J. Hattori Bot. Lab. No. 85 : 245- 265 (Nov. 1998)

ON THE IDENTITY AND POSITION OF PENTAGENELLA FRAGILLIMA, ROCCELLODEA NIGERRIMA, AND SOME RELATED SPECIES (, OPEGRAPHALES)1

2 3 2 GERHARD FOLLMANN , MARGOT SCHULZ , AND BIRGIT WERNER

INTRODUCTION Pentagenella fragillima was described from Chile by Darbishire (1897) and Roccellodea nigerrima from the Galapagos Islands by the same author ( 1932). Because of apparently missing type material, both Roccellaceae were excluded from a cladistical treatment of the family as critical taxa of uncertain position by Tehler (1990), and Weber (1986) listed the second one under "Rejected reports, synonyms, and misapplied names" in his catalogue of Galapagos . On the other side, samples corresponding largely to Darbishire's descriptions of P. fragillima and R. nigerrima growing close to their supposed type localities were repeatedly used for chemo­ taxonomical studies by Huneck & Follmann (1967), Follmann & Huneck (1969), Follmann et al. (1993), etc. and cited in floral inventories and sociological registers (i.a., Follmann 1962, 1964, 1967, 1995, 1997). Meanwhile, the type specimens were located in the herbaria of the National Museum of Natural History at Paris (PC) and of the Department of Botany of the University of Bristol (BRIST), respectively. The following revision, which includes some related taxa, was carried out to bring this unsatisfactory and impeding situation to an end.

MATERIAL AND METHODS For the present check-up, specimens of Roccellaceae preserved in the public herbaria at Boulder (COLO), Bristol (BRIST), Geneva (G), Helsinki (H), London (BM), Lund (LD), Paris (PC), Santa Cruz de Tenerife (TFMC), Stockholm (S), Uppsala (UPS), Vienna (W), and Washington (US) were used. In addition, supplementary field

1 In memory of Otto Vernon Darbishire (1870- 1934) on occasion of the IOOth anm­ versary of the publication of his "Monographia Roccelleorum" (1898) which - despite some misunderstandings and shortcomings concerning the state of knowledge of the period - laid the foundations for many directive biosystematical and geobotanical studies on one of the most peculiar lichen families. 2 Botanical Institute, University of Cologne, Gyrhofstr. 15, D-50931 Cologne, Germany. 3 Institute for Agricultural Botany, University of Bonn, Meckenheimer Allee, D-53115 Bonn, Germany. 246 J. Ha1tori Bot. Lab. No. 85 I 9 9 8 studies were carried out in the Atacama Desert (Chile) and on the Galapagos Islands (Ecuador) by the authors at regular intervals from 1989 to 1997. All lichen samples were subjected to the conventional spot-tests (ST), anatomical (LM), microcrystallographical (MC), thin layer chromatographical (TLC), and high performance liquid chromatographical (HPLC) analyses, if necessary completed by UV spectroscopy (UVS) and melting point determinations (MP). For the anatomical descriptions, freezing microtome sections were used, the corresponding measurements being made with preparations in water. The standard methods for colour reactions, extraction procedures, MC, TLC, and MP described by Huneck & Yoshimura (1996) were followed, but to allow direct comparisons with former results HPLC and UVS were performed according to Follmann et al. ( 1994) with the aid of Beckman's "System Gold" and auxiliary equipment. For co-chromatographical verifications (TLC, HPLC), the Roccellaceae analysed by Follmann et al. (1993) were available, each treatment being repeated at least five times. Finally, all morphological and chemical data were evaluated numerically and confirmed statistically (Forey et al. 1994, Pankhurst 1991 ).

R ESULTS ANO DISCUSSION Pentagenella fragillima Darbishire ( 1897) Roccella fragilissima Montagne in herb. (s.a.). Roccella fragillima (Darbishire) Choisy, comb. inval. ( 1957). As already noted by Darbishire (1898 a), the type collection of Pentagenella fragillima (Herb. C. Montagne, s.a., s.n., PC, actually on loan at KOELN) un­ fortunately consists of a few fragments of a fruticose thallus only, the largest being 35 mm long and in places 5 mm wide, but the others being much smaller. Except Darbishire's revision slip of 1897, there are two labels of Montagne's hand which read as follows: 1. " Rocce/la fragilissima Mont. an R. gracilis Bory?, Coquimbo, Chili, M. Webb" and 2. " Roccellafragilissima Mont. ined. ou a R. taeniata Mont. ined. diversa? Coquimbo in Chile, corn. am. Webb" . R. gracilis Bory is known to be a synonym of R. phycopsis Ach. , and R. taeniata Mont. in herb. belongs to R. portentosa (Mont.) Darb. Consequently, both alternatives can be neglected in virtue of various deviating ch aracters. Deduced from the dimensions of the fragments, the thallus of Pentagenella fragi/lima Darb. might have been 10 to 15 cm high , and the fragile and rigid consistence of the sections points to an upright growth. To judge from a computer simulation, the habit came nearer to that of the African Roccella hypomecha (Ach.) Bory (see Follmann et al. 1994, Fig. 3) than that of the South American Roccel/a portentosa (Mont.) Darb. as insinuated by Darbishire (1898 a). Nevertheless, it should be taken into account that no trace of a holdfast or protothallus could be found, characters included in the generic protologue by the same author (I.e.), but he was certainly right to conclude from the few references at hand that the taxon occurred on rocky ground. Beyond that, the detection of sea salt on the cortex points to a halophytic behaviour typical for most Roccellaceae. G. FOLLMANN et al. : Pentagenella and Roccellodea 247

Fig. I. Type material of Pentagenel/a fragil/ima Darb. with (A) part of largest fragment, note crateriform pycnidia (diam. of branch 1.5 mm) and (B) plate-shaped pseudocarps (diam. of largest one 1.3 mm [phot. B. Werner]).

Wherever possible, the following abbreviated description, focused on essential characters, combines Darbishire's (1897, 1898 a, b) with our own observations: Thallus fruticose, podetia terete (mean diam. 1.2 mm) to applanate (about 0. 7 x 3.0 mm), ash-grey, rough and pruinose, without isidia or soralia, dichotomously branched, extremely fragile (Fig. I); cortex (mean diam. 75 µm) with anticlinally arranged, smooth hyphae (mean diam. 4 µm), in the light outer part more loosely packed, in the darker inner part compact, anastomosing, and interspersed by different types of crystals including calcium oxalate, brownish, granular surface gel only in the inner part; algal laY.er continuous (mean diam. 50 µm), trentepohliaceous phycobionts often deformed, solitary (mean diam. 12 µm) or in short fi laments, covered by hyphal appressoria; medulla light (mean diam. 70 µm), hypomedulla brown, both formed by more or less periclinal strands of conglutinate, verrucose, thick-walled, anastomosing hyphae difficult to distinguish, hypomedullary strands interspersed by brownish, granular, 248 J. Hattori Bot. Lab. No. 85 I 9 9 8

Fig. 2. Type material of Pentagenella fragillima Darb. with discothecia on pseudopodetium-like excrescences, droplets consisting of glue used to fix fragment on herbarium sheet (diam. of ascocarp below droplet 0.4mm [phot. B. Werner]). crystalliferous thallus gel; cortex, medulla, and thalline margin of pseudothecia C+ pink (fugitive), K+ ochre, P+ orange-red, containing erythrin and protocetraric acid (Fig. 3A). Lateral and lamina) ascomata apparently numerous, but supposed larger, thallus-coloured "discocarps" (Fig. 1) in reality plate-shaped vegetative excrescences (mean diam. 1.2 mm) normally bearing several lecanorine pseudothecia (Fig. 2): Roundish, sessile, base barely constricted, plane to subconvex (mean diam. 0.4mm), disk blackish, pruinose, thalline margin relatively thick, crenated, corticate, algiferous; epithecium brown, granular (mean diam. 30 µm), hyphae cylindrical, branched, intertwined; proper exciple a parathecium, dark-brown (mean diam. 45 µm), hyphae slightly clavate; hypothecium brown, not extending down into the medulla (mean diam. 90 µm), hyphae densely covered by crystalloids and granula; hymenium with multiascal locules, pluricarpocentral, hyaline (mean diam. 75 µm), subhymenium G. FOLLMANN et al.: Pentagenella and Rocce/lodea 249

0,20 2 A

0,15

~ ~ 0,10 .0 0 rJJ .0

"' 0,05

0,00

0 10 20 30 40 50 60 70

0' 20 .----rr-..----,--..----,--..----,--..----,--..----,--~--,--.---~ 4 B

0, 15

~ ~ 0,10 .0 0 rJJ 3 .0 "' 0,05

0,00

0 10 20 30 40 50 60 70 retention time [min] Fig. 3. HPLC-profiles (reversed phase) of acetone extracts of type material of (A) Pentagenella fragillima Darb. and (B) lngaderia friabillima Follm. & M. Schulz (I = erythrin, 2 = protocetraric acid, 3 = conpsoromic acid, 4 = psoromic acid [prep. B. Werner]). distinct, light-brown (mean diam. 25 µm), hamathecium paraphysoidal, paraphysoids septate, branched, anastomosing, asci clavate (about 18 x 60 µm), apical cap com­ paratively inconspicuous, amyloid (Kl + blue), eight-spored, ascospores biseriate,. colourless, fusiform, straight, relatively thin-walled, three-septate (about 6 x 19 µm); pycnidia numerous, externally crateriform (ostiole about 50 µm in diam.), corpus globular, black (mean diam. 90 µm), pynoconidia "exobasidial", filiform, curved (about 0.8 x 21 µm). 250 J. Hattori Bot. Lab. No. 85 I 9 9 8

The exceptional brittleness of the dry branches of Pentagenella fragillima Darb. is undoubtedly related to the deposition of unusual quantities of crystalloids and granules in virtually all layers. The first are largely consisting of protocetraric acid determined in situ by crystal form, colour reactions, and solubility in various solvents. According to a quantitative analysis of the acetone extract of a microfragment of P. fragillima, the concentration of the depsidone amounts to approximately 20% d.w. and that of the accompanying depside erythrin to less than 2% d.w. In general, chemosyndromes like this are rare among the Roccellaceae, but are characteristic for the manifold and polymorphous Roccella population of the Galapagos Islands (Follmann et al. 1993, Follmann 1994). Darbishire (1897, 1898 a, b) ignored the noticeable false discocarps of Pentagenella fragillima Darb. which come closer to the cup-shaped pseudopodetia of the sympatric Follmanniella scutellata Peine & Wern. than the sprouting primary pseudothecia of the Caribbean Roccella verruculosa Follm. Regrettably, the scanty type material did not allow a clarification of the·ontogeny of the excrescences and true discothecia under consideration. Two half-sectioned "pseudocarps" found among the fragments might be relics ofDarbishire's activities and perhaps could explain his statement of"colourless hypothecia and parathecia". Other differences between his and our observations, for instance regarding the form and size of the pycnidia, appear to be of minor importance. However, in this context it is indispensable to point to the two misleading figures of a "cross-section of a podetium and an apothecium" (Darbishire 1897, Fig. 6, 1898 a, Fig. 14), where neither the protruding thalline margin nor the dark-brown parathecium nor the clearly contrasted subhymenium and hypothecium are shown, while the paraphysoidal hamathecium is missing completely. All observations considered, the taxonomical meaning of the name Pentagenella (Greek penta = five, Lat. gena -> genella = little cheek, small sidewall) remains unclear: Did the author refer to the relatively thick thalline margin of the discocarps found up to fivefold on the supposed pseudopodetia? Although the foregoing description applies to incomplete fragments of a poor collection, the thallus, ascoma, and pycnidium organization of Pentagenella proves to be widely identical with that of Roccella. Except the pseudopodetium-like excrescences, the only remaining differential character consists in the deviating colouration of the hypothecium and parathecium: brown and dark-brown in Pentagenella, dark-brown or carbonaceous in Roccella. Because these are merely gradual differences, there is no reason to maintain Pentagenella as a separate, monotypic genus; instead it should be placed in the synonymy of Roccella as already proposed by Choisy (1957). However, the species itself has to be uphold because its character combination differs sufficiently from all other Roccella taxa known. Unfortunately, Choisy's (I.e.) earlier combination without basionym, reference, and argumentation is unvalid and unfounded, and so is his introduction of a sect. Pentagenella under Roccella with an incorrect reference and devoid of a diagnosis. As demonstrated, both proposals appear to be justified and are validated here: G. FoLLMANN et al.: Pentagenel/a and Rocce//odea 251

Roccella A. P. de Candolle sect. Pentagenella (Darbishire) Follmann stat. nov. (Bas. Pentagenella Darbishire pro gen., 0. V. Darbishire in Ber. deutsch. bot. Ges. 15: 7; 1897). Spee. typ. Roccella fragillima (Darbishire) Pollmann comb. nov. (bas. Pentagenella fragillima Darbishire, 0 . V. Darbishire in Ber. deutsch. bot. Ges. 15: 5; 1897). Sectio nova generis Roccella, differt a sectione typica excrescentiis disciformibus pseudopodetia similaribus, hypothecio et parathecio infuscatis, thallis acidum proto­ cetraricum continentibus. Irrespective of certain similarities it becomes evident that the Chilean taxon termed Pentagenella fragillima Darb. unaware of the nomenclatural type does not correspond to the species described above. Consequently, the binomial has to be cancelled from the phytochemical literature where it ranges among the lichens with the highest accumulation rate of secondary products ever found (30% d.w. psoromic acid: Huneck & Follmann 1967, Follmann et al. 1993). It should be deleted, too, from ftoristical and phytosociological inventories where it shows up as a characteristic element of the saxicolous and corticolous lichen vegetation of the rocky Central and North Chilean littoral (e.g., in the common Rocce/linetum cerebriformis Follm.: Peine & Werner 1995, Follmann 1997). Although the senior author collected thousands of fruticose Roccellaceae from the Province of Coquimbo and the Atacama Desert in a broader sense from 1960 to 1997, not a single specimen was found approaching P. fragillima s. str. Obviously this became extinct in recent times like Austroplaca johowii (Follm. & Redon) Ryan, Protoroccella minima (Sant.) Follm., or Santessonia cervicornis (Follm.) Follm. (Follmann 1995, 1998). On the contrary, the species mistaken for P. fragillima represents a sporadically frequent endemic of the northern Chilean coast and turned out to be a hitherto undescribed Roccellaceae which is treated hereafter. Ingaderia friabil/ima Follmann & M . Schulz, sp. nov. Species nova fruticulosa, corticicola vel saxicola, ex affinitate lngaderiae pulcherrimae Darb. (Roccellaceae); differt thallo niduliforme, valde friabile, gompho nigro, globoso, forte diffisso, non algifero; ramulis projectis vel erectis, sub ascocarpiis plerumque geniculatis, scabris vel verruculosis, normaliter subcylindricis, interdum inopinate late applanatis, plerumque excrescentiis filiformibus producentibus; strato corticale et medullare subfuscatis, crystalliferis, C-negativo, K+ et P+ luteo, FeCl3 rubro; thallo acidum psoromicum et conpsoromicum continenti; ascocarpiis raris, loculis multiascaliis, monocarpocentralibus, lateralibus, crebrius compluribus, rotundatis vel tortuosis, margine thallino crassiuscula, disco piano vel subconvexo, subnigro, granuloso; epithecio, parathecio et hypothecio fusco, hamathecio paraphysoidali, hymenio hyalino; asci clavatis, octosporis, ascosporis fusiformibus, triseptatis; pycnidiis leviter prominentibus, superne nigris, subglobosis, pycnoconidiis curvatulis. Locus typicus: Chile, Provincia de Antofagasta, Quebrada de Ja Chimba, ad saxa dioritica, 300 m, SW; leg. G. Follmann & M. Schulz (VIII/ 1993), holotypus KOELN 34 896 a, isotypus B 34 896 b. In analogy with Roccella fragillima (Darb.) Follm. the new species is named lngaderia friabillima Follm. & M . Schulz. (Lat. friabilis =crumbly, friable). As well 252 J. Hattori Bot. Lab. No. 85 I 9 9 8 as in R. fragillima the extreme brittleness of the dry thalli of I. friabillima originates from the ample deposition of crystalloids and granules between the hyphae of the vegetative and generative layers, here particularly of the depsidone psoromic acid (determination in situ as described before). According to Huneck & Follmann (1967) and Follmann et al. (1993), the concentration of psoromic acid oscillates between 20 and 30% d.w., and that of the accompanying conpsoromic acid amounts to about 0.3% d.w. (as detected in the acetone extract of a type fragment). The concentration of other phenolic by-products appears to be so low that they may be neglected in this context (Fig. 3). Within the Roccellaceae, the psoromic acid chemosyndrome seems to be restricted to several endemic representatives of the coastal belt of the Atacama Desert where they form characteristic communities (e.g., Ingaderietum graci/limae Follm.), a phenomenon probably related to the harsh ecological conditions. For the following description, Ingaderia friabillima Follm. & M. Schulz was contrasted with the type material of I. pulcherrima Darb. (Herb. C. Montagne, leg. C. Gay 1838, PC, s.n., actually on loan at KOELN) which represents - contrary to the preceding species - a really extensive sample. In view of an uncorrect type citation (Tehler 1990) it seems to be necessary to exhibit the complete labelling of the corresponding herbarium sheet in this context: I. " Evernia ochroleuca Fr. var. crinalis Fr. (Alectoria crinalis Ach.), Chili, Prov. de Coquimbo, M. C. Gay 1838" (scrips. Montagne), 2. " Roccella tinctoria DC. * alectoroides Nyl." (scrips. Montagne), 3. " Jngaderia pulcherrima Darb. (type) 1897" (scrips. Darbishire). The last-named took a duplicate from the original sample and copied the labels literally as given above; this isotype is preserved at BRJST (Herb. 0 . V. Darbishire, s.n., actually on loan at KOELN). A brown-red halo is visible around the type species glued on white paper, a common observation in case of older lichen samples containing larger quantities of lecanoric acid (e.g., Roccella portentosa [Mont.] Darb.). Apart from the incomplete nomenclatural data, the specimen selected by Tehler ( 1990) cannot be taken for the type of I. pulcherrima, if only because of the obviously missing intensive c+ blood-red reaction of the cortex and medulla. At a glance, Jngaderia friabillima Follm. & M. Schulz looks like a stockier and sturdier morphotype of I. pulcherrima Darb., and therefore the abridged description is keyed to the relevant differences between both taxa: Thallus rotund, nest-like (Fig. 4), rigid and extremely friable, light-grey or yellowish grey (mean diam. 5 cm), holdfast blackish, globular, deeply fissured, without phycobionts; branches largely uniform, most of them sprouting directly from the holdfast, projecting or erect, below ascocarps often geniculate, dichotomously ramified, sinuous and disorderly interwoven, normall y subcylindrical (mean diam. 0.5 mm), but sometimes unexpectedly broadened, tortuous lobes (up to 5 x 15 mm) occasionally bearing filiform branchlets which easily break off and might act as isidia, surface rough and (or) verrucose, esorediate; cortex (mean diam. 30 µm) with periclinally arranged hyphae, sometimes single outer ones partly projecting, cortical gel brownish, granular, crystalliferous, algal layer discontinuous, phycobionts trentepohliaceous (mean diam. 12 µm), distributed in paucicellular, elongated groups oriented to the branch axis, surrounding hyphae (mean diam. 2 µm) G. FOLLMANN et al. : Pemagenella and Roccel/odea 253

Fig. 4. lngaderia friabillima Follm. & M. Schulz at the Atacamian type locality (A) on granodioritic rock, note sporadically flattened branch lets (diam. of larger thallus 6 cm) and (B) on columnar cactus (diam. oflarger thallus 5 cm [phot. G. Follmann]). forming appressoria, medulla brownish, comparatively inconspicuous, coalescent, hypomedulla absent, thallus gel like surface gel; cortex and medulla C-negative, K+ and P+ yellow, FeC1 3+ ruby-coloured, containing plenty of psoromic acid and a little of conpsoromic acid (Fig. 3B). Tehler's (1990) revision of several Chilean collections of lngaderia pulcherrima Darb. resulted in the assumption that the two types of ascomata observed belong to lichenicolous fungi. Those lirelliform ascocarps with 7- to 8-septate spores were used by Darbishire ( 1897) for the diagnosis of the species under discussion, and discothecia with 2-septate spores are present in the type material as well , the last also known from f . gracillima (Krempelh.) Feige & Lumbsch. Tehler's (I.e.) interpretation is perfectly corroborated by our re-examination, all the more than f. friabillima Follm. & M. Schulz develops - though rarely - " typical" roccellaceous discocarps: Lateral, often in groups, roundish or undulated (mean diam. 0.8 mm), base slightly constricted, thalline margin pronounced, corticate, without phycobionts, disk plane to subconvex, blackish, pruinose, monocarpocentral with multiascal locules ; epithecium granular, brown (mean diam. 25 µm), parathecium inconspicuous, brown, hypothecium not extending down into the medulla, dark-brown (mean diam. 68 µm), hamathecium paraphysoidal, hymenium hyaline (mean diam. 72 µm), subhymenium delicate, light-brown, asci 254 J. Hattori Bot. Lab. No. 85 1 9 9 8 clavate (about 15 x 60 µm), 8-spored, apical cap amyloid (Kl + blue), overtopped by branched, septate, non-capitate, anastomosing paraphysoids, ascospores biseriate or triseriate, colourless, fusiform, straight, pointed, 3-septate (about 5 x 17 µm); pycnidia solitary, lateral or lamina!, slightly elevated, superficially black, corpus dark-brown, subglobular (mean diam. 0.2 mm), pycnoconidia "exobasidial", filiform, curved (about 0.6 x 20µm). The new taxon raises the species number of Ingaderia to four well delimited entities: Two are fertile(/. friabillima Follm. & M. Schulz, I. gracillima [Darb.] Feige & Lumbsch) and two obviously sterile (I. pulcherrima Darb., I. troglodytica Feige & Lumbsch). The last-named reproduce chiefly by fragmentation instead of soredia; but in view of essential morphological differences they cannot be regarded as secondary species of the former. Chemosystematically, the depsidone-bearing I. friabillima and I. gracillima represent derived taxa, whereas the depside accumulating I. pulcherrima and I. trog/odytica connect the genus to the more elementary lecanoric acid chemosyndrome characteristic for most members of the family. I. friabillima, I. graci//ima, and I. pu/cherrima are known exclusively from southern Pacific South America, while I. troglodytica occupies an outpost in the Atlantic Mediterranean. Concerning Roccellaceae, this most probably relictual disjunction only can be compared with the distribution pattern of Dirina paradoxa (Fee) Tehl. s. lat. extending from Galapagos and the Caribbees via Macaronesia to Socotra (Tehler 1983, 1986). The type locality of lngaderia friabillima Follm. & M. Schulz is situated in the nature reserve Quebrada de la Chimba, a lo ma formation ("fog-oasis") close to the Tropic of Capricorn, Province of Antofagasta, Atacama Desert, North Chile, where it forms luxuriant cushions on steep granodioritic rocks regularly coated by fog-banks from the nearby Pacific Ocean (300 m elev., SW; leg. G. Follmann & M. Schulz [VHI/ 1993), holotype KOELN 34 896 a, isotype B 34 896 b). Additional specimens examined are listed below (sites arranged from north to south): I. Cerro Moreno, Province of Antofagasta, North Chile, southwestern flank of the massif, vertical rock face, abundant, 550-600m elev., SW; leg. G. Follmann (VIH/ 1961 ), KOELN 34 332, 34 334, 34 336, 34 337, 34 359. - 2. Cerro Moreno, Province of Antofagasta, North Chile, south western flank of the massif, exposed cactus skeleton, scarce, 600 m elev., SW; leg. G. Follmann (VIII/1 991), KOELN 34 333. - 3. Quebrada de la Chimba, Province of Antofagasta, North Chile, central canyon, granodioritic rock, abundant, 250m elev., SW; leg. G. Follmann (lII/ 1961), KOELN 7650. - 4. Quebrada de la Chimba, Province of Antofagasta, North Chile, northeastern canyon, ribs of Eulychnia iquiquensis (Schum.) Britt. & Rose, frequent, 200 m elev., SW; leg. G. Follmann (TII/ 196 1), KOELN 7770. - 5. Loma de Paposo north ofTaltal, Province of Antofagasta, North Chile, fog-belt, columnar cacti, frequent, 550 m elev., W; leg. G. Follmann (X/ 1997), KOELN 34 897 . - 6. Coastal formation of Santa Barbara near Huasco, Province of Atacama, North Chile, thorny shrubs, scarce, 20m elev., NW-SW; leg. G. Follmann (lX/ 1962), KOELN 17 515. - 7. Rocky coast of Cachagua, Province of Aconcagua, Central Chile, cactus spines, abundant, 30 m elev., W: leg. G. H. Schwabe (V/ 1941 ), KOELN 25 806. - 8. Coastal strip of El Quisco, Province of Valparaiso, Central Chile, dry shrubs, frequent, 20 m, W; leg. M. Mahu (II/ 1965), KOELN 24 845 . According to the foregoing records, Ingaderia friabillima Follm. & M. Schulz inhabits the coastland between approximately 23 ° and 34° Slat. from North to Central G. FOLLMANN et al.: Pentagenella and Roccellodea 255

Chile and constitutes one of the many endemics among the Roccellaceae of this region considered as primary diversity centre of the family (Follmann 1968, 1998). Like other sympatric Roccellaceae (e.g., Dolichocarpus chilensis Sant., Roccellaria mollis [Hampe] Zahlbr., Roccellina luteola Follm.), the said species does by no means occupy this area of about 1200 km linear extension in a continuous way skipping larger tracts apparently suited for colonization possibly caused by recent ecological alterations. Interestingly, the population density increases in northern direction, while the individual vitality (e .g., height and weight of thalli) appears to be higher in the south. lngaderia friabillima Follm. & M. Schulz was met with as accessory species of the epipetric Roccellinetum cerebriformis Follm. (cover-abundance index +) or of the epiphytic lngaderietum gracillimae Follm. (cover-abundance index 1), two of the most important lichen associations dominated by Roccellaceae (Roccellion portentosae Follm.) of the central and northern Chilean coast (Follmann 1997, 1998). Conse­ quently, the chorological formula ("chorogram") of the new species runs as follows: Neotropical, Andean, subtropical-arid (Atacamian) to mediterranean (Central Chilean), oceanic, planar to colline, disjunct endemic, locally abundant, in communities of the Roccellion portentosae. After our field-notes, the ecological formula ("ecogram") largely coincides with those of other Pacific South American representatives of the family: aerohygrophytic, xeroresistent, ombrophytic, subphotophytic, thermotolerant (stenoclimatic), saxicolous to corticolous, subacidophytic to neutrophytic, halophytic, anitrophytic, toxicophobous, gregarious. Rocce/lodea nigerrima Darbishire ( 1932) Roccel/a nigerrima (Darbishire) Follmann, comb. prov. (1994). Like the type material of Pentagenella fragillima Darb. that of Roccellodea nigerrima Darb. (Herb. 0 . V. Darbishire, acq. 1899, s.n., BRIST, actually on loan at KOELN) consists of a few thallus fragments only. It is labelled " Herb. W. G . Farlow, Roccella tinctoria DC., Galapagos, Hassler Expedition, 1872" (scrips. Farlow), supplemented by Darbishire's hand "Certainly not R. tinctoria DC.: Roccellodea nigerrima Darb. sp. nov., 1932,

Fig. 5. Details of fruiting branches of type material of (A) Roccellodea nigerrima Darb. (diam. of branch 0.9 mm) and (8) Rocce/la obscurissima Follm. & B. Wern. (diam. of branch 2.1 mm [phot. G. Follmann]).

Habit approaching that of Rocce/la galapagoensis Follm. s. !at. (R. portentosa [Mont.] Darb. of previous authors); holdfast hemispherical, warty, corticate, algiferous, externally thallus-coloured, internally black; terete, whitish to yellowish light-grey branches somewhat knotty (Fig. 5A); all layers interspersed by brownish crystalloids and granules of different size (no calcium oxalate found), cortex hyaline, except hyphal tips conglutinate (mean diam. 55 µm), hyphae thick-walled, surface gel yellowish, trentepohliaceous phycobionts often in articulate filaments (mean width of single cells 15 µm), hyphae of algal layer provided with appressoria and haustoria, medulla slightly brownish, hyphae anastomosing and reticularly intertwined, gradually merging into the shiny violet-brown (with naked eye brilliant-black) hypomedulla superficially resembling the black "central cord" of Simonyella variegata J. Stein. and occupying about two thirds of the sectional area of a branch, composed of more or less periclinally oriented strands of a few voluminous, double-walled hyphae (mean exterior diam. 9 µm, lumen about 2 µm), leaving open ample, air-filled, intercellular cavities; all STs negative, but fragments immersed in K for a while showing a yellow halo, dark melanin-like pigment unsoluble in organic solvents; thallus containing galapagin and roccellin (Fig. 6A). G. FOLLMANN et al. : Pentagenel/a and Roccel/odea 257

Darbishire's ( 1932) description of discothecia found on thallus fragments of Roccellodea nigerrima Darb. also harmonizes to a large extent with our observations. Nevertheless, the immersed or slightly raised, reddish brown to blackish ascocarps of irregular or indefinite contour, surrounded by somewhat cracked cortical plectenchyma, and underlayed by the apparently unchanged algal stratum turned out to belong to a lichenicolous . Observed on other Galapagoan Roccellaceae as well, all characters point to an undescribed Arthonia species completely deviating from the only lichenicolous representative of the genus hitherto known to occur on the Galapagos Islands (A . follmanniana Dieder. on Roccella galapagoensis Follm. s. lat.). This presumedly new species will be treated elsewhere together with other parasymbionts of Roccellaceae. Authentic ascomata are absent from the poor type material but were found on additional collections (see below). Discocarps lateral, outline equally irregular (Fig. SA), no true thalline margin formed, juvenile ascocarps covered by rests of ripped up cortex, disk immersed (up to 2 mm in diam.), plane to somewhat elevated, epruinose, black, pluricarpocentral with multiascal locules; epithecium blackish, granular (mean diam. 40 µm), no clearly contrasted parathecium developed, hypothecium carbonaceous (about 250 µm in diam.), not extending down into the medulla, after spore release often thickening while shedding epithecial and hymenial layers, coating large parts of older branches with smooth, deep-black patches, hymenium brown (mean diam. 90 µm), hamathecium paraphysoidal, paraphysoids septate, anastomosing, rarely branched, non-capitate, asci clavate (about 15 x 80 µm), apical cap amyloid (Kl+ blue), 8-spored, ascospores biseriate, yellowish to brownish, fusiform (about 7 x 23 µm), straight, pachydermous, 3-septate; pycnidia (after type material) solitary, lateral, relatively large (up to 0.5 mm in diam.), superficially black, immersed (ostiole about 40 µm in diam.), corpus subglobular, dark-brown, pycnoconidia "exobasidial'', filiform, curved (about 0.8 x 15µm) . The first chemical analysis of a lichen sample designated Roccellodea nigerrima Darb. from San Cristobal, Galapagos, yielded psoromic acid (Follmann & Huneck 1969), the second from Rabida, Galapagos, protocetraric acid (Follmann et al. 1993). In accordance with the studies at hand, the material from San Cristobal represents the externally similar new Roccella obscurissima Follm. & B. Wern. treated in the following, that from Rabida a dark morphotype of R. galapagoensis Follm. s. lat. Consequently, the details included in phytochemical inventories under R. nigerrima have to be replaced by galapagin and roccellin. As both these chromones and depsidones are known from the R. ga/apagoensis complex too, neither the equipment with secondary products nor the vegetative and generative thallus organization justify an eventual maintenance of a separate genus Roccellodea. That is why the provisional combination proposed by Follmann (1994) is validated here. Roccella nigerrima (Darbishire) Follmann, comb. nov. (Bas. Roccellodea nigerrima Darbishire, 0 . V. Darbishire in Ann. Crypt. exot. 5: 157; 1932). As explained before, the type locality of Roccella nigerrima (Darb.) Follm. on 258 J. Hattori Bot. Lab. No. 85 I 9 9 8

0,20 I 6 A

0, 15 .... - 5

0,00 - \ I l J. I ~ • ""- ' 0 10 20 30 40 50 60 70

0 ,20 .---,,--,..----,---,..----,---,..----,---,..----,---,..----,---~--.---~~ 4 B 7 0, 15

~ ~ 0,10 .Q 0 9 CJ) .Q 8 C1l 0,05

0,00

0 10 20 30 40 50 60 70 retention time [min] Fig. 6. HPLC-profiles (reversed phase) of acetone extracts of type material of (A) Roccellodea nigerrima Darb. and (B) Roccella obscurissima Follm. & B. Wern. (4 = psoromic acid, 5 = roccellin, 6 = galapagin, 7 = unidentified phenolic product, 8 = lecanoric acid, 9 = schizopeltic acid [prep. B. Werner]).

San Salvador, Galapagos, cannot be stated with absolute certainty. In any case, the species must be graded as rare, because in the largest collections of Galapagoan Roccellaceae available (Follmann & Werner KOELN, Sanchez-Pinto TFMC, Weber COLO) only two more specimens of R. nigerrima s. str. came across: 1. Galapagos, Isla Pinzon, on volcanic rocks, apparently scattered, 90 m elev. ; leg. L. Sanchez-Pinto (IV/1990), KOELN 34 898, TFMC 6600. - 2. Galapagos, Isla Seymour Norte, on volcanic rocks, apparently gregarious, 10 m elev.; leg. L. Sanchez-Pinto (IV / 1990), G . FOLLMANN et al.: Pentagenel/a and Roccellodea 259

KOELN 34 899, TFMC 6609. The latter were used for the description of authentic ascocarps. With regard to the high conformity of the essential chorological, ecological, and sociological attributes of the three Galapagoan Roccella taxa traced here, these are detailed jointly under R. margaritifera B. Wern. & Follm. Roccella obscurissima Follmann & B. Werner, sp. nov. Species nova fruticulosa, saxicola, ex affinitate Roccellae nigerrimae (Darb.) Follm. (Roccellaceae); differt thallo maiore, solutiore, pendulo, suffiavo vel subfusco, gompho piano, interne fusco vel subnigro, ramulos principali directe generanti, ramulis basalibus modice subdichotome divisis, ad extremum scopuliformibus; phycobiontibus irregulariter dispositis, hypomedulla fusca, hyphis pachydermaticis reticulate intricatis; cortici C+ rosea, K+ suffiava, medulla K+ lutea, KC+ rubrofusca, P+ rubra versus rubrofusca, FeCl 3 + rubra; thallo acidum lecanoricum, protocetraricum et schizo­ pelticum continenti; ascocarpiis plus minusve rotundatis, sine margine thallina, disco in juventute pruinoso, in adulescentia nigro, parathecio subnigro, hamathecio fusco bene evoluto, hymenio alteriore, pycnidiis minoribus. Locus typicus: Insulae Galapagos, Pinzon, ad saxa basaltica, 20 m, S; leg. L. Sanchez-Pinto (IV/ 1990), holotypus KOELN 34 029, isotypus TFMC 5070. In view of various similarities to Roccella nigerrima (Darb.) Follm. the new taxon is named R. obscurissima Follm. & B. Wern. (Lat. obscurissimus = very dark), especially on account of the large black patches on the branches caused by uncovered hypothecia and the unusual gloomy aspect of the thalli. However, despite all resemblances to R. nigerrima and an apparently higher degree of variability of R. obscurissima both species can be distinguished by a series of deviating characters: Thallus larger, more loose, yellowish or brownish, holdfast robust, plane, internally dark-brown to blackish, packed by main branches (up to 20cm long), in the beginning projecting, later pendent, basal parts (up to 2.5 mm in diam.) not often sub­ dichotomously ramified, upper parts (up to 0.5mm in diam.) sometimes broom-like divided (Fig. 7A); cortex often scarred or areolate, comparatively thin (mean diam. 40 µm), trentepohliaceous phycobionts (mean diam. of single cells 11 µm) in roundish colonies of varying size, hypomedulla well developed, brown to dark-brown, composed of thick-walled, more or less reticularly arranged hyphae (mean diam. 5 µm) leaving open interfibrillar spaces; cortex C+ pink, K+ yellowish, medulla K+ yellow, KC+

brown-red, P+ red to brown-red, FeC1 3+ red; thallus containing lecanoric, psoromic, and schizopeltic acid plus an unidentified phenolic compound (Fig. 6B), psoromic acid being the main product (about 12% d.w.). Ascomata numerous, lateral, at least when young roundish (fullgrown up to 2.7mm in diam.) and often pruinose, cortex lifted by developing pseudothecia but no true thalline margin formed (Fig. 5B), unchanged edge of fracture long time visible. as light ring around adult jet-black, immersed disks; epithecium blackish (mean diam. 52 µm), parathecium distinct, dark-brown to blackish (mean diam. 46 µm), hypothecium carbonaceous (about 300 µm in diam.), hymenium brown (mean diam. 102 µm), hymenial strands funnel-shaped (outer diam. about 45 µm), asci resembling 260 J. Hattori Bot. Lab. No. 85 I 9 9 8 those of Roccella nigerrima (Darb.) Follm., ascospores brownish, broad fusiform, poles slightly rounded (about 10 x 28 µm); pycnidia black, immersed, globular (mean diam. 65 µm, ostiole about 15 µm in diam.), pycnoconidia filiform, curved (about 0.8 x I 0 µm). Additional specimens examined (islands arranged from north to south): I. Galapagos, Isla San Salvador, Bahia Sullivan, eastern coast, hard volcanic rocks, rare, 20 m elev.; leg. P. Siebrecht (ll/ 1992), KOELN 35 009. - 2. Galapagos, Isla Rabida, on volcanic rocks, apparently scattered, 30 m elev.; leg. L. Sanchez-Pinto (IV / 1990), KOELN 34 034, TFMC 5072. - 3. Galapagos, Isla Pinzon, caldera rim, northern slope, lava cliffs, about 300 m elev.; leg. D. Cavagnaro (II/ 1964), COLO 188 439 (L-40 760). - 4. Galapagos, Isla Santa Fe, summit of the island, volcanic rocks, about 250 m elev.; leg. C. B. Koford (I- ITI/ 1964), COLO 188 445 (L-40 754). - 5. Galapagos, Isla Santa Fe, on volcanic rocks, probably close to the coast, apparently scattered; leg. R. Orr (l- III/ 1964), COLO 190 242 (L-41 005). - 6. Galapagos, Isla Santa Fe, northern coast, on basalt, apparently scattered, 50m elev.; leg. L. Sanchez-Pinto (IV/ 1990), KOELN 35 001, TFMC 6593 . - 7. Galapagos, Isla Floreana, south side, on base of lava talus, together with Roccella margaritifera B. Wern. & Follm.; leg. W. A. Weber & J. Lanier (IV/ 1976), COLO 297 015 (L-63 037). After this, Roccella obscurissima Follm. & B. Wern. appears to be more widespread on the Galapagos Islands than R. nigerrima (Darb.) Follm., but at none of the known localities it reaches a higher frequency. The different island populations show a certain degree of variation with regard to the strength of colour reactions, concentration of secondary products, thallus colour and form, or anatomical dimensions. Such minor intraspecific deviations, often observed with Roccellaceae of other habitats as well, may be caused by differences in the history of settlement or period of isolation on the single islands and do not deserve taxonomic consideration. Roccella margaritifera B. Werner & Pollmann, sp. nov. Species nova fruticulosa, saxicola, ex affinitate Roccellae galapagoensis Follm. (Roccellaceae); differt gompho amplio, piano vel subgloboso, interne nigro, thallo multiforme, absentia ascomatorum et pycnidiorum, soraliis numerosis, magnis, breve pedunculatis, capitatis, crasse granulatis, basi profunde immersa, soraliis nonnunquam isidia digitata, leviter ramificata formantibus; hypomedulla fusca, hyphis pachydermati­ cis !axe intricatis; cortici C+ rosea, medulla K+ ft ava, P+ rufa, soraliis P+ suffiavis; thallo erythrinum et acidum protocetraricum continenti. Locus typicus: Insulae Galapagos, San Cristobal, Loberia prope Puerto Baquerizo, ad saxa basaltica, !Orn, SW; leg. L. Sanchez-Pinto (III/ 1991), holotypus KOELN 34 900, isotypus TFMC 6616. The fragments of a second species contained in the original capsule of Roccella nigerrima (Darb.) Follm. (see above) correspond completely to the new R. margaritifera B. Wern. & Follm., but for obvious reasons another collection was selected as type material. The specific epithet points to the all-round cover of the branches by eye-catching soralia giving - at least to the pendulous form (Fig. 7B) - the appearance of a string of pearls (Lat. margaritifer = pearl bearing, producing). According to our anatomical and chemical comparisons, it is closely related to the fertile (primary) R. galapagoensis Follm. s. str. and can be taken for the sorediose (secondary) counterpart of this. Therefore, the following description is focused on substantial differences G. FOLLMANN et al. : Pemagene/fa and Roccel/odea 261

Fig. 7. (A) Type material of Roccella obscurissima Follm. & B. Wern. with witches' broom-like ramification of apical part (diam. of broken branchlet with dark medulla in the centre 0.8 mm) and (B) type material of R. margaritifera B. Wern. & Follm., note cloddy soralia (length of largest branch 8 cm [phot. G. Follmann]). between both taxa. Thalli polymorphous, varying from pendent to projecting and upright types (pendent ones up to 20 cm long, others normally not surpassing 10 cm), holdfast extensive, flattened to hemispherical, externally corticate and thallus-coloured, internally black, most branches originating directly from the holdfast, often tortuous, upper parts not or sparsely ramified (mean diam. 2 mm, largest diam. measured 4 mm), if present secondary branchlets here and there pectinate (mean diam. 0.4 mm), neither ascomata nor pycnidia present; except lowest parts, tips, and branchlets main branches on all sides densely covered with relatively large, normally capitate, partly subdivided soralia (up to l mm in diam., if compound up to 2 mm, Fig. 7B), these coarse-grained, marginate, pedunculate, stalk constricted, light globular base deeply extended into the voluminous, non-fibrillate, brown hypomedulla, soralia sometimes producing digitate, rarely ramified isidia; cortex C+ pink, medulla K+ yellow, P+ orange-coloured, soralia P+ yellowish; main secondary products erythrin and protocetraric acid. Selected additional specimens studied (islands arranged from west to east and from north to south, only one characteristic sample being mentioned for each island): I. Galapagos, Isla Darwin, on lava cliffs, apparently scattered; leg. D. Cavagnaro (l ~ HI / 1964), COLO 190 243 262 J. Hattori Bot. Lab. No. 85 I 9 9 8

(L-41 004). - 2. Galapagos, Isla Wolf, near top of the island, on lava rock cliffs, west-facing exposure; leg. D. Cavagnaro (I- IIl/ 1964), COLO 190 239 (L-41 008). - 3. Galapagos, Isla Genovesa, volcanic rocks, apparently abundant, 10- 30m elev .; leg. H. K. Svenson (IV/ 1930), KOELN 34 774. - 4. Galapagos, Isla Isabela, cinder cone above Tagus Cove, on rocks, apparently gregarious; leg. W. A. Weber & J. Lanier (V/ 1976), COLO 297 033 (L-63 288). - 5. Galapagos, San Salvador, close to Bahia Sullivan, stream of lava, apparently gregarious, 40 m elev.; leg. L. Sanchez-Pinto (III/ 199 1), KOELN 35 008, TFMC 6623. - 6. Galapagos, Isla Bartolome, exposed rock cliff at west end of the island, on projecting rock points, abundant, 15m elev.; leg. L. H. Pike (IV/ 1971), COLO 255 647 (L-55 219). -7. Galapagos, Isla Rabida, coastal formation, on volcanic rocks, apparently gregarious, 30 m elev.; leg. L. Sanchez-Pinto (IV/ 1990), KOELN 34 031, TFMC 5071. - 8. Galapagos, Isla Baltra, volcanic cliffs close to the main landing, scattered, 20 m elev., SW; leg. G. Follmann & B. Werner (11 / 1994), KOELN 34 772. - 9. Galapagos, Isla Pinzon, volcanic rocks close to the shore, apparently abundant, !Orn elev.; leg. L. Sanchez-Pinto (IV/ 1990), KOELN 35 007, TFMC 6602. - 10. Galapagos, Isla Santa Cruz, vicinity of Bahia Academia, volcanic rocks, 10 m elev.; leg. W. A. Weber (l- lll/ 1994), COLO 185 684 (L-40 036). - 11. Galapagos, lslote Plaza Norte opposite to the east coast of Santa Cruz, lava plateau, together with other Roccella species; leg. W. A. Weber (I- III/ 1964), COLO 190 302 (L-40 889). - 12. Galapagos, Islote Plaza Sur opposite to the east coast of Santa Cruz, lava plateau, together with other Race/la species; leg. W. A. Weber (l- lll/ 1964), COLO 190 383 (L-40 883). - 13. Galapagos, Isla Santa Fe, west side, close to summit, volcanic rocks, about 200 m elev.; leg. C. B. Koford (l- IH / 1964), COLO 188 444 (L-40 755). - 14. Galapagos, Isla San Cristobal, Loberia south of Puerto Baquerizo, basaltic rocks, abundant, 30 m elev., W; leg. G. Pollmann & B. Werner (lll/ 1994), KOELN 34 923. - 15. Galapagos, Isla Floreana, north coast, Punta Cormoran, basaltic rocks, apparently gregarious, !Orn elev.; leg. L. Sanchez-Pinto ([V/ 1990), KOELN 35 006, TFMC 6608. - 16. Galapagos, Islote Campeon opposite to the northeastern coast of Floreana, south side, bare lava talus, associated with R. galapagoensis Follm. s. str.; leg. W. A. Weber & J. Lanier (IV / 1976), COLO 297 015 (L-63 037). - 17. Galapagos, Isla Espafiola, Punta Suarez, volcanic rocks, apparently scattered; leg. T. Papenfuss (I- III/ 1964), COLO 190 366 (L-40 879). During the examination of many more samples than listed above, not a single thallus was met with bearing ascomata and soralia simultaneously. Previous mis­ understandings certainly were caused by broken branches of fertile and sorediose individuals kept together in a single envelope, especially in older collections. This also happened with the type material of Roccella galapagoensis Follm. from Santa Cruz (KOELN 25 153, Lich. exs. COLO 11 2), the protologue of which mentions among other attributes "Ramulis sorediosis ... apothecia numerosa". Regardless of a current revision of the copious Galapagoan Roccella population as a whole, based on comparative DNA analyses, the required emendation of the primary species as being esorediose, provided with a brown medulla, and so forth is executed here: Roccella galapagoensis Follmann emend. Follmann (Bas. Roccel/a galapagoensis Follmann, G. Follmann in Nova Hedwigia 15: 337; 1968). A detailed discussion of the distinctions between, and the relations among, the Galapagoan Roccella taxa, all of which most probably can be traced back to R. portentosa (Mont.) Darb. from the Pacific coast of Chile and Peru, is reserved to the G. FoLLMANN et al.: Peniagenella and Roccellodea 263 before-mentioned study which will include a determination key, too. If only single specimens of Roccella margaritifera B. Wern. & Follm. are inspected, the different growth forms varying from slender, long pendent, slightly winding to stocky, crooked, octopod-shaped thalli can easily be taken for separate species. However, since continuous transitions occur, these morphotypes are not distinguished taxonomically for the present. Within certain limits, the content of secondary metabolites also appears to be variable: Due to a HPLC screening of about l 00 samples from all islands mentioned in the preceding list, the depside erythrin and the depsidone protocetraric acid represent constant constituents. These may be supplemented by roccellic acid (aliphatic acid), acetylportentol and portentol (cycloaliphatic com­ pounds), lecanoric acid (depside), and galapagin and roccellin (chromones) in varied, mostly low concentrations and differing combinations. The distributive patterns con­ cerning morphotypes, ecotypes, and island populations are analysed currently. All the afore-said lichen substances are known from Roccella galapagoensis Follm. s. str. too (Follmann 1968, Follmann et al. 1993), another indication of the interrela­ tions between the two members of the assumed species-pair R. galapagoensis/ R. margaritifera. The only non-Galapagoan representative with a widely identical spectrum of secondary products seems to be R. lirellina (Darb.) Follm. from the Peruvian Pacific coast with erythrin and protocetraric acid as major and lecanoric acid, roccellic acid, and roccellin as minor metabolites. Beyond that, the sterile R. lirellina, though morphologically distinct, develops soralia similar to those of R. margaritifera. Therefore, it should be considered in the discussion on the origin of the Roccella population on the geologically young Galapagos Islands. Except the four Roccella species treated before, there is only one more known with certainty from the archipelago at present, R. babingtonii Mont. (Follmann 1994, Weber 1986). The first ones are all saxicolous and endemic, the second one is preferably corticolous and widespread in the Neotropics. Even if not collected on all islands and islets as yet, R. margaritifera B. Wern. & Follm. constitutes by far the commonest representative of its genus on all kinds of volcanic rocks, followed by R. galapagoensis Follm., R. obscurissima Follm. & B. Wern., and R. nigerrima (Darb.) Follm. Supplied with a relatively wide ecological ampli tude, the first acts together with R. galapagoensis as characteristic component of a pioneer association on cinder, lava, basalt, tuff, etc. which was provisionally named Roccelletum galapagoensis Follm. on occasion of an overview on the lichen vegetation of the "Enchanted Islands" (Follmann & Werner 1995). A detailed sociological description of this lichen community will be published elsewhere. The joint "chorogram" of the preceding Roccella species can be formulated as follows: Neotropical, Equatorial, Galapagoan, oceanic, arido-humid, planar to colline, neoendemic, scarce to abundant, community-forming. The corresponding "ecogram" based on our field-notes appears to be typical "roccellaceous": aero­ hygrophytic, xeroresistant, ombrophobous, subphotophytic, thermotolerant (steno­ climatic), saxicolous, neutrophytic, halophytic to subhalophytic, anitrophytic, toxico­ phobous, normally gregarious. 264 J. Hattori Bot. Lab. No. 85 I 9 9 8

ACKNOWLEDGMENTS The authors are much indebted to Dr. habil. S. Huneck (formerly Halle a. d. S., Germany), who was the first to elucidate specific secondary products of Chilean and Galapagoan Roccella species, and to the directors and curators of the herbaria at Boulder (COLO), Bristol (BRIST), Geneva (G), Helsinki (H), London (BM), Lund (LD), Paris (PC), Santa Cruz de Tenerife (TFMC), Stockholm (S), Uppsala (UPS), Vienna (W), and Washington (US) for the loan of Roccella collections in their care and the patience they had regarding the return of these. Moreover, the senior author gratefully acknowledges the receipt of financial support from the Association of Promoters of the University of Cologne and of travel grants provided by the University of Cologne.

SUMMARY Analyses of type material of the long-time critical lichen taxa Pentagenella fragillima Darb. (Atacama Desert) and Roccellodea nigerrima Darb. (Galapagos Islands) resulted in the new combinations Roccella fragil/ima (Darb.) Follm. and R. nigerrima (Darb.) Follm. In addition, two externally simil ar lichen species repeatedly mistaken for P. fragi//ima or R. nigerrima are described as new: lngaderia friabillima Follm. & M. Schulz (Atacama Desert) and Roccella obscurissima Follm. & B. Wern. (Galapagos Islands). R. margaritifera B. Wern. & Follm. (Galapagos islands) proved to be the sorediose (secondary) counterpart of the fertile (primary) R. galapagoensis Follm. Finally, Roccella DC. sect. Pentagenella (Darb.) Follm. stat. nov. is proposed. All taxa mentioned are characterized morphologically and chemically as well as chorologically, ecologically, and sociologically.

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