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Climacium Dendroides from Cofre De Perote, a High-Elevation Tropical Montane Site in Veracruz, Mexico

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Climacium Dendroides from Cofre De Perote, a High-Elevation Tropical Montane Site in Veracruz, Mexico The Bryologist 107(3), pp. 368 372 Copyright q 2004 by the American Bryological and Lichenological Society, Inc. Climacium dendroides from Cofre de Perote, A High-Elevation Tropical Montane Site in Veracruz, Mexico ANGELA E. NEWTON Department of Botany, Natural History Museum, London SW7 5BD, United Kingdom; e-mail: a.newton@nhm. ac.uk EFRAIÂN DE LUNA Instituto de EcologõÂa, A.C., Ap. Postal 63, 91000 Xalapa, Veracruz, MeÂxico; e-mail: [email protected] Abstract. Climacium dendroides (Hedw.) F. Weber & D. Mohr is reported new to Mexico and in the ®rst known locality in the tropics, at the elevation of 3,000 m, on the ancient shield volcano of Cofre de Perote. Variation in branching of the costa is assessed in this plant and in material of C. dendroides worldwide. Additional taxa of interest from this locality on Cofre de Perote include Aulacomnium palustre and Polytrichum commune (both new to the state of Veracruz). Keywords. `Boreal' mosses, Climacium, distribution, Mexico, montane tropics. The ancient shield volcano of Cofre de Perote is 20 m across, and consisted of a series of small located in the Neovolcanic Axis of Mexico, at swamps connected by the small stream. Climacium 198329 N, 978309 W, and reaches an elevation of was growing on low hummocks to one side of the 4,200 m. The upper regions of the mountain have stream, and the plants were deeply embedded in the extensive Pinus-Abies forests with a rich ¯ora of surrounding mosses and other plants. Additional mosses, ferns, and cycads, that has been the focus moss species in the immediate area included Po- of much botanical study in the past, including the lytrichum commune Hedw. (De Luna 2348, Newton visit by Baron Alexander von Humboldt in 1803. 4427 ) and Aulacomnium palustre (Hedw.) The moss ¯ora has been studied by De Luna (1988) Schwaegr. (De Luna 2349, Newton 4428 ), both and Delgadillo (1984). At least 173 species of new records for the state of Veracruz and very rare- mosses are listed from the coniferous forest and ly recorded in Mexico (Sharp et al. 1994). Exten- alpine zones. Dominant species on soil include sive areas of Aulacomnium were found in an adja- Hypnum revolutum, Rhytidium rugosum, and Thui- cent, much larger swampy area surrounded by large dium peruvianum. The most frequent epiphytic stands of Juncus and various sedges, but a search mosses are Entodon jamesonii, Fabronia ciliaris, for Climacium in this area was not successful. Fol- Neckera chlorocaulis, Orthotrichum pycnophyllum, lowing severe drought in the summer of 1998 large Platygyrium fuscoluteum, Pylaisiella falcata, Tor- areas of forest on Cofre de Perote were damaged tula amphidiacea, T. fragilis, and Zygodon ehren- by ®re. Fortunately the area in which the Clima- bergii. Open sites are colonized by Symblepharis cium occurs was not affected by the ®re and the vaginata, Zygodon obtusifolius, Didymodon rigi- continued presence of the plants in this locality was dulus, and several species of Bryoerythrophyllum, con®rmed (De Luna) in November 2003. Leptodontium, and Ptychomitrium. It is remarkable The Climaciaceae consist of one genus, Clima- that several new taxa or new records for Mexico cium, with four or ®ve species, and was discussed have been discovered on this volcano: Philonotis by Horton and Vitt in 1976. Climacium dendroides corticata (Crum & Grif®n 1981), Bryoerythrophyl- (Hedw.) Web. & Mohr is known from northern and lum andersonianum (Zander & Sharp 1981), Lep- central Europe and Asia, the South Island of New todontiella apiculata, Pleurozium schreberi (De Zealand, and North America, where it is found Luna 1988), and Leptopterigynandrum austro-al- across the north of the continent above ca 458 N, pinum (De Luna 1986). and in the western mountains as far south as New Plants of Climacium (Newton 4429, De Luna Mexico. Climacium americanum Brid. occurs in 2347, 2358 ) were collected in May and June 1998 eastern North America from the Great Lakes and on the northeastern side of Cofre de Perote at an New York south to Florida and the Gulf Coast. Cli- elevation of 3,000 m (10,200 ft.) in a swampy hab- macium japonicum Lindb. is known from Japan. itat in an open shallow stream valley surrounded by Climacium acuminatum Warnst. was described Pinus spp. The area was fairly small, approximately from North America but was not mentioned by 0007-2745/04/$0.65/0 2004] NEWTON & DE LUNA: CLIMACIUM DENDROIDES 369 Horton and Vitt (1976) and is little known. A ®fth basal spurs or forking of the costa (Fig. 2). Al- taxon is variously recognized as a species, Clima- though only present in ca 1/5 of the stipe leaves, cium kindbergii (Ren. & Card.) Grout, as a variety this feature occurs in leaves along the length of the of C. americanum, or as a form not worthy of tax- stipe, so is not due to heteroblastic changes along onomic rank (Horton & Vitt 1976). For the pur- the stipe. Material from North America and Europe poses of this paper, this latter taxon is regarded as in the herbarium of the Natural History Museum part of Climacium americanum. No specimens of (BM) was examined to determine whether this fea- the genus have been reported previously from lo- ture occurs elsewhere. Considerable variation in the calities in tropical latitudes, with all previous rec- length and stoutness of the costa was seen, often ords north of ca 278 N (North America, Europe, between stems in a single collection. In several col- Asia, and Japan) or south of 408 S (New Zealand). lections short auxillary costae were seen, originat- In North America, the distributions of C. den- ing adjacent to but separate from the principle cos- droides and C. americanum overlap in the north- ta. In a small number of specimens very occasional eastern part of the continent, and the two species forks or spurs were found to be present. It therefore intergrade to some extent in the characters used to seems that some variation in the form of the costa differentiate them, including shape of stipe and is widespread in Climacium dendroides, but has not branch leaves, size of auricles, and length:width ra- been reported in the standard ¯oras. tio of the upper lamina cells (Horton & Vitt 1976). The possible relationship of the Climaciaceae to Although C. americanum extends farther south than the Hylocomiaceae was suggested by Crum and C. dendroides, it is primarily a species of lower Anderson (1981), but the difference in costa mor- elevations, whereas C. dendroides is primarily a phology (long and single versus short and double, boreal-montane species. respectively) was raised as a possible objection by The plants found on Cofre de Perote conform these authors. The presence of forked or spurred closely to the description of C. dendroides as given costae in some specimens of Climacium dendroides by Horton and Vitt (1976), and match specimens might be construed as morphological support for from North America and Europe. Since the nearest this taxonomic placement. known locality for C. dendroides is in New Mexi- Given the high elevation and extensive conifer- co, at 358509 N, 1058409 W(Crosby 16529 MO ous forests on this and other mountains in the neo- http://mobot.mobot.org/W3T/Search/most.html) volcanic axis, it may be hardly surprising that this our report represents a signi®cant range extention. moss assemblage, more typical of boreal habitats, The plants are glossy golden-green, with ®la- should be found on Cofre de Perote. A signi®cant mentous paraphyllia forming a dense stem tomen- number of such ``boreal'' species have previously tum and erect, rather strongly plicate and concave been recorded from this area by Delgadillo (1984): branch leaves. The stipe leaves are up to 2.6 mm Didymodon rigidulus Hedw. var gracilis (Hook. & long by 1.8 mm wide, with obtuse apices abruptly Grev.) Zand.; D. rigidulus var icmadophila (C.M.) narrowed to a slightly curved or re¯exed apiculus Zand.; Grimmia af®nis Hornsch.; G. fuliginosa C. to 0.18 mm long (Fig. 1), and with a single or var- M.; G. fusco-lutea Hook.; G. incurva Schwaegr.; iously spurred or forked costa to the middle or up- Paraleucobryum enerve (Thed.) Loeske; Rhytidium per lamina. The branch leaves are up to 2.3 mm rugosum (Hedw.) Kind.; Tayloria splachnoides long by 1.25 mm wide, deeply plicate when wet Hook.; Zygodon viridissimus (Dicks.) Brid. and by and fairly abruptly narrowed to a short acute and De Luna (1986 and 1988): Timmia megapolitana distinctly serrate apex, with a long, stout, single Hedw. subsp bavarica (Hessl.) Brassard and Pleu- costa reaching into the upper lamina. Auricles in rozium schreberi (Brid.) Mitt. Further exploration both types of leaves are present but weakly devel- of the area and examination of existing collections oped, and with distinct patches of in¯ated thin- can be expected to reveal additional range exten- walled alar cells. Upper lamina cells range from sions. short to long rhomboid, up to 50 mm long, with an The presence of these taxa may represent chance average length:width ratio of 8:1 and a range of 4: colonization by long-distance dispersal of spores 1 to 10:1. According to Horton and Vitt (1976) up- into a suitable habitat, although given the rarity (at per leaf cells of C. dendroides vary from (6)7:1 to least currently) of sporophyte production in species 13:1, whereas in C. americanum the range is from such as Rhytidium rugosum this may be unlikely. 3±7(8):1. Although the average length:width ratio Another possibility is that these distributions may in the Mexican plant falls in the overlap between re¯ect a relictual status of individual species or (as these two species, it is evident that the range more in this case) small communities that have main- closely matches that of C.
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