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DIPLONEURON (Musei: HOOKERIACEAE) Joum. Hal/ori Bot. Lab. No. 61: 45- 64 (Dec. 1986) A REVISION OF THE GENERA PILOTRlCHIDIUM AND DIPLONEURON (MUseI: HOOKERIACEAE) BRUCE H. ALLEN1 AND MARSHALL R. CROSByl ABSTRACT. Pilotrichidium is a genus of two species (P. callicostatum and P. antillarum) with strictly neotropical. primarily geographically distinct, distributions: P. callicostatum north to Mexico, southwest to Ecuador, southeast to Venezuela, and in the Caribbean on the islands of Trinidad and Jamaica; P. antillarum throughout the islands of the Caribbean (except Trindiad and Jamaica) and in Honduras. The affinities of Pilotrichidium lie with the Hookeriopsoid group of the Hookeriaceae, specifically with Diploneuron, S('hi~omitrium, Thamniopsis, and Hookeriopsis. Pilotrichidium brunnescens and P. ant ilia rum var. com­ planatum are syn. novoof P. antillarum, and P. dialOmophilum is transferred to the genus Schizomitrium as S. diatomophilum (e. MiiIL) Alien & Crosby comb. novo Diploneuron, a monotyp ic genus (D. connivens) known only from Jamaica and Cuba is distinguished by its unique double costae. The costae become marginal approximately one-half the distance from the leaf base and coalesce at the apex into a thickened subulate point. The genus is best positioned in the hookeriopsoid group of the Hookeriaceae, and is closely allied to Pilotrichidium. The genus Pilotrichidium was established by Bescherelle (1876) to emphasize the distinctness of a Trinidadian species of Muller's (1851) Hookeria section Callicostata: H. callieostata. Bescherelle maintained Pilotrichidium in the Hookeriaceae, but distanced his new genus from Hookeria on the basis of its leaf cells, which he considered to strongly resemble those found in Pilotrichum (also placed by Bescherelle in the Hookeriaceae). In the same paper he described two additional Carribbean spe­ cies in Pilotrichidium (P. antillarum and P. brunnescens). Bescherelle's treatment of Pilotrichidium was later adopted by Jaeger and Sauerbeck (1879). In 1907, Brotherus re-emphasized the similarity between Pilotrichidium and Pilotriehum when he removed both genera from the Hookeriaceae and placed them in the Pilotrichaceae. Pilotriehidium was subsequently maintained in this position (Fleischer 1923, Brotherus 1925, Crum & Steere 1957, Crum & Bartram 1958) until Crosby (1969) transferred the genus back to the Hookeriaceae. Most recent treatments of Pilotrichidium have agreed (Crosby 1974, Vitt 1984, Walther 1983) in placing the genus in the Hookeriaceae; however, Miller (1971) placed Pilotrichidium in a recon­ stituted Pilotrichaceae. Crosby's earlier paper (1969) was concerned primarily with the disposition of the Pilotrichaceae, which he found to be (in Brotherus's sense) an unsupportable segregate of the Hookeriaceae. Nevertheless, he did give several reasons for considering Pilotrichidium and Pilotriehum as being not closely related. In 1974, Crosby amplified this view when he proposed retaining Pilotriehidium in the Hookeriaceae while I Missouri Botanical Garden, P. O. Box 299. St. Louis, MO. 63166. 46 Journ. Haltori Bot. Lab. No. 61 1 9 8 6 transferring Pilotrichum into the Daltoniaceae. Crosby's rationale for this decision was based on both gametophytic and sporophytic evidence given in his 1969 paper. Gametophytically, the genera differ in a number of features. In Pilotrichidium the plants are never dendroid, the leaves are stongly flattened, pseudoparaphyllia are absent, the calyptrae are naked, the stems and branches have a outer layer of enlarged, hyaline epidermal cells, the costae project dorsally at the apex as spines, and the leaf cells, when papillose are pluripapillose. In contrast species of Pilotrichum are commonly dendroid, the leaves arch out from all sides of the stems, pseudoparaphyllia are present, the calyptrae are pilose (with both uniseriate and multiseriate filaments), stems and branches have an epidermis of 4-5 rows of small, thick-walled, reddish­ brown cells, costae are dorsally crested, and leaf cells, when papillose, have moderate to high, single apical papillae. It is, however, in their sporophytic differences that the strongest evidence for a lack of close relationship between these two genera lie . In Pilotrichidium a broad annulus is present, the exothecial cells are collenchymatous, and the exostome is of the hookeriaceous type. The capsule in Pi/otrichum, on the other hand, lacks an annulus, the exothecial cells are evenly thickened, and the exostome is of the daltoniaceous type. The differences between the hookeriaceous and daltoniaceous exostomes are of a fundamental importance that has generally failed to be recognized. In the hookeriaceous exostome the teeth are broadly triangular at base, the lower two­ thirds of the dorsal lamellae have closely spaced horizontal striae; and secondary wall deposition at the base of the dorsal (outer) surface is considerably greater than the deposition on the ventral (inner) surface. The significance of the last fea­ ture of the hookeriaceous exostome lies in a functional, and therefore evolutionarily important, consequence: the exostome are inflexed when wet and outflexed when dry. In the daltoniaceous exostome, on the other hand, the teeth are linear in shape, the dorsal lamellae are either densely papillose or smooth, and secondary deposition at the base of the ventral (inner) surface is more extensive than the secondary deposition on the dorsal (outer) surface. In the daltoniaceous peristome the action of the exostome teeth is opposite that of the hookeriaceous exostome: inflexed when dry, outflexed when wet. The affinity of Pilotrichidium to the Hookeriaceae is indicated by several sporo­ phytic and gametophytic. features diagnostic of family: the presence of a hookeria­ ceous peristome, collenchymatous exothecial cells, mitrate calyptrae, and strong double costae, and the absence of both pseudoparaphyllia and differentiated alar cells. Within the Hookeriaceae the affinities of the genus lie with what has been termed the Hookeriopsoid group (Crosby 1974), specifically Diploneuron, Schizomitrium (= Callicostella), Hookeriopsis and Thamniopsis. Diploneuron, a monotypic genus known from Cuba and Jamaica, has been closely aligned with Pilotrichidium ever since it was first described (Bartram 1936). The two are indeed extremely close, differing only in two gametophytic features: the outer epidermal layer of cells, while consisting of large, inflated cells, is not distinctly B. H. ALLEN & M. R. CROSBV: A revision of Pi/o/richidium & Dip/oneuron (Hookeriaceae) 47 different from the inner epidermal layers of cells, and the costae are unique "in that the costae become marginal in the upper portion of the leaf and finally coalesce near the apex to form a long-excurrent spine" (Crosby 1969). The further evaluation of the affinity of Pilotrichidium to the Hookeriopsoid group of the Hookeriaceae is hampered because the two largest and most critical genera in the group (Schizomitrium and Hookeriopsis) are in desperate need of taxonomic revision at the generic level. Nevertheless, it is clear to us that· Pilotrichidium is close to Schizomitrium and that Pilotrichidium may represent only a weak segregate of the latter. The two can be distinguished by four gametophytic features. In Pilotrichidium the leaf margins are absolutely entire, the leaf cells are extremely incrassate and irregular in shape, the stern has an outer epidermal layer of large, hyaline cells, and the costae, which are exceedingly strong, project as dorsal spines. By contrast, in Schizomitrium the leaf margins are serrate to dentate above, the cell walls thin to moderately thick-walled and regular in shape, the stem epidermis consists of small , thick-walled, colored cells, and the weaker costae do not project as dorsal spines. The relationship of Pilotrichidium to Hookeriopsis is difficult to evaluate. The genus Hookeriopsis is both large and extremely variable. There are, for example, some species with incrassate and irregularly shaped leaf cells; some with a stem epidermis of large, hyaline cells; some with strong costae that project as dorsal spines; and some with entire leaf margins. No species of Hookeriopsis, however, combines all of these features. Until Hookeriopsis is revised it is impossible to determine exactly how close Pilotrichidium is to that genus. Likewise, Thamniopsis exhibits many features found in Pilotrichidium: stern epidermis of large, hyaline cells, strong costae, entire-margined leaves and thick-walled cells. The two differ in the regularly shaped leaf cells, non-projecting costae, and dendroid habit of Thamniopsis. The closeness of Pilotrichidium to Thamniopsis can perhaps be seen in that the latter genus is in turn similar to some members of Hookeriopsis. So much so that Robinson (1967) considered Hookeriopsis and Thamniopsis congeneric. SYSTEMATIC TREATMENT Pilotrichidium Besch. Ann. Sci. Nat. Bot. ser. 6, 3: 243. 1876. Type. Hookeria callicos/ala C. Mull. , Syn. 2: 216. 1851. Plants medium to large, dark green, yellow-green, to reddish-brown, dull, rigid. Stems creeping or erect ascending, irregularly to subpinnately branched. Stems in cross-section with a si ngle row of enlarged, thin-walled, hyaline epidermal cells, central strand absent. Paraphyllia absent. Pseudoparaphyllia absent. Leaves frequently dimorphic and extremely variable in shape: oblong-ovate to rectangular-oblong; distinctly but shortly rounded to the insertion, apex truncate, broadly obtuse to long acute. Cells firm-walled throughout, those at the margins occasionally forming an indistinct
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