The Family Cladoniaceae (Lecanorales) in the Galapagos Islands

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http://dx.doi.org/10.11646/phytotaxa.129.1.1

The Family Cladoniaceae (Lecanorales) in the Galapagos Islands

A. YÁNEZ-AYABACA1,2, T. AHTI3 & F. BUNGARTZ1* 1Biodiversity Assessment, Charles Darwin Foundation (AISBL), Puerto Ayora, Santa Cruz, Galapagos, Ecuador, *Corresponding author: F. Bungartz, phone: +593-5 2526146/47 ext. 218, fax: +593-5 2527013 ext. 103, email: [email protected] 2Universidad Central del Ecuador, Quito, Ecuador, email: [email protected] 3Botanical Museum, Finnish Museum of Natural History, P.O. Box 7, FI-00014 University of Helsinki, Finland, email: [email protected]

Abstract

As part of an ongoing comprehensive inventory of the Galapagos lichen flora, all species in the Cladoniaceae from the archipelago have been revised using both historic and recent collections. A total of twenty-six species is reported here, one species of Cladia and twenty-five Cladonia species. One species, Cladonia bungartzii, is described as new to science; seven are records new to Ecuador and the Galapagos: Cladonia corymbosula, C. polyscypha, C. pulverulenta, C. pyxidata, C. aff. sphacelata, and C. strepsilis. Four species have previously been reported from Ecuador, but are new to Galapagos: C. cartilaginea, C. chlorophaea, C. dactylota, and C. grayi. Eight species previously reported cannot be confirmed here. Detailed descriptions are presented for all species. They include diagnostic characteristics to distinguish similar species. An identification key to all Galapagos Cladoniaceae is provided. A brief discussion highlights the importance of baseline inventories and uses the Galapagos Cladoniaceae as a case study to discuss important aspects of lichen biogeography in Galapagos.

Key words: Census of Galapagos Biodiversity, Galapagos Lichen Inventory, taxonomy, Cladonia, Cladia, identification key, Ecuador, South America, Cladonia bungartzii sp. nov.

Introduction

The Cladoniaceae are a large family of about 500 species of lichen forming fungi world-wide. Most are relatively large and conspicuous lichens, some even dominate vegetation communities such as the extensive mats of reindeer lichens in the arctic tundras. In the tropics they can also become quite abundant, several species are conspicuous elements for example of the Andean páramo (Ahti 2000). In Galapagos, most species of the Cladoniaceae are generally more common if not restricted to the humid highlands; here Cladonia mats develop on relatively young lava flows with little soil development, but exposed to high humidity, the typical mist of the garúa season. Many species also grow on old logs, rotting wood, often at the basis of trees and as epiphytes also among the dense mats of liverworts and bryophytes. Relatively few species are found in the transition zone or even the dry Galapagos lowlands. Galapagos lichens are today relatively well known. The first inventories were summarized in the 1960's (Weber 1966, Weber & Gradstein 1984, Weber et al. 1977), culminating in a preliminary list of 196 species; followed by a few updates (Elix & McCarthy 1998, Weber 1993). In 2006 the Charles Darwin foundation started the first comprehensive inventory of non-vascular plants and plant-like organisms and during consecutive visits to now 14 different islands almost 14,000 lichen specimens have been collected, all deposited in the herbarium of the Charles Darwin Research Station (CDS). Since 2005 several publications have appeared, treating many larger species groups as well as some otherwise interesting and spectacular reports and many new species were described (Aptroot & Bungartz 2007, Aptroot & Sparrius 2009, Aptroot et al. 2008, Bungartz 2008, Bungartz et al. 2008, 2010, Tehler et al. 2009).

Accepted by Thorsten Lumbsch: 12 July 2013; published: 5 Sept. 2013 1 As a result of this field work and our taxonomic revisions of the material collected, it is now obvious that at least twice as many lichen species occur throughout the archipelago as previously listed in the most recent update of Weber’s original checklist (Elix & McCarthy 1998). Nevertheless, we are still far from understanding the ecology and distribution of Galapagos lichens. They are generally part of lesser known groups of species of the archipelago and despite improved knowledge about their occurrence, little remains known about their general biogeography, i.e., where these species originated and how they reached these isolated islands. Little is also known about the ecological role and habitat preferences of individual species. Galapagos is one of the very few tropical archipelagos, where scientists have suggested that much of its original biodiversity remains intact (Snell et al., 2002). Species inventories are therefore particularly relevant as a necessary pre-requisite to objectively assess biodiversity and thus overcome taxonomic bias otherwise inherent to conservation management (Bungartz et al. 2012, Clark 2002, Dunn 2005, Dunn et al. 2009, Régnier 2009). In conservation biology species inventories thus principally address two objectives: (1) they establish a baseline for species identification and thus the necessary framework for ecological studies, and (2) they provide biogeographic context to better understand how species are distributed, which ones are rare and restricted, and which ones common and widely distributed. While the first objective, a better baseline for species identification, is addressed with this publication by providing an identification key and short diagnostic descriptions specifically focusing on the Galapagos Cladoniaceae. The second aspect, however, remains much more challenging. Despite an extensive monograph on Neotropical Cladoniaceae (Ahti 2000), detailed knowledge on species distribution in this group remains fragmentary. Although only one species is described here as new and presumably endemic, another seven species are reported for the first time from Ecuador (though not confirmed from the mainland), and eight previous reports must be rejected for the archipelago. It is obvious that our understanding of the general distribution in this group thus remains insufficient to speculate, how species of Cladoniaceae reached the Galapagos. It is also not possible, at this stage, to objectively assess species rarity of lichens throughout South America. This highlights the urgency to extended similar studies to the continent. Nevertheless, this inventory of Galapagos Cladoniaceae can be discussed in the context of Galapagos biogeography, where several important trends about Galapagos lichen diversity, and in particular species endemism, emerge.

Material and Methods

The Galapagos Archipelago comprises more than oceanic 123 islands, islets and large rocks that emerged from the sea as a result of volcanic hot spot activity. Fourteen islands are somewhat arbitrarily recognized because of their size as the principal islands. As island groups they are typically associated with numerous smaller islands, rocks and islets within close proximity (Snell et al. 1996). The climate of Galapagos is unusually dry and principally dominated by the sea currents, with a hot and cool season and prevailing winds from the south and southeast. Largely as a result of climate, prevailing winds and elevation, five major vegetation zones can be distinguished: coastal, dry, transition, humid, and high altitude dry zone (Bungartz et al. 2010, Tye et al. 2002). On the southern side of the islands the humid zone is typically much more extensive, whereas it is absent from low islands and typically not well developed on the leeward sides of higher islands. Only on the highest volcanoes of Isabela Island the humid zone forms a transition into a high altitude dry zone above the cloud inversion layer (see, Trueman & d'Ozouville 2010, fig. 1). As part of the Galapagos Lichen Inventory the following islands have been visited, and all vegetation zones with their principal lichen habitats have been surveyed: Isabela (Volcán Sierra Negra, Volcán Alcedo, Volcán Darwin), Santiago (incl. Rábida, Bartolomé), Santa Cruz (incl. Santa Fé, Plaza Sur, Plaza Norte, Roca Gordon, Pinzón), Pinta, Española, Floreana, and San Cristóbal. Herbarium collections of this inventory are deposited at CDS; specimens from many historic collections have also been examined (B, CAS, COLO, FH, H, S). All specimens were examined with a Zeiss Stemi DV4 dissecting microscope and a Zeiss Imager A1 compound microscope equipped with differential interference contrast. Macrophotos were taken with a Nikon D300 or D7000, 62 mm Nikkor Micro Lens and R1C1 macro

2 • Phytotaxa 129 (1) © 2013 Magnolia Press YÁNEZ-AYABACA ET AL. flash directly in the field, or using a Novoflex macro-table to take images of herbarium specimens; for photographic magnifications higher than 1:1 an extension tube or Novoflex bellows was used. For microphotos the compound microscope is equipped with a phototube for the Nikon D300. Photos in the laboratory were taken with Nikon Camera Control Pro 2; all photos are databased with the program IDimager 5 using the Darwin Core XML schema to embed collection and identification information as XMP metadata (http://owl.phy.queensu.ca/~phil/exiftool/TagNames/DarwinCore.html). Photos were processed with Photoshop CS4. Secondary metabolites were examined from a selection of specimens using standardized thin-layer chromatography (Orange et al. 2001, 2010). Instead of the conventional upright TLC tanks a horizontal HPTLC developmental chamber was used as described by Arup et al. (1993). TLC plates were interpreted with the computer program WINTABOLITES (Mietzsch et al. 1994), and photographed for permanent record (Egan 2001). Due to the large amount of specimens examined, collection data for only few representative examples are included here. Where available, at least one specimen per surveyed island (and, in the case of Isabela, the island’s different main volcanoes) has been listed. Due to better accessibility to the specimens, further emphasis is given to collections not deposited at CDS. Detailed collection information of all Galapagos specimens used in this study can be downloaded from the CDF Collections Database online at http:// www.darwinfoundation.org/datazone/collections/.

Results

A total of twenty five species of Cladonia and one species of Cladia are reported here for the Galapagos Islands. Six species constitute new records for Ecuador, C. corymbosula, C. pyxidata, C. polyscypha, C. pulverulenta, C. aff. sphacelata, and C. strepsilis. Four species represent new records for Galapagos, C. cartilaginea, C. chlorophaea, C. dactylota, and C. grayi. One species, Cladonia bungartzii, is described here as new.

Key to species of the Cladoniaceae in the Galapagos Islands

1. Primary thallus granulose, evanescent, never squamulose; podetia generally 4–12 cm tall, always densely branched, typically forming cushions or mats (“reindeer lichens”), always ascyphose, lacking soredia, granules or microsquamules ...... 2 - Primary thallus squamulose, generally persistent, rarely evanescent; podetia present or not, if present, usually 1–3 (– 5) cm tall, simple or branched, scyphose or blunt; commonly sorediate, granulose or microsquamulose...... 5 2. Main thallus corticate, not originating from ascocarpous hyphae (pseudopodetia), with ellipsoid perforations, deep brown or pale yellowish brown, 4–5 cm tall...... Cladia aggregata - Main thallus ecorticate or with discontinuous cortex, originating from ascocarpous hyphae (true podetia), lacking perforations, whitish gray or yellowish gray, not distinctly brown, typically > 5 cm tall ...... 3 3. P+ orange red (fumarprotocetraric acid); podetia apically branched, with brownish to blackening necrotic tips; pycnidia with red jelly ...... C. arcuata - P– (fumarprotocetraric acid absent); podetia apically branched, but tips ±concolorous with the podetia; pycnidial hyaline jelly never red ...... 4 4. Principal axes clearly differentiated because of their anisotomic branching pattern; ramifications dichotomous, rarely trichotomous, generally sparsely branched ...... C. arbuscula subsp. boliviana - Principal axes indistinct because of their isotomic branching pattern; ramification principally trichotomous, very rarely also dichotomous, densely branched ...... C. confusa a. Podetia yellowish to greenish gray, with usnic acid...... C. confusa f. confusa b. Podetia ash gray with brownish gray tips, lacking usnic acid ...... C. confusa f. bicolor 5. Podetia absent; primary thallus dominant ...... 6 - Podetia present; primary thallus persistent, but not dominant ...... 8 6. Medulla C+ green (strepsilin) ...... C. strepsilis - Medulla C− (strepsilin absent) ...... 7 7. Squamules thick, short, laciniate; surface rugulose, often cracked, typically epruinose, rarely pruinose; lower side not cottony, always lacking soredia ...... C. corymbosula

THE FAMILY CLADONIACEAE IN GALAPAGOS Phytotaxa 129 (1) © 2013 Magnolia Press • 3 - Squamules thin, coralloid, elongate to laciniate; surface smooth, not cracked, pruinose; lower side cottony, coarsely granular-sorediate ...... C. nana 8. Apothecia red, pycnidia containing red jelly ...... 9 - Apothecia brown, pycnidia containing hyaline jelly ...... 11 9. Stereomes formed of loosely intertwined hyphae, often with longitudinal cracks, terete to ± flattened; surface mostly corticate, with few decorticated areas, white to pale yellow ...... C. bungartzii - Stereomes of compact, dense hyphae, typically remaining intact, mostly terete; surface ecorticate, greenish gray or pale yellow to brown...... 10 10. Podetia always with dark brown to blackened, necrotic base; densely covered in microsquamules and corticate granules, lacking true soredia ...... C. didyma - Podetia in parts sometimes brownish, but never becoming melanotic (blackened); coarsely sorediate, with scarce microsquamules, corticate granules absent...... C. macilenta 11. Podetia terminating in cups...... 12 - Podetia blunt or subulate, not forming cups ...... 23 12. Podetia occasionally terminating in narrow, open cups (funnels; resembling scyphi, but open inside and always lacking a bottom); typically present only in well developed podetia ...... C. scholanderi - Podetia with narrow to broadened cups that are always closed inside (true scyphi) ...... 13 13. Podetia growing into short, stout, broad cups, not branched, but sometimes proliferating from the rim of the cup and then forming tiers ...... 14 - Podetia elongate stalks, typically terminating in narrow cups, but a few podetia also with blunt tips; podetia often ramified, occasionally as proliferations from the rim of the cups, but not forming distinct tiers...... 17 14. Medulla UV+ whitish blue (grayanic acid); surface, where corticate, verruculose-granular, always microsquamulose, typically lacking soredia or, if soredia present, only forming inside the scyphus ...... C. grayi - Medulla UV– (grayanic acid absent); surface, where corticate, smooth, sometimes flaking of as microsquamules or developing into granules or soredia ...... 15 15. Podetia lacking true soredia but with abundant, corticate granules; surface aereolate-corticate, basally often peeling off and forming schizidia ...... C. pyxidata - Podetia with true, ecorticate soredia, sometimes intermixed with corticate granules; surface mostly ecorticate, sometimes smoothly corticate at the base, but not peeling off, not forming schizidia ...... 16 16. Podetia granular sorediate; soredia of ±uniform size, very rarely intermixed with few, scarce microsquamules; scyphi predominantly simple, rarely proliferating ...... C. chlorophaea - Podetia mostly farinose sorediate, but intermixed with abundant granules and microsquamules; scyphi commonly proliferating ...... C. subsquamosa 17. Podetia with tuberculate soralia; P+ golden yellow (psoromic acid) ...... C. dactylota - Soralia not distinctly delimited; P+ orange or red (fumarprotocetraric acid) or yellow (thamnolic acid), but not golden yellow (never with psoromic acid)...... 18 18. Primary thallus dominant, with large, broad, recurved, rhiziniate squamules; podetia small, densely covered with pin-shaped microsquamules ...... C. ceratophylla Primary thallus not dominant, of small, erhiziniate squamules; podetia lacking pin-shaped microsquamules ...... 19 19. Podetia irregularly corticate throughout; cortex patchy and flaking off (schizidiate), becoming densely sorediate; scyphi, when well developed, very irregular and typically strongly ramified (irregular proliferations) ...... C. aff. ramulosa - Podetia almost entirely lacking a cortex or cortex only covering the lower half to two thirds of the length of the stalk; scyphi not strongly ramified ...... 20 20. Podetia disintegrating and dying of at their base (typically brown to blackened, i.e., necrotic), further up the stalk with an intact cortex ...... 21 - Podetia rarely discolored, occasionally with brown areas, but not blackened, smoothly corticate only at their base, or almost entirely ecorticate ...... 22 21. No more than the lower half of the podetium corticate; ecorticate surface moderately granulose and microsquamulose; farinose soredia scarce; always lacking schizidia; tips always scyphose...... C. polyscypha - Up to two thirds of the podetium corticate; ecorticate surface densely covered by a mixture of soredia, granules, microsquamules and schizidia as a result of cortex disintegration; tips blunt, rarely scyphose ...... C. pulverulenta 22. Podetia whitish gray, rarely pale brown, ecorticate or inconspicuously corticate only at the very base, typically densely covered with isidioid microsquamules, P+ red (fumarprotocetraric acid) ...... C. subradiata - Podetia ash gray or brown in parts, corticate at the base of the stalk and just below the scyphi, with scarce granules and microsquamules not isidioid, but developing true, ecorticate soredia; P+ yellow (thamnolic acid, no fumarprotocetraric acid)...... C. granulosa* (*not confirmed for Galapagos, included here for its similarity to C. subradiata and C. pulverulenta)

4 • Phytotaxa 129 (1) © 2013 Magnolia Press YÁNEZ-AYABACA ET AL. 23. Medulla C+ green (strepsilin), K–, P− (squamatic acid), primary thallus dominant, forming cushions, rarely also developing podetia (not yet observed in Galapagos material) ...... C. strepsilis

- Medulla C−, K+ yellow or brownish, P+ yellow or orange to red (thamnolic or fumarprotocetraric acid), always forming slender, well developed podetia, primary thallus inconspicuous ...... 24 24. Podetia conspicuously yellowish green, with usnic acid; of long, branched stalks with perforate axils; typically with at least a few, well developed podetia terminating in narrow, ± indistinct funnels, but rarely funnels absent; 4–14.5 cm tall ...... C. scholanderi - Podetia not conspicuously yellow, whitish green or greenish gray, lacking usnic acid; branched stalks with closed axils; always lacking funnels; typically less than 5 cm tall ...... 25 25. Podetia nearly always with closely aggregated “turban-like” apothecia; surface ecorticate, densely granular sorediate, lacking microsquamules (previously misidentified as C. peziziformis, which, however, has corticate, squamulose podetia) ...... C. corymbosula - Podetia always with blunt tips, rarely producing apothecia, single, not aggregated in “turban-like” conglomerates; surface ecorticate or in parts corticate, with or without soredia, granules or microsquamules ...... 26 26. Podetia covered in microsquamules that towards the tip develop into fine, corticate granules and towards the base merge with the larger primary squamules ...... C. corniculata - Podetia where a transition from basal squamules to microsquamules into granules cannot be observed...... 27 27. Podetia typically densely microsquamulose; occasionally with few, corticate granules, but always lacking soredia ...... 28 - Podetia with few microsquamules; always granulose to farinose sorediate...... 30 28. Podetia arising from inflated primary squamules (phyllopodiate); stalks typically unbranched to sparsely branched; tips not forming branchlets; surface granulose, but granules not developing into microsquamules; P+ red (fumarprotocetraric acid) ...... 29 - Podetia not arising from inflated primary squamules (not phyllopodiate); stalks moderately to repeatedly branched; tips divided in 2–6 branchlets; surface granulose, granules with age typically becoming larger and branching into microsquamules; P+ yellow (thamnolic acid) ...... C. aff. sphacelata 29. Primary thallus dominant, of ±erect, elongated, strap-shaped (laciniate) squamules, their lower side cottony, coarsely granular-sorediate; podetia rare, typically corticate ...... C. nana - Primary thallus not dominant, of ±erect, stout and broadened, esorediate squamules; podetia mostly ecorticate, if cortex present, restricted to its base ...... C. corymbites 30. Podetia completely ecorticate, moderately covered with a mixture of corticate granules and ecorticate soredia...... C. cartilaginea - Podetia mostly corticate or cortex restricted to the base of the podetium and below and inside the scyphi...... 31 31. Cortex covering up to two thirds of the podetium, otherwise ecorticate and densely covered by a mixture of soredia farinose, granules and schizidia; P+ orange-red (fumarprotocetraric acid) ...... C. pulverulenta - Cortex restricted to the base of the podetium and below and inside the scyphi, otherwise ecorticate and only moderately sorediate, but lacking granules and schizidia; P+yellow (thamnolic acid) ...... C. granulosa* (*not confirmed for Galapagos, included here for its similarity to C. subradiata and C. pulverulenta)

Cladia aggregata (Sw.) Nyl. (1870: 167) (Figs. 1a–b)

Primary thallus not seen, evanescent, of lobulate-papillate squamules (according Ahti, 2000); pseudopodetia erect, forming cushions or dense mats, dark brown or pale yellowish, 4–4.5 cm tall, hollow, terete to ± flattened or angulate, some parts, when older, densely branched; branching anisotomic, dichotomous; surface corticate, glossy or matt; usually abundantly perforate; perforations frequently ellipsoid; pycnidia with hyaline jelly; apothecia not seen. Spot tests and chemistry: P−, K− or + yellow, C−, KC−, UV−; barbatic and 4-O-demethylbarbatic acid (chemotype I). Distribution and ecology: Currently know only from the humid zone in the highlands of Isabela and Santa Cruz; on bare, sunny, exposed soil, often along trails and in reindeer lichen heaths. Notes: The cushions or dense mats of this species may at first be mistaken for a reindeer lichen but, the genus Cladia is easily recognized because it has completely corticate pseudopodetia with abundant ellipsoid perforations.

THE FAMILY CLADONIACEAE IN GALAPAGOS Phytotaxa 129 (1) © 2013 Magnolia Press • 5 According to Ahti (2000) the fertile pseudopodetia typically represent the thickest branches. Fertile material was, however, not encountered among the Galapagos specimens and this observation cannot be confirmed here. Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Cerro Azul, S-slope above Iguana Cove, crater on SE-slope, 800 m, humid zone, 22 June 1976, Sipman, H.J.M. L-58 (L, COLO 297902). Santa Cruz Island, SE of El Puntudo, W of Mt. Crocker, 1972, Weber L-55403 (DUKE, H, TUR), saddle between Mt. Crocker and El Puntudo, 750–800 m, 1976, Weber & Lanier, Lich. Exs. COLO 497 (DUKE, H, TUR, U, QCA, FH 197181, COLO 297764), NE-slope of El Puntudo, 0°38’39.10”S, 90°20’7.90”W, 813 m, humid zone, on open soil among Cladonia spp. 10 Aug 2008, Bungartz, F. 8151 (CDS 40797), along the trail from Media Luna to El Puntudo, at a small stream crossing the trail, 0°39’4”S, 90°20’5”W, 690 m, humid zone, bare ground along the footpath, 28 Jan 2006, Bungartz, F. 3971 (CDS 27901).

Cladonia arbuscula subsp. boliviana (Ahti) Ahti & DePriest (2001: 500) (Figs. 1c–d)

Primary thallus evanescent; podetia forming cushions, slender, whitish gray, up to 12 cm tall, sparsely branched; branching pattern anisotomic, dichotomous, rarely trichotomous, principal axes easily distinguishable; axils occasionally perforate; tips typically slightly brown; surface ecorticate, barely verruculose; algal clumps (glomerules) often observed on the surface of old podetia; pycnidia with hyaline jelly; apothecia brown. Spot tests and chemistry: P−, K−, C−, KC−, UV−; all Galapagos material examined contains only usnic acid (chemotype II of Ahti 2000). Distribution and ecology: Currently known only from a single collection from the humid zone of Isabela Island; forming lichen heaths in sunny, wind- and rain-exposed habitat among ferns (Pteridium arachnoideum, Pernettya howellii, Lycopodium sp.). Notes : The species is similar to C. confusa f. bicolor, but that species differs by thinner, more densely branched podetia and a ramification pattern where no principal axes can clearly be distinguished. The axil perforation observed in the Galapagos material of C. arbuscula subsp. boliviana is not mentioned by Ahti (2000). Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Alcedo, 1970, Prichard (H, LSU), Volcán Sierra Negra, close to the southern crater rim, along the trail to Alemania, 0°51’12.69”S, 91°8’40.5”W, 1055 m, humid zone, on soil,16 Aug. 2008, Bungartz, F. 8338 (CDS 40984).

Cladonia arcuata Ahti (1961: 73) (Figs. 1e–f)

Primary thallus evanescent; podetia forming cushions, mostly whitish gray but apical branches darkening, becoming distinctly brown towards their tips, 4−5 cm tall; branching pattern anisotomic, dichotomous, occasionally trichotomous, main axes distinguishable; axils close or rarely perforate; apical branches usually deflexed, tips sometimes melanotic; surface ecorticate, slightly verruculose; pycnidia with red jelly; apothecia with pale brown jelly. Spot tests and chemistry: P+ orange red, K− or + yellow, C−, KC−, UV−; fumarprotocetraric acid. Distribution and ecology: Known only from Isabela, Santa Cruz, and San Cristóbal Island; from the humid zone extending down into the upper transition zone; on the ground in open vegetation like sparse forests or open scrub, fern-sedge grasslands and reindeer lichen heaths; on soil and amongst plant debris and bryophytes, also on rock. Notes : Cladonia arcuata has a similar morphology to C. confusa f. bicolor, however that species does not have pycnidia with red jelly and always reacts P−, because it lacks fumarprotocetraric acid.

6 • Phytotaxa 129 (1) © 2013 Magnolia Press YÁNEZ-AYABACA ET AL. FIGURE 1. a–b Cladia aggregata (Bungartz 3971). a bifurcate apices of the pseudopodetia (scale 5 mm); b close-up with characteristic oval perforations (scale 3 mm). c–d Cladonia arbuscula subsp. boliviana (Bungartz 8338). c general thallus aspect (scale 5 mm); d podetial apices showing the predominantly anisotomic dichotomous branching pattern (scale 5 mm). e–f Cladonia arcuata (Bungartz 7495); e general thallus aspect (scale 3 mm); f podetial apices showing the predominantly isotomic dichotomous branching pattern (scale 2 mm).

THE FAMILY CLADONIACEAE IN GALAPAGOS Phytotaxa 129 (1) © 2013 Magnolia Press • 7 Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Cerro Azul, S slope above Iguana Cove, 750 m, humid zone, exposed rocky place in fern-sedge vegetation, 22 June 1976, Sipman, H.J.M. L-35 (COLO 297926); Volcán Darwin, southwestern slope, above Tagus Cove, 0°13’34”S, 91°19’21.4”W, 840 m, transition zone, on soil between plant debris, 13 Nov 2007, Bungartz, F. 7495 (CDS 37986); Volcán Sierra Negra, close to the southern crater rim, along the trail to Alemania, 0°51’12.69”S, 91°8’40.5”W, 1055 m, humid zone, on plant debris, 16 Aug 2008, Bungartz, F. 8341 (CDS 40987). San Cristóbal Island, Cerro San Joaquín, 0°53’49.5”S, 89°30’47.7”W, 691 m, humid zone, among mosses on the ground, 24 Aug 2008, Truong, C. 1512 (CDS 39823). Santa Cruz Island, 200 m, transition zone, 1 Apr 1964, Horneman, S. (COLO 316717).

Cladonia bungartzii Yánez-Ayabaca & Ahti sp. nov. MycoBank no. 804595 Diagnosis: A C. macilenta et C. didyma stereomate non compacte instructa differt. Holotype:—ECUADOR. GALAPAGOS: Pinta Island, on top of the highest point of the island, 0°35’3”N, 90°45’12”W, 625 m, humid zone, low and dense vegetation of ferns, grasses (Cyperus andersonii), and Lycopodium sp., open N- exposed, soil semi-shaded by ferns and Lycopodium sp; wind- and rain-exposed, on soil, 26 Feb 2007, Bungartz, F. 5744 (CDS 33396) (Figs. 2a–b)

Primary thallus subpersistent, thin, of short squamules with a corticate upper side and densely granular- sorediate lower side, epruinose; podetia common, stout, 0.5–2cm tall, composed of densely interwoven hyphae, but not compact, and frequently with longitudinal cracks, pale yellow, simple or slightly branched, mainly near the tips; surface mostly corticate-verruculose, sometimes with denuded areas where the cortex disintegrates and there farinose sorediate; lacking microsquamules; pycnidia not seen; apothecia often small, with bright red jelly. Spot tests and chemistry: P+ yellow, K+ yellow, C−, KC−, UV−; thamnolic and traces of didymic acid. Distribution and ecology: Known only from the humid zone of Pinta Island, where the species grows on soil of the fern-sedge grasslands (e.g., Cyperus andersonii, Lycopodium sp.). Notes : In Galapagos Islands, this species can be confused with C. macilenta, both species are densely sorediate, however C. macilenta has ecorticate podetia with few microsquamules. Cladonia didyma is another species that also has red apothecia, but it is different of C. bungartzii because its podetia are ecorticate covered with abundant microsquamules. Both C. macilenta and C. didyma have compact, dense stereomes that typically lack longitudinal cracks that are very characteristic of C. bungartzii. Additional specimen examined (paratype): ECUADOR. GALAPAGOS: Pinta Island, on top of the highest point of the island, 0°35’3”N, 90°45’12”W, 625 m, humid zone, on open soil, 26 Feb 2007, Bungartz, F. 5749 (CDS 33403).

Cladonia cartilaginea Müll. Arg. (1880: 260) (Fig. 2c)

Primary thallus evanescent or subpersistent, of laciniate squamules, esorediate, epruinose; podetia common, whitish gray, elongate, 0.5–1.5 (–2) cm tall, unbranched to slightly branched; axils closed; tips blunt, ascyphose; surface completely ecorticate; moderately covered with granules, ecorticate soredia and scarce microsquamules; macrosquamules confined to the base of the podetia; pycnidia with hyaline jelly; apothecia not seen. Spot tests and chemistry: P+ orange red, K−, C−, KC−, UV−; fumarprotocetraric acid. Distribution and ecology: A new record for Galapagos; currently known only from Santa Cruz Island. Previously reported from high mountains of Prov. Pichincha in Ecuador (Ahti 2000). In Galapagos apparently restricted to the humid zone, where the species has been found on a variety of subtrates: among bryophytes on the ground, as an epiphyte on native trees like Scalesia pedunculata or introduced trees like Cinchona pubescens, and even in the crevices of the windows of an old abandonned car.

8 • Phytotaxa 129 (1) © 2013 Magnolia Press YÁNEZ-AYABACA ET AL. Notes: The species is morphologically and chemically extremely similar to C. subradiata, but the podetia of C. subradiata are characterized by a surface densely covered of isidioid microsquamules and granulose soredia. Podetia of C. cartilaginea, in contrast, are overall only sparsely covered by granules interspersed occasionally by few microsquamules. Cladonia corniculata is also quite similar it shares the ecorticate podetia and same propagules, but unlike C. cartilaginea it is typically more abundantly branched, esorediate and shows a gradual transition from large, laciniate basal squamules to crowded, finely dissected microsquamules along the podetium that eventually become granulose towards the tip. Selected specimens examined: ECUADOR. GALAPAGOS: Santa Cruz Island, tras del Puntudo, ex finca de Don Benito, 0°38’23.18”S, 90°19’57.24”W, 732 m, sobre corteza, 28 Dec 2006, Nugra, F. 240 (CDS 33156), vicinity of Academy Bay, La Copa (= Media Luna), 15 Feb 1964, Weber, W.A. 426 (COLO 193443).

Cladonia ceratophylla (Sw.) Spreng. (1827: 271) (Fig. 2d)

Primary thallus persistent with large and broad, branching squamules, forming large laciniate apically recurved lobes that have a lower surface with scarce, brown, marginal rhizines; esorediate, epruinose; podetia common, greenish gray, short to elongate, 0.8–2 cm tall, unbranched or slightly branched; tips blunt or, when fertile, typically forming narrow scyphi; surface corticate at the base but soon becoming decorticated, sometimes with few, corticate patches remaining; generally lacking corticate granules, but typically densely covered with convex, elongate, unbranched microsquamules, very rarely also with few ecorticate soredia; pycnidia with hyaline jelly; apothecia with brown jelly. Spot tests and chemistry: P+ orange red, K− or K+ yellow, C−, KC−, UV−; fumarprotocetraric acid and traces of atranorin. Distribution and ecology: Known from Isabela, San Cristóbal, Santa Cruz, and Santiago Island; a common species in the humid zone, either growing directly on the ground in open fern-segde grassland, often amongst bryophytes, or sometimes as epiphyte on trunks, branches and even twigs of both on native and introduced tree species. Notes : The species can easily be recognized by its large and broad, rhizinate squamules and the very characteristic shape of its microsquamules, which are stout, elongate, convex lobules that remain unbranched and are thus somewhat pin-shaped. They do not emerge erect from the podetial surface but typically cover the podetial surface densely overlapping, almost like tiles. Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Alcedo, outer SE- exposed slope, ca. 100 m below the crater rim, 0°27’4”S, 91°5’50”W, 1066 m, humid zone, on soil, 6 Mar 2006, Aptroot, A. 64860 (CDS 31436), ca. 500 m below the crater rim; Volcán Cerro Azul, S-slope above Iguana Cove, 700 m, humid zone, on rock, 22 June 1976, Sipman, H.J.M. L-41 (COLO 297920), Sipman, H.J.M. L-57 (COLO 297903); Volcán Sierra Negra, along dirt road from Puerto Villamil to crater of Sierra Negra, farmland, 0°50’38”S, 91°3’45.5”W, 550 m, humid zone, on bark, 9 Sept 2007, Bungartz, F. 6862 (CDS 36311), South side of Sierra Negra crater, trail to Alemania, 0°50’57.5”S, 91°7’41.3”W, 1020 m, humid zone, on soil, 16 Aug 2008, Herrera-Campos, M.A. 10709 (CDS 40447), 0°50’0”S, 91°10’0”W, 800 m, humid zone, on dead wood, 18 Apr 1990, Sánchez-Pinto, L. 5040, 5048 (B 60 0173249, 60 0173257). San Cristóbal Island, bordering lake at El Junco, humid zone, steep slope bordering the lake, on rock, 21 May 1976, Lanier, J. (COLO 298436). Santa Cruz Island, between summit of Mt. Crocker and El Puntudo, 700 m, 1976, Weber & Lanier, Lich. Exs. COLO 502 (DUKE, H, M, TUR); vincinity Academy Bay, 1964, Weber L-40507 (COLO, M, UPS); saddle between Puntudo and Mt. Crocker, 0°38’41”S, 90°19’57”W, 720 m, humid zone, on soil, 18 Apr 1976, Weber, W.A. (CDS 10845); Bellavista, near parking place for trail to Media Luna, 0°40’10”S, 90°19’22”W, 400 m, humid zone, on soil, 27 May 2005, Aptroot, A. 63145 (CDS 29875), along trail from Media Luna to El Puntudo, 0°39’37.79”S, 90°20’0.9”W, 682 m, humid zone, on soil, 28 Dec 2005, Bungartz, F. 3274 (CDS 26913) at the base of the eastern slope below the summit of El Puntudo, 0°38’42”S, 90°20’14”W, 780 m, humid zone, on soil, 28 Feb 2006, Ziemmeck, F. 760 (CDS 27889), sobre corteza, 25 Jan

THE FAMILY CLADONIACEAE IN GALAPAGOS Phytotaxa 129 (1) © 2013 Magnolia Press • 9 2007, Nugra, F. 356 (CDS 35111), cerca la vía sector Los Gemelos, 0°37’33.39”S, 90°23’0.7”W, 611 m, zona húmeda, sobre corteza, 11 Oct 2006, Nugra, F. 146 (CDS 32800), trail between Bella Vista and Media Luna, 550 m, humid zone, on open place in the trail, 18 June 1976, Sipman, H.J.M. L-22 (OSC 53685), south slope of the mountain, 520 m, humid zone, on ground, 9 Apr 1930, Svenson, H.K. 202C (FH 197248, 197249), saddle between summit of Mount Crocker and El Puntudo, 700 m, humid zone, fern-sedge zone, on ground, 18 Apr 1976, Weber, W.A. (FH 197199).

Cladonia chlorophaea (Flörke ex Sommerf.) Spreng. (1827: 273) (Figs. 2e–f)

Primary thallus persistent, of small, compact squamules that easily erode and in some specimens remain only as a few corticate granules, rarely eroding into ecorticate soredia, epruinose; podetia common, grayish green, short; 0.5–1 (1.5) cm tall, always broadly scyphose and shaped like a cone, i.e., gradually narrowing towards their base; scyphi predominantly simple, very rarely proliferating; surface corticate only at the base, soon becoming completely decorticated; densely sorediate-granulose, the soredia generally of uniform size along the entire surface of the podetium; scyphus rarely with few corticate granules inside; old podetia commonly with large denuded areas lacking soredia; pycnidia hyaline; apothecia not seen. Spot tests and chemistry: P+ orange red, K−, C−, KC−, UV−; fumarprotocetraric acid. Distribution and ecology: Newly reported from the Galapagos Islands. Known from Floreana, Isabela, Pinzón, San Cristóbal, Santa Cruz, and Santiago Island; a common species in the humid and transition zone, rarely also in the dry zone; typically on the ground, often on plant debris or rotten wood. Notes : The species can be identified by its abundant, granulose soredia of a more or less uniform size across its podetial surface. Cladonia subsquamosa is very similar but generally has farinose soredia of less uniform size interspersed with granules and microsquamules. At first glance the species also resembles C. pyxidata, which, however, generally has broader scyphi with a corticate surface disintegrating into corticate granules, not forming ecorticate soredia. Selected specimens examined: ECUADOR. GALAPAGOS: Floreana Island, Cerro Alieri, 1°17’18”S, 90°26’60”W, 380 m, zona húmeda, rama de Macraea laricifolia, sobre corteza, 27 Mar 2006, Simbaña, W. 572 (CDS 32407), caldera of Cerro Pajas, trail at the end of road leading up to crater rim, 1°17’54.29”S, 90°27’22.5”W, humid zone, forest, on soil, 2 Jan 2010, Hillmann, G. GAL-61 (CDS 44839). Isabela Island, Volcán Alcedo, in the crater near fumaroles, 0°27’1”S, 91°7’19”W, 780 m, transition zone, on rock, 7 Mar 2006, Aptroot, A. 64790 (CDS 31365); Volcán Cerro Azul, S slope above Iguana Cove, crater on SE-slope, 800 m, humid zone, vegetation dominated by ferns and mosses, 22 June 1976, Sipman, H.J.M. L-56 (COLO 297904); Volcán Darwin, southwestern slope, above Tagus Cove, 0°13’27”S, 91°19’19.5”W, 874 m, transition zone, on soil and plant debris, 15 Nov 2007, Bungartz, F. 7727 (CDS 38231), southwestern slope, above Tagus Cove, 0°13’11.4”S, 91°19’14.1”W, 955 m, transition zone, on soil, 14 Nov 2007, Bungartz, F. 7624 (CDS 38126); Volcán Sierra Negra, El Mango, on the E-side of the dirt road, 0°53’1.7”S, 91°0’50.8”W, 162 m, transition zone, on soil, 15 Aug 2008, Bungartz, F. 8222 (CDS 40868); Volcán Alcedo, on the crater rim near the hut, 0°26’33”S, 91°5’31”W, 1100 m, humid zone, on rock, 3 July 2006, Aptroot, A. 65255 (CDS 31841). Pinzón Island, along the trail going up from Playa Escondida, N- to W-facing cliff above a crater, 0°36’29”S, 90°40’14”W, 318 m, transition zone, on rock, 16 Feb 2006, Bungartz, F. 3659 (CDS 27477). San Cristóbal Island, rim of crater to the NW of Media Luna, inland from the NW-coast, 0°43’51”S, 89°18’55”W, 149 m, transition zone, on rock, 22 Apr 2007, Bungartz, F. 6304 (CDS 34516). Santa Cruz Island, at base of barranco on old trail, 20 m, dry zone, on rock, 11 Apr 1976, Weber, W.A. (CDS 10807, QCA), trail from above Mina Granillo Rojo, leading south towards Cerro Crocker, 0°37’45.79”S, 90°22’0.4”W, 682 m, humid zone, on soil, 29 May 2008, Bungartz, F. 8001 (CDS 39036), above Mina Granillo Rojo, on the N-side of the island, 0°37’7.5”S, 90°21’55.5”W, 607 m, transition zone, on rock, 8 July 2008, Clerc, P. 08-43 (CDS 39897). Santiago Island, summit of Cerro Gavilán, inner N- and NE-exposed crater rim, 0°12’20”S, 90°47’3”W, 840 m, humid zone, on rock, 23 Mar 2006, Aptroot, A. 65693 (CDS 32285).

10 • Phytotaxa 129 (1) © 2013 Magnolia Press YÁNEZ-AYABACA ET AL. FIGURE 2. a–b Cladonia bungartzii (holotype, Bungartz 5744); a general thallus aspect (scale 2 mm); b close-up of the podetia (scale 1 mm); c general thallus aspect of Cladonia cartilaginea (Nugra 240, scale 5 mm); d general thallus aspect of Cladonia ceratophylla (Bungartz 5604, scale 5 mm); e–f cone-shaped, granulose-sorediate podetia of Cladonia chlorophaea; e ( Bungartz 8223, scale 5 mm); f ( Bungartz 6614, scale 5 mm).

Primary thallus evanescent; podetia forming cushions, 4–12 cm tall; densely branched; branching pattern mainly isotomic trichotomous, rarely dichotomous, therefore not forming distinct principal axes; axils frequently perforate; tips slender; surface ecorticate, slightly verruculose; algal clumps (glomerules) visible in old podetia; pycnidia with hyaline jelly; apothecia pale brown, uncommon. Notes: Cladonia confusa is the most common reindeer lichen in Galapagos. Superficially similar species are C. arbuscula subsp. boliviana and C. arcuata. Because of its growth form, Cladia aggregata could also be mistaken for a reindeer lichen. For differences see the notes for these species. Two forms can be distinguished according to their color and the presence, respective absence of usnic acid:

Cladonia confusa f. bicolor (Müll. Arg.) Ahti & DePriest (2001: 501) (Figs. 3a–b)

Podetia ash gray to brownish gray, often with dark brown tips. Spot tests and chemistry: P−, K−, C−, KC−, UV+ bright greenish white; with perlatolic acid only. Distribution and ecology: Know from Fernandina, Isabela, San Cristóbal, Santa Cruz Island; less common than C. confusa f. confusa, but both forms often growing side by side; most specimens collected from the humid zone, but occasionally also found in the transition zone, or rarely even in the dry zone; when well developed forming extensive reindeer lichen heaths. Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Darwin, Tagus Cove, 800 m, humid zone, on large masses 1 ft or more in diam. on lava, 24 Mar 1906, Stewart, A. 430 (CAS-DS 640479), Cerro Beagle, 15 June 1984, Luong, T.T. (CDS 10886), southwestern slope, above Tagus Cove, 0°13’43.29”S, 91°19’47.3”W, 724 m, transition zone, on soil, 12 Nov 2007, Ertz, D. 11797 (CDS 37156). Santa Cruz Island, at the base of the eastern slope below the summit of El Puntudo, 0°38’42”S, 90°20’14”W, 780 m, humid zone, on soil, 28 Feb 2006, Bungartz, F. 3986 (CDS 27916), summit area NW of La Copa (= Media Luna), 830 m, humid zone, in bottom of a small crater, 18 Feb 1964, Horneman, S. [distributed as Weber, Lich. Exs. no. 106 (COLO 185828, LSU, M, NY)].

Cladonia confusa f. confusa (Figs. 3a, c–d)

Podetia yellowish to green gray, with pale brown tips. Spot tests and chemistry: P−, K−, C−, KC+ yellow, UV+ bright greenish white; perlatolic and usnic acids (perlatolic acid rarely absent and then UV–, see notes below). Distribution and ecology: Known from Fernandina, Isabela, Pinta, Pinzón, San Cristóbal, Santa Cruz, and Santiago Island. The most common Galapagos reindeer lichen. Often the dominant taxon in lichen heaths of the humid highlands, where it can form gigantic cusions up to several meters in diameter. Few specimens have also been collected in humid habitats at lower altitudes, in the transition or even dry zone. Notes: The large mayority of specimens of C. confusa f. confusa contains perlatolic acid and thus fluoresces a bright greenish white under UV light. This is not the case, however, for some specimens that also morphologically differ from typical C. confusa f. confusa. These specimens that lack perlatolic acid contain an unidentified substance that forms a pale spot at Rf 1−2 in solvent A. Morphologically the latter two chemotypes are very similar and extremely difficult to distinguish. The atypical chemotype generally appears to have ± finer, but relatively compact podetia, almost exclusively dichotomously ramified, with a rather “bumpy” surface, covered with broad, relatively flat, closely adjoining packets of algae, giving the podetia a overall rather roughened surface aspect (see inset of Fig. 3d).

12 • Phytotaxa 129 (1) © 2013 Magnolia Press YÁNEZ-AYABACA ET AL. In contrast, typical Cladonia confusa f. confusa (Fig. 3c) is characterized by mostly trichotomous ramifications, although specimens frequently also have dichotomously ramified podetia. In fact the majority of the terminal branches are dichotomous, only further down ramifications become increasinly trichotomous. Overall these typical specimens also have a more slender, less stout, less densely ramified appearance, though individual podecia are generally slightly thicker, i.e., generally somewhat broader in diameter. The different surface seems to be perphaps the most diagnostic morphological character of the two chemotypes: typical C. confusa f. confusa has packets of granular algae dispersed on an arachnoid stereome. Its surface appears overall rather smooth, though somewhat cottony and less compact, and as if peppered with granular packets of algae wrapped in hyphae. The other chemotype has a much more uneven surface and seems to lack the granular packets of algae; instead the packets of algae seem to be more closely incorporated into the stereome surface, causing the “bumpy” appearance. Nevertheless, how consistently these characters indeed correlate with the two chemotypes is quite diffcult to assess. To some extent the differences appear quite transient and we therefore hesitate to describe a new species here based on these rather inconspicous characters. Selected specimens examined: ECUADOR. GALAPAGOS: Fernandina Island, W-side, 335 m, transition zone, 15 Feb 1964, Cavagnaro (COLO 193423); green strip on SW slope, 4 Feb 1964, Cavagnaro 26 (duplicate ex US to H, dupl. to CDS). Isabela Island, Volcán Alcedo, outer SE-exposed slope, ca. 2.5 km below the crater rim, 0°26’20”S, 91°4’35”W, 784 m, transition zone, on rock, 7 Mar 2006, Bungartz, F. 4293 (CDS 28365), rim of caldera of Volcán Alcedo, S-side of crater on way to the fumaroles, 700 m, humid zone, on ground, 11 May 1976, Weber, W.A. (COLO 297127); Volcán Cerro Azul, S slope above Iguana Cove, 750 m, humid zone, 22 June 1976, Sipman, H.J.M. L-36 (COLO 297925, H, M, U, US), 700 m, humid zone, on rock, 22 June 1976, Sipman, H.J.M. L-43 (COLO 297918); Volcán Darwin, Cerro Beagle, dry zone, 15 June 1984, Luong, T.T. (CDS 10884), mountain E of Tagus Cove, 910 m, humid zone, Snodgrass, R.E. (CAS-DS 682613), southwestern slope, above Tagus Cove, 0°13’43.29”S, 91°19’47.3”W, 724 m, transition zone, on soil, 12 Nov 2007, Ertz, D. 11822 (CDS 37181); Volcán Sierra Negra, South side of Sierra Negra, trail to Alemania, 0°50’57.5”S, 91°7’41.3”W, 1020 m, humid zone, on soil, 16 Aug 2008, Herrera-Campos, M.A. 10694 (CDS 40432, 10702, 40440), close to the southern crater rim, along the trail to Alemania, 0°51’12.69”S, 91°8’40.5”W, 1055 m, humid zone, on soil and plant debris, 16 Aug 2008, Bungartz, F. 8345 (CDS 40991). Pinta Island, on top of the highest point of the island, 0°35’3”N, 90°45’12”W, 625 m, humid zone, on soil, 26 Feb 2007, Bungartz, F. 5742 (CDS 33393), E-slope of the highest crater, on highest rim of highest crater, 650 m, humid zone, on rock, 10 July 1976, Sipman, H.J.M. L-140 (COLO 297820, H, U). Pinzón Island, from the NE-coast to the highest summit, 350 m, transition zone, on rock, 2 July 1976, Sipman, H.J.M. L-95 (COLO 297865). San Cristóbal Island, Cerro San Joaquín, 0°53’49.5”S, 89°30’47.7”W, 691 m, humid zone, 24 Aug 2008, Herrera-Campos, M.A. 450 (CDS 43341), Volcán Santo Tomás, borde del cráter 0°50’0”S, 91°2’0”W, 1 Feb 1994, Follmann, G. 35302 (B-KOELN 60 0173576), bordering lake at El Junco, humid zone, on rock, 21 May 1976, Lanier, J. (COLO 298430), Wreck Bay, Stewart, A. 431 (339) (CAS-DS 640456, FH 197384), Anonymous collector 4759 (FH 197386), NE-slope of Cerro San Joaquín, shortly below the summit, 0°53’50.79”S, 89°30’49.7”W, 693 m, humid zone, on bryophytes, 24 Aug 2008, Bungartz, F. 8586 (CDS 41232). Santa Cruz Island, below El Puntudo, 720 m, humid zone, on soil, 18 Apr 1976, Weber, W.A. (CDS 10835), near Puntudo, 0°38’41”S, 90°20’13”W, 750 m, humid zone, on soil, 27 May 2005, Aptroot, A. 63203 (CDS 29934), along the trail from Media Luna to El Puntudo, at a small stream crossing the trail, 0°39’4”S, 90°20’5”W, 690 m, humid zone, on soil, 28 Feb 2006, Ziemmeck, F. 734 (CDS 27863), at the base of the eastern slope below the summit of El Puntudo, 0°38’42”S, 90°20’14”W, 780 m, humid zone, on soil, 28-Feb-2006, Bungartz, F. 3985 (CDS 27915), due N of Academy Bay, 610 m, humid zone, 20 Feb 1964, De Roy, A. (distributed by Weber as Lich. Exs. no. 105 to COLO 185829, H, M, NY, US, CAS-DS nos. 629519, 681394).

THE FAMILY CLADONIACEAE IN GALAPAGOS Phytotaxa 129 (1) © 2013 Magnolia Press • 13 FIGURE 3. a–c Cladonia confusa. a thalli of both forms growing side by side, upper pale greenish yellow thallus of C. confusa f. confusa, lower, white thallus with brownish tinge of C. confusa f. bicolor (Bungartz 3985, 3986, scale 10 mm); b close-up of C. confusa f. bicolor, the epithet bicolor justified by the more brownish, sun-exposed parts and the whitish pale underside of the podetia (Bungartz 3986, scale 3 mm); c close-up of the UV+ chemotype of C. confusa f. confusa, the inset shows a predominantly trichotomous branching pattern and an arachnoid granular surface of the podetia (Nugra 261, scale 3 mm); d close-up of the UV– chemotype of C. confusa f. confusa, the inset shows a predominantly dichotomous branching pattern with a uneven podetial surface (Cavagnaro 26, scale 3 mm); e Cladonia corniculata (Sipman L-45, scale 5 mm); f Cladonia corymbites (Bungartz 8334, scale 3 mm).

Primary thallus subpersistent, of small, suborbicular squamules, esorediate, epruinose; podetia common, whitish gray, brown with age, but not becoming necrotic, elongate; 1–3 cm tall; generally dichotomously branched near its apices; axils close; tips blunt or obtuse, ascyphose; surface completely ecorticate; granulose, with scarce microsquamules that towards the tip merge into fine granules and towards the base of the podetium become larger and more laciniate squamules; surface of old podetia commonly denuded, esorediate; pycnidia with hyaline jelly; apothecia not seen. Spot tests and chemistry: P+ orange red, K-, C-, KC-, UV-; fumarprotocetraric acid. Distribution and ecology: Currently known only from Isabela and Santa Cruz Island; known only from the humid zone where the species appears to be moderately common, often growing with other, morphologically similar Cladonia species and amongst bryophytes, in fern-sedge grasslands, over plant debris or even as epiphyte on trunks or branches of both native or introduced trees and shrub. Notes: Cladonia corniculata can be confused with C. cartilaginea or C. subradiata. For comparison see the notes under C. cartilaginea. Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Alcedo, on the crater rim near the hut, 0°26’33” S, 91°5’31” W, 1100 m, humid zone, on soil, 7 Mar 2006, Aptroot, A. 65241 (CDS 31827). Volcán Sierra Negra, near parking place at start of foot path to the crater, 0°49’47.5”S, 91°5’19.80”W, 939 m, humid zone, on bark, 9 Aug 2007, Bungartz, F. 6805 (CDS 36236); Volcán Cerro Azul, S slope above Iguana Cove, W, 700 m, humid zone, on soil, 22 June 1976, Sipman, H.J.M. L-45 (COLO 297916), S-slope above Iguana Cove, crater on SE-slope, 800 m, humid zone, 22 June 1976, Sipman, H.J.M. L-55a (COLO 297905); Volcán Alcedo, on the crater rim near the hut, 0°26’33”S, 91°5’31”W, 1100 m, humid zone, on bark, 7 Mar 2006, Aptroot, A. 65215 (CDS 31801). Santa Cruz Island, along trail from Media Luna to El Puntudo, 0°39’9.80”S, 90°19’59.29”W, 724 m, humid zone, on bark, 10 Aug 2008, Clerc, P. 08-105 A (CDS 39959), cerca la vía sector Los Gemelos, 0°38’2.10”S, 90°23’37.89”W, 661 m, zona húmeda, sobre corteza, 4 Jan 2007, Nugra, F. 266 (CDS 33182), above Mina Granillo Rojo, on the N-side of the island, 0°37’7.5”S, 90°21’55.5”W, 607 m, transition zone, on rock, 7 Aug 2008, Clerc, P. 08-44 (CDS 39898), vicinity of Academy Bay, on trail to La Copa (= Media Luna), humid zone, on bark, 15 Feb 1964, Weber, W.A. 68 (COLO 190028).

Cladonia corymbites Nyl., in Polakowsky (1877: 225) (Fig. 3f)

Primary thallus persistent to evanescent, laciniate squamules, esorediate, epruinose; podetia common, 1–2 cm tall, phyllopodiate, unbranched to slightly divaricately branched; axils closed with age, but in young podetia often open; tips blunt, ascyphose; surface mostly ecorticate, cortex restricted to the base, densely covered with microsquamules, often intermixed with macrosquamules along the length of the podetium, esorediate, egranulose; pycnidia and apothecia not seen. Spot tests and chemistry: P+ orange red, K-, C-, KC-, UV-; fumarprotocetraric acid. Distribution and ecology: Currently know only from Floreana Island (Ahti 2000); apparently a rare species, but possiby overlooked amongst morphologically similar taxa, known only from the humid highlands, on thin soil over rock, rarely as epiphyte. Notes : Cladonia corymbites can be confused with C. cartilaginea, but that species forms only very few microsquamules and its macrosquamules are always restricted to its base. In contrast, C. corymbites is typically densely covered with both micro- and macrosquamules along the entire length of its podetia. Cladonia corymbosula is another very similar species, but it forms podetia that are densely sorediate- granulose and always lack microsquamules. Unlike C. corymbites, both C. cartilaginea and C. corymbosula are not phylopodiate, i.e., their squamules are persistent and not gradually being transformed into podetia.

THE FAMILY CLADONIACEAE IN GALAPAGOS Phytotaxa 129 (1) © 2013 Magnolia Press • 15 Selected specimens examined: ECUADOR. GALAPAGOS: Floreana Island, 300 m, 1976, Gradstein L- 62998 (US). Isabela Island Volcán Sierra Negra, close to the southern crater rim, along the trail to Alemania, 0º51'12.7”S, 91º8'40.5”W, humid zone, on soil, 16 Aug 2008, Bungartz, F. 8334 (CDS 40980).

Cladonia corymbosula Nyl. (1876a: 560) (Fig. 4a–b)

Primary thallus persistent, of short-laciniate, thick squamules, surface scarcely pruinose, rugulose and cracked, esorediate; podetia rare, grayish green, small, 0.6–1.7 cm tall, terete to ± flattened, hyphae of the stereome relatively loose (not compact) and frequently with longitudinal cracks; tips clavate, unbranched to moderately branched, ascyphose, typically bearing closely aggregated, “turban-like” apothecia; surface ecorticate, densely sorediate-granulose; microsquamules absent; pycnidia not seen; apothecia pale brown, globular and closely aggregated and thus resembling a “turban”. Spot tests and chemistry: P+ orange red, K-, C-, KC-, UV-; fumarprotocetraric acid. Distribution and ecology: A new record for Ecuador and the Galapagos Islands. Currently known from Isabela, Santiago, and Santa Cruz Island; known from the humid zone only, on rock or thin soil; possibly quite rare, but the basal squamules lacking the characteristic podetia might be overlooked. Notes : The species can easily be recognized by its relatively short podetia topped by closely aggregated, “turban-like” apothecia. The species could be mistaken for C. peziziformis, which has similarly aggregated apothecia. Cladonia peziziformis, however, has corticate podetia, its cortex peeling off in relatively large squamules. In contrast, podetia of C. corymbosula are ecorticate, and typically sorediate; they always lack squamules. Despite previous reports, Cladonia peziziformis cannot be confirmed for the Galapagos and the reports are almost certainly based on misidentifications of C. corymbosula. Material lacking podetia is not uncommon and can often be identified if compared to the squamulose primary thalli of specimens that bear podetia. It is generally difficult, however, to distinguish clear cut characters that help identify these specimens when the characteristic podetia are missing. Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Alcedo, on the crater rim near the hut, 0°26’33”S, 91°5’31”W, 1100 m, humid zone, on rock, 3 July 2006, Aptroot, A. 65262 (CDS 31848). Santa Cruz Island, near Los Gemelos craters, 0°36’31”S, 90°22’4”W, 350 m, humid zone, on soil, 31 May 2005, Aptroot, A. 63384 (CDS 30130). Santiago Island, summit of Cerro Gavilán, inner N- and NE- exposed crater rim, 0°12’20”S, 90°47’3”W, 840 m, humid zone, on rock, 23 Mar 2006, Aptroot, A. 65721 (CDS 32313).

Cladonia dactylota Tuck. (1859: 201) (Fig. 4c–d)

Primary thallus persistent, of abundant, usually ± erect and rather thick squamules, typically densely cottony sorediate on their lower surface, epruinose; podetia common, greenish gray, sometimes with dark brown necrotic parts, often elongated, 0.8–1.5 (–2.5) cm tall, unbranched to slightly branched; axils closed; tips commonly with narrow and irregular scyphi that sometimes present small marginal proliferations, rarely ascyphose and then acute; surface completely corticate and densely sorediate, soredia farinose, developing into distinctly delimited tuberculose soralia, generally forming bellow the scyphi, less commonly also along the podetial stalk; macrosquamules absents from the podetia, restricted to the basal squamules of the primary thallus; pycnidia with hyaline jelly, apothecia with brown jelly. Spot tests and chemistry: P+ golden yellow, K-, C-, KC-, UV-; psoromic acid and traces of 2’-O- demethylpsoromic and fumarprotocetraric acid. Distribution and ecology: New to Galapagos; reported here from Isabela, Pinta, San Cristóbal, Santa Cruz, and Santiago Island; a common species in the humid zone, sometimes also found in the transition zone; on soil or rock and often among plant debris.

16 • Phytotaxa 129 (1) © 2013 Magnolia Press YÁNEZ-AYABACA ET AL. Notes: The species can easily be recognized by its tuberculose soralia with abundantly farinose soredia in combination with a very characteristic P+ golden yellow spot test reaction caused by psoromic acid. The primary thallus of the species is also very distinct with its stout, ± erect squamules that have a cottony sorediate lower surface. The species can thus even be identified if its podetia are missing. Sepected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Alcedo, on crater rim NW of hut at highest point, 0°25’51”S, 91°5’16”W, 1190 m, humid zone, on soil, 5 May 2006, Aptroot, A. 64830 (CDS 31405). Pinta Island, on top of the highest point of the island, 0°35’3”N, 90°45’12”W, 625 m, humid zone, on rock and plant debris, 26 Feb 2007, Bungartz, F. 5748 (CDS 33402). San Cristóbal Island, NE-slope of Cerro San Joaquín, shortly below the summit, 0°53’50.79”S, 89°30’49.7”W, 693 m, humid zone, on bryophytes, 24 Aug 2008, Bungartz, F. 8587 (CDS 41233). Santa Cruz Island, near Puntudo, 0°38’41”S, 90°20’13”W, 750 m, humid zone, on soil, 27 May 2005, Aptroot, A. 63169, 63202 (CDS 29900, 29933). Santiago Island, Coscojo, 0°13’12”S, 90°45’45”W, 725 m, transition zone, on soil, 24 Mar 2006, Aptroot, A. 65567 (CDS 32155).

Cladonia didyma (Fée) Vain. (1887: 137) (Fig. 4e)

Primary thallus persistent, of crenulate squamules, esorediate, epruinose; podetia common, grayish green to brown, with necrotic dark brown base, short to elongate, (0.8) 1–2.5 (5) cm tall; unbranched to scarcely branched; axils closed; tips blunt or more often with several closely aggregated convex, bright red apothecia, ascyphose; surface completely ecorticate with abundantly olive green to glaucescent microsquamules and granules, rarely becoming farinose sorediate; pycnidia not seen; apothecia with red jelly. Spot tests and chemistry: P+ yellow, K+ yellow, C-, KC-, UV-; thamnolic and didymic acids (chemotype II sensu Ahti 2000, no other chemotypes observed in Galapagos). Distribution and ecology: Know from Isabela, San Cristóbal, Santa Cruz, and Santiago Island; by far the most common red-fruited species, common and possibly restricted to the humid highlands, on a variety of substrates like soil, rock or frequently also as epiphyte, typically amongst bryophytes; both on native and introduced trees. Notes : In Galapagos C. didyma may be confused with C. macilenta; both have generally greenish to brown green, ecorticate podetia with microsquamules, soredia and granules. However, C. macilenta is typically very densely sorediate and its podetia typically bear very few if any microsquamules. In contrast, C. didyma is typically densely microsquamulose and true ecorticate soredia are extremely rare. Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Alcedo, outer SE- exposed slope and crater rim, 0°27’29”S, 91°7’19”W, 1089 m, humid zone, on wood, 5 Mar 2006, Aptroot, A. 65102 (CDS 31684), upper NNW-exposed slope inside the crater, 0°27’27”S, 91°7’23”W, 1055 m, humid zone, on bark, 5 Mar 2006, Bungartz, F. 4092 (CDS. 28056), Sipman, H.J.M. L-46 (COLO 297915); Volcán Sierra Negra, 0°50’0”S, 91°10’0”W, 800 m, humid zone, on soil, 18 Apr 1990, Sánchez-Pinto, L. 5047 (B), Villamil, 150 m, dry zone, 6 July 1906, Stewart, A. 428 (336) (COLO 255412). San Cristóbal Island, 1905- 1906, Stewart 341 (MSC), Lago El Junco, 0°53’0”S, 89°28’0”W, humid zone, on soil, 1 Mar 1994, Follmann, G. 34995 (B-KOELN 60 0173603). Santa Cruz Island, vicinity of Academy Bay, on trail to La Copa (= Media Luna), 15 Feb 1964, Weber L-40271 (H, M, US, COLO 190028), near Puntudo, 0°38’41”S, 90°20’13”W, 750 m, humid zone, on soil, 27 May 2005, Aptroot, A. 63206 (CDS 29937), along the side of a little path to El Puntudo, 0°38’55”S, 90°20’4”W, 698 m, humid zone, on rock, 28 Dec 2005, Bungartz, F. 3301 (CDS 26956), El Puntudo, 0°44’33”S, 90°18’12.6”W, 694 m, zona húmeda, sobre corteza, 17 July 2007, Nugra, F. 412 (CDS 36161), summit of the island between El Puntudo and Cerro Crocker, 700 m, humid zone, 16 Apr 1976, Weber, W.A. (COLO 296977), path from Media Luna to El Puntudo, near El Puntudo, 0°39’8.59”S, 90°20’2.8”W, 684 m, humid zone, on bryophytes, 28 Oct 2010, Yánez-Ayabaca, A. 1537 (CDS 45030). Santiago Island, Munecho rock outcrop, 0°12’35”S, 90°46’57”W, 860 m, humid zone, on rock, 23 Mar 2006, Aptroot, A. 65503 (CDS 32092).

THE FAMILY CLADONIACEAE IN GALAPAGOS Phytotaxa 129 (1) © 2013 Magnolia Press • 17 FIGURE 4. a–b Cladonia corymbosula. a basal squamules (Bungartz 4129, scale 5 mm); b podetia with turban-like apothecia (Aptroot 65255, scale 3 mm); c–d Cladonia dactylota; c atypical, because unusually well-developed podetia (Aptroot 64643, scale 5 mm); d close–up of characteristic squamules (Aptroot 64667, scale 5 mm). e predominantly microsquamulose podetia of Cladonia didyma (Bungartz 4109, scale 5 mm); f tiered podetia of Cladonia grayi (Bungartz 8344, scale 5 mm).

Primary thallus subpersistent, of crenulate squamules, esorediate, epruinose; podetia common, greenish gray but with some brown parts, not melanotic; 0.6–1.5 cm tall, unbranched; always scyphose; scyphi moderately widened, sometimes branching and proliferating from the margin; surface along the stalk initially corticate, but towards the rim soon becoming verruculose–granular, often intermingled with scarce microsquamules; both granules and ecorticate soredia develop inside the cup; pycnidia hyaline; apothecia not seen. Spot tests and chemistry: P+ orange red, K−, C−, KC−, UV+ whitish blue; fumarprotocetraric and grayanic acid. Distribution and ecology: New to Galapagos; known from Isabela and Santa Cruz Island; moderately common and restricted to the humid zone, typically on soil or over rocks with thin soil layer, often among plant debris or bryophytes. Notes: Easily recognized by its UV+ whitish blue reaction caused by grayanic acid. Superficially similar to C. subsquamosa, but that species differs in its secondary chemistry (fumarprotocetraric acid instead of grayanic acid) and its podetial surface is always more densely covered with propagules (microsquamules, granules and soredia). Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Sierra Negra, South side of Sierra Negra crater, trail to Alemania, 0°50’57.5”S, 91°7’41.3”W, 1020 m, humid zone, on soil, 16 Aug 2008, Herrera-Campos, M.A. 10700 (CDS 40438). Santa Cruz Island, near Puntudo, 0°38’41”S, 90°20’13”W, 750 m, humid zone, on soil, 27 May 2005, Aptroot, A. 63195 (CDS 29926).

Cladonia macilenta Hoffm. (1796: 126) (Fig. 5a)

Primary thallus subpersistent, of crenulate squamules, esorediate or sometimes with soredia, epruinose; podetia common, greenish gray or pale yellow, occasionally becoming brown (melanotic), but not necrotic, short to elongate; 0.8–2.5 cm tall, simple to slightly branched, tips obtuse or acute; ascyphose; surface mostly ecorticate, with cortex restricted to the base of the podetium and immediately below the apothecia; densely covered with farinose soredia, scarce microsquamules, lacking granules; pycnidia with bright red jelly, apothecia closely aggregated, with bright red jelly. Spot tests and chemistry: Medulla P+ yellow K+ yellow, C−, KC−, UV−; thamnolic and didymic acid (chemotype I). Distribution and ecology: Currently known only from Isabela, Pinta, Pinzón, San Cristóbal, Santa Cruz, and Santiago Island; a relatively rare species, known from the humid zone only, on soil or rotten wood, less commonly on bark. Notes: The species differs from C. didyma by consistently producing abundant soredia, but scarcely any microsquamules. Cladonia didyma instead has densely microsquamulose podetia, and although its squamules can be minute, almost granular, this species only very rarely forms true soredia. Cladonia bungartzii is similar to both species. Like C. macilenta it is densely sorediate, but has podetia that are mostly corticate. This cortex, although not restricted to the base but present along the length of the podetium, does not remain intact where soredia develop. The stereomes of C. bungartzii are generally not as compact as those of C. macilenta and C. didyma, instead they are composed of relatively loosely interwoven hyphae and unlike the other two re-fruited species, podetia of C. bungartzii are therefore characterized by distinct longitudinal cracks of the cortex and stereome. All Galapagos specimens contain thamnolic and didymic acid and specimens previously identified as C. macilenta var. bacillaris do not correspond to this taxon. Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Alcedo, outer SE- exposed slope and crater rim, 0°27’29”S, 91°7’19”W, 1089 m, humid zone, on wood, 5 Mar 2006, Aptroot, A.

THE FAMILY CLADONIACEAE IN GALAPAGOS Phytotaxa 129 (1) © 2013 Magnolia Press • 19 65100 (CDS 31682), upper NNW-exposed slope inside the crater, 0°27’27”S, 91°7’23”W, 1055 m, humid zone, on bark, 3 May 2006, Bungartz, F. 4093 (CDS 28057); Volcán Darwin, southwestern slope, above Tagus Cove, 0°13’43.29”S, 91°19’47.3”W, 724 m, transition zone, on soil, 11 Dec 2007, Bungartz, F. 7470 (CDS 37957); Volcán Sierra Negra, close to the southern crater rim, along the trail to Alemania, 0°51’12.69”S, 91°8’40.5”W, 1055 m, humid zone, on plant debris, 16 Aug 2008, Bungartz, F. 8342 (CDS 40988), close to Volcán Chico, along the trail, 0°46’57.79”S, 91°5’59.4”W, 944 m, transition zone, on rock, 14 Aug 2008, Truong, C. 1280 (CDS 39591). Pinta Island, on top of the highest point of the island, 0°35’3”N, 90°45’12”W, 625 m, humid zone, on soil, 26 Feb 2007, Bungartz, F. 5743 (CDS 33395). Pinzón Island, E-facing side of a valley on the W-slope of the highest mountain, 0°36’49”S, 90°40’14”W, 294 m, transition zone, on soil, 16 Feb 2006, Bungartz, F. 3606 (CDS 27424). San Cristóbal Island, in encanada near El Junco, humid zone, on bark, 21 May 1976, Lanier, J. (COLO 298441). Santa Cruz Island, vicinity of Academy Bay, La Copa (= Media Luna), humid zone, 15 Feb 1964, Weber, W.A. 426 (COLO 193442), along trail from Media Luna to El Puntudo, 0°39’9.80”S, 90°18’59.3”W, 674 m, humid zone, on bryophytes and bark, 8 Oct 2008, Bungartz, F. 8142 (CDS 40788). Santiago Island, along the trail from Bucanero to Jaboncillos, ca. 1 km below the summit, Cerro Gavilán, 0°11’45”S, 90°47’20”W, 680 m, transition zone, on wood, 22 Mar 2006, Aptroot, A. 65433 (CDS 32019).

Cladonia nana Vain. (1894: 23) (Fig. 5b)

Primary thallus persistent, thin, coralloid, of elongate to laciniate squamules, corticate above and cottony ecorticate below, the lower side covered with coarsely granular-soredia; epruinose, podetia typically very few or absent, if present, short, 0.5–1.5 (–2) cm tall; phyllopodiate; simple to slightly branched; terete to flattened; tips blunt; surface initially corticate, but soon granular-areolate, peeling off as abundant microsquamules, lacking soredia; pycnidia with hyaline jelly, apothecia with brown jelly. Spot tests and chemistry: P+ red; K− or + light brown, C−, KC−, UV−; fumarprotocetraric and traces of protocetraric acid. Distribution and ecology: Known from Floreana, Isabela, Pinta, Santa Cruz, and Santiago Island; possibly the most common species dominated by squamules, often forming extensive matts, most common in the humid and upper transition zone, few collections from the dry zone; on a wide variety of substrates, soil, rotten wood, or as epiphyte. Notes : The dominant coralloid primary thallus of abundant elongated to strap-shaped squamules with small or even no podetia cannot be confused with any other Galapagos species. Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Alcedo, on the crater rim near the hut, 0°26’33”S, 91°5’31”W, 1100 m, humid zone, on bark, 7 Mar 2006, Aptroot, A. 65201 (CDS 31787), on soil, 7 Mar 2006, Aptroot, A. 65239 (CDS 31825); Volcán Cerro Azul, S-slope above Iguana Cove, 800 m, humid zone, on steep rocky bank, 24 June 1976, Sipman, H.J.M. L-84 (COLO 297876). Volcán Sierra Negra, top of eastern crater rim, 0°48’22.30”S, 91°5’15.8”W, 991 m, humid zone, on wood, 8 Sept 2007, Bungartz, F. 6801 (CDS 36220). Pinta Island, along the trail up to the summit from the S-coast, 0°34’47”N, 90°45’8”W, 493 m, humid zone, on bark, 26 Feb 2007, Bungartz, F. 5778 (CDS 33451). Pinzón Island, in the valley on the W-slope of the highest mountain, 0°36’41”S, 90°40’11”W, 310 m, dry zone, on detritus, 16 Feb 2006, Aptroot, A. 64102 A (CDS 30663). Santa Cruz Island, vincinity of Academy Bay, La Copa (= Media Luna), 1964 Weber, W.A. 425 (L-40633; H; COLO 192156), Bellavista, near parking place for trail to Media Luna, 0°40’10”S, 90°19’22”W, 400 m, humid zone, on wood, 27 May 2005, Aptroot, A. 63134 (CDS 29864), above the quarry Mina Granillo Rojo, off the main road to the channel, on the N-side of the island, 0°37’5.79”S, 90°21’59.1”W, 617 m, transition zone, on soil, 21 Oct 2007, Bungartz, F. 7137 (CDS 37622). Santiago Island, along the trail from the caseta in La Central to La Bomba (at the coast), cerro ca. 1 km NE of the caseta and on the W-side of the trail, 0°14’10”S, 90°44’41”W, 664 m, transition zone, on rock, 25 Mar 2006, Bungartz, F. 4855 (CDS 29051), summit of Cerro Gavilán, inner N- and NE-exposed crater rim,

20 • Phytotaxa 129 (1) © 2013 Magnolia Press YÁNEZ-AYABACA ET AL. 0°12’20”S, 90°47’3”W, 840 m, humid zone, on rock, 23 Mar 2006, Aptroot, A. 65700 (CDS 32292), Aptroot, A. 65711 (CDS 32303), area around the entrance of the lava tunnel at La Central, 0°14’23”S, 90°45’8”W, 667 m, humid zone, on rock, 24 Mar 2006, Bungartz, F. 4850 (CDS 29026).

Cladonia polyscypha Ahti & L. Xavier in Ahti et al. (1993:61) (Fig. 5c–d)

Primary thallus persistent though poorly developed, of small squamules, esorediate, epruinose; podetia common, whitish gray, becoming brown with age (melanotic), blackened and dying off at the base (necrotic), elongate; 1.5–4 cm tall; simple to branched; axils closed; tips always with very narrow scyphi; surface corticate at the base but cortex sometimes reaching up to about 1/2 of the stalk; young podetia generally more extensively corticate; typically moderately microsquamulose-granulose, but scarcely sorediate, the soredia, when present, farinose; pycnidia with hyaline jelly, apothecia with brown jelly. Spot tests and chemistry: P+ red, K−, C−, KC−, UV−; fumarprotocetraric acid. Distribution and ecology: New to Ecuador and the Galapagos; known from Isabela, and Santa Cruz Island. Ahti (2000) reported this species only from eastern South America, but it seems to be more widespread. Moderately common in Galapagos, but known only from the humid zone, where it grows mostly on soil or rocks, amongst plant debris and bryophytes, less common also as epiphyte, typically at the base of larger trunks. Notes : This species can easily be confused with C. aff. ramulosa, but that species is more strongly corticate, the cortex typically flaking of as microsquamules, these eventually disintegrating into ecorticate soredia. Cladonia pulverulenta also is very similar, but its podetia are more densely covered by soredia, granules, microsquamules, and schizidia. For a detailed discussion of thallus morphology see C. pulverulenta. Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Darwin, southwestern slope, above Tagus Cove, 0°13’34”S, 91°19’21.4”W, 840 m, transition zone, on soil, 13 Nov 2007, Bungartz, F. 7496 (CDS 37987). Santa Cruz Island, Steve Divine's Farm at the end of Tortoise Road, off the main road to Baltra, Tortoise Territory, 0°40’8”S, 90°24’17”W, 364 m, humid zone, agricultural area, on rock, 23 Feb 2006, Aptroot, A. 64514 (CDS 31086), cerca la vía sector Los Gemelos, 0°37’33.39”S, 90°23’0.7”W, 611 m, zona húmeda, sobre corteza, 10 Nov 2006, Nugra, F. 143 (CDS 32797) above the quarry Mina Granillo Rojo, off the main road to the channel, on the N-side of the island, 0°37’5.79”S, 90°21’59.1”W, 617 m, transition zone, on soil, 21 Oct 2007, Bungartz, F. 7139 (CDS 37624).

Cladonia pulverulenta (L. Scriba ex Sandst.) Ahti (2000: 145) (Fig. 5e)

Primary thallus persistent, of elongate squamules, upper side sometimes coarsely pruinose along the margin or covering the entire squamules, esorediate; podetia common, greenish gray or pale yellow, with dark brown necrotic basal parts, slender, elongate; 0.8–2.5 cm tall, simple to sparsely branched, tips blunts, rarely scyphose; scyphi, if present, narrow; surface corticate at the base with cortex extending up to 1/3 or 2/3 of the stalk, typically covered by granules, microsquamules, schizidia and farinose soredia as a result of cortex disintegrations; pycnidia hyaline, apothecia with brown jelly. Spot tests and chemistry: P+ red, K−, C−, KC−, UV−; fumarprotocetraric acid. Distribution and ecology: New record for Ecuador and Galapagos. Known from Isabela, San Cristóbal, Santa Cruz, and Santiago Island. Ahti (2000) reported this species only from the West Indies and Central America, but it is more widespread, having been confused with C. ochrochlora. A common species in Galapagos; widely distributed through the humid zone, rarely also found in the transition or dry zone, on the ground or as epiphyte.

THE FAMILY CLADONIACEAE IN GALAPAGOS Phytotaxa 129 (1) © 2013 Magnolia Press • 21 FIGURE 5. a the predominantly sorediate podetia of Cladonia macilenta (Bungartz 7758, scale 3 mm); b highly dissected, minute squamules with very short podetia, characteristic of Cladonia nana (Aptroot 64501, scale 5 mm); c–d Cladonia polyscypha. c general growth aspect (Bungartz 3970, scale 5 mm); d close-up with podetia terminating in narrow scyphi (Aptroot 64514, scale 5 mm); e podetia of Cladonia pulverulenta (Aptroot 64518, scale 5 mm); f broad, trumpet-shaped podetia with coarse corticate granules characteristic of Cladonia pyxidata (Aptroot 64846, scale 3 mm).

22 • Phytotaxa 129 (1) © 2013 Magnolia Press YÁNEZ-AYABACA ET AL. Notes: This species is very similar to C. aff. ramulosa; C. granulosa, and C. polyscypha; all have a very similar general morphology of relatively tall and slender podetia with blunt or narrowly scyphose tips. Only C. polyscypha is consistently scyphose and never has blunt apices. Cladonia aff. ramulosa, when well developed, has very irregular cups, sometimes even ± broadened and typically with abundant proliferations. As a result these specimens of C. aff. ramulosa have an overall much more “disheveled look”, but this species is highly variable and less well developed material is often hard to distinguish from the other species. All other species, if scyphose, are generally characterized by very narrow cups that are just barely wider than the stalk and their scyphi have no, or only extremely short proliferations. Cladonia aff. ramulosa is also the only species that is extensively corticate with few denuded areas, where the cortex flakes off forming microsquamules. All other species are not as extensively corticate. Both, C. pulverulenta and C. polyscypha have podetia that become increasingly blackened and die off at their base (necrotic), a character not observed in the other two species. In C. pulverulenta basal cortication typically covers at least 2/3 of the stalk and only then begins to disintegrate into irregular parts peeling off as schizidia. Its surface also develops microsquamules, granules, or even farinose soredia. In contrast, C. polyscypha is less corticate, the cortex does not peel or flake off but develops into few distinct microsquamules or granules, soredia are rare. These similar species all contain fumarprotocetraric acid, only C. granulosa contains thamnolic acid instead. Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Sierra Negra, along dirt road from Puerto Villamil to crater of Sierra Negra, farmland, 0°51’39.89”S, 91°1’41.60”W, 330 m, humid zone, on wood, 9 Sept 2007, Bungartz, F. 6928 (CDS 36427), Villamil, 120 m, dry zone, rocks and dead wood, 6 July 1905, Stewart, A. 428 (336) (CAS-DS 640540); Volcán Alcedo, outer E-exposed slope just below the crater rim, 0°25’17”S, 91°5’8”W, 1077 m, humid zone, on bark, 8 Mar 2006, Aptroot, A. 65127 (CDS 31709). San Cristóbal Island, sector of the "Gotera de agua", trail to Cerro Pelado, 0°51’40.60”S, 89°27’37.20”W, 397 m, transition zone, on bark, 23 Aug 2008, Truong, C. 1493 (CDS 39804). Santa Cruz Island, near Horneman farm, humid zone, on bark, 20 Aug 2008, Truong, C. 1341 (CDS 39652). Santiago Island, along the trail from Cerro Gavilán to La Central, 0°13’2”S, 90°46’33”W, 890 m, humid zone, on soil, 24 Mar 2006, Bungartz, F. 4830 A (CDS 29004).

Cladonia pyxidata (L.) Hoffm. (1796: 121) (Fig. 5f)

Primary thallus persistent, of ascending squamules; esorediate, epruinose; podetia common, grayish green to brown, short; 0.4–1 cm tall; with broad scyphi shaped like a trumpet, i.e., with a wide rim tapering to a thin stalk; cups mostly simple but very rarely proliferating from their margin, forming additional scyphi; surface corticate at the base of the scyphi, becoming aereolate to granulose, but not sorediate, typically with large, corticated granules inside the cup; rarely with schizidia; esorediate and always lacking microsquamules; pycnidia with hyaline jelly; apothecia with brown jelly. Spot tests and chemistry: P+ orange red, K−, C−, KC−, UV−; fumarprotocetraric acid. Distribution and ecology: New to Ecuador and Galapagos; found on Isabela, and Santiago Island. Although C. pyxidata in widespread in the world, it was not reported from most of the Andes by Ahti (2000), although the very similar calciphilous species C. pocillum was found in those areas. Among the “pixie cup lichens” of Galapagos, C. pyxidata is more common than C. grayi, but much rarer than the common C. chlorophaea and the very common C. subsquamosa. Unlike the other superficially similar species, collections of C. pyxidata are quite often also found at the lower altitudes of the dry and transition zone and the species thus appears not as restricted to humid habitat. Notes : Several species of Cladonia with broadly cup-shaped podetia can be distinguished in Galapagos; C. chlorophaea, C. subsquamosa, C. grayi and C. pyxidata. They differ by the shape of their cups, surface cortication, development of propagules and chemistry.

THE FAMILY CLADONIACEAE IN GALAPAGOS Phytotaxa 129 (1) © 2013 Magnolia Press • 23 Cladonia pyxidata is the only species that does not form soredia, but instead only produces relatively large, corticate granules. It forms trumpet-shaped cups with a wide rim and more abruptly tapering stalk. Cups formed by C. chlorophaea are not as broad, generally more cone-shaped and thus gradually tapering towards its base. This species forms farinose soredia of more or less uniform size. Cups of C. subsquamosa are very similar but the species produces microsquamules, granules and soredia, all rather heterogeneous in size. When well developed specimens of C. subsquamosa are abundantly proliferating forming tiers of cups. Such repeated proliferations are not common in the other species but can occasionally be observed in C. grayi, which, however, is reliably distinguished by its conspicuous UV+ whitish blue reaction due to the presence of grayanic acid, a secondary metabolite absent from all other species. Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Alcedo, outer SE- exposed slope, ca. 500 m below the crater rim, 0°27’13”S, 91°5’46”W, 1035 m, humid zone, on bark, 3 June 2006, Aptroot, A. 64846 (CDS 31422). Santiago Island, summit of Cerro Gavilán, inner N- and NE-exposed crater rim, 0°12’20”S, 90°47’3”W, 840 m, humid zone, on rock, 23 Mar 2006, Aptroot, A. 65699 (CDS 32291), along trail to summit above Santiago Bay lava flow, 0°12’35”S, 90°47’5”W, 895 m, humid zone, on rock, 30 Apr 1971, Pike, L.H. 2722 (OSC 54939).

Cladonia aff. ramulosa (With.) J.R. Laundon (1984: 225) (Fig. 6a)

Primary thallus persistent, of ascendant and crenulate squamules, esorediate, epruinose; podetia common, slender, whitish gray or greenish gray sometimes with brown areas but not necrotic, elongate; 0.6–4 cm tall, simple; axils close; tips initially blunt but soon developing into narrow, very irregular, shallow and typically proliferating scyphi; surface mostly corticate but with small denuded areas; patches of the cortex flaking off as microsquamules, these eventually disintegrating into ecorticate soredia, typically becoming densely sorediate, but lacking corticate granules; pycnidia not seen; apothecia with brown jelly. Spot tests and chemistry: P+ red, K−, C−, KC−, UV−; fumarprotocetraric acid. Distribution and ecology: Currently known from Isabela, Pinta, San Cristóbal, and Santa Cruz; common throughout the humid zone, on soil or rock, often among plant debris and bryophytes. Notes : Specimens of C. aff. ramulosa can be extremely similar to C. pulverulenta and poorly developed specimens cannot always be identified with certainty; see the diagnostic differences discussed there. Although Cladonia ramulosa s.str. was reported by Ahti (2000) for South America, it may generally be questioned whether this species even occurs in this continent as preliminary molecular studies suggest that most South American specimens previously identified as C. ramulosa actually belong to at least one, possibly several distinct, yet still undescribed species. Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Cerro Azul, S-slope above Iguana Cove, 300 m, humid zone, on wood, 22 June 1976, Sipman, H.J.M. L-39 (COLO 297923); Volcán Sierra Negra, trail climbing up to Sierra Negra crater, 0°49’41.39”S, 91°5’30.10”W, 967 m, humid zone, on wood, 14 Aug 2008, Herrera-Campos, M.A. 10548 (CDS 40284); El Mango, on the E-side of the dirt road, 0°53’1.7”S, 91°0’50.79”W, 162 m, transition zone, on soil, 15 Aug 2008, Bungartz, F. 8185 (CDS 40831). Pinta Island, E-slope of the highest crater, on highest rim of highest crater, 650 m, humid zone, on rock, 10 July 1976, Sipman, H.J.M. L-139 (COLO 297821); E-slope of the highest crater, 550 m, humid zone, in open mossy Zanthoxylum forest, nearly vertical SE-exposed rock face, on rock, 10 July 1976, Sipman, H.J.M. L-138 (COLO 297822). San Cristóbal Island, Wreck Bay, SE-side of the main mountain, 600 m, humid zone, on rock and moist earth, 6 July 1906, Stewart, A. 426 (COLO 255411); Cerro Colorado, enclosure for Calandrinia galapagosa near the viewpoint on the top, 0°54’58”S, 89°26’5”W, 130 m, transition zone, on rock, 29 Apr 2007, Bungartz, F. 6737 (CDS 34981) Santa Cruz Island, base of El Puntudo, humid zone, summit grassland, large boulders strewn about, on rock, 16 June 1972, Weber, W.A. (COLO 256035), along trail from Media Luna to El Puntudo, 0°39’9.80”S, 90°19’59.29”W, 724 m, humid zone, on bark, 10 Aug 2008, Clerc, P. 08-105B, 08-125A (CDS 46961, CDS 39979), vicinity of Academy Bay, La Copa (= Media Luna), humid zone, 15 Feb 1964, Weber, W.A. 86, 96, 426 (COLO 195014, 192067, 193444).

Primary thallus subpersistent, of small, narrow squamules, esorediate, epruinose; podetia common, conspicuously yellowish green to pale yellow green, elongate, slender, 3–10 (–14.5) cm tall; unbranched to sparsely branched from the rim of narrow, open funnels; axils perforate; tips of funnel proliferations subulate; surface verrucose-arachnoid, densely granulose, but occasionally with schizidia and microsquamules; farinose soredia rare; pycnidia hyaline; apothecia not seen. Spot tests and chemistry: P+ yellow; K+ yellow, C−, KC−, UV−; thamnolic and usnic acids. Distribution and ecology: Known only from Isabela and Santa Cruz Island; known only from the humid zone, typically in fern-sedge grassland, often also part of reindeer heaths. Notes : Cladonia scholanderi is easily recognized by its long, slender, elongate, conspicuously yellowish green podetia. It is also the only Galapagos species with “bottomless” cups, i.e., cups that are not scyphose but are open funnels. Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Alcedo, outer E- exposed slope just below the crater rim, 0°25’17”S, 91°5’8”W, 1077 m, humid zone, basalt outcrops, on rock, 8 Mar 2006, Aptroot, A. 65172 (CDS 31756). Volcán Sierra Negra, South side of Sierra Negra crater, trail to Alemania, 0°51’17.60”S, 91°8’55.2”W, 924 m, humid zone, on soil/rocks, 16 Aug 2008, Clerc, P. 08-237, 08- 193, 08-196 (CDS 40091, 40047, 40050), Herrera-Campos, M.A. 10707, 10704 (CDS 40445, 40442), Truong, C. 1251 (CDS 39562), Bungartz, F. 8346, 8347 (CDS 40992, 40993). Santa Cruz Island, 700 m, 1976, Weber & Lanier L-63343 (COLO, H), eastern slope below the summit of El Puntudo, 0°38’42”S, 90°20’14”W, 780 m, humid zone, on soil, 28 Feb 2006, Aptroot, A. 64673 (CDS 31247), saddle between Mount Crocker and El Puntudo, 700 m, humid zone, on rock, 18 Apr 1976, Weber, W.A. (COLO 297078).

Cladonia aff. sphacelata Vain. (1887: 456) (Fig. 6c)

Primary thallus persistent, of laciniate squamules, esorediate, epruinose; podetia common, grayish green becoming deep brown in parts (melanotic), elongate, 1–5 cm tall; moderately branched; branching pattern irregular anisotomic, mainly dichotomous, rarely trichotomous; axils closed; tips often divided in 2–6 branchlets; ascyphose; surface ecorticate, apically initially with granules becoming larger and branching into microsquamules, eventually developing into narrow and laciniate basal squamules; esorediate; pycnidia with hyaline jelly; apothecia not seen. Spot tests and chemistry: P+ yellow, K+ yellow, C−, KC−, UV−; thamnolic acid (according Ahti (2000), the spot test reactions and chemistry correspond to chemotype I; no other chemotype has so far been confirmed for the Galapagos). Distribution and ecology: New to Ecuador and Galapagos; known from Isabela and Santa Cruz Island; all specimen from the humid zone, growing on the ground in fern-sedge vegetation. Notes: This species is characterized by repeatedly branched podetia with tips often divided in 2–6 (–10) branchlets and a surface densely covered in narrow squamules. The identification of this species is somewhat provisional. Ahti (2000) reported it only from Brazil. However, since then Ahti & Sipman (2013a, 2013b) have reported it from the Guiana Highlands, even segregating two new species off it in the same area. The identity of the Galapagos material is thus not completely clear and molecular studies might help to elucidate it. Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Sierra Negra, South side of Sierra Negra crater, trail to Alemania, 0°51’17.60”S, 91°8’55.2”W, 924 m, humid zone, on rock, 16 Aug 2008, Clerc, P. 08- 249, 08-198, 08-197 (CDS 40103, 40052), 40051), Bungartz, F. 8349 (CDS 40995). Santa Cruz Island, along trail from Media Luna to El Puntudo, 0°39’0”S, 90°20’2.9”W, 740 m, humid zone, on bark, 8 Oct 2008, Clerc, P. 08-118, 08-125B (CDS 39972, 46960), eastern slope below the summit of El Puntudo, 0°38’42”S, 90°20’14”W, 780 m, humid zone, on soil, 28 Feb 2006, Aptroot, A. 64672 (CDS 31246).

THE FAMILY CLADONIACEAE IN GALAPAGOS Phytotaxa 129 (1) © 2013 Magnolia Press • 25 FIGURE 6. a general growth aspect of Cladonia aff. ramulosa (Aptroot 65546, scale 5mm); b pale yellow, elongate podetia of Cladonia scholanderi (Clerc 08-196, scale 10mm); c podetia of Cladonia aff. sphacelata (Bungartz 8349, scale 5 mm). d thick, crenate, and grayish green, esorediate squamules of Cladonia strepsilis (Bungartz 4134, scale 3 mm); e slender podetia of Cladonia subradiata (Bungartz 8162, scale 5 mm); f Cladonia subsquamosa, cone-shaped cups with marginal proliferations, podetia generally covered in a variety of propagules like true soredia, granules, microsquamules, even squamules (Nugra 21, scale 3 mm).

Primary thallus persistent, abundant, dominant, forming dense cushions of thick, crenate, and grayish green squamules, esorediate, epruinose; podetia not observed among the Galapagos material. Spot tests and chemistry: P+ yellow; K−; C+ green, KC+ green; baeomycesic acid, strepsilin. Distribution and ecology: New to Ecuador and the Galapagos; known from Isabela and Santa Cruz Island. The nearest earlier records are from Colombia and Guatemala (Ahti 2000). In Galapagos a rare species, known only from the humid zone; found on the ground, on soil or rock, rarely also as epiphyte. Notes : The distinct cushions of thick, crenate squamules are not easily mistaken for other species of Cladonia and specimens cannot possibly be misidentified because of the conspicuous C+ green spot test reaction caused by strepsilin, a substance not known from any other Cladonia species. Specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Alcedo, outer SE-exposed slope, ca. 100 m below the crater rim, 0°25’36”S, 91°5’12”W, 1146 m, humid zone, on rock, 6 Mar 2006, Bungartz, F. 4134 (CDS 28165). Santa Cruz Island, eastern slope below the summit of El Puntudo, 0°38’42”S, 90°20’14”W, 780 m, humid zone, on soil, 28 Feb 2006, Aptroot, A. 64681 (CDS 31255).

Cladonia subradiata (Vain.) Sandst. (1922: 230) (Fig. 6e)

Primary thallus persistent or subpersistent, of thin, laciniate squamules, esorediate or with scarce granules on the lower side, epruinose; podetia common, whitish gray or grayish green to brownish, elongate, slender, 1–2 cm tall, simple to sparsely branched; tips blunt to scyphose in mature stalks; scyphi very narrow, essentially of the same diameter as the podetium; surface mostly ecorticate or basally barely corticate, typically very densely covered with isidioid microsquamules and sorediate-granulose, sometimes with few macrosquamules at the base of the podetium; pycnidia with hyaline jelly; apothecia with brown jelly. Spot tests and chemistry: P+ orange, K−, C−, KC−, UV−; fumarprotocetraric acid. Distribution and ecology: Known from Isabela, San Cristóbal, Pinzon and Santa Cruz Island; possibly the most common species with long, slender podetia; in the humid highlands often abundant, rarely also in the upper transition zone; often epiphytic on trunk and large branches of native and introduced trees, rarely on soil and then often near the bases of trees. Notes : Cladonia subradiata, when well developed, is best recognized by its terete, slender virtually entirely ecorticate podetia that are typically densely covered with isidioid microsquamules and granulose soredia. Specimens could then be confused with C. granulosa, C. aff. ramulosa, C. polyschypa or C. pulverulenta (see C. pulverulenta), but unlike all these species, podetia of C. subradiata are almost entirely devoid of any well developed cortex. Especially old, mature well developed podetia of Cladonia corniculata can be very similar to C. subradiata, a species that is generally more branched, totally decorticate and has microsquamules that show a transition from large, basal laciniate squamules to microsquamules and finally fine granules towards the tip. Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Sierra Negra, South side of Sierra Negra crater, trail to Alemania, 0°50’57.5”S, 91°7’41.29”W, 1020 m, humid zone, on fern leaves, 16 Aug 2008, Herrera-Campos, M.A. 10695 (CDS 40433), Volcán Alcedo, outer SE-exposed slope, ca. 500 m below the crater rim, 0°27’13”S, 91°5’46”W, 1035 m, humid zone, on bark, 6 Mar 2006, Aptroot, A. 64854 (CDS 31430), Villamil, 150 m, 6 July 1906, Stewart, A. (336) 428 (FH 197407), near parking place at start of foot path to the crater, 0°49’47.5” S, 91°5’19.80” W, 939 m, humid zone, on wood, 8 Sept 2007, Bungartz, F. 6822 (CDS 36253). San Cristóbal Island, Cerro Partido along trail from entrance to Cerro Pelado to El Ripioso, 0°51’23”S, 89°27’37”W, 376 m, transition zone, on rock, 28 Apr 2007, Bungartz, F. 6597 (CDS 34817), SE-slope of Cerro San Joaquín, shortly below the summit, 0º53'52.4”S, 89º30'49.9'W, humid zone, on bark & bryophytes, 24 Aug 2008, Bungartz, F. 8579 (CDS 41225). Santa Cruz Island,

THE FAMILY CLADONIACEAE IN GALAPAGOS Phytotaxa 129 (1) © 2013 Magnolia Press • 27 vicinity of Academy Bay, between first barranca and Bella Vista, transition zone, on rock, 15 Feb 1964, Weber, W.A. 176 (L-40294, COLO 193500, US), Los Gemelos, ca. 100 m S of the craters, along the road, 0°37’38.89”S, 90°23’12.4”W, 618 m, humid zone, on bark, 16 June 2010, Yánez-Ayabaca, A. 1488 (CDS 44918), along trail from Media Luna to El Puntudo, 0°39’9.80”S, 90°18’59.29”W, 674 m, humid zone, on bark, 10 Aug 2008, Bungartz, F. 8143 (CDS 40789), vicinity of Academy Bay, La Copa (= Media Luna), 500 m, humid zone, 31 Jan 1964, Itow, S. 6 (COLO 192183). Pinzón Island, along the trail going up from Playa Escondida, SW-slope of the top, 0º36'36”S, 90º40'11”W, dry zone, on cactus, 16 Feb 2006, Aptroot, A. 64070 (CDS 30631).

Cladonia subsquamosa Kremp in Warming (1874 [1873]: 336) (Fig. 6f)

Primary thallus subpersistent, laciniate squamules, esorediate, epruinose; podetia common, grayish green, sometimes brown with age, 0.8–2.5 (–3) cm tall, always scyphose and shaped like a cone; cups, when well developed, abundantly and repeatedly proliferating (forming 1-2 tiers); surface slightly corticate at the base, otherwise decorticated, basally typically ±finely squamulose, becoming microsquamulose, densely granulose or farinose sorediate; pycnidia with hyaline jelly; apothecia with brown jelly. Spot tests and chemistry: P+ red, K−, C−, KC−, UV−; fumarprotocetraric acid. Distribution and ecology: Currently know from Floreana, Isabela, Pinzón, San Cristóbal, and Santa Cruz Island. The most common of the Galapagos “pixie cup” lichens; most common and often abundant in the humid zone, less common in the transition and rarely in the dry zone; on a wide variety of substrates (rock, soil, among plant debris and bryophytes, as epiphyte on trunks, branches, twigs and even fern fronds). Notes: The cone-shaped cups of C. subsquamosa much resemble C. chlorophaea but unlike this species, cups of C. subsquamosa typically proliferate and the podetia are covered in a variety of propagules (true soredia, granules, microsquamules, even squamules). In C. chlorophaea propagules are all farinose ecorticate soredia of relatively uniform size and the cups do typically not proliferate. Cladonia pyxidata and C. grayi are also similar, but the first species has broader scyphi that do generally not proliferate and coarser, corticate granules, and the latter has a verruculose-granular surface and instead of fumarprotocetraric acid (P+ red, UV−) contains grayanic acid (P−, UV+ bluish white). For a more detailed discussion refer to the notes of C. pyxidata. Selected specimens examined: ECUADOR. GALAPAGOS: Isabela Island, Volcán Alcedo, on the crater rim near the hut, 0°26’33”S, 91°5’31”W, 1100 m, humid zone, on bark, 3 July 2006, Aptroot, A. 65236 (CDS 31822), on soil, 7 Mar 2006, Aptroot, A. 65240 (CDS 31826) Volcán Darwin, southwestern slope, above Tagus Cove, 0°13’28.19”S, 91°19’17.9”W, 872 m, transition zone, on plant debris, 15 Nov 2007, Bungartz, F. 7760 (CDS 38266). Volcán Sierra Negra, mirador El Mango, SE side of island, 0°53’1.39”S, 91°0’48.4”W, 161 m, transition zone, on bryophytes, 15 Aug 2008, Herrera-Campos, M.A. 10582 (CDS 40319). Floreana Island, Asilo de la Paz, Cerro Wittmer, trail in between cliffs, 1°18’50”S, 90°27’13.8”W, humid zone, on detritus, 3 Jan 2010, Hillmann, G. GAL-91, 93, 11 (CDS 44888, 44860, 44871). Pinzón Island, along the trail going up from Playa Escondida, N- to W-facing cliff above a crater, 0°36’29”S, 90°40’14”W, 318 m, transition zone, dry zone, on rock, 16 Feb 2006, Bungartz, F. 3661 (CDS 27479), lava, on rock, 16 Feb 2006, Aptroot, A. 64004 (CDS no. 30565). San Cristóbal Island, Wreck Bay, 610 m, humid zone, on soil, 6 July 1905, Stewart 429 (337) (CAS-DS 640539), 1906, Stewart 7340 (FH), NE-slope of Cerro San Joaquín, shortly below the summit, 0°53’50.79”S, 89°30’49.7”W, 693 m, humid zone, on soil, 24 Aug 2008, Bungartz, F. 8582 (CDS 41228), Cerro Colorado, enclosure for Calandrinia galapagosa near the viewpoint on the top, 0°54’58”S, 89°26’5”W, 130 m, transition zone, on rock, 29 Apr 2007, Bungartz, F. 6736 (CDS 34980), 15 May 2006, Jaramillo, P. 2876 C (CDS 38791). Santa Cruz Island, along trail from Media Luna to El Puntudo, 0°39’37.79”S, 90°20’0.9”W, 682 m, humid zone, on soil, 28 Dec 2005, Bungartz, F. 3271 (CDS 26910), near Puntudo, 0°38’41”S, 90°20’13”W, 750 m, humid zone, on wood, 27 May 2005, Aptroot, A. 63164 (CDS 29895), between Academy Bay and Bella Vista, 150 m, humid zone, 24 Jan 1964, Weber, W.A. 139 (COLO

28 • Phytotaxa 129 (1) © 2013 Magnolia Press YÁNEZ-AYABACA ET AL. 192784), vicinity of Academy Bay, 120 m, transition zone, on wood, 15 Feb 1964, Itow, S. 29 (COLO 192154), between Academy Bay and Bellavista, 150 m, on rock, 24 Jan 1964, Weber, W.A. (FH 197403).

Erroneous records of species to be excluded

Erroneous and doubtful records are corrected in the most recent update to the checklist of the Galapagos lichens (Bungartz et al. 2011): Previous reports of Cladina sandstedei (Abbayes) Ahti in Weber (1986) refer to Cladonia arcuata. Identifications of Cladonia balfourii Cromb. sensu Weber (1986) are misidentifications of Cladonia chlorophaea (see Ahti 2000). Identifications of Cladonia coccifera (L.) Willd. as reported in previous versions of Bungartz et al. (2011) refer to Cladonia corymbosula; Cladonia granulosa, also previously erroneously included in this list is based on a misidentification. Reports of C. polycarpoides Nyl. and C. subcariosa (Weber 1986) are based on misidentifications of C. dactylota, and C. subradiata has previously erroneously been reported as C. subulata (Weber 1986).

Doubtful records

Material in CDS of a specimen collected and identified by A. Aptroot as Cladonia rappii A. Evans is extremely poor and fragmentary, consisting of few squamules and two broken-off podetia. The material is superficially similar to that of C. pulverulenta, a species that also contains fumarprotocetraric acid. However, the specimen is too poor to be reliably identified as either one of the two species. Thus the record of C. rappii cannot be confirmed here.

Discussion

The Galapagos archipelago is well known for its historical role in the development of Darwin’s theory of evolution. These oceanic islands, of volcanic origin and thus never connected to the South American continent, have famously been called a natural laboratory of evolution, where environmental factors set the stage for highly dynamic evolutionary processes: The islands are geographically extremely isolated, ca. 1000 km from the South American mainland, not immediately surrounded by other islands. They were discovered very late (1535 by the Spanish bishop Fray Tomás de Berlanga), even then not colonized for many decades and later only sparsely inhabited for several centuries. Biogeographic isolation of the islands thus remained largely intact until the industrial age. The archipelago is characterized by considerable geomorphologic diversity, the islands build a platform of volcanoes that rise from the deep sea, with some volcanoes barely breaking the sea surface and others reaching up to 1700 m above sea level. Variation in tropical and Antarctic currents that reach the islands result in significant seasonal variation, this climatic variation is further augmented by the unpredictability of erratic catastrophic climate events, commonly referred to as El Niño. As a result of all these factors, two principal tendencies can be observed about Galapagos biodiversity: (1) in comparison with mainland South America, overall Galapagos diversity is unusually low, with species groups on the continent ten or even hundredfold more numerous; (2) in most taxonomic groups endemism is unusually high, i.e., percentages of species only known from the Galapagos, or sometimes even restricted to a particular islands, are often exceptionally high. Generally lichens are no exception. Species of lichens are clearly more diverse on the continent then in Galapagos. With an estimated 8–10% of all lichen species endemic to the Galapagos, they are less unique than other groups or organisms (ca. 49% of all native vascular plants are endemic to Galapagos, Jaramillo et al. 2012), but considering their efficient propagules even rates of endemism that, at first glance, appear less striking, nevertheless represent an unusually high degree of species diversification in Galapagos. Even though species endemic to a few or single islands are generally unknown, quite a considerable number of lichens are considered endemic to Galapagos.

THE FAMILY CLADONIACEAE IN GALAPAGOS Phytotaxa 129 (1) © 2013 Magnolia Press • 29 In this context, the unusually low percentage of endemic Cladoniaceae reported here is clearly unexpected (one species of a total of twenty-six, i.e., only 3%). This low proportion of endemism, nevertheless, needs to be put in context. Endemism of lichens varies considerably across different species groups. Of the five species of Roccella, all but one (80%) are endemic to Galapagos (Tehler et al. 2009). Four of fifteen species (26%) in the genus Ramalina are considered endemic (Aptroot & Bungartz 2007). In the Collemataceae, however, not a single species is known to be endemic (Bungartz 2008). Of forty-two species of Graphidaceae reported from Galapagos by Bungartz et al. (2010) only two were considered endemic; even with the recent discovery that one report was erroneous and instead represents an endemic species (Lücking et al. 2013), the degree of endemism in Graphidaceae does not change considerably: three out of forty-two species (7%). In Bulbothrix (manuscript submitted for publication) one out of four species is possibly endemic, in the Teloschistaceae as many as seven of nineteen species (26%) could be endemic (unpublished data). Of the eight species of Basidiolichens reported from Galapagos, only one is considered endemic (Yánez-Ayabaca et al. 2011), but unpublished data indicate another species might be endemic as well (25%). Lichens are a very diverse group of organisms across a large taxonomic spectrum of fungi, many not closely related, lichenization having occurred repeatedly during the evolution of fungi (Gargas et al. 1995, Lutzoni et al. 2001). It should not be surprising thus, if their biology and ecological adaptations, and in this particular case, distribution strategies and efficiency of their propagules, also prove to be different. At first glance lichen propagules do not, however, appear to be fundamentally different across different groups of lichenized fungi. Generally two dispersal strategies can be observed: (1) sexual reproduction by spores, where, with few exceptions, individual spores are not distributed jointly with the lichen photobiont and establishment of the lichen symbiosis depends on availability of a suitable photobiont; (2) asexual reproduction from lichen hyphae bundled with photobiont cells are jointly distributed in a variety of diaspores that all essentially represent some variation of minute thallus fragments. It can be assumed that lichens with relatively large dispersal units, i.e., propagules that are heavy would be at a disadvantage in reaching the islands. Thus, trends observed in other species groups, where inefficient dispersal mechanisms result in overall lower diversity, but higher rates of endemism should be confirmed also for lichens. With many relatively large groups of Galapagos lichens still requiring further revision (Physciaceae, Parmeliaceae, Lecanoraceae, etc.), it nevertheless remains difficult to see such particular patterns emerge. Cladoniaceae often reproduce by thallus fragments, which are relatively large, on the other hand some species reproduce by ascospores which would be at a disadvantage, if their photobiont were not available. However, Asterochloris (formerly included in Trebouxia), the genus of green algae with which Cladoniaceae form lichen symbioses (Peršoh et al. 2004) is the most common photobiont in the family. Statistically percentages are less meaningful for smaller groups. For example, the taxonomic status of at least three Cladonia species reported here is not yet completely resolved: Preliminary molecular results indicate that Cladonia aff. ramulosa in South America is most likely not the same taxon as in temperate climates (Ahti, unpublished data). The chemical variation of C. confusa f. confusa is poorly understood, its chemical and to some extend even morphological variation is here not assigned a taxonomic rank, but if examined with molecular tools might even be confirmed as a new species. Finally, the specimens here assigned to Cladonia aff. sphacelata, if examined with molecular tools, might as well represent a distinct Galapagos species. It is possible thus that some of the species in the Galapagos Cladoniaceae, although morphologically very close to their continental counterparts, must eventually be considered distinct, yet cryptic species. Assuming that, for example, two of these three taxa were in fact endemic undescribed species, the percentage of endemism in the Cladoniaceae would rise to as much as 13%. In other groups the scenario is similar. If an additional two endemic species of Roccella were to be discovered the rate of endemism in that group jumps from 80 to 85%. For the same 5% increase three more endemic species of Graphidaceae would have to be discovered.

30 • Phytotaxa 129 (1) © 2013 Magnolia Press YÁNEZ-AYABACA ET AL. Nevertheless, the tendency that lichen diversity is generally lower than in other taxonomic groups seems to be confirmed with more and more studies and an overall of trend of 8–10% endemism appears to emerge across all Galapagos lichens, curiously this rate is not significantly different from the endemism already reported in one of the first checklist of Galapagos lichens (Weber 1986); even if today almost three times as many lichens are now known from the archipelago, the percentage of species considered endemic in relation to the total number of species has not much changed. This assertion emphasizes several important aspects for conservation of the Galapagos lichen flora: (1) Because endemism in lichens appears to be generally lower than in many other species groups, any assessment of species rarity must take into account that endemic lichen species are indeed very unique. (2) Paradoxically even species of lichens distributed across large geographic areas can locally become increasingly rare, in particular species adapted to very specific habitats. Even in Galapagos local extinctions (e.g., Usnea angulata) have been reported (Bungartz et al. 2011). Endemism must consequently not be the only criterion assessed when evaluating the conservation status of lichens. (3) Inventory surveys that improve knowledge on lichen distribution in general are urgently needed on a much larger scope. Sampling effort needs to be increased, ideally be combined with molecular methods to better access lichen biogeography to better understand the phenomenon of cryptic endemism. (4) Better understanding of the dispersal characteristics of different lichen propagules would help significantly in elucidating, how lichens are distributed, why certain species are rare and endemic, and how local extinctions in an ever more fragmented environment can be prevented.

Acknowledgements

We thank Frauke Ziemmeck for managing the cryptogam collection at CDS, helping with collecting, data entry and curating of specimens. Successive Directors of Science at the Charles Darwin Foundation have supported this project: Alan Tye, Mark Gardener, and Rodolfo Martínez. We are further indebted to the Galapagos National Park, especially its technical director Washington Tapia for support and specimen export permits. The Census of Galapagos Biodiversity and the CDF Checklist of Galapagos Species is supported by several grants to the Charles Darwin Foundation (donors cited at http://www.darwinfoundation. org/datazone/ checklists/). A checklist of Galapagos lichens is regularly updated and available at http:// www.darwinfoundation.org/datazone/checklists/lichens, where contributing scientists are acknowledged. The lichen inventory continues to receive funds from The Paul and Bay Foundations and the Erwin Warth Stiftung. In 2010 an international lichen workshop was held in Galapagos, supported by two National Science Foundation (NSF) projects entitled “Neotropical Epiphytic Microlichens - An Innovative Inventory of a Highly Diverse yet Little Known Group of Symbiotic Organisms” (DEB 0715660 to The Field Museum; PI Robert Lücking) and “Phylogenetic Diversity of Mycobionts and Photobionts in the Cyanolichen Genus Dictyonema, with Emphasis on the Neotropics and the Galapagos Islands” (DEB 0841405 to George Mason University; PI James Lawrey, subcontract to the Charles Darwin Foundation, local coordinator Frank Bungartz). Finally, we much appreciate the thorough review of this manuscript by an anonymous reviewer. This publication is contribution number 2071 of the Charles Darwin Foundation for the Galapagos Islands.

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