Cyathea

Lehnert, Marcus & Kessler, Michael, 2018, Prodromus of a fern flora for Bolivia. XX. Cyatheaceae, Phytotaxa 334 (2), pp. 118-134 : 123-125

Key to the Bolivian species of Cyathea

1 Lamina pinnnate to pinnate-pinnatisect ............................................................................................................................................ 2

– Lamina 2-pinnate-pinnatifid to 3-pinnate ......................................................................................................................................... 4

2 Plants with thin, erect trunk; fronds notably hairy ..................................................................................................... C. bipinnatifida

– Plants without erect trunk; fronds glabrous or with very few appressed hairs adaxially ................................................................. 3

3 Pinnae incised to 1/2 towards the costae, with lobes rounded, obtuse or acute; veins irregularly foming areoles.............. C. uleana

– Pinnae incised to 1/4 toward the costae, the lobes rounded to truncate; veins always forming areoles ......................... C. subincisa

4 Sori exindusiate .................................................................................................................................................................................. 5

– Sori indusiate .................................................................................................................................................................................. 17

5 Fertile veins of the central pinnae regularly simple ........................................................................................................................... 6

– Fertile veins of the central pinnae forked ......................................................................................................................................... 8

6 Pinnules basally cordate, basal veins of a segment originating from costule, connivent to or ending blindly below a sinus; paraphyses of the same length as or longer than the sporangia ............................................................................................................ C. ulei

– Pinnules basally truncate to cuneate; basal veins of a segment originating from midvein, not connivent to a sinus; paraphyses shorter than sporangia ....................................................................................................................................................................... 7

7 White petiole scale margins with some scattered, dark brown marginal teeth; lamina apex usually gradually reduced; sori ± medial ..................................................................................................................................................................................... C. dombeyi

– White petiole scale margins without brown marginal teeth; lamina apex ± abruptly reduced; sori supramedial to submarginal ....... ........................................................................................................................................................................................... C. pungens

8 Pinnae opposite to subopposite; lamina thin-textured with deeply serrate segments separated polygonal sinuses........ C. conjugata

– Pinnae alternate; lamina thin-textured to coriaceous with entire to crenate segments separated by rounded to triangular sinuses .... ............................................................................................................................................................................................................ 9

9 Costae with long prickles to 5 mm long, these very pointed, almost needle-like ........................................................ C. microdonta

– Costae without long prickles, smooth to muricate or rarely with few longer prickles but then these thick and blunt .................... 10

10 Distal parts of petiole scales with differentiated margins reduced to minute dark teeth, scales mostly light brown to whitish (on trunks and transition to petiole with dark brown center).................................................................................................. C. poeppigii

– Distal parts of petiole scales with differentiated magins, the margins without darker teeth, scales entirely orange-brown to castaneous or with slightly paler margins .................................................................................................................................................... 11

11 Trunks absent, or rarely to 1.5 m; laminae deltate; paraphyses much longer than the sporangia, often broken off, remaining basal cell of the paraphyses not reddish brown ......................................................................................................................................... 12

– Trunks erect, commonly 2 m or taller; laminae ovate; paraphyses shorter than the sporangia or of the same length, if longer then basal cell of the paraphyses reddish brown ...................................................................................................................................... 13

12 Plants of usually warm, dry areas like grassy slopes and sunny rock outcrops; petiole scales narrowly lanceolate, decidedly contorted, ± uniformly reddish-brown with thin white margins easily abraded and often missing........................................... C. villosa

– Plants of cool, wet cloud forests and subparamos; petiole scales long ovate to lanceolate, only weakly contorted, castaneous with broad yellowish white margins...................................................................................................................................... C. zongoensis

13 Petiole scales uniformly orange to reddish-brown ........................................................................................................................... 14

– Petiole scales dark brown to castaneous with lighter colored margin.............................................................................................. 15

14 Croziers covered in white mucus; petioles and rachises soft-hairy on both sides, with spreading green wings (often shrunken and appressed in dried material).......................................................................................................................................... C. mucilagina

– Croziers not covered in white mucus; petioles and rachises only hairy adaxially, without spreading green wings ....... C. phalerata

15 Fronds climbing in understory (rarely hanging free from cliffs), supported over high branches and dangling in distal parts, trunk mostly> 10 cm diam.; pinnae and pinnules more than 2 cm broad, long-stalked .............................................................. C. gibbosa

– Fronds spreading freely, patent-arching from relatively thin trunk usually <10 cm diam.; pinnae and pinnules less than 2 cm broad, sessile to short-stalked...................................................................................................................................................................... 16

16 At least some petiole scales with discernable white margins; petiole scurf an amorphous ephemeral layer of pale brown to whitish squamules; fronds axes abaxially green to yellowish brown, glabrous to glabrescent; veins abaxially without spreading hairs ....... ........................................................................................................................................................................................... C. dombeyi

– Petiole scales without white margins; petiole scurf a fine persistent layer of dark reddish squamules; fronds axes abaxially green to yellowish brown, glabrous to glabrescent, veins abaxially with at least some spreading hairs................................... C. lasiosora

17 Indusia hemitelioid ........................................................................................................................................................................... 18

– Indusia meniscoid to sphaeropteroid................................................................................................................................................ 23

18 Petioles inermous; scales at petiole bases and trunk apices ovate-lanceolate, not tapering at tip.................................................... 19

– Petioles with many spines of variable size; scales at petiole bases and trunk apices lanceolate, or if base ovate then tapering at tip .......................................................................................................................................................................................................... 20

19 Petioles ususally without scales; sori and indusia hidden by small scales inserted at the receptacles; largest pinnules pinnatisect and sessile........................................................................................................................................................................... C. arnecornelii

– Petioles ususally with many persistent scales reaching onto the rachis; indusia and sori not hidden, no scales inserted at the receptacle; largest pinnules sometimes fully pinnate and long-stalked ..................................................................................... C. vilhelmii

20 Petiole scales absent (or few and easily shed), but scurf well developed, white-brownish when dry, darker in fresh material; indusia mostly subsphaeorpteroid but fragmenting to scale-like remnants, variable ................................................................... C. planadae

– Petiole scales present and usually persistent, but scurf usually absent; indusia all of the same size and orientation, not fragmenting but larger ones may split in half ....................................................................................................................................................... 21

21 Indusia to 1 mm long, scale-like, arching over the sori; sori supramedial to inframarginal; segment margins crenulate ................... ............................................................................................................................................................................................. C. andina

– Indusia to 0.5 mm long, often hidden under the sori, appressed to lamina or weakly ascending; sori medial to supramedial; segment margins dentate to serrate................................................................................................................................................................. 22

22 Leaf axes and veins with many hairs; hairs on costules to 2 mm long, when stretched out reaching or surpassing the first pair of sori; petiole scales brown with broad whitish margins, never of reddish color ..................................................... C. leucolepismata

– Leaf axes and veins with scattered hairs; hairs on costules shorter, not reaching sori; petiole scales brown with narrowly light-colored margins, sometimes nearly concolorous and reddish.................................................................................................. C. lindigii

23 Indusia meniscoid to cyatheoid (mostly remnants of larger sphaeropteroid indusia) ...................................................................... 24

– Indusia sphaeropteroid...................................................................................................................................................................... 26

24 Laminae 2-pinnate-pinnatifid; ultimate segments dentate; indusia irregularly fragmenting from cyatheoid to meniscoid or sometimes hemitelioid shape; petiole spiny ............................................................................................................................. C. planadae

– Laminae 2-pinnate; ultimate segments (= pinnules) coarsely crenate; indusia keep shape, remaining entire or as persisting fragments; petiole smooth....................................................................................................................................................................... 25

25 Laminae long elliptic, densely hairy on both sides on and between the veins; costae alate between the pinnules; indusia small discs hidden under sori ...................................................................................................................................... C. bipinnatifida × delgadii

– Laminae deltate to spade-shaped, hairy only on the veins, densely to moderately adaxially, only scarcely abaxially; costae not alate; indusia cyatheoid to subsphaeropteroid.................................................................................................................... C. bettinae

26 Petiole scales entirely whitish to pale yellowish brown or bicolorous with margins of this color, strongly contrasting with brown to dark brown center......................................................................................................................................................................... 27

– Petiole scales dark orange to dark castaneous or shiny black, concolorous or concordantly bicolorous with the narrow margins brownish, orange or yellowish, or rarely in few scales the margins also whitish ............................................................................ 33

27 Laminae and leaf axes abaxially with many minute crested, dark castaneous scales; petiole scales orange brown with whitish margins to dark brown with yellowish margins; segments adaxially glabrous ................................................................ C. carolihenrici

– Laminae and leaf axes with different indument of larger concolorous and/or bicolourous scales; petiole scale color varying from dark brown to blackish with white margins to brown with white to pale brown margins; segments sparsely pubescent adaxially.... .......................................................................................................................................................................................................... 28

28 Laminae with more hairs adaxially than abaxially (especially towards the margins); small flattish concolorous scales of different hues of brown on segments abaxially; petiole with scurf of small brown scales............................................................................. 29

– Laminae with fewer hair adaxially than abaxially; abaxially scales uniformly white or bicolorous dark brown/white; petiole with scurf of crested ivory or stramineous scales..................................................................................................................................... 31

29 Petioles very short; pinnae gradually tapering towards the base, sometimes with a gap between the last two pairs; or if petioles long and without basal pinnae, then laminae broad triangular and plants without notable trunks.......................................... C. parvifolia

– Petioles long, without remote proximal pinnae; laminae broadly elliptic to ovate.......................................................................... 30

30 Trunks slender, 4–6 cm diam., without old petioles; pinnules to 4 cm long and generally less than 1 cm wide; petioles smooth to tuberculate .......................................................................................................................................................................... C. bettinae

– Trunks stout, 8–12 cm diam., with old petioles at least in small plants (less than 2 m height); pinnules to 8 cm long and generally more than 1 cm wide; petioles muricate to aculeate............................................................................................. C. austropallescens

31 Petiole scurf well developed, long lasting, with some larger flattish scales to 4 mm long; laminar scales predominantly bicolorous, some uniformly white or brown ........................................................................................................................................ C. ruiziana

– Petiole scurf scattered, often sparse or abraded, with squamules to 1 mm long; laminar scales white to very light brown............ 32

32 Scales of petiole scurf and laminar indument without dark marginal teeth; laminae ususally with black spots ..... C. dintelmannii

– Petiole scurf and laminar indument with at least some whitish flat scales with some dark marginal teeth; laminae usually without black spots ....................................................................................................................................................................... C. straminea

33 Pinnules long triangular with free basal segments; many small (<0.5 mm), castaneous, strongly dissected squamellae on frond axes, petioles, and croziers ......................................................................................................................................... C. carolihenrici

– Pinnules linear to oblong with adnate basal segments, or if free then green-alate between segments; squamellae on frond axes, petioles, and croziers to 1.0 mm, tan to brown, not strongly dissected............................................................................................ 34

34 Petioles shiny brown, smooth to tuberculate, scurf soon shed, very fine; largest pinnules to 7.0 × 1.1–1.8 cm and notably short-stalked; laminae abaxially glabrous or with few whitish squamules .................................................................................. C. lechleri

– Petioles not shiny and/or muricate to spiny (may be tuberculate basally), with brown scurf persistent to caducous, or with hairs; largest pinnules more than 8.0 × 2.0 cm or if smaller then sessile; laminae abaxially with orange, brown, or tan squamules, and few to many hairs on the veins abaxially ................................................................................................................................................ 35

35 Veins densely hairy on both surfaces; laminae hairy between the veins; pinnules rarely exceeding 80(–95) × 15(–18) mm ....... 36

– Veins only sparsely hairy adaxially, or if abundantly hairy on the veins then no hairs between the veins; largest pinnules at least 85 × 15 mm (generally>100 × 20 mm) ................................................................................................................................................ 39

36 Petiole scales long lanceolate, concolorous orange, rarely yellowish or dull orange-brown; laminae hairy between the veins only abaxially...........................................................................................................................................................................................37

– Petiole scales lanceolate to broadly lanceolate, concolorous brown or with somewhat lighter margins; laminae normally hairy between the veins on both surfaces..................................................................................................................................................38

37 Segments 2–3 mm wide, with margins entire or inconspicuously crenulate; rachises densely hairy or, if hairs abraded, notably scabrous with persistent hair bases; petioles dull brown to yellowish .................................................. C. delgadii (pubescent form)

– Segments 3–4 mm wide, margins conspicously crenulate to crenate; rachises sparsely hairy, not scabrous with persistent hair bases, though sometimes muricate with little spines; petioles blackish to purpureous ......................................... C. delgadii (typical form)

38 Petioles not hairy, or only distally so on the adaxial side; petioles with few to many long prickles; pinnae alternate...... C. herzogii

– Petioles hairy on both sides; petioles without long prickles, inermous or sparsely muricate; pinna ± opposite in lower half of the lamina ................................................................................................................................................................................. C. obnoxia

39 Petioles dark brown with paler, white to tan scurf; basal petiole scales appearing almost concolorous shiny black, margins narrow and often abraded (distal petiole scales with broader paler margins, some of them whitish); basal sinuses narrow, acute; segments falcate with acute, finely serrate tips; laminae and veins with scattered pale squamules and almost no hairs .......... C. dintelmannii

– Petioles blackish to stramineous, scurf equally dark or darker, mostly orange to reddish brown; basal sinuses wider and obtuse to rounded; segments not or weakly falcate, acute to obtuse, crenulate to crenate tips; laminae and veins with orange brown squamules and few to many hairs............................................................................................................................................................ 40

40 Petiole scales concolorous orange to orange-brown, never streaked; scurf caducous ....................................................... C. delgadii

– Petiole scales bicolorous dark castaneous to shiny black with orange margins to nearly concolorous orange-brown, sometimes with blackish streaks in the central portion; scurf persistent.................................................................................................................... 41

41 All petiole scales narrowly lanceolate, with orange-brown margins confluent at the tips, dark center either well defined or appearing washed out from the base to tip; persistent scales may reach up all along the rachis, here with bleached white margins; petiole scurf squamules to 2 mm long; short hairs sometimes copious on costules abaxially but bveins almost glabrous............................. ...................................................................................................................................................................................... C. catacampta

– At least basal petiole scales broadly lanceolate with ovate body and attenuate tip, dark color of the sharply defined darker center reaching tip; petiole scales persistent only at petiole base, rarely few larger scales also retained in axils between rachises and costae; petiole scurf with squamuellae to ± 1 mm long; veins abaxially with some hairs, ususally hairier than costules........................ ........................................................................................................................................................................................ C. squamipes