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The genus Cladonia (Lichenized Ascomycota) from Restinga vegetation of Espírito Santo state, Brazil: Supergroups Cladonia and Cocciferae

Abstract

This paper deals with 14 species of the genus Cladonia, occurring at Restinga vegetation of Espírito Santo state, with eight belonging to the Supergroup Cladonia and six to the Supergroup Cocciferae. Cladonia corallifera, C. crustacea, C. subminiata, C. pityrophylla, and C. polyscypha are new records for the state. An identification key, comments and illustrations are also provided.

Key words:
Atlantic rainforest; Cladoniaceae; dimorphic lichens; lichenized fungi; taxonomy

Resumo

Neste trabalho são tratadas 14 espécies do gênero Cladonia ocorrentes em vegetação de Restinga no estado do Espírito Santo, sendo oito pertencentes ao supergrupo Cladonia e seis ao Supergrupo Cocciferae. Cladonia corallifera, C. crustacea, C. subminiata, C. pityrophylla e C. polyscypha são novos registros para o estado. São fornecidas chave de identificação, ilustrações e comentários.

Palavras-chave:
Mata atlântica; Cladoniaceae; liquens dimórficos; fungos liquenizados; taxonomia

Introduction

The Restinga vegetation occupies the Quaternary sandy plains, being floristic and structurally associated with the Atlantic rainforest (Suguio & Tessler 1984Suguio K & Tessler MG (1984) Planícies de cordões litorâneos Quaternários do Brasil: origem e nomenclatura. In: Lacerda LD, Araujo DSD, Cerqueira R & Turc B (orgs.) Restingas: origem, estrutura e processos. CEUFF, Niterói. Pp. 15-25.; Araujo 1992Araujo DSD (1992) Vegetation types of sandy coastal plains of tropical Brazil: a first approximation. In: Seeliger U (eds.) Coastal plant communities of Latin America. Academic Press, San Diego. Pp. 337-347.; Martin et al. 1997Martin L, Suguio K, Dominguez JML & Flexor JM (1997) Geologia do quaternário costeiro do litoral norte do Rio de Janeiro e do Espírito Santo. CPRM/FAPESP, Belo Horizonte. 112p.) and is characterized by the phytophisiognomic gradient from the beach towards the continent, ranging from grasslands to shrubs and finally trees, intermingled or not by floodplains (Silva & Somner 1984Silva JG & Somner GV (1984) A vegetação da restinga na Barra de Maricá, RJ. In: Lacerda LD, Araujo DSD, Cerqueira R & Turcq B (orgs.) Restingas: origem, estrutura e processos. CEUFF, Niterói. Pp. 217-225.). Brazilian Restinga woods exhibit an abundant and diverse lichen biota, with Cladonia P. Browne among its most representative components (Marcelli 1998Marcelli MP (1998) History and current knowledge of Brazilian lichenology. In: Marcelli MP & Seaward MRW. Lichenology in Latin America: history, current knowledge and applications. CETESB, São Paulo. Pp. 25-45.).

Data about this genus in Brazilian coastal environments can be found in Ahti et al. (1993)Ahti T, Stenroos S & Xavier Filho L (1993) The lichen family Cladoniaceae in Paraiba, Pernambuco and Sergipe, northeast Brazil. Tropical Bryology 7: 55-70., Ahti & Marcelli (1995)Ahti T & Marcelli MP (1995) Taxonomy of the Cladonia verticillaris complex in South America. In: Farkas EE, Lücking R & V Wirth. Scripta Lichenologica Lichenological Papers dedicated to Antonín Vezda. Bibliotheca Lichenologica 58: 5-26., Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362., Gumboski & Eliasaro (2011Gumboski EL & Eliasaro S (2011) Cladonia litoralis (Cladoniaceae), a new species from southern Brazil. The Bryologist 114: 665-667.; 2012aAhti T & Hammer S (2002) Cladonia . In: Nash III TH, Ryan BD, Gries C & Bungartz F (eds.) Lichen flora of the Greater Sonoran Desert Region. Vol. 1. Arizona State University, Tempe. Pp. 131-158.,b)Gumboski EL & Eliasaro S (2012b) Espécies de Cladonia P. Browne (Cladoniaceae, Ascomycota) dos Supergrupos Cocciferae, Crustaceae e Perviae em Restingas e costões rochosos dos estados do Paraná e de Santa Catarina, Brasil. Acta Botanica Brasilica 26: 619-631. and Gumboski et al. (2013)Gumboski EL, Beilke F & Eliasaro S (2013) Cladonia dunensis sp. nov. from southern Brazil, with notes on the genus in beach dune environments. Mycotaxon 124: 333-340.. Records of Cladonia species from Espírito Santo State are relatively few, with only 23 species so far reported by Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362., whereas to the neighbor Rio de Janeiro State, with a similar area, vegetation and geology, the same author registered over 100 species.

The Supergroup Cladonia has a quite diverse thallus morphology, with unbranched to very branched podetia, although always with closed axils and brown apothecial discs. The presence of depsides and depsidones such as atranorin and fumarprotocetraric acids are a diagnostic character for this group (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.; Stenroos et al. 2002Stenroos S, Hyvonen J, Myllys L, Thell A & Ahti T (2002) Phylogeny of the genus Cladonia s.lat. (Cladoniaceae, Ascomycetes) inferred from molecular, morphological and chemical data. Cladistics 18: 237-278.). In contrast, Supergroup Cocciferae is characterized by the red apothecial discs, and occasionally by a yellowish medulla, as well as the production of dibenzofurans and β-orcinol depsides such as didymic, barbatic, squamatic and thamnolic acids (Huovinen et al. 1989Huovinen K, Ahti T & Stenroos S (1989) The composition and contents of aromatic lichen substances in Cladonia, section Cocciferae. Annales Botanici Fennici 26: 133-148.; Stenroos et al. 2002Stenroos S, Hyvonen J, Myllys L, Thell A & Ahti T (2002) Phylogeny of the genus Cladonia s.lat. (Cladoniaceae, Ascomycetes) inferred from molecular, morphological and chemical data. Cladistics 18: 237-278.). For the Restinga vegetation of Espírito Santo, only six species of Supergroup Cladonia and three of Cocciferae were previously known (Ahti et al. 1993Ahti T, Stenroos S & Xavier Filho L (1993) The lichen family Cladoniaceae in Paraiba, Pernambuco and Sergipe, northeast Brazil. Tropical Bryology 7: 55-70.; Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.).

In order to increase our knowledge of the genus Cladonia in the southeastern coastal areas of Brazil, we investigated the diversity of this genus species present along the coast of Espírito Santo State, characterizing their morphology, chemistry, and ecology.

Materials and Methods

This survey was conducted in Restinga vegetation areas along the coast of Espírito Santo State, with boundaries established by the coordinates 18º28'S and 21º15'S (Fig. 1). The collections were made in eight localities (Tab. 1). According to Köppen's classification, the climate is tropical hot and humid (Aw), with hot and rainy summers and dry winters (Pereira & Araujo 1995Pereira OJ & Araújo DSD (1995) Estrutura da vegetação de entre moitas da formação aberta de Ericaceae, no Parque Estadual de Setiba, Espírito Santo. In: Esteves FA (ed.) Oecologia brasiliensis: estrutura, funcionamento e manejo de ecossistemas brasileiros. Vol.1. UFRJ, Rio de Janeiro. Pp. 245-257.).

Figure 1
Espírito Santo state map with collection sites highlighted. 1. ISP; 2. EPACB; 3. RB; 4. MNPJ; 5. PCVSP; 6. EPAS; 7. ARB; 8. NB.

Table 1
Collection sites localities and coordinates.

The specimens were collected and preserved following the standard methods in lichenology (Brodo et al. 2001Brodo IM, Sharnoff SD & Sharnoff S (2001) Lichens of North America. Yale University Press, New Haven, London. 795p.), and deposited at the Herbarium of the Federal University of Paraná (UPCB). Morphological and anatomical analyses were performed under dissecting and compound microscopes using standard microscopic techniques. For the identification of lichen substances, spot tests (K, C, and KC), thin layer chromatography (TLC) with the solvent system C, as well as observation under UV light were conducted (Orange et al. 2001Orange A, James PW & White FJ (2001) Microchemical methods for the identification of lichens. British Lichen Society, London. 111p.; Elix 2014Elix JA (2014) A catalogue of standardized chromatographic data and biosynthetic relationships for lichen substances. 3rd ed. Published by the author, Canberra. 362p.).

Results and Discussion

A total of 14 Cladonia species belonging to the Supergroups Cladonia and Cocciferae were identified, five being new records for the state. The six species belonging to Cocciferae Supergroup identified are: Cladonia corallifera, C. crustacea, C. subminiata (new records for the state), as well as C. didyma, C. macilenta, and C. secundana [previously recorded by Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.]. From Supergroup Cladonia the eight species recognized are: C. pityrophylla and C. polyscypha, two new records for the state. The remaining taxa, C. cartilaginea, C. clathrata, C. furfuracea, C. subradiata, C. subsquamosa and C. verticillaris had already been cited by Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362. from the state.

    Key to Cladonia species of Supergroups Cladonia and Cocciferae in the Restinga vegetation of Espírito Santo
  • 1. Apothecial disc and pycnidia red, hypothallus yellowish (Supergroup Cocciferae).............................2

  • 1'. Apothecial discs and pycnidia brown, hypothallus brown (Supergroup Cladonia)..............................7

    • 2. Primary thallus persistent, predominant, podetia short (up to 1 cm) and often scarce..................3

    • 2'. Primary thallus somewhat evanescent, podetia fairly long (usually longer than 1 cm) and abundant.........................................................................................................................................5

      • 3. Primary squamules sorediate at margins...............................................4. Cladonia crustacea

      • 3'. Primary squamules esorediate................................................................................................4

        • 4. Podetia yellowish green, phyllopodial (a flat squamule that gradually rolling up at the margins so that forms a podetium)............................................11. Cladonia subminiata

        • 4'. Podetia browned, laminal to marginal on primary squamules......................................... ...................................................................................................10. Cladonia secundana

          • 5. Podetia scyphose.................................................................3. Cladonia corallifera

          • 5'. Podetia subulate.....................................................................................................6

            • 6. Podetia cylindrical, with farinose soredia, sparsely squamulose..................... ......................................................................................7. Cladonia macilenta

            • 6'. Podetia thickened at the base, with granulose soredia, granules, and usually with many microsquamules..............................................5. Cladonia didyma

              • 7. Primary thallus persistent, longer than podetia........................................ ...........................................................................8. Cladonia pityrophylla

              • 7'. Primary thallus somewhat evanescent, smaller than podetia.................8

                • 8. Podetia with verticillate scyphi (proliferating from the center).....9

                • 8'. Podetia not proliferating from the center.....................................10

                  • 9. Podetia short (up to 3 cm), scyphi clathrate (e.g., perforate), margins dentate.......................................2. Cladonia clathrata

                  • 9'. Podetia tall (> 3 cm), scyphi not clathrate, margins lacerate (elongated lobes with sharp or irregular edges).................................................................................14. Cladonia verticillaris

                    • 10. Podetia regularly scyphose...........................................11

                    • 10'. Podetia ascyphose to narrowly scyphose.....................12

                      • 11. Podetia short (up to 1.5 cm), with fairly wide scyphi (up to 8 mm wide).........13. Cladonia subsquamosa

                      • 11'. Podetia elongate (1-2.5 cm), with narrow scyphi (up to 2.5 mm wide)..................9. Cladonia polyscypha

                        • 12. Podetia esorediate, with reflexed to isidioid squamules throughout, usually with scyphoid to scyphose apex..........6. Cladonia furfuracea

                        • 12'. Podetia sorediate, ascyphose to narrowly scyphose when developed............................13

                          • 13. Podetia with farinose soredia, usually with microsquamules around the base, with an arachnoid white medulla, with a subulate or scyphoid apex when developed............ ...........................12. Cladonia subradiata

                          • 13'. Podetia with granulose soredia, phyllidiate, without medulla, never scyphose.............. ...........................1. Cladonia cartilaginea

1. Cladonia cartilaginea Müll. Arg., Flora 63 :260 (1880). Fig. 2a

Figure 2
a. Cladonia cartilaginea. b. Cladonia clathrata. c. Cladonia corallifera. d. Cladonia crustacea. e. Cladonia didyma. f. Cladonia furfuracea. (a. C.A.V. Fraga Jr 412; b. C.A.V. Fraga Jr 594; c. C.A.V. Fraga Jr 598; d. C.A.V. Fraga Jr 451b; e. C.A.V. Fraga Jr 698b; f. C.A.V. Fraga Jr 316). Scale bars = 1 cm.

Descriptions: Vainio (1894)Vainio EA (1894) Monographia Cladoniarum Universalis. II. Acta Societatis pro Fauna et Flora Fennica 10: 1-498., Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362..

Spot tests: K+ yellow; C-; KC-; UV-.TLC: fumarprotocetraric and homosekikaic acids.

Selected specimens examined: Presidente Kennedy, NB, 15.I.2015, C.A.V. Fraga Jr 271 (UPCB). Vila Velha, MNPJ, 24.II.2015, C.A.V. Fraga Jr 412, 433a (UPCB).

Cladonia cartilaginea is defined by the subulate, ecorticate podetia, with a hard stereome, covered mainly by granulose soredia and phyllidia (Vainio 1894Vainio EA (1894) Monographia Cladoniarum Universalis. II. Acta Societatis pro Fauna et Flora Fennica 10: 1-498.; Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.). In this study we found specimens with fumarprotocetraric (C.A.V. Fraga Jr 433a) or with homosekikaic acid (C.A.V. Fraga Jr 412). Fumarprotocetraric acid is quite common in this species. However, the occurrence of this compound associated with homosekikaic acid has been detected only in Brazilian samples (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.).

This species is similar to C. subradiata, which also occurs on the State coast. However C. subradiata occasionally exhibits narrow scyphi (1-2 mm wide) when well-developed, and a white medulla and conspicuous microsquamules around the base of the podetia (Ahti & Kashiwadani 1984Ahti T & Kashiwadani H (1984) The lichen genera Cladia, Cladina and Cladonia in Southern Chile. In: H Inoue (eds.) Studies on cryptogams in Southern Chile. Kenseisha, Tokio. Pp. 125-151.). C. cartilaginea is also similar to C. corymbites Nyl. and C. corniculata Ahti & Kashiw., but the former one has crenulated microsquamules (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.), while the second one has larger (2-6 cm) and farinosely sorediate, branched podetia (Ahti & Hammer 2002Ahti T & Hammer S (2002) Cladonia . In: Nash III TH, Ryan BD, Gries C & Bungartz F (eds.) Lichen flora of the Greater Sonoran Desert Region. Vol. 1. Arizona State University, Tempe. Pp. 131-158.).

The analyzed specimens often occur on sandy soil, together with mosses in the shades on the shrubs borders.

2. Cladonia clathrata Ahti & L. Xavier, Trop. Bryol. 7: 58 (1993)Ahti T, Stenroos S & Xavier Filho L (1993) The lichen family Cladoniaceae in Paraiba, Pernambuco and Sergipe, northeast Brazil. Tropical Bryology 7: 55-70.. Fig. 2b

Descriptions: Ahti & Marcelli (1995)Ahti T & Marcelli MP (1995) Taxonomy of the Cladonia verticillaris complex in South America. In: Farkas EE, Lücking R & V Wirth. Scripta Lichenologica Lichenological Papers dedicated to Antonín Vezda. Bibliotheca Lichenologica 58: 5-26., Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362..

Spot tests: K+ yellow; C-; KC-; UV-.TLC: Fumarprotocetraric acid.

Selected specimens examined: Conceição da Barra, ISP, 27.I.2015, C.A.V. Fraga Jr 512 (UPCB); EPACB, 28.I.2015, C.A.V. Fraga Jr 594 (UPCB). Guarapari, PCVSP, 10.II.2015, C.A.V. Fraga Jr 347, 360 (UPCB). Itaóca, ARB, 16.I.2015, C.A.V. Fraga Jr 728 (UPCB). Vila Velha, MNPJ, 24.II.2015, C.A.V. Fraga Jr 456 (UPCB).

Cladonia clathrata belongs to the C. verticillaris complex, and is recognized by its small size (up to 3 cm), the distinctly clathrate (i.e., perforated) scyphi and by the felty surface of podetia (Ahti et al. 1993Ahti T, Stenroos S & Xavier Filho L (1993) The lichen family Cladoniaceae in Paraiba, Pernambuco and Sergipe, northeast Brazil. Tropical Bryology 7: 55-70.; Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.). It is similar to C. rappii A. Evans, which differs by not having the perforations in the scyphi, and also has a strong melanotic base (Ahti & Marcelli 1995Ahti T & Marcelli MP (1995) Taxonomy of the Cladonia verticillaris complex in South America. In: Farkas EE, Lücking R & V Wirth. Scripta Lichenologica Lichenological Papers dedicated to Antonín Vezda. Bibliotheca Lichenologica 58: 5-26.). The ascospores of C. clathrata measure 8-10 × 3 µm and the primary squamules have a distinct crenulate cortex, but these characters were still unreported. This species is present almost all along the State coast, mainly on sand in exposed areas, or borders of brushwood, also growing mixed with mosses of the family Dicranaceae.

3. Cladonia corallifera (Kunze) Nyl., Flora 57: 70 (1874). Fig. 2c

Descriptions: Stenroos (1989c)Stenroos S (1989c) Taxonomy of Cladonia coccifera group. 2. Annales Botanici Fennici 26: 307-317., Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362..

Spot tests: K+ yellow; C-; KC-; UV-.TLC: rhodocladonic, thamnolic, decarboxythamnolic, didymic, 3α-hydroxybarbatic, squamatic and usnic acids plus unidentified compounds with Rfc approximately 34, 38 and 56.

Selected specimens examined: Conceição da Barra, ISP, 27.I.2015, C.A.V. Fraga Jr 509, 575 (UPCB); EPACB, 28.I.2015, C.A.V. Fraga Jr 598 (UPCB). Guarapari, PCVSP, 10.II.2015, C.A.V. Fraga Jr 297, 346 (UPCB); EPAS, 2.III.2015, C.A.V. Fraga Jr 672, 661, 669 (UPCB). Itaóca, ARB, 16.I.2015, C.A.V. Fraga Jr 725, 730 (UPCB). Presidente Kennedy, NB, 14.I.2015, C.A.V. Fraga Jr 221 (UPCB). Vila Velha, MNPJ, 24.II.2015, C.A.V. Fraga Jr 430 (UPCB).

Cladonia corallifera is characterized by a yellowish short podetia (up to 1.5 cm), that is regularly scyphose, granulose sorediate, often also squamulose, and by a diverse secondary chemistry. It occurs mainly in sandy environments of fluvial origin of northern Brazil (Stenroos 1989cStenroos S (1989c) Taxonomy of Cladonia coccifera group. 2. Annales Botanici Fennici 26: 307-317.; Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.).

Cladonia mollis Ahti & Sipman and C. prancei Ahti can be confused with C. corallifera as they occur in similar environments, and also have an ecorticate, scyphose podetia, with farinose to granulose soredia. However, both species have usually larger podetia (1-3 cm), with narrower scyphi (1-4 mm) (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.; Ahti & Sipman 2013Ahti T, Stenroos S & Xavier Filho L (1993) The lichen family Cladoniaceae in Paraiba, Pernambuco and Sergipe, northeast Brazil. Tropical Bryology 7: 55-70.).

Cladonia corallifera is morphologically very similar to C. coccifera, differing primarily in the secondary metabolites. C. coccifera produces porphyrilic acid and zeorin, which are absent in C. corallifera (Stenroos 1989aStenroos S (1989a) Taxonomy of Cladonia coccifera group. 1. Annales Botanici Fennici 26: 157-168.,c; Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.).

Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362. recorded C. coccifera from Restinga vegetation of Espírito Santo state, although in the same paper most specimens were mentioned from highlands, except the collections from the states of Espírito Santo and Bahia. We did not find C. coccifera in our study and, unfortunately the specimens mentioned by Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362. were not located in herbarium SP neither in H (M.P. Marcelli, personal communication) and in H (T. Ahti, personal communication). Thus, we could not confirm its occurrence in the Restinga vegetation of Espírito Santo state. We are inclined to believe that the specimen referred as C. coccifera in Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362. could possibly be related to C. corallifera, since in our investigation, this species is quite abundant along the collection sites and, C. coccifera is mostly reported for highlands in Brazil.

The majority of the specimens analyzed correspond to the chemotype II (usnic and thamnolic acids), and only one to the chemotype I (usnic, thamnolic and didymic acids) mentioned by Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.. However, in our samples, 3α-hydroxybarbatic and decarboxythamnolic acids are often present, though have not yet been reported. Despite showing a wide variation in the composition of its secondary metabolites, we did not notice any significant morphological correlation with the chemistry.

In the field, C. corallifera was found on white sand or on dry wood, in well-lit environments such as in open shrub formations, forming yellowish colonies, growing alone or together with C. polystomata Ahti & Sipman, C. subradiata, C. didyma and C. polyscypha. It is mainly reported for northern Brazil (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.), being a new record for Espírito Santo State.

4. Cladonia crustacea Ahti, Fl. Neotrop. Monogr.78: 191 (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.. Fig. 2d

Description: Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362..

Spot tests: K+ yellow; C-; KC-; UV-.TLC: thamnolic and decarboxythamnolic acids plus an unidentified compound with Rfc approximately 47.

Selected specimens examined: Vila Velha, MNPJ, 24.II.2015, C.A.V. Fraga Jr 440a, 439a, 451b (UPCB).

Cladonia crustacea is defined by the abundantly sorediate margins of the squamules, forming a subcrustose coverage, sparsely bearing inconspicuous podetia (up to 5 mm tall), and by producing thamnolic and decarboxythamnolic acids (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.).

It is similar to C. ahtii S. Stenroos and C. miniata G. Meyer by having squamules with a sorediate margin, but differs from C. ahtii, which has corticate podetia and from C. miniata, which has orange medulla. Also, both C. ahtii and C. miniata occur in higher altitudinal environments (Stenroos 1989b; Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.). It is also morphologically similar to C. hypoxantha Tuck. However, in this species, the squamules are narrower, and not abundantly sorediate (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.).

It is an inconspicuous species, and is essentially corticolous/lignicolous, forming powdery, and easily overlooked crusts. It was found above 50 cm from the ground in an arboretum in open shrub formation. This is the first record of this species for the Espírito Santo State.

5. Cladonia didyma (Fée) Vain., Acta Soc. Fauna Fl. Fenn. 4: 137 (1887). Fig. 2e

Descriptions: Vainio (1887)Vainio EA (1887) Monographia Cladoniarum Universalis. I. Acta Societatis pro Fauna et Flora Fennica 4: 1-509., Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362..

Spot tests: K+ yellow or -; C-; KC-; UV-; TLC: rhodocladonic didymic, condidymic, subdidymic, thamnolic and decarboxythamnolic acids, plus two unidentified compounds with Rfc approximately 30, 34 and 52.

Selected specimens examined: Conceição da Barra, ISP, 27.I.2015, C.A.V. Fraga Jr 557 (UPCB). Guarapari, PCVSP, 13.II.2015, C.A.V. Fraga Jr 315a (UPCB); EPAS, 23.VII.2015, C.A.V. Fraga Jr 904 (UPCB). Itaóca, ARB, 16.I.2015, C.A.V. Fraga Jr 698b, 701 (UPCB). Presidente Kennedy, NB, 14.I.2015, C.A.V. Fraga Jr 203, 220, 248 (UPCB). Vila Velha, MNPJ, 24.I.215, C.A.V. Fraga Jr 431 (UPCB).

Cladonia didyma is distinguished by its ecorticate and subulate ascyphose podetia, with a granulose to squamulose cover, without soredia (Vainio 1887Vainio EA (1887) Monographia Cladoniarum Universalis. I. Acta Societatis pro Fauna et Flora Fennica 4: 1-509.; Fleig et al. 1995Fleig M, Ahti T & Stenroos S (1995) A família Cladoniaceae (líquens) no Rio Grande do Sul, Brasil. Napaea 11: 1-29.) and also, by the production of barbatic, thamnolic or didymic acids (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.).

According to Gumboski & Eliasaro (2012b)Gumboski EL & Eliasaro S (2012b) Espécies de Cladonia P. Browne (Cladoniaceae, Ascomycota) dos Supergrupos Cocciferae, Crustaceae e Perviae em Restingas e costões rochosos dos estados do Paraná e de Santa Catarina, Brasil. Acta Botanica Brasilica 26: 619-631., C. didyma may comprise a complex of species accordingly to some morphological differences found in the populations studied by them. The specimens included here as C. didyma also exhibit some morphological variations, such a melanotic stereome in some specimens, and farinose soredia with granules and squamules. In addition, the specimen C.A.V. Fraga Jr 557, showed a clearly pellucid central canal with yellow hyphae in podetia section. Thus, a phylogenetic analysis using molecular data are required to elucidate if the different populations of C. didyma comprises or not a complex of species, and also to delimit the values of the taxonomic characters.

Cladonia didyma is very similar to C. macilenta since both species have subulate and ecorticate podetia, also by the production of the same secondary metabolites (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.). However, C. didyma differs from C. macilenta by having a more granulose/squamulose podetia, leaving an exposed stereome. We also observed that the podetia of C. didyma are somewhat more robust than C. macilenta, and usually have a squamule ring below the apothecial discs.

When the apothecial discs and pycnidia are absent, C. didyma can be confused with C. cartilaginea or even with young specimens of C. furfuracea, by presenting an ecorticate surface and subulate podetia. However, both C. cartilaginea and C. furfuracea produce the fumarprotocetraric acid complex, and C. furfuracea is also usually scyphose (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.; Gumboski & Eliasaro 2012aGumboski EL & S Eliasaro S (2012a) Espécies de Cladonia P. Browne (Cladoniaceae, Ascomycota) do Supergrupo Cladonia em Restingas e costões rochosos dos estados do Paraná e de Santa Catarina, Brasil. Hoehnea 39: 315-337.,bGumboski EL & S Eliasaro S (2012a) Espécies de Cladonia P. Browne (Cladoniaceae, Ascomycota) do Supergrupo Cladonia em Restingas e costões rochosos dos estados do Paraná e de Santa Catarina, Brasil. Hoehnea 39: 315-337.).

This is one of the most commonly found Cladonia species along the Espírito Santo Restinga woods, occurring throughout the coast, colonizing sand, termite hills, litter and dead branches. It occurs in open areas or borders of bushes, without accompanying species or together with C. subradiata, C. subsquamosa, and C. furfuracea.

6. Cladonia furfuracea Vain., Acta Soc. Fauna Fl. Fenn. 10: 375 (1894). Fig. 2f

Descriptions: Vainio (1894)Vainio EA (1894) Monographia Cladoniarum Universalis. II. Acta Societatis pro Fauna et Flora Fennica 10: 1-498., Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362..

Spot tests: K+ yellow or -; C-; KC-; UV-; TLC: fumarprotocetraric acid.

Selected specimens examined: Conceição da Barra, ISP, 27.I.2015, C.A.V. Fraga Jr 578 (UPCB); EPACB, 28.I.2015, C.A.V. Fraga Jr 638 (UPCB). Guarapari, PCVSP, 10.II.2015, C.A.V. Fraga Jr 316, 352, 369 (UPCB). EPAS, 2.III.2015, C.A.V. Fraga Jr 660, 664 (UPCB). Itaóca, ARB, 16.I.2015, C.A.V. Fraga Jr 685a (UPCB). Presidente Kennedy, NB, 14.I.2015, C.A.V. Fraga Jr 253b (UPCB). Vila Velha, MNPJ, 24.II.2015, C.A.V. Fraga Jr 421, 435 (UPCB).

This species can be distinguished by the ecorticate, esorediate and usually scyphose podetia, that are 3-6 (-8) cm tall, often with a bare stereome, and covered by some corticated verruculae (Vainio 1894Vainio EA (1894) Monographia Cladoniarum Universalis. II. Acta Societatis pro Fauna et Flora Fennica 10: 1-498.; Fleig et al.1995Fleig M, Ahti T & Stenroos S (1995) A família Cladoniaceae (líquens) no Rio Grande do Sul, Brasil. Napaea 11: 1-29.), and reflexed isidioid squamules, producing fumarprotocetraric acid (Fleig et al. 1995Fleig M, Ahti T & Stenroos S (1995) A família Cladoniaceae (líquens) no Rio Grande do Sul, Brasil. Napaea 11: 1-29.; Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.; Stenroos et al. 2002Stenroos S, Hyvonen J, Myllys L, Thell A & Ahti T (2002) Phylogeny of the genus Cladonia s.lat. (Cladoniaceae, Ascomycetes) inferred from molecular, morphological and chemical data. Cladistics 18: 237-278.).

Cladonia furfuraceoides Ahti & Sipman is very similar, but it is smaller (1-3 cm tall) than C. furfuracea, and exhibits a strongly melanotic base of podetia and elongated podetial squamules (Stenroos et al. 2002Stenroos S, Hyvonen J, Myllys L, Thell A & Ahti T (2002) Phylogeny of the genus Cladonia s.lat. (Cladoniaceae, Ascomycetes) inferred from molecular, morphological and chemical data. Cladistics 18: 237-278.; Ahti & Sipman 2013Ahti T, Stenroos S & Xavier Filho L (1993) The lichen family Cladoniaceae in Paraiba, Pernambuco and Sergipe, northeast Brazil. Tropical Bryology 7: 55-70.). In this study, some specimens also exhibited a melanotic base (ca. 15%), mainly in buried parts. The podetial squamules, range from isidioid to clearly flattened, elongated and reflexed, and the secondary thallus hardly reaches 3 cm tall. Such characteristics approach the description of C. furfuraceoides. However, we believe that these variations can be caused by environmental conditions, since the thallus size of C. furfuracea seems to increase with altitude (Charnei & Eliasaro 2014Charnei AM & Eliasaro S (2014) Espécies de Cladonia com cifas não verticiladas do Supergrupo Cladonia em ambientes de altitude da Serra do Mar no Sul do Brasil. Hoehnea 41: 453-468.), whereas is found smaller in coastal environments. These variations deserve further investigations with support of molecular data to evaluate the range of its phenotypic plasticity and of its taxonomical characters.

Cladonia polystomata Ahti & Sipman is another similar species, also have an ecorticate, esorediate podetia with squamules and both can occur together in field. However, C. polystomata usually has funnels and open axils, also differing by producing thamnolic acid and related compounds (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.).

It is a common species along the Espírito Santo State coast, growing on sand at edge of shrubs, occasionally growing together with C. clathrata and C. subradiata.

7. Cladonia macilenta Hoffm., Deutschl. Fl. 2: 126 (1796). Fig. 3a

Figure 3
a. Cladonia. macilenta. b. Cladonia pityrophylla. c. Cladonia polyscypha. d. Cladonia secundana. e. Cladonia subminiata. f. Cladonia subradiata. (a. C.A.V. Fraga Jr 704b; b. C.A.V. Fraga Jr 335b; c. C.A.V. Fraga Jr 588; d. C.A.V. Fraga Jr 549; e. C.A.V. Fraga Jr 891; f. C.A.V. Fraga Jr 253a). Scale bars = 1 cm.

Descriptions: Vainio (1887)Vainio EA (1887) Monographia Cladoniarum Universalis. I. Acta Societatis pro Fauna et Flora Fennica 4: 1-509., Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362..

Spot tests: K+ yellow; C-; KC-; UV-; TLC: rhodocladonic, barbatic, thamnolic, condidymic, didymic and subdidymic acids.

Selected specimens examined: Itaóca, ARB, 16.I.2015, C.A.V. Fraga Jr 690, 704b, 713 (UPCB).

The diagnostic characters of C. macilenta are the cylindrical, ecorticate and always ascyphose podetia, covered by a farinose soredia (Vainio 1887Vainio EA (1887) Monographia Cladoniarum Universalis. I. Acta Societatis pro Fauna et Flora Fennica 4: 1-509.), and the production of barbatic, didymic and thamnolic acids (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.).

It is morphologically and chemically close to C. didyma, but it is often difficult to interpret their morphological characters (see discussion under C. didyma). The analyzed specimens displayed slender and usually proportionally cylindrical podetia. Also, the presence of a whorl of squamules below the hymenial discs is absent. We noticed that there is a tendency for the apothecial discs to be simple, flattened, and often with a central perforation (not funnel), rather than agglomerate, like in C didyma. It was also observed that there are granular soredia and few granules, but to a lesser extent farinose soredia. C. macilentoides Ahti & Fleig is also very similar to C. macilenta, but it presents a more robust morphology, exhibiting taller podetia (3-5 cm), with an acuminate apex, tending to be more branched (Fleig et al. 1995). In addition, C. macilentoides does not produce both barbatic and didymic acids, which are present in C. macilenta (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.). When sterile, it may be confused with C. subradiata, by its slender, ecorticate and subulate podetia, differing by their secondary metabolites, and by the presence of a scyphoid apex in C. subradiata. Cladonia macilenta produces thamnolic and didymic acids, which are absent from C. subradiata (Fleig et al. 1995Fleig M, Ahti T & Stenroos S (1995) A família Cladoniaceae (líquens) no Rio Grande do Sul, Brasil. Napaea 11: 1-29.).

It is an inconspicuous species that occurs on litter and dead wood, rarely on sand, in fairly shaded areas. It was found mostly alone or intermixed with C. didyma.

8. Cladonia pityrophylla Nyl., Flora 57: 70 (1874). Fig. 3b

Descriptions: Vainio (1894)Vainio EA (1894) Monographia Cladoniarum Universalis. II. Acta Societatis pro Fauna et Flora Fennica 10: 1-498., Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362..

Spot tests: K+ yellow → reddish; C-; KC-; UV-; TLC: fumarprotocetraric acid.

Selected specimens examined: Guarapari, PCVSP, 10.II.2015, C.A.V. Fraga Jr 314, 335b (UPCB). Vila Velha, MNPJ, 24.II.2015, C.A.V. Fraga Jr 386b, 433b, 444a, 452, 457a (UPCB).

Cladonia pityrophylla is characterized by the well-developed primary thallus without rhizines in the margins, by the laminal growing, ecorticate, usually subulate podetia, covered by bullate to globose squamules, and by the production of fumarprotocetraric acid (Vainio 1894Vainio EA (1894) Monographia Cladoniarum Universalis. II. Acta Societatis pro Fauna et Flora Fennica 10: 1-498.; Fleig et al. 1995Fleig M, Ahti T & Stenroos S (1995) A família Cladoniaceae (líquens) no Rio Grande do Sul, Brasil. Napaea 11: 1-29.; Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.).

Cladonia ceratophylla (Sw.) Spreng. is a similar species. However, the podetia do not exhibit globose or bullate squamules, and also can produce atranorin (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.). In the specimens C.A.V. Fraga Jr 444 and 457, we found scarce black rhizines on the margins of squamules, which we consider as a morphological variation, since other morphological and chemical characters fits the description present in Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.. C. latiloba Ahti & Marcelli and C. subcariosa Nyl. are also similar because of the conspicuous squamules (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.), but both are distinguishable by their podetia, which in C. latiloba are clavate and ecorticate, while in C. subcariosa they are corticate. Moreover, neither of them have bullate squamules, and generally produce atranorin, a compound not found in C. pityrophylla.

It is a rare species in the study area, essentially arenicolous, found mostly solitary in shady places, colonizing bare sand around shrubs or even buried. This is the first record of this species for Espírito Santo state.

9. Cladonia polyscypha Ahti & L. Xavier, Trop. Bryol. 7: 61 (1993)Ahti T, Stenroos S & Xavier Filho L (1993) The lichen family Cladoniaceae in Paraiba, Pernambuco and Sergipe, northeast Brazil. Tropical Bryology 7: 55-70.. Fig. 3c

Descriptions: Ahti et al. (1993)Ahti T, Stenroos S & Xavier Filho L (1993) The lichen family Cladoniaceae in Paraiba, Pernambuco and Sergipe, northeast Brazil. Tropical Bryology 7: 55-70., Ahti (2000).

Spot tests: K+ yellow → brownish; C-; KC-; UV-; TLC: fumarprotocetraric acid.

Selected specimens examined: Conceição da Barra, ISP, 27.I.2015, C.A.V. Fraga Jr 487 (UPCB); EPACB, 28.I.2015, C.A.V. Fraga Jr 588, 616 (UPCB). Guarapari, PCVSP, 10.II.2015, C.A.V. Fraga Jr 299 (UPCB); EPAS, 2.III.2015, C.A.V. Fraga Jr 645 (UPCB). Linhares, RB, 26.II.2015, C.A.V. Fraga Jr 468c (UPCB). Itaóca, ARB, 16.I.2015, C.A.V. Fraga Jr 704a (UPCB). Presidente Kennedy, NB, 14.I.2015, C.A.V. Fraga Jr 213c (UPCB). Vila Velha, MNPJ, 24.II.2015, C.A.V. Fraga Jr 424c (UPCB).

Cladonia polyscypha is characterized by the slender and ecorticate podetia, usually with a melanotic, compact corticate sheath around the base (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.), with a very narrow scyphi, and by producing fumarprotocetraric acid (Ahti et al. 1993Ahti T, Stenroos S & Xavier Filho L (1993) The lichen family Cladoniaceae in Paraiba, Pernambuco and Sergipe, northeast Brazil. Tropical Bryology 7: 55-70.).

It is very similar to C. subradiata, which differs by the absence of a melanotic base of podetia and by the more subulate/ascyphose apex (Ahti et al. 1993Ahti T, Stenroos S & Xavier Filho L (1993) The lichen family Cladoniaceae in Paraiba, Pernambuco and Sergipe, northeast Brazil. Tropical Bryology 7: 55-70.). C. ramulosa (With.) J.R. Laundon and C. ochracea L. Scriba are also similar, but C. ramulosa has a more granular soredia and C. ochracea has granules, and in both cases the base of the podetia is neither melanotic nor well corticate (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.; Gumboski & Eliasaro 2012aGumboski EL & S Eliasaro S (2012a) Espécies de Cladonia P. Browne (Cladoniaceae, Ascomycota) do Supergrupo Cladonia em Restingas e costões rochosos dos estados do Paraná e de Santa Catarina, Brasil. Hoehnea 39: 315-337.).

It was mainly found on sand, also occurring in termite hills, organic and clay soil in open areas or edges of brushwood. Found together with C. didyma, C. furfuracea and C. subradiata. This is the first record of this species for the Espírito Santo state.

10. Cladonia secundana Nyl., Flora 57: 71 (1874). Fig. 3d

Descriptions: Stenroos (1989b)Stenroos S (1989b) Taxonomic revision of the Cladonia miniata group. Annales Botanici Fennici 26: 237-267., Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362., Ahti & Sipman (2013)Ahti T, Stenroos S & Xavier Filho L (1993) The lichen family Cladoniaceae in Paraiba, Pernambuco and Sergipe, northeast Brazil. Tropical Bryology 7: 55-70..

Spot tests: K+ yellow; C-; KC-; UV-: TLC: rhodocladonic, didymic, condidymic, subdidymic, fumarprotocetraric and usnic acids, plus an unidentified substance with Rfc 58.

Selected specimens examined: Conceição da Barra, ISP, 27.I.2015, C.A.V. Fraga Jr 549 (UPCB). Guarapari, PCVSP, 10.I.2015, C.A.V. Fraga Jr 301a (UPCB).

Cladonia secundana is characterized by the long and deeply laciniate squamules, that are olive brown in color, with podetia arising laminally or marginally (Stenroos 1989bStenroos S (1989b) Taxonomic revision of the Cladonia miniata group. Annales Botanici Fennici 26: 237-267.), and by the highly variable chemistry, with about 30 compounds reported (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.).

It is close related to C. subminiata, another species of the C. miniata complex. Both have a similar gross morphology and geographical distribution in eastern Brazil. However, in C. subminiata the thallus is more yellowish due the presence of usnic acid, and the podetia are clearly phyllopodial (Ahti et al. 1993Ahti T, Stenroos S & Xavier Filho L (1993) The lichen family Cladoniaceae in Paraiba, Pernambuco and Sergipe, northeast Brazil. Tropical Bryology 7: 55-70.; Stenroos 1989bStenroos S (1989b) Taxonomic revision of the Cladonia miniata group. Annales Botanici Fennici 26: 237-267.). The specimen C.A.V. Fraga Jr 549 presented usnic acid, in addition to didymic, condidymic and subdidymic acids, compounds reported for C. subminiata. Despite this, the specimen showed laminal to marginal podetia, with more brownish coloration, characteristics that morphologically distinguish it from C. subminiata. We accept the aforementioned specimen as C. secundana based in a similar case reported by Stenroos (1989b), which also considers a specimen with a similar morphology as C. secundana, arguing about its highly variable chemistry.

According to Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362., this species has long been confused with C. miniata G. Mey., but is differentiated by the color of the medulla, which is red in C. miniata.

It is an essentially an arenicolous species, found mainly buried in the white sand, leaving few parts exposed. It was recorded alone, forming inconspicuous patches on the soil in open and well-lit areas.

11. Cladonia subminiata S. Stenroos, Ann. Bot. Fenn. 26: 256 (1989). Fig. 3e

Descriptions: Stenroos (1989b)Stenroos S (1989b) Taxonomic revision of the Cladonia miniata group. Annales Botanici Fennici 26: 237-267., Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362..

Spot tests: K+ yellow; C-; KC-; UV-; TLC: Rhodocladonic, didymic, condidymic, subdidymic, barbatic and usnic acids.

Selected specimens examined: Conceição da Barra, ISP, 27.I.2015, C.A.V. Fraga Jr 504, 518, 519, 535 (UPCB); EPACB, 28.I.2015, C.A.V. Fraga Jr 541 (UPCB). Guarapari, PCVSP, 10.II.2015, C.A.V. Fraga Jr 302, 334, 339, 365 (UPCB); EPAS, 2.III.2015, C.A.V. Fraga Jr 891 (UPCB).

Cladonia subminiata is characterized by the conspicuous, ascending and yellowish primary squamules, with constantly phyllopodial podetia, and by the production of usnic and didymic acids (Stenroos 1989bStenroos S (1989b) Taxonomic revision of the Cladonia miniata group. Annales Botanici Fennici 26: 237-267.).

Cladonia subminiata is also included in Cladonia miniata complex, and is very close to C. secundana (see comments under the later species).

This species forms extensive colonies on bare white sand, being highlighted by the red apothecial disc through the white sand. It was found mainly isolated, but was also registered together with C. clathrata. This is the first record of this species for Espírito Santo State.

12. Cladonia subradiata (Vain.) Sandst., Abh. Naturwiss. Vereine Bremen 25: 230 (1922). Fig. 3f

Description: Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362..

Spot tests: K+ yellow → brownish; C-; KC-; UV-; TLC: fumarprotocetraric acid.

Selected specimens examined: Conceição da Barra, ISP, 27.I.2015, C.A.V. Fraga Jr 507, 552 (UPCB); EPACB, 28.I.2015, C.A.V. Fraga Jr 611, 612a (UPCB). Guarapari, PCVSP, 10.II.2015, C.A.V. Fraga Jr 293 (UPCB); EPAS, 23.VII.2015, C.A.V. Fraga Jr 882 (UPCB). Itaóca, ARB, 16.I.2015, C.A.V. Fraga Jr 698 (UPCB). Linhares, RB, 27.I.2015, C.A.V. Fraga Jr 458 (UPCB). Presidente Kennedy, NB, 15.I.2015, C.A.V. Fraga Jr 239b, 253a (UPCB). Vila Velha, MNPJ, 24.II.2015, C.A.V. Fraga Jr 399 (UPCB).

This species is characterized by the simple to little branched, subulate and ecorticate podetia, often with squamules around the base, ascyphose to and narrowly scyphose when mature, and by the production of fumarprotocetraric acid (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.).

It is inconspicuous and easily confused with C. cartilaginea or C. polyscypha (see discussion on these taxa). C. subradiata is also very similar to C. didyma when both are sterile, due its ecorticate and subulate podetia, but C. didyma has a yellow hypothallus, and produces barbatic, didymic and thamnolic acids (Fleig et al. 1995Fleig M, Ahti T & Stenroos S (1995) A família Cladoniaceae (líquens) no Rio Grande do Sul, Brasil. Napaea 11: 1-29.; Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.). In some cases, the latter species can have a darker to melanotic podetia, which do not occur in C. subradiata. Cladonia corniculata Ahti & Kashiw. is also morphologically similar, but never produces scyphi and is reported mainly from highlands (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.).

This is one of the most common species of Cladonia along the Espírito Santo state coast, occurring in a wide variety of substrates, like on sand, wood and termite hills, in open or shaded areas. Usually occurs together with C. subsquamosa, C. didyma and C. polyscypha.

13. Cladonia subsquamosa Kremp. in Warming, Vidensk. Meddel.Dansk Naturhist. Foren. Kjøbenhavn 5: 336 (1874, ‘1873'). Fig. 4a

Figure 4
a. Cladonia subsquamosa. b. Cladonia verticillaris. (a. C.A.V. Fraga Jr 483; b. C.A.V. Fraga Jr 428). Scale bars = 1 cm.

Descriptions: Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362., Ahti & Sipman (2013)Ahti T & Sipman HJM (2013) Ten new species of Cladonia (Cladoniaceae, lichenized Fungi) from Guianas and Venezuela. South America. Phytotaxa 93: 25-39..

Spot tests: K+ yellow → brownish or -; C-; KC-; UV-; TLC: fumarprotocetraric acid.

Selected specimens examined: Conceição da Barra, ISP, 27.I.2015, C.A.V. Fraga Jr 483 (UPCB); EPACB, 28.I.2015, C.A.V. Fraga Jr 633 (UPCB). Guarapari, PCVSP, 10.II.2015, C.A.V. Fraga Jr 357, 374 (UPCB); EPAS, 2.II.2015, C.A.V. Fraga Jr 635 (UPCB); 23.VI.2015, C.A.V. Fraga Jr 885 (UPCB). Itaóca, ARB, 16.I.2015, C.A.V. Fraga Jr 721 (UPCB). Linhares, RB, C.A.V. Fraga Jr 467a, 468b (UPCB). Presidente Kennedy, NB, 15.I.2015, C.A.V. Fraga Jr 235 (UPCB). Vila Velha, MNPJ, 24.II.2015, C.A.V. Fraga Jr 401, 454a (UPCB).

This species is characterized by the ecorticate and scyphose podetia, with a throughout deciduous sorediate layer, that in most cases leaves a bare stereome, and by the production of fumarprotocetraric and quaesitic acids (Ahti & Kashiwadani 1984Ahti T & Kashiwadani H (1984) The lichen genera Cladia, Cladina and Cladonia in Southern Chile. In: H Inoue (eds.) Studies on cryptogams in Southern Chile. Kenseisha, Tokio. Pp. 125-151.; Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.; Ahti & Sipman 2013Ahti T & Sipman HJM (2013) Ten new species of Cladonia (Cladoniaceae, lichenized Fungi) from Guianas and Venezuela. South America. Phytotaxa 93: 25-39.).

Cladonia chlorophaea (Flörke ex Sommerf.) Spreng. is morphologically similar by having an ecorticate and scyphose podetia, however, it is covered mainly by granules rather than soredia, and is almost constantly corticated in the basal region (Bungartz & Ahti 2009Bungartz AR & Ahti T (2009) Cladoniaceae. Flora Liquenológica Ibérica. Vol. 4. Sociedad Española de Liquenología (SEL), Madrid. 111p.). Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362. argues that C. fimbriata (L.) Fr. is also morphologically and chemically similar to C. subsquamosa, but differs by having a more persistent farinose soredia and medulla. Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362. reports the constant presence of quaesitic acid (= convirensic acid) in C. subsquamosa, but all analyzed specimens here produced only fumarprotocetraric acid as a secondary metabolite. The same was noticed by Gumboski & Eliasaro (2012)Gumboski EL & S Eliasaro S (2012a) Espécies de Cladonia P. Browne (Cladoniaceae, Ascomycota) do Supergrupo Cladonia em Restingas e costões rochosos dos estados do Paraná e de Santa Catarina, Brasil. Hoehnea 39: 315-337. and Charnei & Eliasaro (2014)Charnei AM & Eliasaro S (2014) Espécies de Cladonia com cifas não verticiladas do Supergrupo Cladonia em ambientes de altitude da Serra do Mar no Sul do Brasil. Hoehnea 41: 453-468..

It is a widely distributed and common species, being one of the most common species in Espírito Santo Restinga woods. It occurs on sand, wood and termite hills, without accompanying species or with C. subradiata, C. furfuracea and C. polyscypha.

14. Cladonia verticillaris (Raddi) Fr., Lichenogr. Eur. Reform. 465 (1831). Fig. 4b

Descriptions: Ahti & Marcelli (1995)Ahti T & Marcelli MP (1995) Taxonomy of the Cladonia verticillaris complex in South America. In: Farkas EE, Lücking R & V Wirth. Scripta Lichenologica Lichenological Papers dedicated to Antonín Vezda. Bibliotheca Lichenologica 58: 5-26., Ahti (2000)Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362..

Spot tests: K+ yellow → brownish; C-; KC-; UV-; TLC: fumarprotocetraric acid.

Selected specimens examined: Guarapari, PCVSP, 10.II.2015, C.A.V. Fraga Jr 287, 322 (UPCB). Itaóca, ARB, 16.I.2015, C.A.V. Fraga Jr 724, 731 (UPCB). Presidente Kennedy, NB, 15.I.2015, C.A.V. Fraga Jr 230 (UPCB). Vila Velha, MNPJ, 24.II.2015, C.A.V. Fraga Jr 428, 442 (UPCB).

Cladonia verticillaris is characterized by the brownish green podetia, with yellowish to brown or melanotic bases (Ahti & Marcelli 1995Ahti T & Marcelli MP (1995) Taxonomy of the Cladonia verticillaris complex in South America. In: Farkas EE, Lücking R & V Wirth. Scripta Lichenologica Lichenological Papers dedicated to Antonín Vezda. Bibliotheca Lichenologica 58: 5-26.), bearing broad and verticillate scyphi, with lacerate margins that are often squamulose, producing fumarprotocetraric acid (Ahti 2000Ahti T (2000) Cladoniaceae. Flora Neotropica Monograph 78: 1-362.).

Besides C. verticillaris, C. clathrata is the only other species that has verticillate scyphi found in Espírito Santo Restinga vegetation. These species are quite distinct. However, as C. verticillaris exhibits taller podetia (3-15 cm), with lacerate scyphi, instead of being dentate. For additional information about species of the C. verticillaris complex in Brazil see Ahti & Marcelli (1995)Ahti T & Marcelli MP (1995) Taxonomy of the Cladonia verticillaris complex in South America. In: Farkas EE, Lücking R & V Wirth. Scripta Lichenologica Lichenological Papers dedicated to Antonín Vezda. Bibliotheca Lichenologica 58: 5-26. and Charnei & Eliasaro (2013)Charnei AM & Eliasaro S (2013) Verticilate Cladonia species (Lichenized Ascomycota) from high altitude environments of Serra do Mar in Southern Brazil. Hoehnea 40: 87-97..

This is a common species along the Southeast and Northeast Brazilian coast, found growing on bare sand and litter, both in open areas and in borders of brushwood. Usually grows alone, but was found with C. crispatula (Nyl.) Ahti.

Acknowledgements

We are thankful to Daniel Goetze for the great help in field excursion, Dr. Luciana Canêz, Dr. Fernando Bittencourt, Dr. Vagner Cortez, Dr. Robert Lucking and anonymous revisors for their precious considerations about the manuscript; to Instituto Estadual de Meio Ambiente (IEMA), Secretaria de Desenvolvimento Sustentável de Vila Velha (SEMDESU) for the collection licenses; and CAPES (Coordenação de Aperfeiçoamento de Pessoal do Nível Superior) for the financial support.

References

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Publication Dates

  • Publication in this collection
    Oct-Dec 2017

History

  • Received
    20 June 2017
  • Accepted
    18 Sept 2017
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