the genus Coccocarpia (Peltigerales: Coccocarpiaceae) - Instituto ...

Lichenological Contributions in Honour of David Galloway. 

I. Kärnefelt (ed.): Bibliotheca Lichenologica 95:429–457. 

J. Cramer in der Gebrüder Bornträger Verlagsbuchhandlung, Berlin · Stuttgart, 2007. 

A first assessment of the TICOLICHEN biodiversity inventory 

in Costa Rica: the genus Coccocarpia 

(Peltigerales: Coccocarpiaceae) 

Robert LÜCKING 1 , André APTROOT 2 , José Luis CHAVES 3 , Harrie J. M. SIPMAN 4 

& Loengrin UMAÑA 3 

1 Department of Botany, The Field Museum, 1400 South Lake Shore Drive, Chicago, IL 60605-2496, USA; 

email: rlucking@fieldmuseum.org 

2 ABL Herbarium, G.v.d. Veenstraat 107, NL-3762 XK Soest, The Netherlands; 

email: andreaptroot@wanadoo.nl 

3Laboratorio de Hongos, Instituto Nacional de Biodiversidad (INBio), Apdo. 22-3100, Santo Domingo 

de Heredia, Costa Rica; email: jchaves@inbio.ac.cr; lumana@inbio.ac.cr 

4Botanisches Museum Berlin Dahlem, Königin-Luise-Strasse 6-8, D-14191 Berlin, Germany; 

email: h.sipman@bgbm.org 

Abstract: The genus Coccocarpia is treated as part of the ongoing TICOLICHEN biodiversity inventory in 

Costa Rica, including a thorough revision of all the material reported by Dodge and held at the Farlow 

Herbarium (FH). Eighteen species are distinguished, among which four taxa are described as new: 

Coccocarpia gallaicoi Lücking, Chaves & Umaña (with the non-isidiate counterpart C. aff. gallaicoi), C. 

microphyllina Lücking & Aptroot (phyllidiate relative of C. epiphylla), C. neglecta Aptroot & Lücking 

(differing from C. domingensis by its squamiform isidia), and C. prostrata Lücking, Aptroot & Sipman 

(related to C. stellata but with ascending, fruticulose secondary lobes). The name C. guimarana (Vain.) 

Swins. & Krog is reinstated for a species with narrow, isidiate lobes. Two further, possibly undescribed 

species are tentatively identified as C. aff. gallaicoi Lücking, Chaves & Umaña and C. aff. imbricascens Nyl. 

Six taxa, viz. Coccocarpia adnata L. Arvidss., C. dissecta Swinsc. & Krog, C. epiphylla (Fée) Kremp., C. 

filiformis L. Arvidss., C. glaucina Kremp., and C. tenuissima Tuck., are reported for the first time from Costa 

Rica. Coccocarpia adnata and C. glaucina are also new records for the Neotropics, and C. filiformis is new to 

Central America. Analysis of the available material shows ecological differentiation between species: 

Coccocarpia erythroxyli and C. palmicola have a wide ecological amplitude, being found from (semi-) arid to 

perhumid situations and from sea level to the subparamo zone at 3500 m, while Coccocarpia dissecta is 

mostly a lowland species typical of (semi-)arid to humid climates. The other taxa are confined to humid 

climates; while C. guimarana, C. glaucina, C. neglecta, and C. tenuissima seem to prefer lowland to 

submontane zones, the remaining species are chiefly montane, with C. domingensis, C. filiformis, C. pellita 

and C. prostrata reaching the subparamo zone. Coccocarpia as a whole is most abundant and most diverse in 

humid to perhumid, lower montane to montane situations, and the species generally prefer semi-exposed to 

exposed (micro-)habitats. 

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Introduction 

In 2002, we started TICOLICHEN, a project supported by the National Science Foundation 

(NSF) to unravel Costa Ricas tremendous lichen diversity (GÁMEZ et al. 1997; LÜCKING et 

al. 2004; CHAVES et al. 2004; LÜCKING 2005). The goals of this international collaboration 

between The Field Museum in Chicago, the Instituto Nacional de Biodiversidad (INBio) in 

Costa Rica, the Botanischer Garten & Botanisches Museum in Berlin, and various other 

institutions in the United States (University of Wisconsin-Madison, New York Botanical 

Garden), The Netherlands (Adviesbureau voor Bryologie en Lichenologie), and Austria 

(Karl-Franzens-Universität), are to enhance the taxonomic knowledge of tropical lichens at 

both the local and world-wide level, through a comprehensive, printed lichen biota of Costa 

Rica, tentatively scheduled for publication in 2008. Several genera and families have 

already been treated in individual assessments, in order to communicate improvements in 

our taxonomic understanding of tropical lichens (CHAVES et al. 2004; GRUBE et al. 2004; 

LÜCKING et al. 2006a, b; NELSEN et al. 2006; RIVAS PLATA et al. 2006). 

Here we present a preliminary treatment of Coccocarpia Pers., a genus of about 25, 

small-foliose, cyanophilic lichen species mainly found in the humid tropics, but with a few 

taxa extending into the northern and southern hemisphere. Coccocarpia forms the core 

genus of the Coccocarpiaceae, a small family of about five genera now confirmed to belong 

to suborder Collematineae within Peltigerales, together with Collemataceae, Pannariaceae, 

and Placynthiaceae (MIADLIKOWSKA & LUTZONI 2004; MIADLIKOWSKA et al. 2006). 

Coccocarpia was first treated in a modern sense by SANTESSON (1952), HENSSEN (1963) 

and HENSSEN & JAHNS (1974) and eventually monographed more than 20 years ago by 

ARVIDSSON (1983). Since then, remarkably few papers have been published with new 

discoveries. Only two new species were described, both from Venezuela (MARCANO et al. 

1995): Coccocarpia culatensis V. Marcano & A. Morales, related to C. epiphylla, and C. 

duidensis V. Marcano, Galiz & A. Morales, related to C. dissecta. 

An extensive treatment of Coccocarpia in Costa Rica was presented by DODGE (1933), 

who reported 11 names, four species and seven varieties, including the new species C. 

albida Dodge. Unfortunately, his treatment is completely untenable, since a revision of the 

rich material in FH (75 specimens) revealed these names to be a confusing mix of mostly 

C. erythroxyli, C. palmicola, and C. pellita, but also C. adnata, C. dissecta, and C. 

glaucina, and the overlooked new species, C. gallaicoi, described herein. During our 

TICOLICHEN survey, we identified 18 species of Coccocarpia, four of which could not be 

identified with any known species and which are here described as new, including the C. 

gallaicoi dedicated to David Galloway, and a further one which it tentatively identified as 

C. aff. gallaicoi. We also present a key and an ecological analysis of the genus in Costa 

Rica. 

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Material and Methods 

Specimens were mainly studied at The Field Museum (F) in Chicago, the Instituto Nacional 

de Biodiversidad (INBio) in Costa Rica, the Farlow Herbarium (FH) at Harvard University 

in Cambridge, Massachussetts, and the Botanischer Garten & Botanisches Museum in 

Berlin, Germany. At the Farlow Herbarium, we revised all the material of the genus 

reported for Costa Rica by Dodge (1933). Thallus morphology was assessed using a 

LEICA MS5 stereo microscope, and thin hand sections of thalli and apothecia were studied 

under a ZEISS Axioskop 2 compound microscope. Images were taken with a NIKON 

Coolpix 5400 digital camera through a ZEISS W-PI 10x/23 eyepiece directly attached to 

the stereo microscope. 

For the ecological analysis, we assessed environmental parameters for all studied 

collections when available, using three environmental gradients, based on HOLDRIDGE 

(1982) and LÜCKING (1995, 1997): (1) Precipitation regime (arid - semi-arid - subhumid - 

humid - perhumid); (2) altitude (lowland - colline - submontane - lower montane - montane 

- upper montane - subparamo); and (3) light exposure (fully shaded - shaded - intermediate 

- exposed - fully exposed). According to their range within each gradient, the species were 

ordinated within a two-dimensional grid, using precipitation regime and altitude as main 

axis and expressing light exposure by shades of black, grey, and white. 

The Species 

Our survey of recent and historical collections revealed a total of 18 taxa (Table 1) which 

are keyed out below. Nomenclature follows ARVIDSSON (1983) unless otherwise stated; no 

nomenclatural references are given here for names already treated in that monograph. 

Many of the names reported earlier by MÜLLER ARGOVIENSIS (1892, 1894) and DODGE 

(1933) had already been placed into synonymy by ARVIDSSON (1983), and thorough 

revision of the material at FH and G was necessary to elucidate their correct identifications 

(Table 2). 

Judging from their thallus and apothecial morphology, several of the taxa distinguished 

here can be considered primary versus secondary species (Table 3), and molecular 

phylogenetic analysis is required to see whether the species concepts hold in these cases. 

Studies in other groups, such as the Physciaceae, have shown that in many cases, primary 

and secondary species form part of either a single-species complex with variable dispersal 

strategy or several species which all may include both sexual and vegetative forms. 

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Table 1. Species of Coccocarpia recorded for Costa Rica. Detailed collection data, including color images, 

can be retrieved online from The Field Museum's Botany Collections Database at 

http://emuweb.fieldmuseum.org/botany/search_crf.php. 

__________________________________________________________________________________ 

Coccocarpia adnata L. Arvidss. 

Coccocarpia dissecta Swinsc. & Krog 

Coccocarpia domingensis Vain. 

Coccocarpia epiphylla (Fée) Kremp. 

Coccocarpia erythroxyli (Spreng.) Swinsc. & Krog 

Coccocarpia filiformis L. Arvidss. 

Coccocarpia gallaicoi Lücking, Chaves & Umaña sp. nov. 

Coccocarpia aff. gallaicoi Lücking, Chaves & Umaña 

Coccocarpia glaucina Kremp. 

Coccocarpia guimarana (Vain.) Swins. & Krog. 

Coccocarpia aff. imbricascens Nyl. 

Coccocarpia microphyllina Lücking & Aptroot sp. nov. 

Coccocarpia neglecta Aptroot & Lücking sp. nov. 

Coccocarpia palmicola (Spreng.) L. Arvidss. & D. Gall. 

Coccocarpia pellita (Ach.) Müll. Arg. 

Coccocarpia prostrata Lücking, Aptroot & Sipman sp. nov. 

Coccocarpia stellata Tuck. 

Coccocarpia tenuissima Müll. Arg. 

__________________________________________________________________________________ 

The morphology and position of isidia and lobules plays an important role in distinguishing 

species of Coccocarpia. The distinction of isidia, phyllidia, lobules, and other similar 

structures, is not always clearcut in the literature (see discussion in ARVIDSSON 1983: 12), 

and we have here used a combination of morphological and functional definitions (Table 

4). The term "phyllidia" was originally introduced for flattened isidia, but in our opinion 

covers perfectly the marginal isidioid, strongly branched structures found, for example, in 

Sticta species (MCDONALD et al. 2003). The structures have the same color as the thallus 

(while true isidia in Sticta are usually darker) and do not seem to act as dispersal organs, 

but rather increase the thallus surface for enhanced gas exchange. So defined, phyllidia 

differ from both flattened isidia (in function) and accessory lobules (in the absence of 

rhizines). The new species Coccocarpia microphyllina is the first in the genus in which 

such phyllidia are found; all other species either produce isidia in the strict sense or 

accessory lobules. 

Table 2. Revision of names in Coccocarpia published by DODGE (1933) for Costa Rica and on labels of 

specimens in FH, their current nomenclatural status, and taxonomic identification of the material (see also list 

of cited specimens under each taxon entry below). *names not published by Dodge (1933) but used on 

specimen labels, **superfluous new combinations published by DODGE (1933) of names already recombined 

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previously by Vainio and Zahlbruckner, ***unpublished new combinations made by Dodge on specimen 

labels. 

Name Current name Correct identification 

albida Dodge palmicola palmicola (type), glaucina 

cronia (Tuck.) Vain. palmicola palmicola, pellita, dissecta 

var. furfuracea (Müll. Arg.) Vain.* pellita palmicola, erythroxyli 

var. granulosa (Müll. Arg.) Dodge** erythroxyli palmicola 

var. isidiophylla (Müll. Arg.) Dodge** palmicola palmicola, dissecta 

var. isidiosa (Müll. Arg.) Vain.* palmicola dissecta, palmicola, gallaicoi 

var. lividorufa (Meyen & Flotow) Dodge** pellita pellita, palmicola, erythroxyli 

var. prolificans (Malme) Dodge** pellita pellita 

elegans Müll. Arg. stellata stellata 

isidiophylla (Müll. Arg.) ined.*** palmicola palmicola, dissecta 

var. granulosa (Müll. Arg.) ined.*** palmicola palmicola, dissecta, glaucina 

var. prolificans (Müll. Arg.) ined.*** pellita pellita, palmicola 

pellita (Ach.) Müll. Arg. pellita pellita 

var. parmelioides (Hook.) Müll. Arg. erythroxyli erythroxyli, palmicola, glaucina 

var. pyrrhichocarpa Hue erythroxyli erythroxyli 

var. smaragdina (Pers.) Müll. Arg.* smaragdina erythroxyli 

var. strigosa Müll. Arg. erythroxyli erythroxyli, adnata 

tenuissima Müll. Arg.* tenuissima neglecta 

Table 3. Presumed primary and secondary species in the genus Coccocarpia in Costa Rica, separated by type 

of isidia present. 

Thallus type Primary species Secondary species 

cylindrical / flattened isidia squamiform isidia phyllidia 

broadly flabellate C. erythroxyli C. palmicola C. pellita 

linear dissected C. adnata C. dissecta 

squamulose C. aff. gallaicoi C. gallaicoi 

narrowly flabellate C. epiphylla C. microphyllina 

linear spaced C. stellata C. domingensis C. neglecta 

Table 4. Morphological and functional definitions used here for isidia, phyllidia, accessory lobules, and 

similar structures in the genus Coccocarpia. 

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Structure Section Symmetry Position Orientation Rhizines Function Example 

isidia round radiately ± laminal vertical — dispersal palmicola 

(cylindrical) 

symmetrical 

isidia flattened bisymmetrical ± laminal oblique — dispersal domingensis 

(flattened) (± unbranched) 

isidia flattened dorsiventral ± laminal horizontal — dispersal pellita 

(squamiform) (± unbranched) 

phyllidia flattened dorsiventral ± marginal horizontal — ?surface microphyllina 

(± branched) 

increase 

accessory flattened dorsiventral ± marginal horizontal present ?surface several species 

lobules 

increase 

Key to the species of Coccocarpia in Costa Rica 

1 Cylindrical to flattened or squamiform isidia or phyllidia present or thallus ascendent to fruticulose ............2 

1 Isidia absent and thallus flattened-foliose ......................................................................................................12 

2 Thallus with richly branched phyllidia or secondary lobes or ascendent to fruticulose...................................3 

2 Thallus with cylindrical to flattened or squamiform isidia...............................................................................5 

3 Thallus fruticulose, lobes less than 0.1 mm wide...........................................................................C. filiformis 

3 Thallus foliose but with branched marginal phyllidia or partially ascendent secondary lobes, lobes 0.2–1 

mm wide .......................................................................................................................................................4 

4 Lobes flabellate, terminally up to 1 mm wide, marginally with branched phyllidia .............. C. microphyllina 

4 Lobes linear to very slightly flabellate, up to 0.4 mm wide, with numerous thin (0.1–0.3 mm wide), partially 

ascendent and richly branched (especially at tips) secondary lobes........................................... C. prostrata 

5 Lobes broadly flabellate to almost semicircular, (1–)2–7(–12) mm wide........................................................6 

5 Lobes linear to narrowly flabellate, 0.2–1(–1.5) mm wide ..............................................................................9 

6 Isidia distinctly flattened, squamiform ................................................................................................C. pellita 

6 Isidia cylindrical to coralloid............................................................................................................................7 

7 Lobes dark grey when dry, with thin white longitudinal striae, rather short and imbricate, 1–2 mm wide, 

thallus therefore appearing squamulose.......................................................................................C. gallaicoi 

7 Lobes light grey to bluish grey when dry, lacking distinct white striae, elongate, adjacent but usually not 

imbricate, usually 2–7(–12) mm wide, thallus distinctly foliose..................................................................8 

8 Lobes flat; isidia laminal ...............................................................................................................C. palmicola 

8 Lobes slightly to distinctly canaliculate; isidia mostly marginal.....................................................C. glaucina 

9 Lobes mostly 0.5–1 mm wide, adjacent, lacking distinct white striae; usually corticolous ...........................10 

9 Lobes mostly 0.2–0.5 mm wide, leaving distinct interspaces, with thin white longitudinal striae; usually 

foliicolous...................................................................................................................................................11 

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10 Lobes rather thick (70–130 µm), linear, with well-developed layer of black rhizines; ascospores ellipsoid, 

9–14 × 3–4 µm ............................................................................................................................. C. dissecta 

10 Lobes rather thin (50–80 µm), flabellate, with sparse, white rhizines; ascospores not observed 

.................................................................................................................................................C. guimarana 

11 Isidia distinctly flattened, squamiform, horizontal; apothecia usually abundant........................... C. neglecta 

11 Isidia cylindrical to slightly flattened, elongate, erect; apothecia rare ................................... C. domingensis 

12 Lobes broadly flabellate to almost semicircular, (1–)2–7(–12) mm wide....................................................13 

12 Lobes linear to narrowly flabellate, 0.2–1(–1.5) mm wide ..........................................................................15 

13 Lobes elongate, adjacent or partly overlapping but not imbricate, 2–7(–12) mm wide, thallus distinctly 

foliose ...................................................................................................................................... C. erythroxyli 

13 Lobes rather short and imbricate, 1–2 mm wide, thallus therefore appearing squamulose..........................14 

14 Lobes dark grey with white longitudinal striae, apically curved downward.......................... C. aff. gallaicoi 

14 Lobes pale grey and lacking white striae, apically slightly curved upward ....................C. aff. imbricascens 

15 Lobes mostly 0.5–1 mm wide, adjacent .......................................................................................................16 

15 Lobes mostly 0.2–0.5 mm wide, leaving distinct interspaces ......................................................................17 

16 Lobes rather thick, linear, with well-developed layer of black rhizines; surface lacking distinct white 

striae; ascospores ellipsoid, 8–12 × 3–4 µm...................................................................................C. adnata 

16 Lobes rather thin, flabellate, with sparse, white rhizines; surface with thin white longitudinal striae; 

ascospores shortly ellipsoid, 4–6 × 3–4 µm ............................................................................... C. epiphylla 

17 Lobes 0.05–0.1 mm wide, leaving large interspaces................................................................. C. tenuissima 

17 Lobes 0.2–0.5 mm wide, leaving small interspaces ........................................................................ C. stellata 

Coccocarpia adnata L. Arvidss. Fig. 1A 

Notes: Three specimens are assigned here to this taxon, which is newly reported for the 

Neotropics, previously known from the eastern Paleotropics only (ARVIDSSON 1983). One 

specimen (Lücking 15579d) agrees well with paleotropical material, although in bad 

condition and with partly broken lobes. The other three specimens (Dodge 10660, Lücking 

17252e, Sipman 53299) are intermediate between Coccocarpia adnata and the narrowlobed 

form of C. erythroxyli called var. strigosa (see below), sharing the narrow, dissected 

lobes with the first and the rounded lobe tips with the second. These specimens are here 

tentatively referred to C. adnata, but more detailed studies in the C. erythroxyli-complex 

are necessary to sort out their correct taxonomic placement. 

Selected specimens examined: COSTA RICA. Alajuela: Volcán Tenorio National Park, Mar 2004, 

Lücking 17252e (F). Heredia: Potreros near Coleman Finca, Aug 1936, Dodge 10660 (FH; as Coccocarpia 

pellita var. strigosa). Puntarenas: San Vito de Coto Brus, Las Cruces Biological Station, Oct 2004, Sipman 

53299 (B, INB-3993704). Las Tablas Protection Zone, Jun 2002, Lücking 15579d (F). 

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Coccocarpia dissecta Swinsc. & Krog Fig. 1G 

Notes: Previously known from the Paleotropics only (SWINSCOW & KROG 1976; 

ARVIDSSON 1983; DIN et al. 1998), but recently also reported from Paraguay (LLARENS 

2002). Several specimens were observed in the Costa Rican material which match the 

paleotropical populations perfectly. The almost linear, richly dissected lobes, with the black 

rhizines visible laterally along the lobe margins, distinguish this taxon from Coccocarpia 

palmicola. Coccocarpia adnata is identical in thallus morphology but lacks isidia and can 

be considered the primary species of C. dissecta. 

A species closely related to Coccocarpia dissecta, C. duidensis, was described from 

Venezuela by MARCANO et al. (1995). This taxon seems to be rather distinctive, differing 

from C. dissecta not only by its shorter ascospores but also by its chemistry, including 

anthraquinones and a xanthone. 

Selected specimens examined: COSTA RICA. Alajuela: Altamera, s.dat., Holm & Iltis 1529 (FH; as 

Coccocarpia cronia). Cartago: Santiago, Oct 1929, Dodge 4638 (FH; as C. cronia var. isidiosa). 

Guanacaste: Barra Honda National Park, Mar 2004, Will-Wolf 12834f (INB-4031112, WIS). Between Río 

Las Cañas and Río San José, Feb 1930, Dodge & Thomas 6719 (FH; as C. cronia var. isidiosa). Hacienda 

Santamaría, Jan 1930, Dodge & Thomas 7007 (FH; as C. cronia var. isidiosa). Palo Verde National Park, 

Mar 2003, Buck 43929 (INB, NY). Heredia: La Selva Protection Zone, Jun 2002, Lücking 15026a (INB- 

3932456), 15026b (F). Limón: Between Río Cimarrones and Waldeck Farm and lake on Monte Verde Farm, 

Apr 1930, Dodge s.n. (FH; as C. isidiophylla var. granulosa). Gandoca-Manzanillo Wildlife Refuge, Mar 

2004, Lücking 17086h (F). Puntarenas: Along seashore between Río Sandoval and Río Tigre, May 1930, 

Dodge 8057 (FH; as C. isidiophylla). Flood-plain of Río Sándalo, peninsula of Osa, Aug 1936, Dodge & 

Goerger 10048 (FH; as C. cronia var. isidiophylla). Las Tablas Protection Zone, Jun 2002, Lücking 15593b 

(F, INB-3981686). San José: N of Turrúcares, Mar 1930, Dodge & Thomas 8056 (FH; as C. isidiophylla var. 

granulosa). 

Coccocarpia domingensis Vain. Fig. 3A–B 

Notes: A very common foliicolous species, characterized by linear lobes and marginal, 

cylindrical to slightly flattened isidia (SANTESSON 1952; ARVIDSSON 1983). Careful study 

of the available material showed that another species was hidden within this taxon, 

distinguished by its squamiform isidia; it is described below as Coccocarpia neglecta. 

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Fig. 1. Habit of Coccocarpia species. A. Coccocarpia adnata (Lücking 15579d). B. 

Coccocarpia erythroxyli (Lücking 15148a). C. Coccocarpia palmicola (Lücking 

15199a). D. Coccocarpia guimarana (Lücking 15615d). E–F. Coccocarpia pellita 

(Lücking 17252b and Will-Wolf 12735b), with phyllidia (E) and squamulose isidia 

(F). G. Coccocarpia dissecta (Lücking 17086k). H. Coccocarpia glaucina (Chaves 

128). Scale = 2 mm, in E and F = mm. 

437

Selected specimens examined: COSTA RICA. Cartago: Tapantí National Park, Mar 2004, Aptroot 60982 

(ABL, INB-4021820); Apr 2003, Trest 1365 (GZU). Heredia: La Selva Protection Zone, Jul 1997, Lücking 

97-1486 (F); Oct 2002, Lücking 15662 (CR, INB-4030117, USJ), 16001g (B, CR, F, INB-4031113, USJ), 

16002f (F). 

Coccocarpia epiphylla (Fée) Kremp. Fig. 2A–B 

Notes: New to Costa Rica, already listed in our online checklist 

(http://www.fieldmuseum.org/ticolichen/checklist.html). The neotropical species belongs in 

the Coccocarpia stellata group and is rather similar to the latter, differing mainly by its 

broader, flabellate lobes and slightly more elongated ascospores. 

MARCANO et al. (1995) described the new species Coccocarpia culatensis from 

Venezuela, being similar to C. epiphylla but differing by its lichenicolous instead of 

foliicolous growth and its subglobose versus fusiform ascospores. Our material shows that 

C. epiphylla commonly grows on bark as well, and the 'lichenicolous' growth over other 

foliose lichens can certainly not be used as a specific character for C. culatensis. 

ARVIDSSON (1983) gives the ascospores of C. epiphylla as 6–7 × 2–4 µm, but according to 

our observations they are mostly 6 × 4 µm, thus slightly elongate. This variational 

amplitude perfectly includes the specimen representing the type of C. culatensis, and also 

the habit photograph (MARCANO et al. 1995: 220) indicates that it deals with C. epiphylla. 


Aptroot 60512 (ABL, INB-3967666). Cartago: Tapantí National Park, Apr 2003, Will-Wolf 12540l (WIS). 

Guanacaste: Palo Verde National Park, Mar 2003, Lücking 16514d (INB-3987277). Puntarenas: La Amistad 

International Park, Jun 2002, Lücking 15221a (INB-3978438), 15235k (F), 15239h (INB-3976200), 15240ab 

(F). 

Coccocarpia erythroxyli (Spreng.) Swinsc. & Krog Fig. 1B 

Notes: A very common and widespread, quite variable taxon (ARVIDSSON 1983). 

Typically, the apothecia are convex and lack laterally spreading rhizines; their color varies 

from orange to dark red to almost black, apparently depending on the degree of sun 

exposure. ARVIDSSON (1983) reports and illustrates a narrow-lobed form (f. ciliata Müll. 

Arg. and f. strigosa Müll. Arg.) with rhizinate apothecia from Mexico, Ecuador, and Brazil. 

We have observed the same form in Costa Rica. Molecular studies are required in this 

species and also in Coccocarpia palmicola to better assess the enormous morphological 

variation in these taxa. 

Selected specimens examined: COSTA RICA. Alajuela: La Palma de San Ramón, Oct 1929, Brenes 41 

(FH; as Coccocarpia pellita var. parmelioides). Santiago de San Ramon, Dec 1929, Brenes 238 (FH; as C. 

pellita var. smaragdina). Volcán Tenorio National Park, Mar 2004, Aptroot 60451 (ABL, INB-3964128), 

Lücking 17234 (CR), 17251r (INB-4004879). Cartago: E terrace of Río Birris, Oct 1929, Dodge & Thomas 

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4708 (FH; as C. pellita var. smaragdina). Potrero and bananal below railroad, Oct 1929, Dodge 8050 (FH; as 

C. pellita var. pyrrhichocarpa), 8051 (FH; as C. pellita var. strigosa). Tapantí National Park, Jul 2002, 

Lücking 15415 (F), 15459 (INB-3979984); Apr 2003, Lücking 16606b (F, INB-4002728), Sipman 51052 (B, 

INB-3998684); Jul 2002, Will-Wolf 10171b (INB, WIS). Guanacaste: Hacienda Santamaría, Jan 1930, 

Dodge & Thomas 4709 (FH; as C. pellita var. parmelioides), 6813 (FH; as C. pellita var. smaragdina). 

Monte Alto Forest Reserve, Mar 2004, Sipman 52011 (B, INB-4031185), Will-Wolf 12816q (INB-4031186, 

WIS). NE of Tilaran, Pacific slope, Feb 1930, Dodge & Thomas 6559 (FH; as C. cronia var. furfuracea). 

Palo Verde National Park, Mar 2003, Trest 1334b (INB-4031188, WIS). Upper portion of Cañon of Río San 

José, Feb 1930, Dodge & Thomas 6581 (FH; as C. pellita var. parmelioides). Upper slopes of Cerro San José 

de Libano, Feb 1930, Dodge et al. 6684 (FH; as C. pellita var. parmelioides). Heredia: Potreros near 

Coleman Finca, Aug 1936, Dodge 10184 (FH; as C. cronia var. lividorufa). Limón: Between Río Cimarrones 

and Waldeck Farm and lake on Monte Verde Farm, Apr 1930, Dodge 7402 (FH; as C. pellita var. 

parmelioides). Gandoca-Manzanillo Wildlife Refuge, Mar 2004, Lücking 17086i (F). Guapiles, Nov 1988, 

Sipman et al. 41893 (B, CR). Puntarenas: La Amistad International Park, Jul 2002, Lücking 15245a (F, INB- 

3976210), 15258 (F), Sipman 48037b (B, INB-3944629). Las Cruces Biological Station, Oct 2004, Sipman 

53280 (B, INB-3993688). Las Tablas Protection Zone, Jun 2002, Lücking 15148a (F, INB-3974792), Will- 

Wolf 10061g (INB-4031193, WIS). Manuel Antonio National Park, Nov 1988, Sipman 42335 (B, CR). Near 

Río Sandalo, Apr 1930, Dodge 7742 (FH; as C. pellita var. prolificans). Puerto Jimenez de Osa (Golfo 

Dulce), Apr 1930, Brenes 825a (FH; as C. pellita var. parmelioides). San José: Chirripó National Park, Llano 

Bonito, Nov 1988, Kappelle & Monge 3430 (B, CR). Los Santos Forest Reserve, Jul 2002, Lücking 15349 

(F), Will-Wolf 10156c (WIS). 

Coccocarpia filiformis L. Arvidss. Fig. 3H 

Notes: Previously known from South America and Madagascar (ARVIDSSON 1983; 

APTROOT 1990). The species at first glance resembles a Polychidium, but is more robust, 

and the tiny ascendent lobes are actually dorsiventral and produce sparse rhizines. If 

present, the often rhizinate apothecia immediately indicate the taxonomic affinities of this 

lichen. 

Selected specimens examined: COSTA RICA. Alajuela: San Carlos, Boca Tapada, Chaves 2241 (INB- 

3942439), Chaves 2259 (INB-3942457), Chaves 2321 (INB-3942540).Volcán Tenorio National Park, Mar 

2004, Aptroot 60470 (ABL, INB-3974792), Lücking 17252c (F, INB-4004881, USJ). Cartago: Tapantí 

National Park, Jul 2002, Will-Wolf 10209b (WIS). Guanacaste: Volcán Tenorio National Park, Mar 2004, 

Aptroot 60652 (ABL, INB-3968066). Heredia: La Selva Protection Zone, Oct 2002, Lücking 16002f (F). 

Puntarenas: Terraba Valley, Clavera, Potrero Grande savannas, Jul 2002, Sipman 48159b (B, INB-3946555). 

439

Fig. 2. Habit of Coccocarpia species. A–B. Coccocarpia epiphylla (Lücking 

15239h and s.n.). C–D. Coccocarpia gallaicoi (holotype). E. Coccocarpia aff. 

gallaicoi (Lücking 15240a). F. Coccocarpia aff. imbricascens (Chaves 2556). G– 

H. Coccocarpia microphyllina (Sipman 48010l and holotype). Scale = 2 mm, in D 

= 1 mm. 

440

Fig. 3. Habit of Coccocarpia species. A–B. Coccocarpia domingensis (Lücking 

1486 and Lücking 462). C–F. Coccocarpia neglecta (isotype). F. Coccocarpia 

tenuissima (holotype). H. Coccocarpia filiformis (Lücking s.n.). Scale = 1 mm. 

441

Coccocarpia gallaicoi Lücking, Chaves & Umaña sp. nov. Fig. 2C–D 

Species nova similis Coccocarpia epiphylla sed lobis brevioribus isidiis instructis differt. 

Typus: COSTA RICA. San José: South wall of canon of Río Virilla between El Brazil and Santa Ana, May 

1930, Dodge 7777 (FH – holotype; originally identified as Coccocarpia cronia var. isidiosa). 

Thallus corticolous, foliose-squamulose, covering irregular areas of up to 50 mm diam. 

Lobes thin, short, ascendent, 1–2 mm wide, broadly flabellate, shallowly branched, densely 

imbricate and giving the thallus a squamulose appearance; lobe apices curved downward. 

Upper surface dark blue-green when moist, dark grey when dry, with thin, longitudinal, 

white striae. Lower surface black, with numerous, black, unbranched, 0.1–0.2 mm long 

rhizines forming a mat, not or scarcely protruding from under the lobe margins. Lobe 

surface isidiate. Isidia laminal, numerous, erect, cylindrical, partly branched, 0.2–0.4 × 

0.08–0.1 mm, creme-colored but apices dark grey. Apothecia not observed. Pycnidia not 

observed. 

Chemistry: No substances detected by TLC. 

Notes: This very conspicuous new species was hidden under the name Coccocarpia 

cronia var. isidiosa in the Dodge herbarium at FH. While the latter is a synonym of C. 

palmicola (ARVIDSSON 1983), the present specimen turned out to have exactly the same 

thallus morphology as the non-isidiate material named C. aff. gallaicoi below. Both belong 

in the C. epiphylla group, but are characterized by rather short lobes that give the 

impression of a squamulose thallus. Superficially most similar is C. imbricascens, but that 

species has a pigmented medulla and more delicate, light grey lobes and lacks isidia (see 

below). 

Distribution and Ecology: So far only known from the type locality in a montane 

rainforest. 

Etymology: We name this new species after our colleague and friend David, the 

subject of this splendid compilation, on the occasion of his 65th birthday. To avoid the 

usual 'gallowayi', already found in several lichen names, we looked up the meaning of the 

name Galloway and came up with 'foreign Gael'. The Gaels are an ethno-linguistic group in 

Ireland, Scotland and the Isle of Man, speaking one of the Gaelic languages and 

presumably originating from Gallaecia, a Roman province that comprised a territory in the 

north-west of Hispania (present-day Galicia in Spain and northern Portugal). The Romans 

gave the name Gallaecia to that province after the tribe of the 'Gallaicoi', who had been 

their foremost enemy in the region, and we choose that name as epithet for your new 

species in honour of David Galloway. 

442

Coccocarpia aff. gallaicoi Lücking, Chaves & Umaña Fig. 2E 

Notes: This taxon keys out as Coccocarpia imbricascens in Arvidsson's (1983) key and 

also fits the description, but differs in its dark grey, more robust lobes with distinct white 

striae and apical margins curved downward, and its white medulla, among other characters. 

It has exactly the same lobe morphology as C. gallaicoi (see above), but lacks isidia and 

represents the non-isidiate counterpart of that species. However, since there is only one 

collection, we opted for not describing this taxon formally. 

Selected specimens examined: COSTA RICA. Puntarenas: La Amistad International Park, Jun 2002, 

Lücking 15240aa (F). 

Coccocarpia glaucina Kremp. Fig. 1H 

Notes: Previously known from the eastern Paleotropics only (ARVIDSSON 1983; 

MAKHIJA et al. 1999). The collections from Costa Rica fit the paleotropical material well; 

they can be distinguished from Coccocarpia palmicola by the canaliculate lobes with 

mostly marginal isidia. 

Selected specimens examined: COSTA RICA. Alajuela: San Carlos, Boca Tapada, Chaves 2240 (INB- 

3942438). Limón: Finca Castilla, Jul 1936, Dodge & Goerger s.n. (FH; as Coccocarpia albida). W bank of 

Río Siquirres, Dec 1929, Dodge et al. 5595 (FH; as C. isidiophylla var. granulosa). Puntarenas: Flood-plain 

of Río Sandalo, peninsula of Osa, Aug 1936, Dodge & Goerger 9902 (FH; as C. pellita var. parmelioides). 

Monteverde, San Gerardo Biological Station, Chaves 128 (INB- 3762776). 

Coccocarpia guimarana (Vain.) Swinsc. & Krog Fig. 1D 

Notes: The present material, three small but well-developed thalli from the same 

locality, was first identified as Coccocarpia dissecta, due to the comparatively narrow 

lobes. However, a detailed study of its morphology showed that this taxon is distinct in its 

narrowly flabellate, very thin lobes which lack a pronounced hypothallus. It is thus more 

similar to C. epiphylla and narrow- and small-lobes forms included by ARVIDSSON (1983) 

in C. palmicola. The latter author interprets these forms, exemplified by the type material 

of C. guimarana, as ecotypes of C. palmicola typical of lowland rainforest. However, we 

have collected very typical C. palmicola in Costa Rican lowland rainforest and even in dry 

and coastal areas, and they agree well with the montane forms in their broad, rounded 

lobes. The photographs of C. palmicola and C. guimarana depicted by ARVIDSSON (1983: 

73), as well as the present material, suggest that the narrow- and small-lobed form is 

specifically distinct, and following SWINSCOW & KROG (1976), the name C. guimarana is 

here tentatively reinstated, pending further study of this difficult complex of species. We 

have not seen the type material of C. cronia var. microphyllina Vain., but acording to the 

443

statements by ARVIDSSON (1983: 74), this might represent a synonym of C. guimarana 

sensu SWINSCOW & KROG (1976). 

Distribution and Ecology: Pantropical, apparently typical of lowland rainforest with a 

more or less pronounced dry season. 

Selected specimens examined: COSTA RICA. Puntarenas: Carara National Park, Jul 2002, Lücking 

15615d (F, INB-3980902, USJ). 

Coccocarpia aff. imbricascens Nyl. Fig. 2F 

Notes: ARVIDSSON (1983) gives only the type as belonging to this species, with 

unknown locality data and thus unknown distribution. In the meanwhile, the taxon has been 

reported from South America (Venezuela, Guyana, Brazil) and the eastern Paleotropics 

(Thailand) and thus seems to be possibly pantropical (MARCANO et al. 1995). However, as 

discussed under Coccocarpia aff. gallaicoi, the identification of this taxon is not 

straightforward and the aforementioned citations might include misidentifications. We have 

studied the type specimen of C. imbricascens and found that it has a yellow pigmented 

medulla, a fact not mentioned by ARVIDSSON (1983). The present material agrees perfectly 

with the type, except that the medulla is unpigmented. Since yellow pigments (both cortical 

and medullary) are rare within Coccocarpia, the present material seems to be specifically 

different from C. imbricascens, but is to scanty to allow a formal description. Both taxa are 

characterized by their very short, rounded, imbricate lobes, which give the thallus an almost 

squamulose appearance, similar to C. gallaicoi and C. aff. gallaicoi. 

Selected specimens examined: COSTA RICA. Guanacaste: Volcán Tenorio National Park, Alto Masis, 

Mar 2006, Chaves 2556 (INB-4010098). 

Coccocarpia microphyllina Lücking & Aptroot sp. nov. Fig. 2G–H 

Species nova similis Coccocarpia epiphylla sed lobulis phyllidiis marginalis instructis differt. 

Typus: COSTA RICA. Puntarenas: La Amistad International Park, Jun 2002, Lücking 15212a (F – holotype). 

Thallus corticolous, foliose, starting as round colonies but covering irregular areas of up 

to 20 mm diam. Lobes thin, flat, 0.4–0.7 mm wide but tips more or less flabellate and up to 

1 mm wide, anisotomically branched and dissected, leaving rather large interspaces; lobe 

apices slightly ascending but tips curved downward. Upper surface greenish grey when 

moist, plumbeous to bluish grey when dry, with thin, longitudinal, white striae. Lower 

surface black, with numerous, black, unbranched, 0.1–0.2 mm long rhizines forming a mat, 

not or scarcely protruding from under the lobe margins. Lobe margins phyllidiate. Phyllidia 

numerous, ascending, elongated, branched, 0.1–0.4 × 0.05–0.1 mm, forming a 

characteristic dissected rim. Apothecia rare, up to 3 mm diam; disc orange-brown to red- 

444

own, more or less flat and with indistinct, concolorous margin, with scattered white, 

stellately arranged, 0.3–0.5 mm long rhizines attaching the apothecia to the thallus. 

Ascospores broadly ellipsoid to subglobose, 5–7 × 4–6 µm. Pycnidia not observed. 


Notes: This new species is most similar to Coccocarpia epiphylla in thallus characters, 

but differs in the presence of marginal, branched, dorsiventral phyllidia. So far, phyllidia as 

defined here (see above) were unknown in the genus Coccocarpia (ARVIDSSON 1983), but 

similar structures are known from other Peltigerales, e.g. in Sticta carolinensis 

(MCDONALD et al. 2003). Coccocarpia guimarana is also similar in thallus morphology, 

but differs in having laminal, cylindrical isidia. 

Coccocarpia prostrata, another new species described below, is superficially similar in 

having ascending lobes that resemble the phyllidia of C. microphyllina. However, these 

lobes are clearly of primary origin, while in the latter, the phyllidia are secondary lobules 

produced marginally along the primary lobes. Also, the main lobes in C. prostrata are 

linear, rather than flabellate, and narrower. 

Distribution and Ecology: Known from a few collections in wet montane rainforest. 

Selected specimens examined: COSTA RICA. Cartago: Irazu National Park, July 2002, Sipman 48327 

(B, INB-3986727). Tapantí National Park, Apr 2003, Trest 1375e (INB-4031197, WIS). Guanacaste: Novasa 

Wind Power Project, Mar 2004, Will-Wolf 12915k (INB-4031198, WIS). Puntarenas: La Amistad 

International Park, Jun 2002, Sipman 48010l (B, INB-3927515), 48020e (B, INB-3927547). Puntarenas: La 

Amistad International Park, Jun 2004, Chaves 1737 (INB-3939332) 

Coccocarpia neglecta Aptroot & Lücking sp. nov. Fig. 3C–F 

Species nova similis Coccocarpia domingense sed isidiis squamiformis differt. 

Typus: COSTA RICA. Cartago: Jicotea near Turrialba, Jul 1991, Lücking 1243a (CR – holotype; ABL, F, B, 

INB, USJ – isotypes). 

Thallus foliicolous, foliose, in round colonies of up to 30 mm diam. Lobes thin, flat, 

0.2–0.4 mm wide, linear, isotomically branched and dissected, leaving rather large 

interspaces; lobe tips curved downward. Upper surface blue-green when moist, plumbeous 

to bluish grey when dry, with thin, longitudina, white striae. Lower surface pale, with 

numerous, white, unbranched, tapering, 0.3–0.5 mm long rhizines not forming a mat but 

protruding from under the lobe margins and easily visible from above. Lobes isidiate. Isidia 

marginal and laminal, numerous, flattened, slightly raised, semicircular to somewhat 

elongated, sometimes crenulate, 0.1–0.2 × 0.07–0.15 mm, in mature thalli covering most of 

the thallus interior. Apothecia abundant, originating on the lobe margins but soon wider 

than the individual lobes, up to 3 mm diam; disc orange brown, more or less flat with a 

thin, slightly raised, concolorous margin, becoming immarginate and convex when old, 

445

with numerous white, stellately arranged, 0.3–0.5 mm long rhizines attaching the apothecia 

to the thallus. Ascospores globose, 4–5 µm diam. Pycnidia not observed. 


Notes: Until recently, this new taxon was not properly recognized, hence the epithet 

neglecta, but considered to represent an extreme form of Coccocarpia domingensis, 

another commonly foliicolous species with thin, linear lobes and isidia, However, the isidia 

in C. domingensis are of a different nature, being cylindrical to somewhat flattened but 

always elongated and erect rather than horizontal. Also, apothecia are much less common 

in C. domingensis. Coccocarpia stellata is also similar but completely lacks isidia or 

lobules. The three species can often be found side by side on the same leaves, especially 

Citrus leaves in humid, warm conditions. 

The isidia of Coccocarpia neglecta can be confused with secondary lobules but, for 

example contrary to the phyllidia in C. microphyllina, clearly represent diaspores and thus 

must be interpreted as isidia, Very similar isidia are known from C. pellita. 

Distribution and Ecology: A common foliicolous species in exposed situations in 

areas of wet tropical rainforest from sea level to 2000 m; also known from Ecuador 

(reported as Coccocarpia domingensis in Lücking, Lich. Fol. Exs. 194). 

Selected specimens examined: COSTA RICA. Heredia: La Selva Biological Station, Jun 1997, Lücking 

97-989 (F); Jul 1997, Lücking 97-1358 (F). Limón: Finca Hamburg, Aug 1936, Dodge & Goerger 10271 

(FH; as Coccocarpia domingensis). Forest along new tramline beyond bridge over Río Parismina on Finca 

Siam, Jul 1936, Dodge & Goerger 9530 (FH; as C. tenuissima). Puntarenas: La Amistad International Park, 

July 2002, Sipman 48039 (B, INB-3944635). 

ECUADOR. Pichincha: Guajalito Biological Station, May 1996, Lücking 96-292 (F). 

Coccocarpia palmicola (Spreng.) L. Arvidss. & D. Gall. Fig. 1C–D, 4A–H 

Notes: This is the most common species in the genus, with a wide morphological and 

ecological amplitude (ARVIDSSON 1983). Our observations both in the field and in the 

laboratory confirm ARVIDSSON's statements that lowland forms tend to be more adnate and 

often more narrow-lobed, while montane populations are usually loosely attached and 

broad-lobed. We also observed that the lowland forms (Fig. 4A–D) mostly have blue-black 

rhizines and rather compact, clavate isidia, while the montane populations (Fig. 4E–H) 

feature white rhizines and irregularly elongate, often almost budding, cylindrical isidia. 

Since intermediate forms exist, we follow ARVIDSSON's concept and keep all these forms 

under a single species, but it will be interesting to study this intriguing variation with 

statistical and molecular tools based on a large specimen sampling. 

Selected specimens examined: COSTA RICA. Alajuela: Alto de La Palma de San Ramón, Jan 1930, 

Brenes s.n. (FH; as Coccocarpia isidiophylla var. prolificans). Alto del Mondongo del San Ramon, Jan 1930, 

446

Brenes s.n. (FH; as C. cronia). Cerros de Pata de Gallo a San Rafael de San Ramón, Dec 1929, Brenes 215 

(FH; as C. isidiophylla var. granulosa). La Palma de San Ramón, Oct 1929, Brenes 104 (FH; as C. cronia 

var. isidiophylla). San Pedro de San Ramón, Feb 1930, Brenes 459 (FH; as C. isidiophylla var. granulosa). 

Volcán Poás, Puente del Río Poás, Sep 1950, Dodge s.n. (FH; as C. cronia var. isidiophylla). Vicinity of Los 

Chiles, Río Frio, Aug 1949, Holm & Iltis 1506 (FH; as C. cronia). Volcán Tenorio National Park, Mar 2004, 

Aptroot 60418 (ABL, INB-3963861). Lücking 17215 (F, INB-4004852), 17252d (CR, F, INB-4004883, 

USJ), Nelsen 3760 (INB-4031204, WIS), 3773 (INB-4031205, WIS), Sipman 51770 (B, INB-4031206), 

Trest 1539 (INB-4031207), 1553b (INB-4031208). Cartago: Cangreja village, Oct 2002, Lizano et al. 49 

(CR, INB, USJ). CATIE agricultural research centre near Turrialba, Nov 1988, Sipman & Döbbeler 42392 

(CR). Cerro Carpintera, Nov 1929, Dodge & Thomas 4763 (FH; as C. cronia var. isidiosa). Finca Coliblanco, 

Oct 1929, Dodge s.n. (FH; as C. cronia). Hills S of Agua Caliente, Mar 1930, Dodge & Thomas 7087 (FH; as 

C. cronia var. isidiophylla). Paraiso village, Jul 2002, Lücking 15441c (INB-3980085). Río Macho village, 

Apr 2003, Grube 11764 (CR, GZU), Trest 1427b (INB-4031209), 1428a (INB-4031210, WIS). Santiago, 

May 1930, Dodge 8054 (FH; as C. isidiophylla var. granulosa). Tapantí National Park, Apr 2003, Lücking 

16562 (INB-4002568). Guanacaste: Along Río Liberia near Hacienda Santamaria, Jan 1930, Dodge & 

Thomas 8055 (FH; as C. isidiophylla var. granulosa). Barra Honda National Park, Mar 2004, Aptroot 60825 

(ABL, INB-4020489), 60845 (ABL, INB-4020712), Trest 1606f (INB-4031211). Hacienda Santamaría, Jan 

1930, Dodge & Thomas 6987 (FH; as C. cronia var. furfuracea), 6900 (FH; as C. cronia var. isidiosa). 

Lomas de Barbudal Biological Reserve, Mar 2003, Trest 1345a (GZU), Trest 1346c (CR, WIS), Will-Wolf 

12529d (WIS). Monte Alto Forest Reserve, Mar 2004, Aptroot 60787 (ABL, INB-4020406). NE of Tilaran, 

Atlantic slope, Feb 1930, Dodge & Thomas 6556 (FH; as C. isidiophylla var. granulosa). Palo Verde 

National Park, Mar 2003, Sipman 51008a (B, INB-3987468), 51021 (B, INB-3987487), Will-Wolf 12511a 

(CR, WIS). S of Liberia, Jan 1930, Dodge & Thomas 6582 (FH; as C. cronia var. granulosa). Valley below 

Tilaran, Feb 1930, Dodge & Thomas 8256 (FH; as C. cronia var. isidiophylla). Volcán Tenorio National 

Park, Mar 2004, Will-Wolf 12792c (INB-4031212). Heredia: Carrizal village, Jul 2002, Sipman 48370 (B, 

INB-3987180). Cerro Central de Zurqui, Dec 1929, Dodge et al. 6089 (FH; as C. cronia var. isidiophylla). La 

Selva Protection Zone, Jun 2002, Lücking 15022b (F, INB-3932449); Oct 2002, Lücking 16002f (F). Volcán 

Barva, March 1985, Sipman 20613 (B, CR). Limón: Braulio Carrillo National Park, Mar 2003, Grube 11228 

(GZU). Finca Castilla, Jul 1936, Dodge & Goerger 9172 (FH; as C. cronia var. granulosa). Gandoca- 

Manzanillo Wildlife Refuge, Mar 2004, Lücking 17086g (CR, USJ), 17093c (F), 17095b (CR), 17102h 

(INB), Sipman 51691 (B, IBN), 51691a (B, INB). Hamburg Farm, Mar 1930, Dodge & Neverman 7401 (FH; 

as C. albida). Cerere Biological Reserve, Mar 2004, Trest 1510b (INB), Will-Wolf 12719g (INB). 

Puntarenas: Corcovado National Park, Apr 2003, Sipman 51152b (B, CR), Will-Wolf 12575e (INB). Floodplain 

of Río Sándalo, peninsula of Osa, Sep 1936, Dodge & Goerger 10426 (FH; as C. cronia var. 

granulosa), 10164 (FH; as C. cronia var. isidiophylla), 10162 (FH; as C. cronia var. isidiophylla). Floodplain 

of Río Terrones, peninsula of Osa, Aug 1936, Dodge & Goerger 10331 (FH; as C. cronia var. 

isidiophylla). La Amistad International Park, Jun 2002, Lücking 15186a (F, INB-3975917), 15199a (CR, F, 

INB-3976003), 15307 (INB-3976366). Las Cruces Biological Station, Oct 2004, Sipman 53292 (B, INB- 

3993697). Las Tablas Protection Zone, Jun 2002, Lücking 15136a (INB-3974770), Sipman 47948b (B, INB- 

3924211), Trest 488 (INB). San José: Along road from San Jerónimo to Alto La Palma, Nov 1988, Sipman et 

al. 41583 (B, CR). Cerro de la Muerte, along Panamerican Highway, March 1985, Sipman & Chaverri 20744 

(B, CR). Finca Guayabillos, Jul 1936, Dodge & Goerger 9240 (FH; as C. cronia var. granulosa). Forest 

along Río Paquita, Aug 1936, Dodge & Goerger 9759 (FH; as C. cronia var. lividorufa), 9757 (FH). Los 

Santos Forest Reserve, Apr 2003, Grube 11804 (GZU, USJ); Jul 2002, Lücking 15551 (F), Trest 640a (INB- 

4031273). N of Turrúcares, Mar 1930, Dodge & Thomas s.n. (FH; as C. pellita var. parmelioides). Valley of 

San Gerardo de Dota, July 1989, Wijtzes 1090 (B). 

Coccocarpia pellita (Ach.) Müll. Arg. Fig. 1E–F 

447

Notes: Coccocarpia pellita is characterized by is flattened, squamiform isidia, as 

opposed to the cylindrical isidia in C. palmicola (ARVIDSSON 1983). In the material 

observed by us, we found two different types of isidia: semicircular and entire to 

marginally incised (Fig. 1F), and elongate-branched, resembling phyllidia (Fig. 1E). 

ARVIDSSON (1983) accepts this as variation within the species, which we follow here, but 

this matter requires attention since no such range of variation is observed in the other 

isidiate species of Coccocarpia. 

Selected specimens examined: COSTA RICA. Alajuela: La Palma de San Ramón, Oct 1929, Brenes 38a 

(FH; as Coccocarpia isidiophylla var. prolificans), 43 (FH; as C. pellita var. lividorufa). Piedades de San 

Ramón, Jan 1930, Brenes 385 (FH; as C. isidiophylla var. prolificans). Volcán Tenorio National Park, Mar 

2004, Aptroot 60482 (ABL, INB-3964245), Lücking 17235 (F, INB-4004859), 17252b (CR, F, USJ), Nelsen 

3779 (INB-4031292, WIS), 51781 (B, INB-4030472), Trest 1562 (INB-4031316), Will-Wolf 12735b (INB- 

4031317, WIS). Cartago: Camino al Río Diquís, Apr 1960, Dodge s.n. (FH; as C. cronia). E terrace of Río 

Birris, Oct 1929, Dodge & Thomas 4710 (FH; as C. cronia var. prolificans). La Chonta, Nov 1988, Sipman et 

al. 41788 (B, CR). Río Macho village, Apr 2003, Grube 11735 (GZU, INB). Tapantí National Park, Apr 

2003, Nelsen 3607a (INB-4031318, WIS). Valley of Río Reventazon, near Interamerican Institute of 

Agricultural Sciences, s.dat., Holm & Iltis 1473 (FH; as C. cronia var. prolificans). Guanacaste: Along Río 

Liberia near Hacienda Santamaria, Jan 1930, Dodge & Thomas 8053 (FH; as C. isidiophylla var. prolificans). 

Hacienda Santamaría, Jan 1930, Dodge & Thomas 8082 (FH; as C. cronia var. prolificans), 6999 (FH; as C. 

cronia var. prolificans). NE of Tilaran, Pacific slope, Feb 1930, Dodge & Thomas 6557 (FH; as C. pellita 

var. lividofusca). Limón: Finca Castilla, Jul 1936, Dodge & Goerger 9211 (FH; as C. cronia var. lividorufa), 

9249 (FH; as C. cronia var. lividorufa), 9213 (FH; as C. prolificans). W bank of Río Siquirres, Dec 1929, 

Dodge et al. 8058 (FH; as C. pellita var. lividorufa). San José: Along road from San Jerónimo to Alto La 

Palma, Nov 1988, Sipman et al. 41579 (B, CR), 41582 (B, CR). Los Santos Forest Reserve, Jul 2002, 

Lücking 15348 (INB-3976412). 

Coccocarpia prostrata Lücking, Aptroot & Sipman sp. nov. Fig. 5C–H 

Species nova similis Coccocarpia stellata sed lobulis pro parte irregulariter elongatis asdendentibus 

ramificantibusque differt. 

Typus: COSTA RICA. Cartago: Tapantí National Park, Apr 2003, Nelsen 3557d (INB–4031018, holotype; 

WIS – isotype). 

Thallus corticolous, foliose, starting as round colonies but eventually covering irregular 

areas of up to 50 mm across, with lobes in one to several planes. Lobes in first plane thin, 

flat, 0.2–0.4 mm wide, linear, isotomically branched, leaving interspaces, at lobe tips often 

multiply branched; lobe apices more or less ascending but tips curved downward and often 

bifurcate; lobes in second planes thin, irregularly ascending, 0.1–0.3 mm wide, linear, 

strongly isotomically branched and often radiating towards the lobe tips. Upper surface 

blue-green when moist, plumbeous to brownish or greenish grey when dry, with thin, 

longitudinal, white striae. Lower surface pale, with numerous white, unbranched, tapering, 

448

0.2–0.4 mm long rhizines not forming a mat but protruding from under the lobe margins 

and easily visible from above; interior of thalli sometimes with small, rounded adventive 

lobules along the margins of the lobes, which stay in the same plane as the lobes they are 

attached to. Apothecia uncommon, central in the thallus, up to 3 mm diam; disc orangebrown, 

soon immarginate and convex, with a few white, stellately arranged, 0.3–0.4 mm 

long rhizines attaching the apothecia to the thallus. Ascospores globose, 4–5 µm diam. 

Pycnidia uncommon, pale brown, c. 0.05 mm diam, marginal along the lobes. Conidia not 

observed. 

Distribution and Ecology: Apparently a species confined to very wet submontane to 

montane rainforests, often growing between bryophyte mats. Known from various 

collections throughout the Neotropics, including Venezuela (illustration of Coccocarpia 

'domingensis' by ARVIDSSON 1983: fig. 35C). 

Notes: Coccocarpia prostrata belongs in the group of species formed by C. stellata and 

relatives, characterized by narrow, linear lobes. Its lobe morphology is indeed very similar 

to that of C. stellata, but the latter has small, circular thalli of adnate lobes. The thalli of C. 

prostrata often appear almost fruticulose, due to the combination of prostrate main and 

ascending secondary lobes growing in various planes. Indeed, the new species should not 

be confused with C. filiformis, which has an entirely fruticulose thallus that might resemble 

the interior thallus parts of C. prostrata; however, C. filiformis lacks prostrate, linear 

primary lobes. The phyllidoid secondary lobules of C. prostrata also resemble the phyllidia 

of C. microphyllina, another new species described above, but that taxon has flabellate 

primary lobes and the phyllidia are not homologous to the thallus lobes of C. prostrata. 

ARVIDSSON (1983: 49) included at least one specimen of Coccocarpia prostrata in his 

concept of C. domingensis. The two species are easily distinguished, however, since C. 

domingensis has a completely flattened, stellate, distinctly bluish-grey thallus with 

cylindrical to flattened, true isidia. 

Selected specimens examined: COSTA RICA. Alajuela: Grecia Forest Reserve, Jun 2005, Chaves 2445 

(INB-3945752). Volcán Tenorio National Park, Mar 2004, Aptroot 60437 (ABL, INB-3964115). Cartago: 

Along Panamerican Highway ca. 5 km N of Cerro Vueltas, March 1985, Sipman & Chaverri 20671 (B, CR). 

La Chonta, Nov 1988, Sipman et al. 41815 (B, CR). SW-slope of Volcán Turrialba, March 1985, Sipman 

20432 (B, CR). Tapantí National Park, Apr 2003, Buck 44140 (CR, NY), Lücking 16597b (F, INB-4002722), 

Nelsen 3557d (INB, WIS), Trest 1432c (INB, WIS), Will-Wolf 12536b (INB, WIS). Heredia: Braulio 

Carrillo National Park, Oct 2005, Chaves 2456 (INB-3979135). Limón: Hitoy Cerere Biological Reserve, 

Mar 2004, Nelsen 3709 (INB, WIS). Puntarenas: La Amistad International Park, Jun 2002, Lücking 15252a 

(F), Trest 523 (INB), Will-Wolf 10079d (WIS), Jun 2002, Trest 498a (INB). Las Cruces Biological Station, 

Oct 2004, Sipman 53301 (B, INB-3993706). 

U.S.A. Florida: Ocala National Forest, Aug 1985, Harris 18042 (ABL). DOMINICAN REPUBLIC. 

Independencia: Sierra de Boaruco, Jan 1987, Harris 20453 (ABL). La Vega: Armando Bermudez National 

Park, Jan 1987, Harris 19742 (ABL). COLOMBIA. Santuario, Oct-Dec 1989, Wolf et al. T115 (ABL). 

449

VENEZUELA. Mérida: Prado Verde, ceranías de Mérida, Apr 1976, López-Figueiras 13543 (GB, MER). 

FRENCH GUIANA. Maripasoula: Saül, 1986, Montfoort & Ek s.n. (ABL). BRAZIL. Minas Gerais: Caraça, Sep 

1997, Aptroot 40885 (ABL), 41198 (ABL). São Paulo: Campos de Jordão, Sep 1997, Aproot 41793 (ABL). 

BOLIVIA. La Paz: B. Saavedra, 15 km de Charazani hacia Apolo, July 1997, Bach et al. 272B (B). 

450

Fig. 4. Morphotypes of Coccocarpia palmicola. A–D. Dark rhizinate morphotype 

with compact, erect isidia (Lücking 15199a). E–H. Pale rhizinate morphotype with 

fragile, prostrate isidia (E–F, Lücking 17215 and G–H, Lücking 15022b). Scale = 2 

mm, in B, D, and F = 1 mm. 

451

Fig. 5. Habit of Coccocarpia species. A–B. Coccocarpia stellata (Lücking s.n.). C– 

H. Coccocarpia prostrata (C and D, Lücking 15252a and Lücking 16597b; E and 

F, holotype; G and H, Lücking s.n.). Scale = 1 mm. 

452

Fig. 6. Ecology of Coccocarpia species in Costa Rica. The two axes depict the 

main precipitation and altitudinal gradient, while the illumination gradient is 

represented by different shades of white (exposed), grey (semi-exposed) and black 

(shaded). 

Coccocarpia stellata Tuck. Fig. 5A–B 

Notes: Coccocarpia stellata is commonly found on leaves and thin branches in more or 

less exposed situations. It is most similar to C. domingensis and C. neglecta but lacks 

isidia, and its lobes are apically often very slightly wider than those of the two isidiate 

species. C. epiphylla differs from C. stellata by its lobes being about twice as wide and by 

the absence of rhizines projecting laterally from the apothecial margins. The newly 

described C. prostrata has very similar primary lobes but grows much more irregularly and 

also forms additional, subfruticulose, ascending, rather thin and richly branched secondary 

lobes, while C. stellata is always flat and regularly radiating. 

453

Selected specimens examined: COSTA RICA. Alajuela: Volcán Tenorio National Park, Mar 2004, Nelsen 

3762 (INB-4031349, WIS), Sipman 51769 (B, INB-4031350), 51831 (B, INB-4031351), Trest 1553c (CR, 

INB-4031352). Guanacaste: Volcán Tenorio National Park, Mar 2004, Aptroot 60647 (ABL, INB-3968061). 

Heredia: La Selva Protection Zone, Oct 2002, Lücking 16002f (F). Puntarenas: La Amistad International 

Park, Jul 2002, Lücking 15260f (INB-3976254), Nelsen 2116a (WIS), Sipman 48020g (INB-3927549). Las 

Cruces Biological Station, Oct 2004, Sipman 53372 (B, INB-3993874). 

Coccocarpia tenuissima Müll. Arg. Fig. 3G 

Notes: Found only once but the material agrees well with the type (Fig. 3G). 


Aptroot 60525 (ABL, INB-3967677). 

Ecological analysis 

Coccocarpia erythroxyli and C. palmicola are not only the most abundant species but also 

show the widest ecological amplitude, ranging from (semi-)arid to perhumid climates and 

from sea level to the subparamo zone at 3500 m (Fig. 6). This coincides with the statements 

by ARVIDSSON (1983) that these two species are the only ones also found in extratropical 

regions and the only ones that reach altitudes above 4000 m in the Andes. We could not 

confirm the observation of ARVIDSSON (1983) that the two forms of Coccocarpia with 

adnate thallus and blue-black rhizines versus loosely attached thallus and white rhizines 

intergrade along an altitudinal gradient; the typical 'montane' form was also found in the 

lowland rain forest at 50 m altitude. 

According to ARVIDSSON (1983), Coccocarpia pellita is another common and 

widespread taxon at a world-wide level, but at the regional level in our material was 

restricted to (per-)humid climates between 1000 and 3500 m. On the contrary, C. dissecta 

seems to be a lowland species with a range from (semi-)arid to (sub-)humid climates. The 

collections of C. guimarana also originate from lowland subhumid forest. Three other 

species apparently confined to lower altitudes are C. glaucina, C. neglecta, and C. 

tenuissima (Fig. 6). 

Another group of species, being mostly found at mid elevations in (sub-)humid to 

perhumid climates, includes C. adnata, C. epiphylla, C. aff. imbricascens, C. 

microphyllina, and C. stellata (Fig. 6). Coccocarpia gallaicoi possibly also belongs here, 

but the label data on the type material are insufficient to draw meaningful ecological 

conclusions. 

The three remaining taxa, Coccocarpia domingensis, C. filiformis, and C. prostrata, are 

mostly confined to humid climates but show a wide altitudinal range, from the lowland and 

the colline to the upper montane and subparamo zone (Fig. 6). 

454

There is no clear correlation between morphology and ecological preferences, although 

broad-lobed and adnate taxa and those with true, cylindrical isidia (Coccocarpia adnata, C. 

guimarana, C. erythroxyli, C. glaucina, C. palmicola) seem to prefer, or tolerate, drier and 

hotter conditions. On the other hand, most narrow-lobed species and those with phyllidia or 

flattened isidia, as well as the ascendent and fruticulose forms (C. domingensis, C. 

filiformis, C. microphyllina, C. pellita, C. prostrata), are more typically found in humid to 

perhumid, montane climates. 

Coccocarpia as a whole is most abundant and most diverse in humid to perhumid, 

lower montane to montane situations. Localities falling under this category not rarely have 

up to ten or even more taxa. The species also prefer (semi-)exposed microhabitats and are 

usually absent under fully shaded conditions (Fig. 6). 

Acknowledgements 

The Ticolichen biodiversity inventory in Costa Rica is supported by a grant from the National Science 

Foundation (DEB 0206125 to The Field Museum; PI Robert Lücking) and by funds from the World Bank to 

the Instituto Nacional de Biodiversidad (INBio). We greatly appreciate the support of the Sistema Nacional 

de Áreas de Conservación (SINAC) and the Ministerio de Ambiente y Energía (MINAE) in receiving the 

necessary collection permits. Daniela Lizano (Universidad de Costa Rica), Eida Fletes, Enia Navarro, Ronald 

Rodriguez, and Eduardo Alvarado (INBio), Susan Will-Wolf, Marie Trest (University of Wisconsin- 

Madison), as well as Martin Grube (University of Graz, Austria) and William Buck (New York Botanical 

Garden), participated in the field work, and their vaulable support and company are warmly acknowledged. 

For the generous support in organizing the visit to the Farlow Herbarium (FH) to study the Dodge material, 

the first author in indepted to Dr. Donald Pfister and his staff. 

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