Diversity of the Bryoflora of the Tazekka National Park, North-East Morocco

Abstract

The Tazekka National Park, in the Moroccan Middle Atlas, offers a great diversity of flora and fauna that has been the subject of much research. However, bryophytes are the least studied in the area. In this context, the present study aims to establish an exhaustive inventory of the bryoflora of the park. Sampling was carried out along altitudinal transects, taking into consideration 10 stations spread over the entire massif, making a total of 112 records, and the period of collection was spread over two years. The exploration of the park revealed the existence of 112 taxa belonging to 100 mosses, 10 liverworts and 2 hornworts distributed over 32 families and 66 genera. The Pottiaceae family is predominant (26 species). The present study has enabled us to enrich the Moroccan bryoflora with five newly observed taxa: Anomodon attenuatus, Amblystegium serpens, Didymodon nicholsonii, Orthothecium intricatum and Stegonia latifolia var. Latifolia. Moreover, we note the predominance of acrocarpous mosses over pleurocarps, and a dominance of terricolous mosses over the other habitat types. The level of abundance of the species is variable depending on the environmental conditions of the station, showing that two liverworts are dominant and the most widely distributed in this area (Lunularia cruciata and Targonia hypophylla) and four epiphytes belonging to Orthotrichum are very dominant in an only station. This study clarifies the bryological component of the park's ecosystems. This is not without interest considering the role of these plants in the equilibrium of the environments.

Appendix

Taxonomic checklist of bryophytes from the Tazekka National Park collected in the 10 stations and their IES index. (1-Bab Azhar, 2-Bab Kefza, 3-Bab Boudir, 4-Bab Marzouka, 5-Cave of Frioauto, 6-Ras Lma, 7-Oued Amlil, 8-Jbel Tazekka, 9-Ahl Boudris, 10-Boucheffaa, *New for Morocco).

Families Species	Stations										Substrates-Altitudes (m)
	1	2	3	4	5	6	7	8	9	10
MOSSES
Amblystegiaceae
*Amblystegium serpens (Hedw.) Schimp					30						Humid siliceous soil near cascades - 1470
Cratoneuron filicinum (Hedw.) Spruce						183					Waterlogged soil near cascades - 981, 995
Hygroamblystegium tenax (Hedw.) Jenn				6							Waterlogged siliceous soil - 1088
Hygroamblystegium varium (Hedw.) Mönk				6		33					Shaded siliceous soil - 1088, 995
Anomodontaceae
*Anomodon attenuates (Hedw.) Huebener								53			Humid limestone rocks, humid shaded soil and at the base of the cedar - 1866, 2053, 2146
Bartramiaceae
Bartramia pomiformis Hedw			117					29			Schist rock, at the base of the zeen oak and holm oak, limestone rock, siliceous soil with or without shade - 1422, 1444, 1481, 1729, 1824
Bartramia stricta Brid			50	108			92	74	43		Shaded siliceous soil, limestone rock, at the base of the cedar, shale rocks, in a ravine, on barks and branches of the cork oak in a shaded and unshaded environment, on soil exposed to the sun, on trunk of the holm oak - 671, 715, 761, 776, 826, 829, 849, 1087, 1234, 1444, 1581, 1610, 1673, 1866, 1923, 1931
Philonotis caespitosa Jur			83								Waterlogged soil with seepage - 1532, 1587
Philonotis marchica (Hedw.) Brid			67	17					43		Water source in a very humid environment, on fresh soil soaked with seeping water - 1071, 1088, 1532, 1587
Brachytheciaceae
Brachytheciastrum velutinum (Hedw.) Ignatov & Huttunen			67	46			25	29			Limestone rock, dry siliceous soil, at the base and on the trunk of the holm oak in a shady area, on the trunk and at the base of the zeen oak, limestone rock - 761, 853, 1444, 1451, 1480, 1689, 1723
Brachytheciastrum velutinum var. Salicinum (Schimp.) Ochyra & Zarnowiec								6			On cedar bark, under holm oak and zeen oak, on shaded siliceous soil, between cracks in limestone rocks - 1680
Brachythecium rivulare Schimp						183					Siliceous soil soaked with sweat water, calcareous rocks - 995, 1021
Brachythecium rutabulum (Hedw.) Schimp			25					13			Under the holm oak, on schist, on the trunk and at the base of the holm oak, limestone rocks, shaded siliceous soil - 1444, 1678, 1681, 1751
Brachythecium salebrosum (Hoffm. ex F.Weber & D.Mohr) Schimp								12			At the base and on the trunk of the cedar, the shaded limestone rocks - 1923, 2053
Cirriphyllum crassinervium (Taylor) Loeske & M.Fleisch						83	8				Moist soil, near swamps, on limestone rocks near running water - 1021
Homalothecium lutescens (Hedw.) H.Rob	250		42	46				53	86		On the trunk and base of cork and zene oaks, on siliceous soil shaded by a fern grouping, on cedar trunks and limestone rocks - 931, 1045, 1071, 1108, 1157, 1480, 1581, 1610, 1729, 1866
Homalothecium sericeum (Hedw.) Schimp	150		50	46				44	71		Siliceous soil in eagle fern undergrowth, on schist rocks, siliceous soil exposed to the sun, trunk of zeen oak and holm oak, under cedar and limestone rocks - 671, 757, 795, 831, 1480, 1558, 1888, 1923
Kindbergia praelonga (Hedw.) Ochyra						100					Porphyry rocks, damp and shaded limestone rocks in cascades, limestone rocks near running water - 995, 1021
Pseudoscleropodium purum (Hedw.) M.Fleisch						33			29		Slightly moist shaded siliceous soil, limestone rocks, dry siliceous soil, under oleaster, pistacia and Tetraclinis articulata - 715, 1012
Rhynchostegiella tenella (Dicks.) Limpr								41			Slightly moist, shaded siliceous soil on limestone and dry siliceous soil under oleaster, pistacia and Tetraclinis articulata -1765, 1770
Sciuro-hypnum plumosum (Hedw.) Ignatov & Huttunen	75			13							Schist rock, basaltic rock, cork oak wood, limestone soil, bare siliceous soil exposed to the sun - 757, 795, 896, 1045
Scleropodium cespitans (Wilson ex Müll.Hal.) L.F.Koch						24					Near a stream, cascade, on siliceous soil soaked with running water - 1923, 2053
Scleropodium touretii (Brid.) L.F.Koch		100	33	17			33				In a shady and humid environment, under holm oak trees, on the trunk and base of the zeen oak, on siliceous soil, in cracks in limestone rocks, in an agricultural enclave - 896, 1078, 1249, 1480
Scorpiurium circinatum (Bruch) M.Fleisch. & Loeske						133					Limestone rocks, under cedar, on shaded or unshaded siliceous soil, near running water - 995, 1021
Bryaceae
Anomobryum julaceum (Schrad. ex P. Gaertn. et al.) Schimp								7			Soaked earth with seeping water - 1729
Bryum argenteum Hedw			33								Shaded siliceous soil, fresh soil soaked with seeping water - 1422, 1587
Bryum elegans Nees						25		6			Under holm oak, on limestone - 1012, 1923
Bryum radiculosum I.Hagen					40		17				Under holm oak, on limestone and shale rock - 1234, 1470
Bryum sp.			13	8							Siliceous soil, cool, humid and shaded, near the cedar - 848, 1587
Imbribryum alpinum (Huds. ex With.) N. Pedersen			33	17							Fresh soil soaked with water - 1088, 1518
Ptychostomum imbricatulum (Müll. Hal.) Holyoak & N. Pedersen				21	100		108		71		Limestone and schist rocks, on dry siliceous soil - 377, 776, 853, 1234, 1354, 1525
Ptychostomum capillare (Hedw.) Holyoak & N. Pedersen	75		25	17	160	67	125	15			Schist rocks, in the ravine, on trunk and branches of cork oak, in cracks in limestone rocks, on holm oak trunk, dry siliceous soil - 377, 776, 853, 1234, 1354, 1525
Ptychostomum donianum (Grev.) Holyoak & N. Pedersen				17			13	9			Shaded and humid siliceous soil, on trunks and branches of cork oak, schist rocks - 377, 761, 1143, 1770
Ptychostomum pseudotriquetrum (Hedw.) J.R. Spence & H.P. Ramsay			13								Sunny siliceous soil - 1378
Dicranaceae
Dicranella varia (Hedw.) Schimp			33								Schist rock, siliceous soil, on the trunk and at the base of the zeen oak - 1447
Ditrichaceae
Ceratodon purpureus (Hedw.) Brid								18			Moist siliceous soil, under cedar, zeen oak and holm oak - 1723, 1729
Encalyptaceae
Encalypta vulgaris (Hedw.) Hoffm	75	140				100		12			Under the zeen oak, on schist, the trunk and the base of the holm oak, on schist and limestone rocks - 995, 1021, 1157, 1649, 1712, 1762
Fabroniaceae
Fabronia pussila Raddi				42				59	171		Holm oak trunk, on shaded siliceous soil, at the base and on the trunk of cedar, pistacia and cork oak bark - 761, 831, 925, 967, 1581, 1689, 1765, 1832
Fissidentaceae
Fissidens bryoides Hedw						33					Limestone rocks, near the river, in the cascades, on siliceous soil immersed in running water - 1021
Fissidens exilis Hedw					60						Shaded siliceous soil - 1511
Fissidens taxifolius Hedw. Susbp. Taxifolius	100			8		33			29		Limestone rocks, in the gully, grassland near running water and with Eagle Fern stand in cascades, wall, running water, damp siliceous soil - 795, 925, 931, 1021, 1045
Fissidens viridulus (Sw. ex anon.) Wahlend	150							18	29		Soaked siliceous soil, in flowing water, limestone rocks, in the ravine, on the lawn near flowing water and in a stand of eagle fern, in cascades - 795, 925, 1045, 1566, 1581
Fontinalaceae
Fontinalis antipyretica Hedw								26			Shaded siliceous soil under cedar in running water - 1566, 1581
Funariaceae
Entosthodon attenuatus (Dicks.) Bryhn									43		At the edges of limestone walls and on dry siliceous soils - 1354
Funaria hygrometrica Hedw		40									Schistes- 1712
Grimmiaceae
Grimmia decipiens (Schultz) Lindb				13				18			Schist rocks, limestone rocks and under holm oak on sunny siliceous soil - 829, 1466, 1678
Grimmia pulvinata (Hedw.) Sm	100	80					33				Schist rocks, under cork oak trees, on siliceous soil exposed to the sun - 1143, 1157, 1504
Grimmia tergestina Tomm. ex Bruch & Schimp				42			25		29		Schist rocks, limestone rocks, under holm oaks, on Aleppo pine trunks, dry siliceous soil, in cracks in limestone rocks - 720, 757, 896, 1234, 1354
Grimmia torquata Drumm				29							Schist rocks and under holm oak trees, limestone rocks, dry siliceous soil - 404, 487
Grimmia trichophylla Grev	125	100		100	120	100	108	59	143		Schist rock and under the cork oak, on juniper trunk, trunk and bark of the holm oak, limestone rock, shaded siliceous soil - 377, 715, 757, 776, 826, 829, 1012, 1087, 1157, 1234, 1354, 1470, 1498, 1504, 1581, 1678, 1866
Schistidium crassipilum H.H.Blom		80	83					29			Schist rocks, in cracks in limestone rocks, on shaded siliceous soil - 1444, 1560, 1673, 1712, 1923
Hedwigiaceae
Hedwigia ciliata (Hedw.) P.Beauv		100	42					12			Schist, shaded siliceous soil - 1447, 1712, 1842
Hypnaceae
Hypnum cupressiforme Hedw				38			50	15			Schist, basalt rocks, on trunks and branches of cork oak, schist rocks and soil, in cracks in limestone rocks - 671, 720, 1122, 1581
Hypnum cupressiforme var. Lacunosum Brid	125							15			Schist, basalt rock, trunk and branches of cork oak, cedar, on shaded siliceous soil - 795, 1729
Hypnum cupressiforme var. Tectorum Brid								9			Moist shaded siliceous soil, under cedar - 1888
*Orthothecium intricatum (Hartm.) Schimp					60						Limestone rock and shaded siliceous soil - 1511
Leucodontaceae
Antitrichia californica Sull		180	152					24	143		Trunk of holm oak, trunk and branches of mastic and cork oak, trunk of cedar and zeen oak - 831, 1111, 1279, 1445, 1481, 1504, 1639, 1682, 1753
Leucodon sciuroides (Hedw.) Schwägr	125		42	17							Trunk of cork oak and holm oak - 761, 1207, 1445
Nogopterium gracile (Hedw.) Crosby & W.R.Buck			42	50			50	71	86		Trunk and branches of holm oak, on schist in a stand of eagle fern, siliceous soil exposed to sun and humidity, on trunk of cork oak, trunk and base of holm oak and cedar - 829, 931, 1087, 1111, 1444, 1558, 1770, 1824, 2146
Mniaceae
Epipterygium tozeri (Grev.) Lindb				6							Siliceous soil under holm oak -761
Plagiomnium cuspidatum (Hedw.) T.J.Kop					40	200					Porphyritic rocks, humid and shaded limestone rocks, and limestone rocks in flowing water - 995, 1021, 1511
Neckeraceae
Leptodon smithii (Hedw.) F.Weber & D.Mohr		60	33	17				24	57		Holm oak trunk, trunk and branches of pistacia and cork oak, cedar trunks and zeen oak - 761, 831, 1445, 1712, 1814, 1923
Orthotrichaceae
Orthotrichum affine Schrad. ex Brid				21	100			29	71	467	Trunk and base of holm oak, trunk of cedar and pistacia - 408, 967, 1474, 1525, 1544, 1581, 1615, 1765
Orthotrichum anomalum Hedw			33	13	60			24	43	100	Trunk of holm oak, trunk and base of cedar - 761, 1111, 1444, 1525, 1615, 1770, 1931
Orthotrichum cupulatum Hoffm. ex Brid			42	21	100			21		267	Trunk of the holm oak and at the base of the cedar - 967, 1444, 1474, 1525, 1544, 1581, 1723
Orthotrichum lyelli Hook. & Taylor	150		50	25	120			29	71	600	Trunk of cork oak and holm oak, sunny bare siliceous soil, limestone rock, at the base and on the trunk of cedar and zeen oak - 761, 1111, 1207, 1444, 1474, 1525, 1544, 1615, 1689, 1765
Orthotrichum pumilum Sw. ex anon								24			Trunk of cedar - 1866, 2053
Orthotrichum rupestre Schleich. ex Schwägr		120		50				76	71	200	Rocks in the eagle fern stand, at the base of the zeen oak - 408, 761, 967, 1339, 1504, 1581, 1615, 1689, 1765, 1770
Orthotrichum speciosum Ness								6			Trunk of cedar - 1770
Orthotrichum striatum Hedw							33	53			Trunk and base of the cedar, on slightly moist shaded siliceous soil, on limestone rocks - 1290, 1723, 1729, 1866, 2053
Orthotrichum tenelllum Bruch ex Brid			33	38	80			44	43		Trunk of holm oak, cedar and zeen oak - 715, 761, 967, 1339, 1444, 1525, 1689, 1765
Polytrichaceae
Polytrichum juniperinum Hedw			42	25				35			Shaded siliceous soil under zeen oak and at the base of cedar trees - 1422, 1451, 1581, 1729
Pogonatum aloides (Hedw.) P.Beauv		100	83					18			Schist rock, on trunk and base of holm oak, shaded siliceous soil and base of cedar - 1378, 1444, 1692, 1866
Pottiaceae
Barbula unguiculata Hedw		60	33	54		33	33	9	43		Siliceous soil, at the base of the holm oak, schist rock, between the cracks of the limestone rocks - 618, 671, 715, 1012, 1049, 1290, 1498, 1560, 1692
Cinclidotus riparius (Host ex Brid.) Arn								24			Shaded or unshaded siliceous soil, in running water - 1566, 1581
Dialytrichia mucronata (Brid.) Broth						100					On slightly moistened walls - 995, 1021
Didymodon acutus (Brid.) K.Saito				17			25				Between the cracks in the limestone - 404, 1234
Didymodon fallax (Hedw.) R.H.Zander				38	60		50		43		Dry siliceous soil, limestone and schist - 849, 853, 1234, 1354, 1451, 1525
Didymodon insulanus (De Not.) M.O.Hill			25	33		50		12	57		Sunny bare siliceous soil, on limestone and schist rocks - 668, 715, 761, 1012, 1480, 1729
Didymodon luridus Hornsch				50		83	25		57		Calcareous rocks, dry siliceous soil, at the base of the holm oak, oleaster and thuya plants - 377, 668, 715, 1012, 1143, 1451
*Didymodon nicholsonii Culm		60		25			50	9			Schist rock, in cracks in limestone, on siliceous soil - 377, 487, 1087, 1234, 1712, 1762
Dydimodon tophaceus (Brid.) Lisa	100			13			25	9	43		Schist rocks, limestone rocks, shaded siliceous soil - 987, 1050, 1157, 1354, 1753
Didymodon vinealis (Brid.) R.H.Zander				25			33				Dry siliceous soil, between limestone and schist cracks - 404, 849, 1234
Eucladium verticillatum (With.) Bruch & Schimp	37				40						Calcareous soil, shaded siliceous soil - 1045, 1511
*Stegonia latifolia (Schwägr.) Venturi ex Broth								9			Moist siliceous soil - 1824
Syntrichia ruralis (Hedw.) F.Weber & D.Mohr			42	58				53			Limestone rock, schist rock, trunk of Aleppo pine, on trunk and base of holm oak - 720, 967, 1049, 1339, 1480, 1770, 1866
Syntrichia laevipila Brid								9			At the base of the cedar, on the trunk of the holm oak, on limestone rocks - 1729
Syntrichia norvegica F.Weber								4			At the base of the cedar - 1770
Timmiella barbuloides (Brid.) Mönk	100			17			25	9			Schist rock, basalt rocks, siliceous soil, at the base of the cedar, in cracks in limestone rocks - 795, 931, 1087, 1729
Tortella humilis (Hedw.) Jenn								4			Moist siliceous soil - 1765
Tortella squarrosa (Brid.) Limpr		120		75		100	50		86		Schist rock, limestone rock, dry siliceous soil - 487, 640, 715, 850, 1012, 1087, 1649
Tortula inermis (Brid.) Mont			33								At the base of the zeen oak and holm oak, in a shady environment - 1639
Tortula muralis Hedw						50					On wall - 1055, 1137
Tortula subulata Hedw			42	38				44			Sunny limestone rock, dry siliceous soil, on trunk and base of holm oak, cedar, on siliceous soil near limestone - 487, 967, 1480, 1729, 1770, 1866
Tortula subulata Hedw. Var. Subulata								4			Shaded siliceous soil with low humidity under cedar trees - 1682
Tortula vahliana (Schultz) Mont			42	21				29			Trunk of zeen oak, holm oak and at the base of the cedar - 757, 1639, 1770, 1931
Trichostomum brachydontium Bruch	75	100		25							Bare siliceous soil exposed to the sun, under holm oak and cedar - 795, 1108, 1451, 1712
Trichostomum crispulum Bruch		60		17							Shale rock, limestone and siliceous soil - 668, 1712
Weissia controversa var. Crispata (Nees & Hornsch.) Nyholm		80						9			Schist, siliceous soil, under cedar - 1712, 1770
Pterigynandraceae
Pterigynandrum filiforme Hedw								65			At the base of the cedar, on limestone rocks, on trunks and branches of zeen oak - 1770, 1814, 1824, 2146
LIVERWORTS
Arnelliaceae
Gongylanthus ericetorum (Raddi) Ness				25							Shaded siliceous soil - 931, 1108
Aytoniaceae
Reboulia hemisphaerica (L.) Raddi			75					121	86		At the base of zeen oak trees, in a stand of eagle fern, on siliceous soil, on a cork oak trunk - 925, 1111, 1378, 1422, 1447, 1498, 1610, 1678, 1680, 1681, 1739, 1753, 1762, 1842
Corsiniaceae
Corsinia coriandrina (Spreng.) J.Lindb	38										Moist and shady soil, schistose rocks - 1045
Fossombroniaceae
Fossombronia angulosa (Dicks.) Raddi	150			17				12			In a ravine by a river, on shady ground soaked with seeping water - 795, 931, 1045, 1108, 1751
Fossombronia foveolata J.Lindb								12			In a ravine near running water, under cedar, on shaded or unshaded siliceous soil - 1566, 1581
Frullaniaceae
Frullania dilatata (L.) Dumort								53			Limestone rocks, on trunk and base of cedar, on shaded siliceous soil - 1931, 2053, 2146
Lunulariaceae
Lunularia cruciata (L.) J.Lindb	250		133	179	40	300	83	106	243		In rock crevices, on the trunk and branches of cork oak and schist rocks, siliceous soil exposed to the sun, schist and basalt - from 404 to 1842
Pelliaceae
Pellia neesiana (Gottsche) Limpr						167					Moist limestone rock, shaded, on walls near cascades - 995, 1021
Ricciaceae
Riccia lamellosa Raddi					20			18			In cedar forest on dry, unshaded siliceous soil - 1581, 1751
Targioniaceae
Targionia hypophylla L	350		100	142	120	83	100	106	200		In cracks in limestone rocks, schist rocks, on siliceous soil and under the feet of holm oak and thuya - from 404 to 1842
HORNWORTS
Anthocerotaceae
Phaeoceros laevis (L.) Prosk	38			19					57		Shaded siliceous soil soaked with seeping water - 795, 931, 1071, 1088, 1108
Phymatoceros bulbiculosus (Brot.) Stotler, W. T. Doyle & Crand.-Stotl			13								Moist siliceous soil, on the trunk and at the base of the zeen oak - 1480
Number of species by station	21	17	37	53	18	22	25	64	30	5