0007-2745/05/$0.75/0

The Bryologist 108(4), pp. 520 525Copyright q 2005 by the American Bryological and Lichenological Society, Inc.

Phloeopeccania anemoides, a New Lichen Species from Baja California Sur andSinaloa, Mexico

MATTHIAS SCHULTZ

Biozentrum Klein Flottbek und Botanischer Garten, Arbeitsbereich Systematik der Pflanzen, Universitat Hamburg,Ohnhorststr. 18, D-22609 Hamburg, Germany; e-mail: schultzm@botanik.uni-hamburg.de

BURKHARD BUDEL

Pflanzenokologie und Systematik, FB Biologie, TU Kaiserslautern, Postfach 33049, D-67653 Kaiserslautern, Ger-many; e-mail: buedel@rhrk.uni-kl.de

Abstract. The new cyanobacterial lichen species Phloeopeccania anemoides is described fromBaja California Sur and Sinaloa, Mexico. The species colonizes open rock surfaces on volcanicboulders in rocky slopes. A key to the known North American species of Phloeopeccania is pro-vided.

Keywords. Phloeopeccania anemoides, Lichinaceae, Mexico, Sonoran Desert.

During a joint field trip organized by T. Nash IIIwithin the Sonoran Desert Lichen Flora Project toBaja California in 1988/89 a supposedly unde-scribed cyanobacterial lichen species was discov-ered by the second author and C. Wetmore. Sub-sequently, the species was collected at several ad-ditional localities by the first author as well as byP. L. Nimis and M. Tretiach (Trieste). More materialwas discovered among unidentified collections ofSonoran Desert Lichinaceae made by T. Nash III.In spite of a similar thallus anatomy and an externalresemblance with members of the genus AnemaNyl. ex Forssell anatomical studies of the apothe-cial development suggested that the new lichenshould be referred to Phloeopeccania J. Steiner.Whereas in Phloeopeccania coiled ascogonia arisein groups of three to five, subsequently forming aspheroid ascomatal initial (Schultz & Budel 2002:52, fig. 7G), the ascogonia are formed beneath pyc-nidia (pycnoascocarp) in Anema. Similarly, the sep-aration of the new genus Digitothyrea P. Moreno &Egea from Thyrea A. Massal. by Moreno and Egea(1992) was based on differences in the type of as-comatal ontogeny. Phloeopeccania is a small genusin the Lichinaceae. The growth form varies fromcrustose to squamulose and subfruticose. The thal-lus anatomy is reticulate. The mycobiont forms aloose network of short, roundish to somewhat an-gulose hyphae. The cells of the photobiont are largeand have thick gelatinous sheaths. Three species arecurrently recognized (Henssen 1979, 1990).Phloeopeccania pulvinulina J. Steiner occurs insouthern Arabia and Socotra (Schultz 1998; Schultz& Mies 2003), Iran (Henssen & Jørgensen 1990),the Canary Islands and North America (Henssen &

Jørgensen 1990). Phloeopeccania hispanica Wil-lems & Henssen is known from southeastern Spain(Henssen 1990) and Socotra (Schultz & Mies2003). Phloeopeccania australiensis Henssen ishitherto only known from the type locality in Aus-tralia (Henssen 1990). A fourth species, Phloeo-peccania major Henssen ined. is rather common onlimestone in New Mexico and Texas. It was rec-ognized as new by Aino Henssen, but has not beenpublished yet. The first author is aware of anotherapparently undescribed species of the genus fromsouthern Arabia. For separation of the North Amer-ican species of Phloeopeccania a key is providedin this paper. Only one species of Anema, A. pro-digulum (Nyl.) Henssen is known from NorthAmerica so far (Schultz 2002). Anema dodgei Her-re does not belong to the genus, and its identitywill be clarified elsewhere.

MATERIAL AND METHODS

Cryotome sections were mounted using lactophenolcottonblue. Micrographs were taken using a Zeiss Axios-kop; macrographs were taken using Zeiss Luminar 3.5/40mm and Canon 2.8/35 mm lenses mounted on bellows,Nikon F3 and Canon T70 cameras and electron flash.

RESULTS AND DISCUSSION

PHLOEOPECCANIA ANEMOIDES M. Schultz & Budel,sp. nov.

Thallus nigricans squamulosus umbilicatus vel stipitatusparvus homoiomerus, squamulis rotundis crenatis velrosulatis 0.25–2 mm. Cyanobacteria unicellularia 12.5–20 mm gelatina crassa flavo-brunnea. Apothecia lam-inalia semi-immersa vel sessilia parva 0.25–0.5(20.6)mm, discus apertus depressus, margo thallinus promi-nentus rotundus vel sinuosus, excipulum proprium ab-

2005] 521SCHULTZ & BUDEL: PHLOEOPECCANIA

sunt, hymenium hyalinus in iodo coerulescens, asci cla-vati leptodermatici in iodo non tincti octospori, ascos-porae simplicae hyalinae ellipsoidea 10–12.5 3 5–7.5mm. Pycnidia immersa, conidia ellipsoidea ca. 3–5 31.5 mm.

TYPE: MEXICO. Baja California Sur, along Rte. 1, 68km NE of Ciudad Insurgentes in pass through Sierra laGiganta, hillside and rock ledges in thorn forest with Aca-cia, Lycium, Prosopis, 400 m, 258369 N, 1118209 W, 8January 1989, B. Budel [20062b] & C. Wetmore [63927,63959] (ASU–holotype; hb. B. Budel, hb. M. Schultz &MIN–isotypes); Baja California Sur, Sierra La Giganta,dirt road to Agua Verde, Rancho Viejo ca 7 km SE ofRte. 1, Sonoran Desert scrub, N-facing rocky slope, onSE-exposed rock faces, volcanic conglomerate, 300 m,258339430 N, 1118169030 W, 18 March 2003, M. Schultz16262a (ASU, hb. M. Schultz– paratypes).

Illustration: FIGURES 1 & 2.Thallus blackish, dull to slightly glossy, squa-

mulose-umbilicate. Squamules 0.25–2 mm wide,roundish, margin entire at first, becoming crenateto somewhat lobulate or effigurate and then resem-bling small rosettes (FIGS. 1A–C), mature squa-mules bulging, composed of laterally compressed,densely aggregated lobules (0.2–0.3 mm thick) withflattened tips, attached by small umbilicus, some-times shortly stipitate. Surface of the squamulessmooth at first, becoming irregularly tessellate withage. Thallus anatomy homoiomerous, but strands ofhyphae formed in basal and central parts of thesquamules (FIGS. 2A–B), these hyphal strands grad-ually merging into the umbilicus. Mycobiont cellsshort to elongate, angulate to roundish, ca 5–10 32.5–5 mm; photobiont cells large, 12.5–20 mm in-cluding yellowish brown, K-, ca. 2–3 mm thick ge-latinous sheath (FIG. 2B). Apothecia semi-immersedto sessile, circular or compressed with age, 0.25–0.5(20.6) mm wide with dark red, depressed discs;thalline margin thick, 75–125 mm wide (FIG. 2B),persisting, prominent, smooth or becoming irregu-larly crenate, circular to finally often compressed oreven sinuose; proper margin virtually absent andnegligible, hymenium hyaline, IKI1 blue, to 125mm high, subhymenium hyaline, inversely cone-shaped, sometimes elongate as a stipe, IKI1 blue,epihymenium faintly yellowish-brown, K-, paraph-yses distinctly septate, branched and anastomosing,apical cells to 3 mm wide. Asci subclavate, thin-walled, IKI-, 8-spored; ascospores simple, hyaline,broad ellipsoid, 10–12.5 35–7.5 mm, thin walled.Ascomatal ontogeny starting with free ascogones ingroups of 3–4, later becoming surrounded by in-terwoven hyphae forming a spheroid ascomatal ini-tial. Pycnidia immersed, globose, to 0.125 mmwide; wall simple or eventually convolute; conidiasmall, ellipsoid, ca. 3–5 3 1.5 mm. Chemistry: Nosecondary products detected.

Substrate and ecology. The species grows inSonoran Desert thorn scrub habitats. It prefers

steep, shaded volcanic boulders in rocky slopes atmid elevations, but it was also found on exposedpebbles on ground ca. 50 m above sea level.

Distribution. The new species is common inthe Sierra la Giganta in Baja California Sur, Mex-ico. The species seems to be absent from northernparts of Baja California and it has not been foundin the desert regions of southern California and Ar-izona. However, there is a record from rocky slopesN of Los Mochis in Sinaloa and the species mayalso occur in suitable habitats in Sonora. Interest-ingly, Phloeopeccania anemoides is replaced by P.major in hot Chihuahuan Desert habitats in south-ern New Mexico and western Texas. This may beexplained by the different substrate preferences.Phloeopeccania major, a yet undescribed but verycommon species in the Chihuahuan Desert region,is exclusively found on dry limestone boulders,whereas P. anemoides prefers acidic substrata likevolcanic rock. Since acidic rock types are widelydistributed in the Sonoran Desert Regions of BajaCalifornia and limestone is most common in north-ern parts of the Chihuahuan Desert, both speciesseem to form a pair of vicarious species.

Notes. Phloeopeccania pulvinulina is distin-guished from the new species by the formation ofsmall cushions composed of short, densely aggre-gated, cylindrical lobules (FIG. 3A–B; Schultz &Budel 2002: 46, fig. 4G) and asci usually contain-ing 16–24 spores. Furthermore, the squamules areoften surrounded by a hyaline gelatinous sheath(Schultz & Budel 2002: 48, fig. 5B). Phloeopec-cania major differs from P. anemoides in theroundish, convex squamules with down-curvedmargins and eventually indented tips. In Phloeo-peccania hispanica the squamules are not effigurateat the margin and not divided into compressed, ag-gregated lobules. Finally, Phloeopeccania hispani-ca usually has asci with 16–32 spores. Phloeopec-cania anemoides externally resembles rosette-shaped species of Anema, Phylliscum Nyl. ex A.Massal. and Paulia Fee. However, Anema andPhylliscum differ from Phloeopeccania in the typeof ascomatal ontogeny (pycnoascocarps) and thethallus hyphae are more elongated and angulose.Paulia and Phloeopeccania are similar in ascomataldevelopment. However, Paulia lacks an excipulumproprium and usually has thick walled ascospores.Paulia myriocarpa (Zahlbr.) Henssen was recentlyreported from Baja California Sur (Schultz & Tre-tiach 2002) and it may occur together with Phloeo-peccania anemoides in suitable thorn scrub habi-tats. Paulia myriocarpa forms regularly tessellate,roundish squamules with distinctly effigurate mar-gins. However, the apothecia are smaller than inPhloepeccania anmoides and they lack a prominentthalline margin.

522 [VOL. 108THE BRYOLOGIST

FIGURE 1. Phloeopeccania anemoides, external morphology. — A. (left) Three mature, bulging squamules withirregularly tessellate surface, (center) young, flat squamule with effigurate or rosulate margin (holotype). — B. (top)Young squamules, (bottom) mature squamules, apothecia with bulging thalline margin (isotype, hb. M. Schultz). — C.Young, flat squamules forming rosettes with effigurate margin, apothecia with prominent thalline margin and depresseddiscs (Schultz 16106b). (bars 5 1 mm).

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FIGURE 2. Phloeopeccania anemoides, thallus and apothecial anatomy. — A. Mature, sessile apothecium, thallinemargin prominent, proper exciple lacking (arrow), subhymenium inversely cone-shaped and elongate into thin centralstrand (holotype). — B. Reticulate hyphal arrangement with large photobiont cells (holotype). (bars A 5 100 mm, B5 50 mm).

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FIGURE 3. Phloeopeccania spp. — A & B. Phloeopeccania pulvinulina. — A. Squamules very small and appearinggranulose, apothecia very small, discs punctiform (arrow) (Schultz 16045k). — B. Unusually large, hypertrophic,bulging cushions (Wetmore 63890). — C. Phloeopeccania major: squamules small, of roughly equal width and heigth,tips becoming indented, hiding the deeply sunken apothecial discs (Schultz 16301a). (bars 5 1 mm).

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The following key treats the three species ofPhloeopeccania known from North America andnorthern Mexico.

KEY TO THE SPECIES

1. Squamules forming cushions 0.1–1 mm wide;cushions small, flat, irregularly granulose (FIG.3A) or roundish, slightly to sometimes distinctlybulging (FIG. 3B), composed of densely aggregat-ed, short to elongate, cylindrical lobules, 0.1–0.15mm wide, 0.2–0.5 mm long; apothecia 1 to sev-eral per cushion; asci usually 16–24-spored, rarely8-spored; widely distributed in the American SW,on acidic and calcareous rock ------------- P. pulvinulina

1. Thallus forming small squamules or rosettes, notcomposed of short cylindrical lobules; asci 8-spored ---------------------------------------------------------------------------- 2

2. Squamules small, 0.25–0.9 mm wide, convex,margins bent downwards, not effigurate, sur-face smooth but eventually becoming some-what rimose and indented at the tips (FIG.3C); apothecia usually single, disc deeplysunken, punctiform to irregularly compressed;restricted to limestone boulders in hot Chi-huahuan Desert habitats ---------------------------------

---------------------------------------------------------------------- P. major2. Squamules 0.25–2 mm wide, flat to bulging,

rosette-shaped with effigurate margins andeventually with tessellate surface (FIG. 1A–C); apothecia usually several per squamule,discs open, margin prominent, often sinuose;restricted to acidic rock in Baja California Surand Sinaloa -------------------------------------- P. anemoides

Additional specimens examined.—Phloeopeccania ane-moides: MEXICO. BAJA CALIFORNIA SUR: Sierra la Gi-ganta, Cerro Adamn at rd. 1, ca 50 km S of Loreto, Sono-ran Desert scrub, N-facing hill slope, shaded, volcanicconglomerate, ca 500 m, 258379 N, 111816.59 W, 2 Mar1999, Schultz 16106b (hb. Schultz); W of Sierra la Gi-ganta, ca 20 km NE of Ciudad Insurgentes, MagdalenaRegion of Sonoran Desert, exposed pebbles on ground, 50m, 25820932.50 N, 1118379210 W, 19 Mar 2003, Schultz16269a (hb. Schultz). SINALOA: 38 km N of Los Mochis,hill beside cemetery, Konglomeratfels, NO-exponiert, 50–60 m, 268079 N, 1098029 W, 12 Feb 1993, Budel 20138e(hb. Budel).

Phloeopeccania major: U.S.A. NEW MEXICO: Dona AnaCo., Bishop Cap Hills SE of Las Cruzes, Chihuahuan De-sert scrub, rocky NE slope, on 458 inclined limestone,boulders, 1,550 m, 328129 N, 1068369 W, 28 Mar 2003,Schultz 16301a (hb. Schultz).

Phloeopeccania pulvinulina: U.S.A. ARIZONA: Marico-pa Co., S end of South Maricopa Mountain Wilderness,

N of hwy 8, exit 140, N-facing cliff, shaded, sligthlysloped rhyolithic rock surface, ca. 520 m, 328539 N,1128229 W, 14 Feb 1999, Schultz 16045k (hb. Schultz).MEXICO. BAJA CALIFORNIA SUR: Along rte. 1, 31 km Wof San Ignacio at junction of road to Punta Abreojos, onNE slope of a cerro with Bursera microphylla, Lemaireo-cereus thurberi, Lophocereus schottii, Pachycereus prin-glei, elev. 180–200 m, 278209 N, 1138079 W, 7 Jan 1989,Wetmore 63890 (MIN).

ACKNOWLEDGMENTS

The authors would like to thank T. Nash III (Tempe)for field trip invitations and various generous help. Specialthanks to C. Wetmore (St. Paul) and R. Worthington (ElPaso) for additional study material. The authors are grate-ful to E. Brodo (Ottawa) and T. Nash III for improvingthe original text. This study was funded by DFG (SCHU1493/2). The first author received travel funds fromDAAD.

LITERATURE CITED

HENSSEN, A. 1979[1980]. Problematik der Gattungsbe-grenzung bei den Lichinaceen. Berichte der DeutschenBotanischen Gesellschaft 92: 483–506.

———. 1990. Lichenes cyanophili et fungi saxicolaeexsiccati, fasc. II, nos. 26–50. Marburg.

——— & P. M. JøRGENSEN. 1990. New combinations andsynonyms in the Lichinaceae. Lichenologist 22: 137–147.

MORENO, P. P. & J. M. EGEA. 1992. Digitothyrea, a newgenus in the family Lichinaceae. Lichenologist 24:215–228.

SCHULTZ, M. 1998. Studies on lichens from southern Yem-en Arabian Peninsula. Lichenologist 30: 293–297.

———. 2002. Anema, Pp. 97–98. In T.H. Nash III, B.D.Ryan, C. Gries & F. Bungartz (eds.), Lichen Flora ofthe Greater Sonoran Desert Region. Lichens Unlimit-ed, Arizona State University, Tempe, Arizona.

——— & B. BUDEL. 2002. Key to the genera of the Lich-inaceae. Lichenologist 34: 39–62.

——— & B. MIES. 2003. The saxicolous and terricolous,cyanobacterial lichens of Socotra (Indian Ocean).Nova Hedwigia 77: 73–97.

——— & M. TRETIACH 2002. Paulia, Pp. 329–331. InT.H. Nash III, B.D. Ryan, C. Gries & F. Bungartz(eds.), Lichen Flora of the Greater Sonoran Desert Re-gion. Lichens Unlimited, Arizona State University,Tempe, Arizona.

ms. received March 20, 2005; accepted June 20, 2005.