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Notes on the Dicnemonaceae (Musci): 11. the Status of Dicnemon Rugosum and Werneriobryum Geluense

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Notes on the Dicnemonaceae (Musci): 11. the Status of Dicnemon Rugosum and Werneriobryum Geluense Journ. Hallori BOI. Lab. No. 61. 273- 280 (1986) NOTES ON THE DICNEMONACEAE (MUSCI): 11. THE STATUS OF DICNEMON RUGOSUM AND WERNERIOBRYUM GELUENSE BRUCE H. ALLEN ABSTRACT. The dicranaceous pcristomes. unicellular spores, and lack of both leaf nematogens and leaf rhizoids in Dicnemon rugosum and Werneriobryurn geluense make the placement of these species in the Dicnemonaceae untenable. Conversely, the presence of leaves with a hyaline border, well differentiated alar cells, smooth, elongated and porose leaf cells, dense tomentum, and polyseteous condition all indicate that these species are properly placed in the genus Dicranoloma. D. rugosum and D. gelURnse belong to an atypical element of Dicranoloma. Species belonging to this element are also present in New Zealand, Australia, and ew Guinea. Erroneously attributed to Australia, Dicranoloma rugosum is endemic to Tahiti; D. geluense is endemic to the high mountains of eastern New Guinea. A Systematic Account of Dicranoloma rugosum Dicranoloma rugosum was originally described by Hooker (1818) as a species of Leucodon . At the time Leucodon represented a group of mosses ..... having the peristome of a Trichostomum or Didymodon (i.e. single), and lateral fruit (i .e. pleurocarpous) .. ,. (Hooker 1818). Hooker considered the species closely allied to Leucodon calycinus (=Dicnemon calycinum) and due to the mitriform calyptra of L. calycinus provisionally established the genus Lyellia for the two species. It was not until 1824, however, that Schwaegrichen transferred the two species from Leucodon. By then the name Lye/fia had been validly published by Brown (1819) for another genus of mosses and Schwaegrichen placed both species into the genus Dicnemon. Most subsequent authors have maintained this species in Dicnemon (Muller 1849; Jaeger, 1872; Paris 1896, 1904; Brotherus 1901 , 1924; Whittier 1976; Miller, Whittier& Whittier 1978). Muller (190 I), after examining the type of Eucamplodon, transferred a Tahitian species he earlier had described as Dicnemon banksii into Eucamptodon. M uller gave no other reason for his transfer, and shortly thereafter Salmon (1902) placed Dicnemon banksii into the synonomy of D. rugosum. Alien (1985) in a revision of the Dicnemonaceae considered the family highly derived and characterized by (among other features) the presence of a complex and strongly modified peristome; massive, endosporically germinating protonemata (so­ called " multicellular spores"); and numerous leaf nematogens from which leaf rhizoids arise. Hooker's L. rugosum, however, has a typical dicranaceous peristome, unicellular, exosporically germinated spores, and lacks both leaf nematogens and leaf rhizoids. These features make the placement of L. rugosum in the Dicnemonaceae untenable. 1 Missouri Botanical Garden, P. O. Box 299, St. Louis, MO 63166. 274 Journ. Hattori Bot. Lab. No. 61 1 9 8 6 As indicated by its dicranaceous peristome, erect habit, well difTerentiated, inflated alar cells, and smooth, elongated and porose leaf cells this species belongs in the Dicranaceae. Within the Dicranaceae, its strongly sheathing perichaetial leaves are reminescent of Holomitrium. However, the presence of leaves with a hyaline border of elongated cells, its robust size, dense tomentum, smooth elongate, porose leaf cells, polyseteous condition and the lack of strongly sheathing vegetative leaf bases all indicate the genus Dicranoloma. Furthermore, while many species of Dicranoloma lack sheathing perichaetialleaves, some species of the genus in New Zealand have more or less sheathing perichaetial leaves (D. plurisetum, D. robustum, and D. menziesii). Within Dicranoloma, D. rugosum belongs to an atypical element that macroscopi­ cally is very similar to the Hawaiian Dicranum speirophyllum . This element of Dicranoloma is also present in New Zealand (D. obesifolium), Australia (D. eucamplo­ dontoides), and New guinea (D . undulatifolium and D. geluense). There is some controversy surrounding the type locality of Dicranoloma rugosum . The species was originally described by Hooker from a "New Holland" (i.e. Australia) collection sent to Turner by Dickson. Mitten (1875), however, questioned the correctness of Dickson's reported localities in general: "Dickson obtained specimens of mosses from the voyages of trading vessels, and localities were then not so precisely mentioned, so that any gathered during the voyage were liable to be reported as if gathered in the country the vessel had returned from; and there are some of the species which Dickson in this manner procured, of which the native country has yet to be discovered." Salmon (1902) noting that this "conspicuous" moss had never been recollected in Australia and citing Mitten's comments regarded Dickson's New Holland locality for this species with suspicion. Finally, on a note accompanying an isotype of Dicranoloma rugosum in BM there is a discussion of Dickson 's localities that ends: "J . D . got the specimen from an East India man, & did not know she (sic) had called at Madeira. Nor did he know that the Leucodon rugosus specimen came from Tahiti or Fiji & not from New Holland." Aside from the type collection, this species is still known only from Tahiti. Despite its restricted distribution, Dicranoloma rugosum is far from a stenotypic species. Collections are extremely variable in plant size, leaf length, the degree in which the leaves are rugose, the strength of the hyaline border and leaf serrations, capsule shape and length, and the number of setae per perichaetium. Plants that are extremely robust, have long narrow rugose leaves that are distinctly denticulate have been called Dicnemon giganteum. M tiller (1858) described Dicnemon banksii for a Tahitian moss that differed from D. rugosum in a number of vegetative characters, most prominent of which were its denticulate leaf and non-rugose leaves. Bescherelle (1895) was thefirst to question the distinctness of these two mosses, noting that in all Tahitian collections he had examined the leaves were more or less rugose (as in D. rugosum) and had denticulate margins (as in D. banksii). Salmon (1902) examined the types of both species, and found them to be identical. More specifically, Salmon noted that the type of Dicnemon banksii was" . decidedly rugose (if first moistened and then allowed to dry naturally B. H. ALLEN: Notes on the Dicnemonaceae (Musci): Il 275 without being subjected to pressure) .. " and that the leaf margin of D. rugosum was, contrary to Hooker and Schwaegrichen's description, not entire but rather, H ••• ir­ regularly denticulate, in exactly the same manner as in M liller's D. banksii." All au­ thors since Salmon have placed Dicnemon banksii under the synonymy of Dicnemon rugosum. Unfortunately, the type of Dicnemon banksii, which was deposited at B, appears to have been destroyed. Taxonomic Treatment Dicranoloma rugosum (Hook.) Alien, comb. novo Leucodon rugosum Hook., Musei Exotici 1, t. 20. 18 18 . (basionym). Dicnemon rugosum (Hook.) Schwaegr., Spec. Muse. Suppl. 2(1): 127. 1824. Eucnemis rugosa(Hook.) Brid., Bryol. Univ. 2: 217. 1827. comb. inval. Type. Australia (sic). In Nova Hollandia. Turn. Herb. a Dicksonia missus (holotype, BM; isotypes, 'BM, G, MANCH, NY). Dicranum densifolium Mohr, Ann. Bot. 2: 246. 1806. nom. nudo (vide e. Muller 1858). Dicnemon banksii e. Mull., Bot. Zeit. 16: 16 1. 1858. Dicnemos banksii e. Mull ex Beseh., Ann. Sci. at. Bot. ser. 7, 20: 18. 1894. or/hogr. var. Eucamptodon banksii (e. Mull.) e. Mull., Gen Muse. Fr.: 248 . 1901. Type. Tahiti. Ex insula Tahiti retulit Banks (type, B; apparently destroyed). Dicnemon giganteum Schimp. in Salm., J. Bot. 40: 7. 1902. nom. inval. in synom . Dicnemon procerrimum Schimp. in Salm., J. Bot. 40: 7. 1902. nom. inval. in synom. Plants medium to large, 4 - 9cm high, more or less shiny, yellow-brown to yellow-green, caespitose. Stems red, densely tomentose; in cross-section 2 - 3 rows of small, thick-walled, red­ brown epidermal cells, a cortex of larger, thinner-walled, reddish-yellow cells and a central strand of small thin-walled hyaline cells. Branches infrequent, identical to stems. Leaves 4 - 6 mm x 1.5 mm, very dense, rigid, imbricate-appressed to erect-spreading, more or less transversely rugose, ovate-lanceolate to lanceolate, concave or plane; margins entire and plane below, more or less denticulate and incurved above, variably bordered by 2 - 7 rows of long, narrow, hyaline cells; apex acute or shortly acuminate; costa single, E-type (see Kawai 1968), percurrent, strong throughout (80 - 100 J.lm wide at base). Cells narrowly elongate, longer and narrower toward the margins, shorter and wider toward the costa; upper cells long-rhomboidal to rhomic, distinctly porose; alar cells orange-yellow to yellow-brown, conspicuously enlarged. Dioicous. Male plants dwarfed on female plants. Sexually mature males 1.0 - 1.3 mm long; leaves lanceolate, ecostate, 0.4 - 1.0 mm long, cells elongate-rhomboidal, more or less thick-walled, not porose, alar cells weakly differentiated. Perichaetia terminal, becoming laterally displaced; outer leaves 3 - 4 mm long, sheathing, abruptly long setaceous-acuminate, costa present; inner leaves 1 PLATE I. Distribution of Dicranoloma rugosum in Tahiti. 276 Journ. Hattori Bot. Lab. No. 61 1 9 8 6 8 - 12 mm long, closely sheathing the seta, long setaceous above. Archegonia numerous, 120 mm long, paraphyses absent. Capsules exserted or shortly emergent. Setae 8 - 15 mm long, red or yellow, I - 3 per perichaetium; in cross-section 3 - 4 layers of small, thick-walled
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