The Bryologist 108(1), pp. 47 49 Copyright ᭧ 2005 by the American Bryological and Lichenological Society, Inc.

Anoectangium handelii (Bryopsida, Pottiaceae) in the New World

RICHARD H. ZANDER Missouri Botanical Garden, P.O. Box 299, St. Louis, MO 63166-0299, U.S.A. e-mail: [email protected]

WILLIAM A. WEBER University of Colorado Museum, Campus Box 265, Boulder, CO 80309, U.S.A. e-mail: [email protected]

Abstract. New to the Americas is Anoectangium handelii Schiffn., which differs signi®cantly from A. aestivum (Hedwig) Mitt. by its small size, axillary propagula, and cauline leaves usually blunt and shorter than the perichaetial leaves. Previously, this species was known only from south- central Eurasia.

Keywords. Anoectangium, Colorado, disjunct, moss, Pottiaceae, south-central Eurasia.

The discovery of a diminutive, propaguliferous ANOECTANGIUM HANDELII Schiffn., Annalen des Naturhis- Anoectangium in Colorado by W. A. Weber and R. torischen Hofmuseums Wien 27: 490. 51±59. 1913. Wittmann prompted a re-examination of study Plants growing in low, dense turf, vivid green slides associated with a synopsis of the Pleuro- above, tan below, commonly encrusted below with weisieae of Middle America (Zander 1977). Three lime. Stems occasionally branching, to 0.4 cm, additional Colorado collections of what are clearly transverse section rounded-pentagonal, scleroder- A. handelii Schiffn. were uncovered, reported ear- mis thin and sharply distinguished from in¯ated lier (Zander 1977) as Molendoa sendtneriana cells of inner cylinder, central strand present; axil- (Bruch, Schimp. & W. GuÈmbel) Limpr. Anoectan- lary hairs of 2±3 cells, basal cells 1±2, ®rm-walled; gium handelii was originally described from Turk- radicles sparse. Leaves crowded, weakly secund, ish Kurdistan, and has been reported from Iraq and weakly spreading when dry, spreading-recurved Turkey (Agnew & VondraÂcÏek 1975), and the Cri- when moist, ligulate to short-elliptic or ovate, 0.4± mean Peninsula, southern Siberia, and the south- 0.5(±0.6) mm long, distal lamina strongly keeled, central Eurasian states north of Iran and Afghani- margins plane, apex usually blunt, occasionally stan in the former Soviet Union (Ignatov & Afonina with 1-celled apiculus; leaf base little differentiated in shape; costa usually ending (1±)3±4 cells before 1992); it was mistakenly synonymized with M. apex, adaxial and abaxial super®cial cells elongate, sendtneriana by Zander (1977). transverse section semicircular, with 2 guide cells Anoectangium handelii is a species with no sa- exposed adaxially, one stereid band, and abaxial lient features aiding identi®cation in the ®eld. It epidermis of thick-walled cells, hydroid strand ab- looks much like Gymnostomum aeruginosum Sm., sent; distal laminal cells subquadrate, often elon- which, however, is found in Colorado in well-wa- gate longitudinally, 5±9 ␮m in width, 1(±2):1, tered places, not xeric deep crevices. Tell-tale mor- sometimes bistratose in patches along margins, pa- phology under the dissecting microscope include pillae low and simple, 3±4 per lumen or apparently the leaves secund when dry, recurved when wet, absent; basal cells weakly differentiated, rising and the lateral perichaetia on short branches. Im- higher along costa, little wider than distal cells, 2± portant details visible with the compound micro- 3:1, usually thick-walled. Propagula uncommon, scope and sectioning are the blunt or very short- ovate, occasionally apiculate by projection, of ca 8 apiculate leaf apex, leaves keeled and appearing cells, born on rhizoids in leaf axils. Apparently di- deeply grooved along the costa, costa usually end- oicous; perichaetia lateral, inner leaves lanceolate, ing before the apex, costal section showing only 0.5±0.7 mm long, laminal cells elongate; perigonia one stereid band and the guide cells apparently ex- not seen. Sporophytes not seen. Laminal color re- posed adaxially, and ovoid or ovate-apiculate pro- action to 2% KOH yellow-orange. pagula borne on rhizoids from the leaf axils. Peri- gonia were not seen, and the species may reproduce Specimens examined.ÐU.S.A. COLORADO. Boulder Co., Hall Ranch Open Space, 1,700 m, 15 May 2004, Weber, vegetatively by plant fragmentation because of Wittmann, & Lehr B-114035 (COLO, MO); W slope Steam- fragile stems. boat Mountain (cuesta), W of Lyons, 1,900 m, May 1971,

0007-2745/05/$0.45/0 48 THE BRYOLOGIST [VOL. 108

FIGURES 1±9. Anoectangium handelii, from American material. Ð 1. Habit, moist. Ð 2±3. Fertile stems. Ð 4. Stem section. Ð 5. Three leaves. Ð 6. Distal leaf and leaf apex. Ð 7±8. Mid-leaf sections. Ð 9. Two gemmae. Scale bars: A ϭ 1 mm (1), B ϭ 1 mm (2±3), C ϭ 0.2 mm (5), D ϭ 50 ␮m (4, 6±9).

Hermann 23589, 23593 (COLO). Larimer Co., Buckhorn short-elliptic to ovate, 0.4±0.5(±0.6) mm long, Creek, Stove Prairie road, NNW of Masonville, April apex usually blunt, often with 1-celled apiculus, 1976, Hermann 27016 (COLO). TURKEY. Western Kur- margins weakly crenulate or irregular near apex, distan, Cataonia, Handel-Mazzetti 2024 (FH, holotype). apex usually blunt, sometimes apiculate, costa ending (1±)3±4 cells below apex; propagula un- KEY TO NORTH AMERICAN common, axillary, ovate, of several multiseriate ANOECTANGIUM SPECIES cells, often tapering to a hyaline, acute to attenu- 1. Stems to 0.4 cm long; leaves short-ligulate or ate apex ------Anoectangium handelii 2005] ZANDER & WEBER: ANOECTANGIUM HANDELII 49

1. Stems usually 1±3 cm long; leaves long-ligulate typic Proterendothrix scolecoidea W. & G. S. West, to broadly lanceolate, 1±2 mm long, margins en- as treated by Geitler (1925: 389), and said to occur tire, apex sharply acute (in America north of Mex- ico), usually with 1±3-celled apiculus or mucro, on moist earth in Africa. costa usually percurrent; propagula absent ------At the station, the layers of rock were almost ------Anoectangium aestivum horizontal and separated by very deep ®ssures. The Molendoa sendtneriana is found in Colorado as moss was growing in the deepest parts of the re- poorly developed forms that morphologically phe- cesses, dif®cult and dangerous (because of rattle- nocopy A. handelii. The former grows in a more snakes) for retrieval. The critical feature of the mi- open habitat, such as vertical cliff faces, and may crohabitat appears to be the direction of exposure further be distinguished by the stem central strand, of the rocks, such that the opening of the crevices broad and somewhat sheathing leaf base, total lack faced southwest, providing protection and a guar- of a leaf apiculus, and two stereid bands or at least antee of some small amount of moisture throughout a layer of epidermal cells adaxial to the guide cells. the year, and permitting the entry of sunlight during In the New World, A. handelii is known only for the winter when the sun lies low on the horizon. a few stations in Colorado. Excepting the 2004 col- This feature is also present in southwestern Colo- lection, the specimens are extremely scant. In May rado, in the Four Corners desert, where growth of 2004, Colorado suffered a severe drought. A lucky Weissia ligulifolia is possible in spite of the almost few days of heavy rains prompted a visit to a re- always dry soil. Phytogeographically, this record cently designated Open Space area on the west side reinforces the thesis of Weber (2004) that the local of a cuesta in the outermost foothills of the Front southern Colorado ¯ora is a relictual one, with nu- Range west of Lyons. Here the slope was less steep merous taxa in vascular and cryptogamic plant than on Steamboat Mountain where small scraps of groups occurring disjunct from Middle Asia. the species (then thought to be Molendoa) had been collected years ago in cracks of the rim-rock. The ACKNOWLEDGMENTS slopes are covered conspicuously by Cercocarpus We appreciate the aid of Ronald C. Wittmann, who has montanus and Opuntia phaeacantha, with scattered helped the junior author in his twilight years accomplish joint exploration of the bryophyte ¯ora of a daunting, rug- trees of Sabina virginiana. A variety of calcareous ged landscape. We also thank Patricia Eckel for her usual and non-calcareous sandstones form ledges of red superb illustration. and white layers. Recent moisture made visible a variety of mosses, including Crossidium squamifer- LITERATURE CITED um, Encalypta vulgaris, Grimmia anodon, Ptery- AGNEW,S.&M.VONDRAÂ CÏ EK. 1975. A moss ¯ora of Iraq. goneurum subsessile, Syntrichia caninervis, Tortula Feddes Repertorium 86: 341±489. obtusifolia, and Weissia ligulifolia. Here also, oc- GEITLER, L. 1925 [1926]. Cyanophyceae, pp. 1±450. In A. Pascher, Die SuÈûwasserÐFlora, Deutschlands, OÈ ster- curring adjacent to tufts of Didymodon rigidulus, reichs und der Schweiz. Heft 12, Gustav Verlag., Jena. were beautiful patches of the continental rarity D. IGNATOV, M. S. & O. M. AFONINA (eds.). 1992. Checklist anserinocapitatus, which has been found only once of mosses of the former USSR. Arctoa, a Journal of before in Colorado. The area immediately adjacent Bryology. 1: 1±85. was occupied by a minute terricolous blue-gray cy- WEBER, W. A. 2004. The Middle Asian Element in the Southern Rocky Mountain Flora of the western United anobacterium in the Oscillatoriaceae that forms rig- States: a critical biogeographical review. Journal of id, short spikes suggestive of a sterile lichen. Each Biogeography 30: 649±685. solitary ®lament is enveloped by a thick, rough ZANDER, R. H. 1977. The tribe Pleuroweisieae (Pottiaceae, sheath, out of which the ®lament protrudes termi- Musci) in Middle America. THE BRYOLOGIST 80: 233± 269. nally. Subject to veri®cation by J. Johansen, John Carroll University, who is culturing the cyanobac- terium, this curious plant appears to be the mono- ms. received Aug. 12, 2004; accepted Sept. 14, 2004.