinctive character of Megalastrum is that the cited below. veins in almost all species end before the lamina margins Results in enlarged tips (hydathodes). This type of vein termina- tion is uncommon among dryopteroid ferns, which typically We recognize seven species, of which three are new have slender vein tips that reach the margin. Some other ( M. masafuerae, M. peregrinum, and M. taafense ). No infraspe- Dryopteridaceae may have conspicuous hydathodes, how- cific taxa are recognized. These species are keyed to, described, ever, including Stigmatopteris C. Chr. (Moran 1991), the spe- and illustrated below. A synopsis of their distributions is cies of the monophyletic subsection Setosa of Elaphoglossum given in Appendix 1. A list of collectors and their numbers is Schott ex J. Sm. (Mickel and Atehortúa 1980; Rouhan et al. given in Appendix 2.

461 462 SYSTEMATIC BOTANY [Volume 35

Six of the seven species recognized are endemic to oceanic A. R. Sm. & R. C. Moran [basionym: Polypodium subin- islands of recent volcanic origin. They are all under the influ- cisum Willd.] ence of the prevailing westerly trade winds. The seventh spe- Plants terrestrial; rhizomes erect to decumbent; petioles cies is found on mainland South America in the Valdivian scaly toward the base, with 4–10 vascular bundles, the two forests of Chile and Argentina, which are temperate rainfor- adaxial bundles enlarged; laminae 1-pinnate-pinnatifid to ests. Strangely, no species of Megalastrum are known from 4-pinnate-pinnatifid, catadromic above the basal pinnae; New Zealand, despite it being in line with the westerly trade basal pinnae inequilateral and more developed on the basi- winds. scopic side or (less commonly) equilateral; rachises, costae, and costules not grooved or only shallowly so adaxially, scaly Megalastrum Holttum, Gard. Bull. Straits Settlem., ser. 3, and pubescent abaxially, densely pubescent on the adaxial 39: 161. 1986.—TYPE: Megalastrum villosum (L.) Holttum surfaces, the hairs whitish, spreading to antrorsely strigose, [basionym: Polypodium villosum L.]. multicellular, if glands present, these ca. 0.1 mm wide, spheri- Dryopteris subgenus Ctenitis C. Chr., group of D. subincisa cal, shiny, yellowish to orangish, sessile to stalked; basal basi- C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. scopic segment of more distal pinnules becoming decurrent Math. Afd. ser. 8, 6: 32, 59. (1919) 1920. Dryopteris subgenus and adnate to the pinna rachises, the vein supplying the seg- Ctenitis C. Chr. sect. Subincisae C. Chr., Index Filic., suppl. ment springing from the pinna rachis instead of the costule; 3: 7. 1934. Ctenitis subgen Ctenitis C. Chr. sect. Subincisae hydathodes (enlarged vein ends) present adaxially; indusia (C. Chr.) Tindale, Contr. New South Wales Natl. Herb. absent or (less commonly) present, circular, brown, firm, in 3(5): 252. 1965.—TYPE: Megalastrum subincisum (Willd.) some species minute and fugacious; x = 41.

Key to the Species of M EGALASTRUM in the Circumaustral Region 1. Hairs of the lamina margins 0.5–0.7 mm long, 4–7-celled, straight; indusia fugacious (sometimes absent), usually visible as a tuft of (1)5–10 erect acicular hairs, 0.5 mm long, emerging from the center of the sorus; rhizomes forming an erect trunk up to 1 m tall (Isla Masatierra, Juan Fernández Islands) ...... M. inaequalifolium 1. Hairs of the lamina margins 0.2–0.4 mm long, 3- or 4-celled, curved, or the lamina margins glabrous; indusia present or absent, provided with indument 0.1 mm long including either cilia, sessile glands, and or 1–2 setae; rhizomes not forming an erect trunk ...... 2 2. Indusia present, 0.3–0.8 mm wide; abaxial lamina surfaces between veins glabrous ...... 3 3. Lamina margins glabrous; laminae abaxially lacking hairs; hairs of the adaxial pinna rachises 0.1–0.3 mm long (Juan Fernández Islands, Isla Masafuera) ...... M. glabrius 3. Lamina margins sparsely ciliate, the hairs restricted to proximal portions of the ultimate segments; laminae abaxially sparsely pubescent along veins; hairs of the adaxial pinna rachises 0.4–0.8 mm long (Juan Fernández Islands, Isla Masafuera) ...... M. masafuerae 2. Indusia fugacious or absent, 0.1–0.4 mm wide; abaxial lamina surfaces between veins glabrous to puberulent and /or glandular ...... 4 4. Adaxial lamina surfaces between the veins sparsely to densely pubescent and glandular, the hairs 0.3–0.5 mm long, the glands evenly distributed (Gough Island) ...... M. peregrinum 4. Adaxial lamina surfaces between the veins glabrous to sparsely pubescent, sparsely glandular or nonglandular, the hairs ca. 0.3 mm long, the glands near the costae only ...... 5 5. Lamina margins flat, ciliate; indusia absent; adaxial laminae with hairs present on the veins and often between the veins, lacking glands (Chile, Argentina) ...... M. spectabile 5. Lamina margins recurved, sparsely ciliate to glabrous; indusia present, fugacious (rarely absent); adaxial laminae lacking hairs, with or without glands ...... 6 6. Lamina surfaces between veins pubescent and glandular abaxially; hairs of the abaxial pinna costules 0.5–0.7 mm long, 5- or 6-celled; adaxial lamina surface nonglandular or with glands restricted to the costae (Tristan da Cunha and Gough Islands) ...... M. aquilinum 6. Lamina surfaces between veins glabrous abaxially; hairs of the abaxial pinna costules 0.2–0.5 mm long; adaxial lamina surface nonglandular (Amsterdam and St-Paul Islands) ...... M. taafense

Taxonomic Treatment quilateral, short-stalked to sessile; pinnules sessile, the basal segments frequently overlapping the pinna rachis, the largest Megalastrum aquilinum (Thouars) Sundue, Rouhan, & pinnule of the basal pinna 6 cm long; pinna rachises abaxially R. C. Moran, comb. nov. Polypodium aquilinum Thouars, glandular, scaly, and densely pubescent, the glands sessile, Esquisse Fl. Tristan d’Acugna 32. 1808. Phegopteris aquilina yellow, hyaline, 0.1 mm wide, the hairs 0.2–0.7 mm long, 3- or Mett. ex Kuhn, Fil. Afr. 121. 1868. Nephrodium aquilinum 4-celled, hyaline, acicular, the cross walls reddish, the scales (Thouars) Hemsley, Rep. Voy. Chall., Bot., 1(2) 167, t. 39. 1.5–2.0 × 0.5 mm, ovate to lanceolate, light brown, translucent, 1885. Dryopteris aquilina (Thouars) C. Chr., Index Filic. entire, the cells isodiametric, adaxially glandular, scaly, and 252. 1905. Ctenitis aquilina (Thouars) Pic. Serm., Webbia densely pubescent, the glands and scales similar to those of 28: 468. 1973.—TYPE: TRISTAN DA CUNHA. without the abaxial side, the hairs 0.2–0.3 mm long, 2- or 3-celled, red- date, du Petit-Thouars, L.-M. A. s. n. (lectotype here desig- dish, blunt-tipped; costules abaxially densely glandular, scaly, nated: P- P00713304!; duplicate P- P00713305!). and pubescent, the glands like those of the pinna rachises, the Plants terrestrial; rhizomes not seen; fronds 0.5–1.0 m long; scales 1.0–1.5 × 0.3–0.5 mm, otherwise similar to those of the basal petiole scales ca. 4.0 × 0.2 cm, linear-lanceolate, light pinna rachises, the hairs 0.5–0.7 mm long, 5- or 6-celled, oth- brown, translucent, minutely denticulate, twisted, the cells erwise similar to those of the abaxial pinna rachises, adaxially elongate; laminae 25–50 cm long, 3-pinnate-pinnatifid at base, the costae densely glandular and pubescent, the hairs 0.5 mm 2-pinnate-pinnatifid medially; basal pinna 18 cm long, inae- long, 4-celled; laminar tissue between the veins abaxially 2010] SUNDUE ET AL.: MEGALASTRUM IN CIRCUMAUSTRAL REGION 463 slightly pale, glandular, glabrous or with scattered erect P. cruciata (Willd.) Holltum, a species that is doubtfully distinct hairs 0.2 mm long, adaxially darker in color, glabrous; veins from P. aubertii (Holttum 1974). After departing from Tristan not visible on either side of the lamina, abaxially glandular, da Cunha, du Petit-Thouars later traveled to Madagascar and pubescent, and provided with scattered 0.2 mm long uniseri- the Mascarenes ( Williams 2003 ) where P. aubertii is present. ate scales, the hairs 0.4–0.6 mm long, 4-celled, adaxially with It seems likely that the specimen at B was collected there. glands along the segment costa, otherwise glabrous; lamina Consequently, if the B material was not collected on Tristan da margins recurved, glabrous or very sparsely ciliate, the hairs Cunha, then it is in conflict with the protologue. Considering 0.2–0.3 mm long, 2- or 3-celled, acicular, restricted to the prox- these points and Recommendation 9A.5 of the ICBN (McNeill imal portions of the ultimate lobes; indusia present or rarely et al. 2006), we chose to designate the specimen at P as the lec- absent, fugacious, 0.1–0.4 mm wide, provided with ca. 0.1 mm totype for this species. long sessile glands, and sometimes with 1–4 ca. 0.1 mm long An illustration of this species appears in Hemsley (1885, setae; n = 41 ( Manton and Vida 1968 ). Figures 1M–R , 2 , 3A . plate XXXIX) based on Carmichael s. n. (K, n. v.). The name Etymology— From the Latin aquilinum, like that of an Polypodium acunhianum Carmichael also appears in Hemsley eagle. Du Petit-Thouars chose this specific epithet because (1885 , p. 167), placed in synonymy, but Carmichael’s manu- he found the “root” (likely the rhizome) similar to the one script (K), was not validly published and the name therefore of “ Pteris aquilina” (= Pteridium aquilinum). This is a strange is considered nom. inval. finding because the “roots” are probably not similar at all. Christensen (1940) and Tryon (1966) remarked on the sim- Although we have not seen the rhizome of M. aquilinum , it is ilarity of Megalastrum aquilinum to M. spectabile from Chile probably erect, like all other species in the genus, not creep- and Argentina, and suggested long distance dispersal as ing as in Pteridium . a means to explain this apparent amphi-atlantic distribu- Distribution and Ecology—Endemic to Tristan da Cunha, tion. Morphologically, the plants have similar lamina cut- Nightingale, Inaccessible, and Gough Islands in the south ting, densely pubescent axes provided with short erect hairs, Atlantic Ocean, where it reportedly grows in dense thick- brown lanceolate scales, and subglabrous to puberulent abax- ets beneath groves of Phylica arborea Thouars (Rhamnaceae), ial lamina surfaces between the veins. Megalastrum aquilinum which is one of the few trees native to the island (Wace and differs from M. spectabile most notably by its entirely glabrous Holdgate 1958 ); 50–600 m. adaxial lamina surfaces, the presence of fugacious indusia Additional Specimens Examined— TRISTAN DA CUNHA. Gough (sometimes absent, see below), sparsely ciliate lamina mar- Island: slopes behind hut, above Glen Beach, [40°20’S, 10°0’W], 50 ft, 30 gins with hairs that are often restricted to the proximal por- Nov 1955, Wace 12 (GH). Inacessible Island: Plateau at west end, [37°18′9”S, 12°40′28”W], 450 m, 26 Feb 1938, Christophersen 2545 (GH, O). [37°18′9”S, tions of the ultimate segments, thicker laminae, and recurved 12°40′28”W], s. d., Mosely et al. s. n. (GH); Base of Denstone Hill, [37°07’S, lamina margins. These last two characters, which were also 12°17’W], ca. 400 m, 14 Oct 1989, Roux 2147 (NBG). Nightingale : At the cited by Christensen (1940) , may be a response to wind dam- peak, [37°07’S, 12°17’W], 300 m, 10 Feb 1938, Christophersen 2217 (O). Trail age, but are consistent in all of the material that we have seen. to lake region, [37°07’S, 12°17’W], 100 m, 12 Feb 1938, Christophersen 2232 (O); streambed along West Road to summit, [37°07’S, 12°17’W], ca. 300 m, Christensen reported (1940) that he did not find an indusium Roux 2092 (NBG); Phylica forest near first pond, [37°07’S, 12°17’W], ca. 200 in M. aquilinum . We have seen one specimen annotated by m, 20 Oct 1989, Roux 2193 (NBG). Tristan da Cunha: Above settlement, Christensen (Christophersen 2545), and confirm that it does not [37°07’S, 12°17’W], 600 m, 21 Dec 1937, Christophersen 23 (O); East of Big have any trace of an indusium, but all other specimens that Point, [37°07’S, 12°17’W], 250 m, 29 Dec 1937, Christophersen 423 (O); Base we have seen do have small fugacious indusia (0.1–0.4 mm of Round Hill, [37°07’S, 12°17’W], 650 m, 14 Jan 1938, Christophersen 1061 (O); Round Hill, [37°07’S, 12°17’W], 800 m, 14 Jan 1938, Christophersen 1093 wide) bearing sessile glands and setae. (O); Up slope about 1 mi S of settlement, [37°07’S, 12°17’W], 600–1000 ft, 3 Megalastrum glabrius (C. Chr. & Skottsb.) Sundue, Rouhan Jan 1937, Dyer 3554 (NY); [37°07’S, 12°17’W], without precise date, Keytel & R. C. Moran, stat. nov. Dryopteris inaequalifolia (Colla) 4713 (NBG); [37°07’S, 12°17’W], 12 Nov 1852, Mac Gillivray 327 (GH); [37°07’S, 12°17’W], 12 Nov 1852, Mac Gillivray s. n. (NY); Above Soggy C. Chr. var. glabrius [as “glabrior ”] C. Chr. & Skottsb., Plain, at tree limit, [37°07’S, 12°17’W], 18 Feb 1938, Mejland 1575 (O). C. Chr., Kongel. Dankse Vidensk. Selsk. Skr. Naturvidensk. Discussion— In addition to the type at P, a du Petit-Thouars Math Afd., ser. 8, 6: 74. 1920. Dryopteris inaequalifo- collection at B is also annotated by Willdenow as type mate- lia (Colla) C. Chr. f. glabrius [as “glabrior ”] C. Chr. & rial (BGBM virtual herbarium, image ID: 260189, barcode: Skottsb., in Skottsb., Nat. Hist. Juan Fernández 2: 19. B-W 19721–01 0; Röpert 2000 ) but it is not clear that this col- 1920. Ctenitis inaequalifolia (Colla) Ching var. glabrius [as lection should be regarded as such. Although some of du “ glabrior ”] (C. Chr. & Skottsb.) Kunkel, Nova Hedwigia Petit-Thouars’ collections were distributed to B ( Stafleu and 9: 261. 1965. Ctenitis inaequalifolia (Colla) Ching f. glabrius Cowan 1976 ), the sheet in question does not bear a label indi- [as “ glabrior”] R. A. Rodri., Duek & R. A. Rodr. Bol. Soc. cating that it was in fact part of du Petit-Thouars’ herbarium. Biol. de Conception 45: 148. 1972. Megalastrum inaequali- The status of the B specimen is critical though, because it is folium (Colla) A. R. Sm. & R. C. Moran var. glabrius [as a different species, which we have identified as belonging to “ glabrior”] (C. Chr. & Skottsb.) R. A. Rodr., Gayana, Bot. Pseudophegopteris . 46: 10. 1989.—TYPE: CHILE. Juan Fernández, Masafuera, The protologue provided by du Petit-Thouars (1808) is i Casas-dalen [valley of the Rio Casas], 11 Feb 1917, brief and might pertain to either a species of Megalastrum C. Skottsberg & I. Skottsberg 466 [as “446”] (lectotype here or Pseudophegopteris. The provenance of the collection designated: UPS!; duplicate: BM!). at B provides some evidence of the name’s application. Plants terrestrial, rhizomes not seen; petioles not seen; Pseudophegopteris is not known from Tristan da Cunha fronds 2-pinnate-pinnatifid medially, 3-pinnate-pinnatisect at ( Christensen 1940 ; Holttum 1969 , 1974 ; Tryon 1966 ; Wace and the base, basal pinnae up to 60 cm long (estimated); medial Holdgate 1958 ; Wace 1960 , 1961 ), suggesting that the speci- pinnae 45 cm long, the base sessile, overlapping the rachis; pin- men at B was collected elsewhere and mislabeled. The B sheet nules 10 cm long, sessile, overlapping the pinna costa; pinna is most likely P. aubertii (Desv.) Holttum, a species named by rachises with a reddish hue, abaxially nearly glabrous, sparsely Desvaux in honor of du Petit-Thouars. Alternately, it could be glandular and sparsely pubescent, the hairs 0.3–0.4 mm 464 SYSTEMATIC BOTANY [Volume 35

Fig . 1. Detail of indument on three cicrumaustral Megalastrum species. A–E. Megalastrum peregrinum (Wace 133, GH) A. Abaxial costule and segments. B. Detail of scale. C. Detail of indument on abaxial costule. D. Adaxial costule and segments. E. Detail of hairs and glands on adaxial lamina surface. F–L. Megalastrum taafense (Jolinon 1071 , P) F. Abaxial surface of pinnule and costule at the juncture with the pinna rachis. G. Detail of the hairs on the abaxial costule. H. Detail of hair from the abaxial costule. J. Abaxial lamina surface. K. Detail of scale from the pinna costa. L. Adaxial lamina surface. M–R. Megalastrum aquilinum ( Christopherson 2545, GH) M. Abaxial surface of pinnule and costule at the juncture with the pinna rachis. N. Detail of indument upon abaxial lamina surface and veins. O. Detail of scale from pinna rachis. P. Adaxial lamina surface. Q. Detail of adaxial costule. R. Detail of indument from adaxial costule. 2010] SUNDUE ET AL.: MEGALASTRUM IN CIRCUMAUSTRAL REGION 465

Fig . 2. Distribution of the circumaustral species of Megalastrum . long, 2–4 celled, hyaline, the cross walls reddish, adaxially mod- Etymology—From the Latin glabri- , glabrous, and the com- erately puberulent, the hairs conspicuously short, 0.1–0.3 mm parative suffix –or/-us, more so. The specific epithet glabrius long, 1–3-celled, the cross walls reddish, the apices blunt; cos- refers to the absence of hairs on either side of the laminae and tules with a reddish hue, abaxially sparsely pubescent and the lamina margins. scaly, the hairs 0.3–0.5 mm long, 2–4-celled, the scales 0.5–0.7 × Distribution and Ecology—Endemic to Isla Masafuera 0.3–0.5 mm, ovate to lanceolate, gold-brown, translucent, (Alexander Selkirk Island) of the Juan Fernández Islands where entire, the cells isodiametric, adaxially moderately pubescent, it is known with certainty only from Quebrada de las Casas. the hairs 0.3–0.5 mm long, 3–4-celled; lamina tissue between Specimens Examined— CHILE. Juan Fernandez Islands: Masafuera, the veins glabrous on both sides, abaxially glaucous, adaxially without precise locality, 3 Mar 1920, Bryan s. n. (OSU); Masafuera Quebrada de las Casas, [33°46’S, 80°47’W], 28 Aug 1908, Skottsberg 731 dark green; veins visible on both sides of the lamina, abaxi- (BM, UPS). ally sparsely provided with appressed, filiform, uniseriate, Discussion— Megalastrum glabrius is known only from reddish, scales, 0.2–0.4 mm long, adaxially glabrous; lamina three fragmentary specimens, two of which were cited margins glabrous; indusia present, 0.3–0.4 mm wide, brown, by Christensen (1920). The specimen at BM, collected by the margin ciliate, the cilia 0.2 mm long, the same color as the Skottsberg, does not have a collection number, but has the indusia. Figures 4A –C, 2 , 5 A. 466 SYSTEMATIC BOTANY [Volume 35

Fig . 3. Lamina dissection of three Megalastrum species. A. Megalastrum aquilinum ( Christopherson 2545). B. Megalastrum peregrinum ( Wace 133 ). C–D. Megalastrum taafense (C. Jolinon 1071 , D. Aubert s. n. ). 2010] SUNDUE ET AL.: MEGALASTRUM IN CIRCUMAUSTRAL REGION 467

Fig . 4. Detail of indument on two circumaustral Megalastrum species. A–C. Megalastrum glabrius ( Skottsberg & Skottsberg 466 , BM) A. Abaxial pinna rachis and pinnules at the juncture with the rachis of the lamian. B. Adaxial pinna rachis and pinnules. C. Detail of indument on adaxial pinna rachis. D. Megalastrum masafuerae (Solbrig et al. 3676 , MICH), Abaxial pinna rachis and pinnules. same date and locality as Skottsberg 731 at UPS. Therefore, lium Colla, Mem. Reale Accad. Sci. Torino 39: 49–50. 1836; we consider it part of that collection. It should be noted that Herb. Pedem. 6: 228. 1836. Dryopteris skottsbergii C. Chr., Skottsberg 731 bis at UPS, identified as M. inaequalifolium , has a Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Math. different collection date and locality. Afd. ser. 8, 6: 15. 1920. Dryopteris inaequalifolia (Colla) In the original publication of the name Dryopteris inaequal- C. Chr., Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. ifolia var. glabrius C. Chr. & Skottsb., the collection number Math. Afd. ser. 8, 6: 74. 1920. Ctenitis inaequalifolia (Colla) “ 446 ” should be considered as a typographical error (instead Ching, Sunyatsenia 5: 250. 1940. Ctenitis inaequalifolia of “466”), because the specimen cited by Christensen (1920) (Colla) Ching fo. inaequalifolia R. A. Rodri., in Duek & clearly corresponds to C. Skottsberg & I. Skottsberg 466 : except R. A. Rodr. Bol. Soc. Biol. de Conception 45:148. 1972.— the collection number, all of the other information on the label TYPE: CHILE. Juan Fernández Island, “in sylvis umbrosis matches, and it bears the corrected identification annotated collium et montium Insulae Juan Fernández” Mar–May 1830, by Skottsberg. C. L. G. Bertero 1550 (lectotype here designated: P ex herb. Megalastrum glabrius differs from its congeners by the Bory barcode: P00600449!; duplicates: BM-000890150!, absence of hairs on either side of the laminae and the lamina NY!, P barcodes: P00600447!, P00600448!, P00600451!, margins, the short-puberulent adaxial pinna rachises, and the P00600672!, WAG). ciliate-margined indusia. Also distinctive are the dark green Polypodium berteroanum Hook., Sp. Fil. 4: 269. 1862, non Spreng. color of the adaxial lamina surfaces, the glaucous abaxial sur- 1827.—TYPE: CHILE. Juan Fernández Islands, “ sylv. faces, and the reddish axes. umbros. coll. Isl. Juan Fernández ”, May 1830, Bertero 1660 (lec- Megalastrum inaequalifolium (Colla) A. R. Sm. & R. C. totype here designated: K, the sheet bearing the barcode: Moran, Amer. Fern J. 77: 128. 1988. Polypodium inaequalifo- K000512194!; duplicate: K, a sheet labeled “1660(bis)” !, 468 SYSTEMATIC BOTANY [Volume 35

Fig . 5. Lamina dissection of two Megalastrum species. A. Megalastrum glabrius (Skottsberg 466 ). B. Megalastrum masafuerae ( Skottberg 448 ).

[without barcode]). Dryopteris villosa (L.) Kuntze var. lamina margins ciliate, the hairs 0.5–0.7 mm long, 4–7-celled; berteroana (Hook.) C. Chr., Ark. Bot. 10(2): 13. 1910. indusia present or rarely absent, fugacious, ca. 0.2 mm wide, provided with (1)5–10 erect cilia, these ca. 0.5 mm long, and Plants terrestrial; rhizomes not seen, reportedly forming emerging from the center of the sorus in a tuft. Figures 6G –M, erect trunks up to 1 m tall, 10 cm diam.; fronds 1.2–1.6 m long; × 2 , 7C–D . petiole scales 3.50 0.25 cm, linear-lanceolate, brown, minutely Etymology— From the Latin inaequal-, uneven, unequal, denticulate, several layers of cells thick at the base, opaque or dissimilar, and –folium, leaf, referring to the inaequilateral partially translucent distally; laminae 0.4–1.0 m long, 3-pin- basal pinnae. nate-pinnatisect; basal pinnae 22–50 cm long, inaequilateral; Distribution and Ecology— Endemic to the Isla Masatierra largest pinnules of the basal pinna 9.5–16.0 cm long, short- (Robinson Crusoe Island) of the Juan Fernández Islands in the stalked to sessile; pinna rachises abaxially glandular, pilose, South Pacific Ocean; 215–660 m. and with scattered scales, the glands sessile, yellow, hyaline, Additional Specimens Examined— CHILE. Juan Fernandez Islands: 0.1 mm wide, the hairs 1.0 mm long, hyaline, catenate, with Without locality or date, Bertero s. n. (P); Masatierra, subida al Mirador reddish cross walls, 5- or 6-celled, the scales 1.5–2.0 × 0.3– de Selkirk desde San Juan Bautista, [33°38’S, 78°51’W], 440 m, 27 Jan 1980, 0.4 mm, lanceolate, light brown, translucent, entire, the cells Marticorena et al. 9011 (OSU); Masatierra, Mirador de Selkirk, [33°38’S, 78°51’W], 575 m, 27 Jan 1980, Maticorena et al. 9026 (OSU); Masatierra, elongate, adaxial pinna rachises densely pubescent, the hairs Quebrada de Villagra, Salto de La Pulga, [33°38’S, 78°51’W], 445 m, 28 Jan ca. 0.6 mm long, 5-celled, hyaline, catenate, antrorse; costules 1980, Marticorena et al. 9037 (MO, OSU); Masatierra, plazoleta del Yunque, abaxially glandular, moderately pilose and with scattered [33°38’S, 78°51’W], 280 m, 2 Feb 1980, Marticorena & Ugarte 9082 (OSU); scales, the glands sessile, yellow, hyaline, ca. 0.1 mm wide, the Masatierra, Cerro Pascua, [33°38’S, 78°51’W], 450 m, 8 Feb 1980, Marticorena et al. 9153 (OSU); Mas Atierra, Cerro Salsipuedas, [33°38’S, 78°51’W], 1,800 hairs ca. 1.0 mm long, hyaline, catenate, with reddish cross × ft, 10 Dec 1965, Meyer 9499 (GH, MO); Mas Atierra, along trail to Comote, walls, 5- or 6-celled, the scales 0.4–0.6 0.2–0.5 mm, ovate, [33°38’S, 78°51’W], 1,000 ft, 25 Dec 1965, Meyer 9661 (GH, MO); s. d., Miller light brown, translucent, entire to erose, the cells isodiamet- 119 (BM); Isla Mas Atierra, trail above Weber’s house on the way to the ric, adaxially densely pubescent, the hairs ca. 0.4 mm long, divide S of El Tranque, [33°38’S, 78°51’W], 400 m, 1 Mar 1939, Morisson hyaline, catenate, antrorse; laminar tissue between the veins 17326 (GH, US); Isla Robinson Crusoe, El Camote, [33°38’S, 78°51’W], 520 m, 21 Jan 1992, Ricci 402 (MO); At rd. to Portezuelo de Villagra, [33°38’S, abaxially glabrous or provided with appressed uniseriate red- 78°51’W], 215 m, 24 Dec 1954, Skottsberg & Skottsberg 12 (NY, S); Mas a dish scales, ca. 0.4 mm long, adaxially glabrous; veins visible Tierra, [33°38’S, 78°51’W], 250 m, 6 Dec 1916, Skottsberg & Skottsberg 65 on both sides of the laminae, abaxially glandular and mod- (P, S, UPS); Mas a Tierra, Salsipuedes, [33°38’S, 78°51’W], 435 m, 20 Dec 1916, erately pilose, the glands sessile, yellow, hyaline, ca. 0.1 mm Skottsberg & Skottsberg 168 (S, UPS); Masatierra, oversta delen av Villagre- dalen, [33°38’S, 78°51’W], 500 m, 10 Jan 1917, Skottsberg & Skottsberg 274 long, the hairs ca. 0.8 mm long, hyaline, composed of 6 cells, (P, S, UPS, US); Masatierra, [33°38’S, 78°51’W], 500 m, 10 Jan 1917, Skottsberg spreading, adaxially with scattered hairs along the midvein, & Skottsberg 275 (BM, P, S); Mas a Tierra, V. Colonial, Monte Maderugo, the other veins glabrous, the hairs ca. 0.4 mm long, appressed; [33°38’S, 78°51’W], 24 Jan 1917, Skottsberg & Skottsberg 339 (S); Mas a Tierra, 2010] SUNDUE ET AL.: MEGALASTRUM IN CIRCUMAUSTRAL REGION 469

Fig . 6. Detail of indument on two circumaustral Megalastrum species. A–F Megalastrum spectabile (A–B Dusén s. n. NY; C–F Solomon & Solomon 4559 , NY). A. Basal petiole. B. Basal petiole scale. C. Abaxial surface of pinna rachis and pinnae at the juncture of the rachis of the lamina. D. Detail of hairs on abaxial rachis of the lamina. E. Detail of hairs and scales on abaxial pinna costa. F. Adaxial surface of pinna rachis and pinnule. G–M Megalastrum inaequalifolium (G–H Meyer 9661, NY; J–M Skottsberg & Skottsberg 12, US) G. Basal petiole. H. Basal petiole scale. J. Abaxial pinna rachis and pinnules. K. Detail of hairs on abaxial pinna costa. L. Adaxial pinna rachis and pinnule. M. Detail of hairs on adaxial pinna costa. 470 SYSTEMATIC BOTANY [Volume 35

Fig . 7. Lamina dissection of two Megalastrum species. A–B. Megalastrum spectabile (A. Skottsberg 921 , B. Hunnewell 15852 ). C–D. Meglastrum inaequalifolium (C. Skottsberg 420 , D. Morisson 17326 ). 2010] SUNDUE ET AL.: MEGALASTRUM IN CIRCUMAUSTRAL REGION 471

[33°38’S, 78°51’W], 22 Aug 1908, Skottsberg 721 (S); Mas a Tierra, Quebrada 20 cm long; pinna rachises abaxially sparsely-glandular, Salsipuedes, [33°38’S, 78°51’W], 660 m, 13 Jan 1917, Skottsberg & Skottsberg sparsely scaly and sparsely pubescent, the scales and hairs 1151 (S, UPS); Masatierra, Valle Colonial, [33°38’S, 78°51’W], 22 Aug 1908, slightly more abundant distally, the glands short-stipitate, ca. Skottsberg 731 bis (UPS); Mas a Tierra, Salto de Pangal, [33°38’S, 78°51’W], × 180 m, 25 Oct 1955, Sparre 75 (S); Masatierra, on path from pueblo San 0.1 mm long, hyaline, the scales 3.0–4.5 0.5–0.6 mm, linear- Juan Bautista to Mirador Selkirk, [33°38’S, 78°51’W], 480 m, 28 Jan 1980, lanceolate, slightly bullate, lustrous, light brown, entire, the Stuessy & Saunders 5014 (OSU); Masatierra, Cerro Agudo, down into sec- cells elongate, the hairs 0.4–0.8 mm long, 3–6-celled, acicular, ond valley on Villagra side, [33°38’S, 78°51’W], 630 m, 12 Feb 1990, Stuessy antrorse, hyaline with reddish cross-walls, the rachises adax- et al. 11644 (OSU); Masatierra, Puerto Frances, up the ridge W of the ascent ridge, then down into the forest and across in La Piña, [33°38’S, 78°51’W], ially nonglandular, without scales, densely pubescent, the 590 m, 23 Jan 1991, Stuessy & Crawford 11963 (OSU); Masatierra, up from hairs like those of the abaxial surface; costules on the abax- La Plazoleta del Yunque on trail to La Carbonera de Torres, [33°38’S, ial surface nonglandular, sparsely pubescent and sparsely to 78°51’W], 440 m, 28 Jan 1991, Stuessy & Soto 12055 (OSU); Masatierra, moderately scaly, the hairs like those of the rachises, the scales down from top of cerro (ridge up from Centinela) into quebrada toward × the Corrales de Molina, [33°38’S, 78°51’W], 640 m, 30 Jan 1991, Stuessy 0.6–2.0 0.2–1.0 mm, ovate, gold-brown, translucent, slightly & Crawford 12129 (OSU); Masatierra, Valle Villagra, below Cerro Agudo, iridescent, the cells mostly isodiametric, the apices acute or [33°38’S, 78°51’W], 350 m, 24 Jan 1997, Stuessy et al. 15396 (WU). filiform, the filiform portion up to 0.5 mm long when pres- Discussion—Colla appears to have published the name ent, the margins erose to entire, sometimes with a few broad Polypodium inaequalifolium in two places at nearly the same teeth or cilia up to 0.2 mm long, adaxially moderately pubes- time, Herb. Pedem. 6: 228. 1836, and Mem. Reale Accad. Sci. cent, the hairs erect to spreading, otherwise similar to those Torino 39: 49. 1836. According to TL-2 (Stafleu and Cowan of the abaxial rachises; lamina tissue between the veins abaxi- 1976), Herb. Pedem 6 was received between November and ally and adaxially glabrous; veins visible on both sides of the December of 1836. A date which volume 39 of the Mem. Reale lamina, the veins glabrous adaxially, abaxially the larger veins Accad. Sci. Torino was received by libraries is not known, sparsely pubescent, the hairs 0.5 mm long, 4-celled, smaller but it has come to our attention that the Memoire was in fact veins moderately provided with 0.2–0.7 mm long, nong- read by Colla to the society during a meeting on the 29th of landular, erect, reddish, filiform, scales, these uniseriate or November 1835. A copy of volume 39 at the library of the rarely bifurcating; lamina margins sparsely ciliate, the hairs Accademia della Scienze in Torino attests to this with an restricted to proximal portions of the pinnules, the hairs 0.3 inscription on the inside cover that reads “exhibitus die 29 mm long, 3-celled; indusia present, 0.4–0.8 mm wide, brown, novembris 1835,” but according to article 29.1 of the ICBN, often folded or lost in mature sori, the margins sparsely cili- effective publication would require that a pamphlet also be ate, the cilia 0.1–0.2 mm long, brown. Figures 4D , 2 , 5B . distributed at the time of the reading ( McNeill et al. 2006 ). Etymology— The specific epithet is derived from the origi- Megalastrum inaequalifolium is distinguished by massive nal name given to Alexander Selkirk Island, Isla Masafuera. trunk-forming rhizomes (reportedly up to 1 m tall, 10 cm Distribution and Ecology— Endemic to the Isla Masafuera diam.), large petiole scales (3.50 × 0.25 cm), pilose pinna (Alexander Selkirk Island) in the Juan Fernández Islands rachises (the hairs 1.0 mm long), veins of the abaxial lam- in the South Pacific Ocean, known only from the Valley of ina surfaces with hairs 0.8 mm long, ciliate lamina margins Quebrada de las Vacas and North branch of the Quebrada (the hairs 0.5–0.7 mm long), and fugacious indusia. The latter Varadero. appear as (1)5–10 erect cilia (0.5 mm long) that emerge as a Additional Specimens Examined— CHILE. Juan Fernandez Islands: tuft from the center of the sorus. Indusia are often missing in Masafuera, North branch of Quebrada Varadero, at the first waterfall, [33°46’S, 80°47’W], 100 m, 3 Feb 1986, Ruiz & Landero 8265 (OSU); mature sori, and in some cases apparently absent altogether. Masafuera, Quebrada de los Vacas, [33°46’S, 80°47’W], 10 Feb 1917, The type of Megalastrum inaequalifolium ( Bertero 1550) is a Skottsberg & Skottsberg 448 (S). mixed collection. Another sheet of this number at BM (no. Discussion— Megalastrum masafuerae is distinguished by 000890149) is M. spectabile . large indusia (0.4–0.8 mm wide) with sparsely ciliate margins, Hooker’s syntypes for Polypodium berteroanum included axes and veins sparsely pubescent abaxially, and nearly gla- both M. inaequalifolium , M. spectabile (Chile, 1831, Anderson brous lamina margins with marginal hairs (if any) restricted s. n. K), and what is most likely M. pleiosorus (Hook. f.) C. V. to the proximal portions of the sinuses. Megalastrum glabrius, Morton from the Galapagos Islands (n.v.). The lectotype cho- also endemic to Isla Masafuera, is indusiate, but its indusia sen here agrees with Christensen’s (1913) application of that are smaller (0.3–0.4 mm wide). Furthermore, it lacks hairs on name as a synonym of M. inaequalifolium . the laminae and has puberulent adaxial axes. Megalastrum Megalastrum masafuerae Sundue, Rouhan & R. C. Moran, sp. inaequalifolium on Isla Masatierra also has indusia, but it dif- nov.—TYPE: CHILE. Juan Fernández Islands, Masafuera, fers by the indusia being fugacious, smaller (ca. 0.2 mm wide), Quebrada Vacas, on slopes of canyon among Myrceugenia , and bearing (1)5–10 erect acicular hairs, as well as by its pilose between rocks, about 60–90 cm tall, no trunk, near brook, axes and evenly ciliate lamina margins that are provided with rare, 28 Nov 1965, O. T. Solbrig, H. E. Moore Jr., & J. Walker 0.5–0.7 mm long hairs. 3676 (holotype: GH!; isotypes: MICH!, US!). Megalastrum peregrinum Sundue, Rouhan & R. C. Moran A Megalastro inaequalifolio indusiis majoribus et lamina mar- sp. nov.—TYPE: TRISTAN DA CUNHA. Gough Island, gine breviter ciliata laminae differt . southern end of Gonydale, frequent between boulders Plants epipetric; rhizomes not seen, reportedly not form- in sheltered hollows and crevices, 1,600 ft., 12 Feb 1956, ing erect trunks; leaves ca. 1.2 m long; scales of the petiole N. M. Wace 133 (holotype: GH!). base 1.5–2.0 × 0.20–0.35 cm, linear-lanceolate, entire, brown, A Megalastro aquilino laminae pagina adaxiali supra dense lustrous, the cells isodiametric to elongate; laminae ca. 0.6 m pubescenti et glandulosa differt . long, 3-pinnate-pinnatifid at base, 2-pinnate-pinnatisect Plants terrestrial; rhizomes erect, ca. 1.5 cm wide; fronds medially; basal pinnae 35–55 × 40 cm, inaequilateral; pinnules 0.35–1.00 m long; basal petiole scales 3.0 × 0.1–0.15 cm, lin- short-stalked to sessile, the largest pinnule of the basal pinnae ear-lanceolate, light brown, translucent, minutely denticulate, 472 SYSTEMATIC BOTANY [Volume 35 twisted, the cells elongate; laminae 30–60 cm long, 3-pinnate volcanic origin, with ages of one million years (Tristan), six at base, 2-pinnate-pinnatifid medially; basal pinna 12–22 cm million years (Gough), eight million years (Inaccessible), and long, inaequilateral, short-stalked; pinnules sessile, the basal 16 million years (Nightingale group) (Wace and Dickson 1965). segments frequently overlapping the pinna rachis, the largest The islands have a temperate flora that is low in diversity, but pinnule of the basal pinna 7–9 cm long; pinna rachises abaxi- rich in ferns. About half of the vascular plant diversity is com- ally glandular, scaly, and densely pubescent, the glands ses- prised of ferns and lycophytes ( Christensen 1940 ; Tryon 1966 ; sile, globose, 0.1 mm long, the hairs 0.2–0.4 mm long, 3- or Wace and Holdgate 1958 ). The islands’ flora is notable for its 4-celled, hyaline, acicular, the cross walls reddish, the scales high endemism: at least half of its species or varieties of ferns 1.5–4.0 × 0.4–1.2 mm, ovate to lanceolate, translucent, light and lycophytes are endemic ( Christensen 1940 ; Tryon 1966 ). brown, slightly bicolorous, the center darker than the edges, The addition of M. peregrinum brings the number of ferns and entire, the cells isodiametric, adaxial pinna rachises not seen; lycophyte species in the island group up to 34. Based on the costules abaxially glandular, scaly, and pubescent, the glands young age of the Tristan da Cunha archipelago and the simi- like those of the costae, the scales 1.5–2.5 × 0.5 mm, ovate to larity of M. peregrinum to the South-American M. spectabile , lanceolate, the apices attenuate, up to 1/3 the length of the an allopatric origin of this species through long-distance dis- scale, otherwise similar to those of the pinna rachises, the persal from South America is likely. A similar long-distance hairs similar to those of the abaxial costae, adaxial costules not dispersal event has been inferred for several species in the seen; lamina tissue between the veins abaxially glandular and “ ciliatum group” of Elaphoglossum on Tristan da Cunha (Vasco sparsely pubescent, the glands sessile, globose, 0.1 mm long, et al. 2009 ). Alternatively, M. peregrinum might have evolved the hairs 0.2–0.3 mm long, 2-celled, erect, adaxially glandular locally on the islands from M. aquilinum , to which it is per- and pubescent, the glands sessile, globose, 0.1 mm long, the haps even more similar morphologically. hairs 0.3–0.5 mm long, 3- or 4-celled, spreading; veins not vis- Megalastrum spectabile (Kaulf.) A. R. Sm. & R. C. Moran, ible on either side of the lamina, abaxially glandular, pubes- Enum. Filic. 121. 1824. Polypodium spectabile Kaulf., cent, and provided with scales along the main vein, the hairs Enum. Fil.: 121. 1824. Phegopteris spectabilis (Kaulf.) Fée, 0.3–0.5 mm long, 3- or 4-celled, adaxially pubescent, the hairs Mém. Foug. 5 Gen. Filic. 243. 1852. Dryopteris spectabilis like those between the veins; lamina margins recurved, cili- (Kaulf.) C. Chr., Kongel. Dankse Vidensk. Selsk. Skr. ate, the hairs 0.2–0.4 mm long, 2- or 3-celled, acicular; indusia Naturvidensk. Math. Afd., ser. 8, 6: 69. 1920. Ctenitis spec- present, 0.3–0.4 mm wide, light brown, fugacious, the sur- tabilis (Kaulf.) Kunkel, Nova Hedwigia 13: 333. 1967.— face and margins with sessile glands, ca. 0.1 mm long, and TYPE: CHILE. 1827, L. A. von Chamisso s. n. (lectotype, sometimes with 1 or 2 setae, ca. 0.1 mm long; spores echinate chosen by Christensen (1920) : P!). ( Tryon and Lugardon 1990 : 460 as Ctenitis aquilina ). Figures 1A–E , 2 , 3B . Polypodium contractum Desv. Mém. Soc. Linn. Paris 6: 242. Etymology—From the Latin peregrinus, foreign, having a 1827.—TYPE: CHILE. “habitat in regno Chilense,” no date, tendency to wander. Referring to the ancestor of this species collector unknown, s. n. (holotype: P!, herb. Desvaux). that must have wandered to these remote islands from South America. Dryopteris spectabilis (Kaulf.) C. Chr. var. philippiana C. Chr., Distribution and Ecology—Endemic to Gough Island in the Kongel. Danske Vidensk. Selsk. Naturvidensk. Math. South Atlantic Ocean, which is part of the Tristan da Cunha Afd., ser. 8, 6: 70. 1920. Ctenitis spectabilis (Kaulf.) Kunkel archipelago; among rocks, on banks, in the open, or under var. philippiana (C. Chr.) R. A. Rodr., in Duek et R. A. Rodr., stands of Phylica arborea ca. 55–500 m. Bol. Soc. Biol. Concepción 45: 148. 1972. Megalastrum Additional Specimens Examined— TRISTAN DA CUNHA. Gough spectabile (Kaulf.) A. R. Sm. & R. C. Moran var. philippi- Island: Transvaal Bay, damp area below Phylica trees, growing on moss anum (C. Chr.) A. R. Sm. & R. C. Moran.—TYPE : CHILE. bank, 40°21’S, 9°53’W, 22 Oct 1989, Cooper G89053 (NBG); Snoek Gat., tus- Valdivia, Krause s. n. (lectotype here designated: NY!). sock grassland and fernbush, [40°20’S, 10°0’S], 22 March 2000, Hänel & Jones CH 021 (NBG); Under Phylica arborea north-west of base, 40°S, 9°W, Plants terrestrial; rhizomes not forming erect trunks; fronds 55 m, 31 Oct 1979, Roux 716 (NBG). up to 1.4 m long; petiole scales 1.0–2.2 × 0.06–0.15 cm, linear- Discussion— Megalastrum peregrinum is distinguished from lanceolate, gold-brown, sparsely and minutely denticulate, the all other species in the circumaustral region by the lamina tis- base of the scale thickened and curved immediately above the sue between the veins adaxially pubescent and glandular. In point of attachment, the apices filiform, somewhat tortuous, other aspects, M. peregrinum is similar to M. spectabile of Chile the cells elongate; laminae 35–60 cm long, 2-pinnate-pinnati- and Argentina, M. aquilinum of Tristan da Cunha and Gough sect to 3-pinnate-pinnatifid at the base, 2-pinnate-pinnatifid Islands, and M. taafense of Île Amsterdam and Île Saint-Paul. In medially; basal pinnae (12)22–28(52) cm long, inaequilateral; addition to the indument of the adaxial laminae, it differs from pinnules short-stalked to sessile, the largest pinnule of the M. spectabile in having recurved lamina margins, a fugacious basal pinna 9–11 cm long; pinna rachises sparsely glandular, indusia, and longer scales (1.5–2.5 mm long vs. 0.5–1.2 mm densely pubescent and moderately scaly both abaxially and long) on the abaxial costules, and from M. aquilinum and. adaxially, the glands ca. 0.1 mm long, sessile, hyaline, yellow- M. taafense by the adaxial indument, and having more densely ish, the hairs 0.3–0.5 mm long, 2–5-celled, acicular, hyaline, ciliate lamina margins and longer scales (1.5–2.5 mm long vs. spreading, the scales 0.8–3.5 × 0.2–0.6 mm, lanceolate, brown, 1.0–1.5 mm long) on the abaxial costules. lustrous, entire to erose, the cells elongate; costules abaxially The South Atlantic islands of the Tristan archipel- sparsely glandular, densely pubescent and moderately scaly, ago, including Tristan da Cunha, the Nightingale group, the glands ca. 0.1 mm long, sessile, hyaline, yellowish, the Inaccessible, and Gough Island, are about 1,750 miles west of hairs 0.1–0.5 mm long, 1–5-celled, acicular, hyaline, spread- South Africa and 2,088 miles east of South America. Gough ing, the scales 0.5–1.2 × 0.1–0.2 mm, lanceolate, brown, adaxi- Island lies 230 miles southeast of the other islands in the ally pubescent, the hairs like those of the abaxial side; laminar Tristan da Cunha archipelago. All the islands are of recent tissue between the veins abaxially glabrous to puberulent, 2010] SUNDUE ET AL.: MEGALASTRUM IN CIRCUMAUSTRAL REGION 473 the hairs erect 0.3 mm long, also provided with appressed 0.4 species, M. caribeum ( Funk & Schlim 282 , Moritz 202 pp ) and mm long reddish uniseriate proscales, adaxially glabrous to M. subincisum ( Funck & Schlim 1575 , Moritz 202 ). By chosing sparsely pubescent, the hairs ca. 0.3 mm long, 3- or 4-celled; the Chamisso collection as a type, Christensen restricted the veins visible on the abaxial side of the lamina, abaxially mod- application of the name. erately pubescent and glandular, the hairs 0.2–0.7 mm long, Length and distribution of pubescence is variable in erect, the glands sessile, yellow, hyaline, adaxially sparsely Megalastrum spectabile, with specimens ranging from glabrous pubescent to moderately pubescent, the hairs 0.3–0.7 mm to puberulent on the lamina surfaces between the veins abaxi- long, appressed; lamina margins ciliate, the hairs 0.3–0.4 mm ally. The length of hairs upon the veins also varies in these long, 3–4-celled; indusia absent. Figures 6A–F , 2 , 7A–B . plants, abaxially ranging from 0.2–0.7 mm long, and adaxi- Etymology— From the Latin spectabilis, spectacular, ally ranging from 0.3–0.7 mm long. Longer haired plants showy. have been recognized as M. spectabile var. philippiana , which Distribution and Ecology— Chile, Argentina, restricted to Christensen described as being smaller, more coriaceous, more Valdivian temperate rain forests; 10–350(–500) m. densely scaly, and having subulate whitish hairs intermixed Specimens Examined—ARGENTINA. Chubut: Lago [illeg.], 2 Feb with the shorter hairs. While none of Christensen’s charac- 1945, Castellanos 114252 (NY). Neuguen: Lago Espejo, 40°37’S, 71°45’W, 20 ters reliably distinguish var. philippiana , it can be identified Mar 1939, Cabrera 5037 (GH). Rio Negro: Neuguen, Lago Nahuel Huapi, Puerto Manzano, 41°0’S, 71°30’W, 12 Mar 1963, de la Sota 2799 (MICH); by being glabrous between the veins abaxially, and by having Bariloche, Puerto Blest Area, west end of Lago Nahuel Huapi, 41°0’S, slightly longer hairs on the veins abaxially (0.5–0.7 mm long). 71°30’W, 5 Nov 1982, Sage s. n. (UC). Unknown: s. d., Weiland s. n. (S). By comparison, var. spectabile is identified by the puberulent CHILE. Aisen: Patagonia occidental, Rio Aisen [Aysen], 26 jan 1897, lamina tissue between the veins abaxially (the hairs 0.3 mm Dusen s. n. (BM, S); Patagonia occidental, [45°24’S, 72°42’W], 15 Jan 1897, Dusen s. n. (NY); Puerto Aisen, [45°24’S, 72°42’W], 17 Feb 1945, Looser long, erect), and relatively shorter hairs on the veins abaxially 5062 (GH). Araucania: Temuco, [38°45’S, 72°40’W], 5 Feb 1906, anon. s. n. (0.2–0.6 mm long). We find that the variation in pubescence (BM); Temuco, [38°45’S, 72°40’W], Jan 1919, Claude-Joseph 885 (US); between these varieties is part of a continuum with numerous Cautin, Llaima, 38°42’S, 71°46’W, 25 Jan 1925, Claude-Joseph 3046 (US); intermediates, and not worthy of recognition. Melleco, Angol, Parque Nacional Contemo, [38°0’S, 70°52’W], 24 Apr Megalastrum spectabile is restricted to the Valdivian temper- 1970, de la Sota 6011 (NY); Lota, [37°5′S, 73°10’W], 5 Mar 1877, Savatier 272 (P); Cautin, Temuco, Cerro Nielol, 38°44’S, 72°40’W, 350 m, 27 Nov ate rainforests of Chile and neighboring Argentina. Most col- 1947, Wall & Sparre 3240 (S). Biobio: Concepción, 37°5′S, 73°15’W, s. d., lections have been made between 36°–42° deg. S, with one anon. 156 (K); Concepción,Talcahuano, Parque Hualpen, 36°46’S, 73°6’W, ( Sparre 2948 ) at 30° S, outside of Ovalle, in the Región de ′ 21 Apr 1970, de la Sota 6028 (US); Concepción, Valparaiso, [37°5 S, Coquimbo, where northern relictual pockets of Valdivian for- 73°15’W], s. d., Gaudichaud 34 (P); Concepción, [37°5′S, 73°15’W], 1841, Hombron s. n. (GH); Concepción, quebrada on Avenida Alemana, 18 est occur. Feb 1951, Junge 2924 (US); Concepción, Collin, cerca de Tome, 36°37’S, Megalastrum taafense Rouhan, Sundue & R. C. Moran, sp. 72°57’W, 24 Jul 1934, Looser s. n. (US); Concepción, Talcahuano, 36°43’S, nov.—TYPE: TERRES AUSTRALES ET ANTARCTIQUES 73°7’W, Apr 1868, Poeppig 45 (P); Arauco, Contulmo, 38°0’S, 73°14’W, 16 July 1975, Zollner s. n. (NY). Coquimbo: Coquimbo, 8 Feb 1948, Files FRANÇAISES. Nouvelle Amsterdam, dans une voûte 574 (S); Ovalle, Bosque de Talinay, [30°35’S, 71°12’W], 500 m, 8 Oct 1947, d’une coulée, ombragée, 12 Jul 1963, A. Lourteig & P. Cour Sparre 2948 (S). Los Lagos: Valdivia, Quitalito, [39°48′50”S, 73°14′45”W], 50 (holotype: P!). 23 Mar 1931, anon. 1430.4 (GH); Valdivia, [39°48′50”S, 73°14′45”W], s. d., Buchtien s. n. (S); Valdivia, [illeg.], [39°48′50”S, 73°14′45”W], 15 Apr 1904, A Megalastro aquilino lamina adaxialiter glabra abaxialiter Buchtien s. n. (S); Valdivia, [39°48′50”S, 73°14′45”W], 1906, Buchtien s. n. inter venas glabra differt . (S); Valdivia, [39°48′50”S, 73°14′45”W], 15 Feb 1904, Buchtien s. n. (BM); Plants terrestrial, rhizomes erect, 2.7 cm diam., not forming Valdivia, Corral, [39°48′50”S, 73°14′45”W], 8 Jul 1930, Gunkel 2467 (GH); large erect trunks, the scales 35.0 × 1.0–1.5(–2.3) mm, linear Llanguihue, Puerto Varas, 41°19’S, 72°59’W, Feb 1926, Gunther & Buchtien lanceolate, thin, brown, translucent, irregularly twisted, sub- 13 (S); Valdivia, [39°48′50”S, 73°14′45”W], 200 m, Mar 1925, Hollerymayer 693 (B, BM, F, GH, MO); Panguipulli, [39°38′40”S, 72°19′50”W], s. d., entire, bearing few teeth distally; fronds up to 80 cm long, pet- Hollerymayer 156c (B); Llanguihue, Peulla, 41°19’S, 72°59’W, 2 Jan 1939, ioles sulcate; petiole bases scaly with scales similar to those Hunnewell 15852 (GH); Chiloe, Piruquina, [42°28’S, 73°48’W], 17 Dec of the rhizome, but shiny dark brown; laminae up to 41 cm ′ 1931, Junge 84 (107) (MO); Llanquihue, Casa Pangue, 41°3 S, 71°52’W, long, lanceate to triangular; 2-pinnate-pinnatisect (rarely trip- Dec 1926, Shannon & Shannon 31 (US); Chiloe, 42°17’S, 73°40’W, 26 Jul 1908, Skottsberg 921 (S); Chiloe, vid mynningen af Rio Pudeto, [42°28’S, innate) at base, 2-pinnate-pinnatifid to 2-pinntate-pinnatisect 73°48’W], 16 Jul 1908, Skottsberg s. n. (UPS); Chiloe, W of rt. 5 ca. 5 km medially; basal pinnae up to 23 cm long, short-stalked, the past Coipomo, ca. 20 km S of Ancud, 41°52’W, 73°50’W, 80–100 m, 15 stalk up to 1.5 cm long, slightly to strongly inequilateral with Jan 1981, Sobel & Strudwick 2738 (NY); Valdivia, Corral, Quebrada La the basal basiscopic pinnules more elongate, the most basal ′ ′ Aguada, [39°48 50”S, 73°14 45”W], 10 m, 3 Feb 1979, Solomon & Solomon basiscopic pinnules (to 15 cm long) 1.4–2.3 times the length 4559 (MO); Osorno, Puyehue, [40°34’S, 73°09’W], 15 Jan 1947, Sparre 2131 (S, US); Osorno, Puyehue, [40°34’S, 73°09’W], 15 Jan 1947, Wall s. n. (NY); of the most basal acroscopic ones, the pinnules on the acro- Osorno, Puyehue, [40°34’S, 73°09’W], s. d., Wall & Sparre 9 (S). Los Rios: scopic side not or only slightly reduced toward the base of the Southern Corral, [39°48′50”S, 73°14′45”W], 15 Feb 1907, anon. s. n. (S); pinnae; pinnules sessile, the basal segments frequently over- ′ ′ [Valdivia], [39°48 50”S, 73°14 45”W], s. d., Bridges 806 (BM, K); Valdivia lapping the pinna rachis; rachises and pinna rachises puber- [illeg.], [39°48′50”S, 73°14′45”W], 15 Feb 1904, Buchtien s. n. (US); Valdivia, Casilla, [39°48′50”S, 73°14′45”W], 1904, Buchtien s. n. (US); Valdivia, ulent to densely pubescent on both surfaces, nonglandular, Panguipulli, 38°48’S, 71°57’W, 200 m, July 1926, Hollerymayer 1933 (MO, the hairs erect, 0.2–0.5 mm long, to 5-celled, sparsely scaly, US, S); Valdivia, [39°48′50”S, 73°14′45”W], 1854, Lechler 501 (BM, K, P, S, the scales up to 4.0 × 1.0 mm, lanceolate, entire, flat (nonbul- UPS); Valdivia, [39°48′50”S, 73°14′45”W], s. d., Philippi s. n. (K, P). late), shiny dark brown with clearer margins; costules abax- Unknown: s. d., anon. s. n. (F); Jan 1919, Claude-Joseph 884 (US); Jan ially pubescent, non-glandular, the hairs erect to spreading, 1921, Claude-Joseph 2660 (US); [illeg.], Dec 1928, Claude-Joseph 5853 (US); 3 Jun 1897, Dusen s. n. (S); s. d., Gay s. n. (P); Southern Chile, s. d., Philippi s. 0.2–0.5 mm long, to 5-celled, sparsely scaly, the scales simi- n. (K); 1 Feb 1940, Schwabe 6 (NY). lar to those of the pinna rachises but shorter, 1.5 × 0.5 mm, Discussion— As originally described, the name P. specta- adaxially puberulent to pubescent, nonglandular, the hairs bile Kaulf. was applied to plants from throughout the west- erect to somewhat antrorse, 0.2–0.5 mm long; laminar tissue ern hemisphere, with syntypes that included two other between veins glabrous on both surfaces; veins 3–6 pairs per 474 SYSTEMATIC BOTANY [Volume 35 pinnule lobe, visible on both surfaces, abaxially pubescent publications of the Accademia. Harutu Fukuda prepared the illustrations. and sparsely glandular, the hairs erect, 0.2–0.5 mm long, usu- Alejandra Vasco assisted with the preparation of the manuscript. This study was funded by a grant to Moran from the United States National ally 2–5-celled, the glands spherical, clear yellowish, shiny, Science Foundation (DEB 0717056). short-stalked, ca. 0.1 mm diam., 1–2-celled, adaxially glabrous; lamina margins slightly recurved, sparsely ciliate, the cilia 0.3 mm long, 2- or 3-celled, fugacious indusia present, Literature Cited inconspicuous, 0.1–0.3 mm wide, provided with ca. 0.1 mm Christensen, C. 1913 . A monograph of the genus Dryopteris , Part I. The long sessile glands, and sometimes with 1–4 ca. 0.1 mm long tropical American pinnatifid-bipinnatifid species. Kongelige Danske setae. Figures 1F–L , 2 , 3C–D . Videnskabernes Sselskabs Skrifter. Naturvidenskabelige og Mathematiske Etymology— From ‘TAAF’, the French acronym for the Afdeling, ser. 7 , 10 : 55 – 282 . Christensen, C. 1920 . A monograph of the genus Dryopteris , Part II. The Territory of the French Southern and Antarctic Lands (French: tropical American bipinnate-decompound species. Kongelige Danske ‘Terres Australes et Antarctiques Françaises’). Amsterdam Videnskabernes Sselskabs Skrifter. Naturvidenskabelige og Mathematiske and Saint-Paul Islands, where the species is endemic, belong Afdeling, ser. 8 , 6 : 3 – 132 . to this Territory. Christensen, C. 1931 . Asiatic pteridophyta collected by Joseph F. Rock. Distribution and Ecology— Amsterdam and St-Paul Contributions from the United States National Herbarium 26: 265 – 337 . Christensen, C . 1940 . The pteridophytes of Tristan da Cunha. In Results Islands, South Indian Ocean. Terrestrial in gullies, in shade of of the Norwegian scientific expedition to Tristan da Cunha 1937–1938 , ed. lava crevices and caves; observed along river and waterfall. E. Christophersen , 6: 1 – 25 . Plants isolated, rare, 50–350 m. Du Petit-Thouars, L.-M. A . 1808 . Esquisse de la flore de l’Isle de Tristan Additional Specimens Examined— TERRES AUSTRALES ET d’Acugna . Paris . ANTARCTIQUES FRANÇAISES. Amsterdam Island: Caverne des Hemsley, W. B . 1885 . Report on the botany of the Bermudas and various Venus(?) de Verre(?), [37°50’S, 77°30’E], 20 Mar 1960, Aubert s. n. (P); other islands of the Atlantic and southern oceans, 2 nd part. Report [37°50’S, 77°30’E], 20 Mar 1960, Aubert s. n. (P); Au bord d’une cavité, près on the Scientific Results of the Voyage of H. M. S. Challenger during the du camp, [37°50’S, 77°30’E], 31 Mar 1958, Cour s. n. (P); Au dessus du bois years 1873–76: under the command of Captain George S. Nares and the late de Phylica , direction le Fourneau, [37°50’S, 77°30’E], 200–300 m, 27 Dec Captain Frank Tourle Thomson. Prepared under the superintendence of Sir 1984, Jolinon 891 (P); Au dessus du bois de Phylica , direction le Fourneau, C. Wyville Thomson and John Murray. Botany . London . [37°50’S, 77°30’E], 200–300 m, 27 Dec 1984, Jolinon 892 (P); Entrecasteaux, Holttum, R. E. 1969 . Studies in the family Thelypteridaceae. The genera rivière à 700 m de la baraque, [37°50’S, 77°30’E], 50 m, 3 Jan 1985, Jolinon Phegopteris, Pseudophegopteris , and Macrothelypteris . Blumea 17: 1 – 32 . 933 (P); Le Grand Tunnel, coulée, [37°50’S, 77°30’E], 110 m, 8 Jan 1985, Holttum, R. E. 1974 . Thelypteridaceae of Africa and adjacent islands . Jolinon 938 (P); Versant des Taureaux Sauvages, [37°50’S, 77°30’E], 332 m, Journal of South African Botany 40: 123 – 168 . 23 Jan 1985, Jolinon 1016 (P); Del Cano, dans la Ravine le long de la cas- Manton, I. and G. Vida . 1968 . Cytology of the fern flora of Tristan da cade, [37°50’S, 77°30’E], 160 m, 27 Jan 1985, Jolinon 1068 (P); Deal Cano, Cunha . Proceedings of the Royal Society of London. Series B. Biological dans la Ravine le long de la cascade, [37°50’S, 77°30’E], 160 m, 27 Jan 1985, Sciences 170: 361 – 379 . Jolinon 1071 (P); De la coulée Heurtin au Faux Sommet, [37°50’S, 77°30’E], Marticorena, C. and R. Rodríguez (eds.). 1995 . Flora de Chile 1. Universidad 250 m, 4 Feb 1985, Jolinon 1101 (P); Mêmes endroits que les deux numéros de Concepción : Concepción . précédents, [37°50’S, 77°30’E], 23 Dec 1974, L’Isle 28 (P); [37°50’S, 77°30’E], McNeill, J., F. R. Barrie , H. M. Burdet , V. Demoulin , D. L. Hawksworth , 12 Dec 1963, Lourteig & Cour 50 (P); Vallée entre les 2 Venus, [37°50’S, K. Marhold , D. H. Nicolson , J. Prado , P. C. Silva , J. E. Skog , J. H. Wiersema , 77°30’E], 350 m, 16 Dec 1963, Lourteig & Cour 78 (P); [37°50’S, 77°30’E], and N. J. Turland . 2006 . International code of botanical nomenclature (Vienna 1 Jan 1962, Noël 29 (P). Saint-Paul Island: [38°43’S, 77°31’E], 11 Nov 1974, Code). Regnum Vegetabile 146. Ruggell : A. R. G. Gantner Verlag. L’Isle 3 (P). Mickel, J. T. and L. Atehortúa . 1980 . Subdivision of the genus Elaphoglossum . Discussion— Given some known mistakes in the labels by American Fern Journal 70: 47 – 68 . Moran, R. C. 1991 . Monograph of the neotropical fern genus Stigmatopteris . L’Isle, Tardieu-Blot (1954) postulated that this taxon, treated Annals of the Missouri Botanical Garden 78: 857 – 914 . by her as Dryopteris aquilina (Thouars) C. Chr., had been Moran, R. C., J. Prado , and P. H. Labiak . 2009a . Megalastrum (Dryopteridaceae) erroneously recorded from Amsterdam. Several recent and in Brazil, Paraguay, and Uruguay. American Fern Journal 99: 1 – 44 . annotated collections attest the presence of this taxon on Moran, R. C., J. Prado , and P. H. Labiak . 2009b . Megalastrum (Dryopteri- Amsterdam. The presence of this species on St. Paul however daceae) in the West Indies . Brittonia 61: 273 – 292 . Röpert, D . (ed.). 2000 - (continuously updated): Digital specimen images at is still based on a single collection ( L’Isle 3). Until more recent the Herbarium Berolinense. - Published on the Internet http://ww2. collections can be made, some doubt remains as to the pres- bgbm.org/herbarium/default.cfm [accessed 12 Feb 2009]. ence of the taxon on that Island. Rouhan, G., J.-Y. Dubuisson , F. Rakotondrainibe , T. J. Motley , J. T. Mickel , Megalastrum taafense resembles M. aquilinum from Tristan J.-N. Labat , and R. C. Moran . 2004 . Molecular phylogeny of the fern genus Elaphoglossum (Elaphoglossaceae) based on chloroplast non- da Cunha in the cutting of the laminae, revolute margins, and coding DNA sequences: contributions of species from the Indian fugacious indusia. However, M. taafense differs by having Ocean area . Molecular Phylogenetics and Evolution 33: 745 – 763 . shorter hairs on the abaxial costules, and glabrous lamina tis- Smith, A. R. and R. C. Moran . 1987 . New combinations in Megalastrum sue between the veins abaxially. It also lacks glands adaxially, (Dryopteridaceae) . American Fern Journal 77: 124 – 130 . whereas M. aquilinum sometimes has sessile spherical glands Stafleu, F. A. and R. S. Cowan . 1976 . Taxonomic Literature, a selective guide to botanical publications and collections with dates, commentaries and types. near the adaxial costae. Vol. I, A–G . 2 nd ed . Utrecht : Bohn, Schletema & Holkem . The brown, nonbullate scales of the abaxial surfaces of the Tardieu-Blot, M.-L. 1954 . Sur les Fougères récoltées par Aubert de la Rüe laminae, and the laminar surfaces glabrous between veins aux îles Kerguelen et Amsterdam. Mémoires de l’Institut Scientifique de on both surfaces easily distinguish Megalastrum taafense from Madagascar. Série B. Biologie Végetale 5: 59 – 64 . Tryon, A. F. 1966 . Origin of the fern flora of Tristan da Cunha . British Fern the three geographically closest tropical taxa: M. lanuginosum Gazette 9: 269 – 276 . (Africa, Madagascar, Mascarene Islands, Comoros), M. canacae Tryon, A. F. and B. Lugardon . 1990 . Spores of the Pteridophyta . New York : (Mauritius and La Réunion), and M. lanatum (La Réunion). Springer-Verlag . Vasco, A., R. C. Moran , and G. Rouhan . 2009 . Circumscription and phy- Acknowledgments . We would like to thank the curators of the fol- logeny of the Elaphoglossum ciliatum group (E. sect. Lepidoglossa , Dry- lowing herbaria for providing loans of material: B, BM, F, GH, K, MICH, opteridaceae) based on cpDNA sequences. Taxon 58: 825 – 834 . NY, MO, NBG, O, OSU, P, S, UC, UPS, US, WU. We would like to thank Wace, N. M. 1960 . Botany of the southern oceanic islands . Proceedings of the Alan Smith for discussions regarding Pseudophegopteris . Roy Gereau Royal Society of London. Series B. Biological Sciences 152: 475 – 489 . assisted with Latin diagnoses. We are thankful to Elena Borgi from the Wace, N. M. 1961 . The vegetation of Gough Island . Ecological Monographs Accademia delle Scienze, Torino, for providing us with information about 31: 337 – 367 . 2010] SUNDUE ET AL.: MEGALASTRUM IN CIRCUMAUSTRAL REGION 475

Wace, N. M. and J. H. Dickson . 1965 . The terrestrial botany of the Tristan numbers in bold type indicate type specimens. Megalastrum aquilinum da Cunha islands In Report of the Royal Society Expedition to Tristan (1), Megalastrum glabrius (2), Megalastrum inaequalifolium (3), Megalastrum da Cunha, 1962 . Philosophical Transactions of the Royal Society of London. masafuerae (4), Megalastrum peregrinum (5), Megalastrum spectabile (6), Series B , 249: 273 – 360 . Megalastrum taafense (7). Aubert s. n. (7). Bertero 1550 (3); 1660 (3). Wace, N. M. and M. W. Holdgate . 1958 . The vegetation of Tristan da Bridges, T. 806 (6). Cabrera, A. L. 5037 (6). Castellanos 114252 (6). L. A. Cunha . Journal of Ecology 46: 593 – 620 . von Chamisso s. n. (6). Christophersen, E. 23, 423, 1061, 1093, 2217, 2232, Williams, R. L. 2003 . French botany in the enlightenment: the ill-fated voyages 2545 (1). Claude-Joseph 884, 885, 2660, 3046, 5853 (6). Cooper, J. G89053 of La Pérouse and his rescuers . Boston : Kluwer Academic Publishers . (5). Cour s. n. (7). de la Sota, R.E 2799, 6011, 6028 (6). du Petit-Thouars, L.-M. A. A. s. n. (1). Dyer, R. A. 3554 (1). Files P., C. 574 (6). Gaudichaud- Appendix 1. Synopsis of distributions. Baupre, C. 34 (6). Gunckel, H. 2467 (6). Gunther, E. 13 (6). Hänel, C. CH 021 (5). Hollermayer, P. A. 693, 1933, 156c (6). Hunnewell, F. W. 15852 (6). South America, Chile/Argentina: M. spectabile. South Pacific Oceanic Jolinon, J.-C. 891, 892, 933, 938, 1016, 1068, 1071, 1101 (7). Junge, E. 2924, Islands, Masatierra: M. inaequalifolium, Masafuera: M. glabrius, M. masa- 84 (107) (6). Keytel, P. 4713 (1). Krause s. n. (6). Lechler, W. 501 (6). L’Isle, fuerae. South Atlantic Oceanic Islands, Tristan da Cunha: M. aquilinum , M. G. 3, 28 (7). Looser, G. 5062 (6). Lourteig, A. 78, 50 (7). Mac Gillivray, J. Gough Island: M. aquilinum, M. peregrinum , Inaccessible Island: M. aqui- 327 (1). Marticorena, C. 9011, 9026, 9037, 9082, 9153 (3). Mejland, Y. 1575 linum , Nightingale Island: M. aquilinum. South Indian Ocean Islands, Île (1). Meyer, F. G. 9499, 9661 (3). Miller 119 (3). Morisson, J. L. 17326 (3). Amsterdam: M. taafense , Île St. Paul: M. taafense. Noël, P. 29 (7). Poeppig, E. 45 (6). Ricci, M. 402 (3). Roux, J. P. 716 (5); 2092, 2147, 2193 (1). Savatier, L. 272 (6). Schwabe, G. H. 6 (6). Shannon, R. C. 31 (6). Skottsberg, C 12, 65, 168, 274, 275, 339 (3); 466 (2); 448 (4); 721, 730 (3); Appendix 2. Index to Collections. 731 (2); 921 (6); 1151, 731 bis (3). Sobel, G. 2738 (6). Solbrig, O. T. 3676 (4). Solomon, J. 4559 (6). Sparre, B. 75 (3); 2131, 2948 (6). Stuessy, T. F. 5014, The numbers in parentheses refer to the corresponding species. 11644, 11963, 12055, 12129, 15396 (3). Wace, N. M. 12 (1); 133 (5). Wall, E. 9, Specimens are listed in alphabetical order by first collector. Collection 3240 (6). Collector unknown s. n. (6).