Afoninia dahurica Ignatovа, Goffinet & Fedosov, 2015

Ignatov, Michael S., Ignatova, Elena A., Czernyadjeva, Irina V., Goffinet, Bernard, Kuznetsova, Oxana I. & Fedosov, Vladimir E., 2015, Afoninia, a new moss genus of Funariaceae from Transbaikalia (East Siberia, Russia), Arctoa 24 (1), pp. 14-20 : 16-20

Afoninia dahurica Ignatovа, Goffinet & Fedosov , gen. et sp. nov.

Figs. 2-12 .

Type: Russia, Southern Siberia , Zabaikalsky Territory , Gazimuro-Zavodskoy District, midstream of Gazimur River , circa 12 km SW of Batakan Settlement, 51°50’46"N, 118°43’48"E, alt. 635 m. Rock outcrops. 23.VII.2012. Coll. O.M. Afonina #3512 (Holotype LE, isotypes MHA, MW) .

Etymology: The species name indicates the collecting locality, Dahuria, a part of Trans-Baikal area, called by Russian colonists after a local population name.

Plants pale green, dull green or yellowish, growing on soil as individual shoots or forming loose tufts. Stems 2.5– 4.0 mm, green or brownish at base, with well-differentiated hyalodermis, abruptly delimited sclerodermis consisting of 1–2 rows of thick-walled red-brown cells, very thin-walled medullar cells and narrow central strand. Lower leaves small and distant, upper leaves enlargered and crowded in distal part of stem, contorted when dry, erectspreading when moist, (1.8–)2.5–3.5 u (0.8–) 1.2–1.6 mm, spatulate, slightly concave, widely acute or obtuse at apex; margins obtusely serrulate in distal 1/2, entire in proximal part, plane; costa moderately strong, slightly narrowing upwards, ending 4–6 cells below leaf apex; distal and median laminal cells 40–60 u (18–)22–28(–40) µm, 4-6- angled, thin-walled, not differentiated or colored at margins, basal laminal cells elongate rectangular, 90–120 u 30– 40 µm, basal marginal cells slightly shorter. Apparently autoicous (smaller male shoot mixed among plants with sporophytes, but seems so easily broken off that no one obvious junction was seen in a limited material available for study). Male shoots slightly smaller than female ones. Androecia terminal, perigonial leaves similar to stem leaves, obtusely serrulate throughout. Paraphyses mostly 5-celled, 3 basal cells narrow rectangular, two upper cells inflated, more than twice wider, both spheric, or terminal cell pyriform, all cells not colored. Seta 1.0– 1.6 mm, light reddish brown. Capsules 2.0– 2.6 mm long, ca. 0.8 mm wide, symmetric, cylindric, erect or slightly inclined, yellowish to light brownish when mature, urn smooth, neck ca. 1/4–1/3 the capsule length, weakly delimited; exothecial cells oblong, ± irregular, rhombic and hexagonal, slightly bulging, with moderately thickened outer walls, with cuneate thickenings of radial walls, at urn mouth in 3 rows transversely rectangular to quadrate, with numerous one-celled stomata at the neck and urn base. Annulus compound, revoluble, consisting of 2 rows of large inflated cells. Peristome double, hardly extending above urn mouth; exostome teeth 16, attached to endostome, 120– 140 µm long, slightly narrowing distally, obtuse, occasionally irregularly notched in upper part, finely papillose throughout on outer surface, with low ventral trabeculae; endostome segments 16, opposite to teeth, ca. 150 µm long, fused at base and forming low basal membrane, segments irregular in shape, slightly wider than exostome teeth, vertically papillose-striolate on outer surface. Operculum flat or weakly convex, indistinctly mamillate, with red rim. Spores 18–21 µm, brownish, finely verrucose. Calyptra cucullate, inflated below, with rostrum ca. 0.9 mm long.

Differentiation. There is no species in the Zabaikalsky Territory that can be confused with Afoninia . The Funariaceae are represented in the area by Funaria hygrometrica and Entosthodon pulchellus , both having asymmetric and inclined capsules and well-developed peristome, while species of Physcomitrum and Entosthodon hungarius , reported from there, are eperisomate.

The species of the clade most closely related to Afoninia , i.e., Mediterranean Funariella curviseta , South American Entosthodon laevis , and Australian E. apophysatus , all have inclined and more or less asymmetric capsules, non-revoluble annulus, and different peristomes: Funariella is eperistomate, Entosthodon laevis has well developed double peristome, E. apophysatus ,in contrast, develops only a rudimentary endostome, whereas Afoninia has a strongly reduced double peristome. Other differences include mitrate calyptrae in both these Entosthodon species versus Afoninia and Funariella , both sharing cucullate calyptra. However, the latter genus is characterized by a short rostrum of the calyptra, whereas in Afoninia the rostrum is long, resembling that of Funaria hygrometrica .

Although the Funaria -like annulus and the long rostrate calyptra may point to affinities of Afoninia to species of Funaria , inferences from DNA data do not support this hypothesis, and similarities with F. aequidens , F. flavicans and F. polaris , must hence reflect homoplasy. Resolving Afoninia outside of Funaria s. str. is consistent with it developing a smooth and symmetric urn, an erect capsule and a peristome only slightly exceeding the level of annulus, all features not known from Funaria .

Entosthodon is very variable and likely will be split in the future into monophyletic entities; however, in any case all its species in modern definition lack a revoluble annulus, and E. attenuatus , the type of the genus has long peristome teeth.

Ecology: Afoninia dahurica was collected in five localities in the eastern part of Zabaikalsky Territory, at altitudes ranging from 635 m to 1170 m a.s.l., on soilcovered rock outcrops, on fine soil in crevice and in niches between rocks, and on bare soil on rocky slope with steppe-like vegetation. In the type locality, in the end of July the plants had both immature and almost ripe capsules and well-preserved male and female inflorescences, while in other places, plants had mostly mature and deoperculate sporophytes despite of approximately the same time of collecting.

Additional specimens examined: ASIATIC RUSSIA: Zabaikalsky Territory : Kalgansky District , Nerchinsky Mt. Range , 10 km from Kalga Settlement to Aleksandrovsky Zavod Settlement, 50°56’37”N, 118°41’23”E, 754 m alt., 26.VII.2012, Czernyadjeva 31-12 ( LE, MHA) ; Kyra District , Sokhondinsky Nature Reserve: Enda River, ca. 49°27’N, 110°51’E, 1070 m alt., 14.VII.2010, Czernyadjeva 19-10 ( LE, MHA) ; Agutsa River , ca. 49°38’N, 111°27’E, 1170 m alt., 18.VII.2010, Czernyadjeva 28-10 ( LE, MHA) ; Gazimuro-Zavodsky District , Pryamoy Mulday River, 52°14’48”N, 119°23’22”E, 664 m alt., 22.VII.2012, Afonina #2812 ( LE, MHA) .

Figures

Figs. 2-6. Afoninia dahurica (from holotype: Russia, Zabaikalsky Territory, Gazimur River, Afonina 3512, LE): 2, 4, 5 – capsule mouth from outside, showing annulus and peristome variation; 3, 6 – peristome attached to spore sac wall from outside: note the teeth tapering towards the bases, so their lower parts are narrower than endostome segments, while distally the teeth and segments have about the same width. Median line is seen on few teeth (arrowed in 2). Scale bars: 100 Pm for 2, 4-5; 50 Pm for 3, 6.

Figs. 7-11. Afoninia dahurica (from holotype: Russia, Zabaikalsky Terri- tory, Gazimur River, Afonina 3512, LE): 7: exostome tooth from outside, show- ing ornamentation; 8: peristome from inside, showing ornamentation of endostome segments (En) on their inner surface and also partly ornamentation of exostome tooth (Ex) from inside; note that anticlinal cell divisions in the inner peristomial layer, which resulted in peristomial formula 4:2:4, are seen only in most basal plates (arrowed), while distally corresponding divisions are not performed, hence the peristome has 4:2:2 pattern distally; 9-10: spores; 11: endostome segment inner surface, showing longitudinal rows of papillae, a close up from 8. Scale bars: 30 Pm for 8, 10 Pm for 7, 10-11; 2 Pm for 9.

Fig. 12. Afoninia dahurica (from holotype: Russia, Zabaikalsky Territory, Gazimur River, Afonina 3512, LE): 1 – habit, wet; 2, 4 – capsules; dry; 3,5 – capsules, wet; 6 – peristome & exothecium at urn edge; 7 – exothecium in the middle part of urn; 8 – exothecium & stomata at urn base; 9 – habit, dry; 10 – stem transverse section; 11 – transverse section of urn wall in the middle part of urn; 12-13, 15-16 – leaves; 14 – upper laminal cells; 17-18 – paraphyses from perigonium; 19 – median laminal cells; 20 – basal laminal cells. Scale bars: 5 mm for 1; 3 mm for 12-13, 15-16; 2 mm for 2-3, 9; 1 mm for 4-5; 100 Pm for 6-8, 10-11, 14, 17-20.