The Family Cladoniaceae (Lecanorales) in the Galapagos Islands family...Cladoniaceae. The second...

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TheFamilyCladoniaceae(Lecanorales)intheGalapagosIslands

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PHYTOTAXA

ISSN 1179-3155 (print edition)

ISSN 1179-3163 (online edition)Copyright © 2013 Magnolia Press

Phytotaxa 129 (1): 1–33 (2013) www.mapress.com/phytotaxa/

Article

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] Central del Ecuador, Quito, Ecuador, email: [email protected] 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

mailto:mailto:[email protected]

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

YÁNEZ-AYABACA ET AL. 2 • Phytotaxa 129 (1) © 2013 Magnolia Press

flash directly in the field, or using a Novoflex macro-table to take images of herbarium specimens; forphotographic 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 microsqua-

mules ............................................................................................................................................................................ 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; pyc-

nidia 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

Phytotaxa 129 (1) © 2013 Magnolia Press • 3THE FAMILY CLADONIACEAE IN GALAPAGOS

http://www.darwinfoundation.org/datazone/coll

- 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 gran-

ules, 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 lack-

ing 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 microsquamu-

lose, 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, some-

times 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 microsquamu-

lose; 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 fumarproto-

cetraric acid)............................................................................................................................................. C. granulosa*

(*not confirmed for Galapagos, included here for its similarity to C. subradiata and C. pulverulenta)


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 soredi-

ate, 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 (fumarpro-

tocetraric 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 moder-

ately 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; surfacecorticate, 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.


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; apotheciawith 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.


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).


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. 804595Diagnosis: 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; apothecianot 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.


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; podetiacommon, 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, WeberL-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


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).


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).


Cladonia confusa R. Sant. (1942: 13)= C. galapagosensis Ahti (1961:46); according to Ahti (2000).

(Figs. 3a–d)

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 R

f 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).


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).


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).


Cladonia corniculata Ahti & Kashiw. (1984: 136)(Fig. 3e)

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. corymbosulaare not phylopodiate, i.e., their squamules are persistent and not gradually being transformed into podetia.


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; surfaceecorticate, 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.


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).


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).


Cladonia grayi G. Merr. ex Sandst. (1929: 1847)(Fig. 4f)

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.


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,


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; podetiacommon, 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; surfacecorticate 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 t

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