Ctenitis in, 1938

Viveros, Raquel Stauffer, Rouhan, Germinal & Salino, Alexandre, 2018, A taxonomic monograph of the fern genus Ctenitis (Dryopteridaceae) in South America, Phytotaxa 385 (1), pp. 448-450 : 448-450

Key to the taxa of Ctenitis in South America

1.

–.

2.

–. Laminae 2–4-pinnate-pinnatifid (Fig. 03C) at base and and medially................................................................................... 2 Laminae 1-pinnate-pinnatifid/pinnatisect (Fig. 03A) at base and medially, to 2-pinnate-pinnatifid at base and 1-pinnate-pinnatifid medially (Fig. 05E) ................................................................................................................................................ 5 Rachises flexuous (Fig. 03B), abaxially lanuginose with short to long-fimbriate scales, tangled among proscales............... ................................................................................................................................................................................ C. flexuosa Rachises more or less straight (Fig. 03C), abaxially not lanuginose, with entire or short-fimbriate scales, without proscales ................................................................................................................................................................................................ 3

A TAXONOMIC MONOGRAPH OF CTENITIS IN SOUTH AMERICA

Phytotaxa 335 (1) © 2018 Magnolia Press • 19

3. Rachises and costae with scales stiff, patent or retrorse, linear or linear-lanceolate with truncate base ( Fig. 19G ) ............... ............................................................................................................................... ............................................. C. grisebachii

–. Rachises and costae with scales flaccid, ascending or appressed, lanceolate or ovate with cordate base ( Figs. 12D , 17B )... ................................................................................................................................................................................................ 4

4. Rachises and costae with scales dark brown to blackish, sometimes pale at edges, iridescent ( Fig. 17B ) ............................. ......................................................................................................................................................... C. equestris var. equestris

–. Rachises and costae with scales uniformly light castaneous or castaneous, not iridescent ( Fig. 12D ) ..................... C. ampla

5. Pinnae incised 1/4–2/3 of the distance between the segment apex and costa (Figs. 05A–C); basal veins from adjacent segments reach the margin at sinus (e.g. Figs. 13A ; 22A ; 24A ) ................................................................................................. 6

–. Pinnae incised more than 2/3 (rarely ca. 1/2) of the distance between the segment apex and costa (e.g. Fig. 05D); basal veins from adjacent segments ending before the margin towards above the sinus or end at margin somewhat or well above the sinus (e.g. Figs. 05E; 12A; 13F)....................................................................................................................................... 9

6. Lamina apex conform (Fig. 04A) or subconform (Fig. 04 B); indusia present ................................................. C. aspidioides

–. Lamina apex confluent (pinnatifid with the most distal pinnae reduced and confluent to the rachis; Fig. 04C); indusia absent...................................................................................................................................................................................... 7

7. Petioles, rachises and costae abaxially with scales stiff, dark brown (Fig. 06C) or castaneous (Fig. 06B); abaxial laminar surface between veins glabrous; costa adaxially without scales .............................................................................. C. nervata

–. Petioles, rachises and costae abaxially with scales flaccid, light castaneous (Fig. 06A) or castaneous (Fig. 06B); abaxial laminar surface between veins with sparse bacilliform trichomes (Figs. 07C; 24B, D); costa adaxially with scales ........... 8

8. Pinnae incised 1/2–2/3 of the distance between the segment apex and costa (Figs. 05B–C); costa abaxially with linear scales ( Figs. 24A–C ) ...................................................................................................................... C. refulgens var. refulgens

–. Pinnae incised 1/4–1/3 of the distance between the segment apex and costa (Fig. 05A); costa abaxially with lanceolate scales ( Figs. 24 D–E ) ..................................................................................................................... C. refulgens var. peruviana

9. Indusia absent or small and inconspicuous .......................................................................................................................... 10

–. Indusia conspicuous ............................................................................................................................................................. 17

10. Sori with a tuft of proscales among sporangia (Fig. 08E) ................................................................................... C. fenestralis

–. Sori without a tuft of proscales among sporangia.................................................................................................................11

11. Rachis and costal scales ovate ( Fig. 12B ) .................................................................................................................. C. abyssi

–. Rachis and costal scales linear (e.g. Figs. 12H ; 13D ), linear-lanceolate (e.g. Fig. 15E ) or lanceolate (e.g. Fig. 13H )....... 12

12. Lamina 2-pinnate-pinnatifid basally, 1-pinnate-pinnatisect to 1-pinnate-pinnatifid medially (Fig. 05E); basal veins from adjacent segments end before margin, towards above the sinus (illustrated in Viveros & Salino 2017)................................. .................................................................................................................................................................. C. megalastriformis

–. Lamina 1-pinnate-pinnatisect to 1-pinnate-pinnatifid basally and medially; basal veins from adjacent segments reach the margin at sinus (e.g. Fig. 15D ), or somewhat or well above the sinus (e.g. Fig. 15A ; 17 C–D ; 19A ) ................................ 13

13. Segments margin glabrous ................................................................................................................................................... 14

–. Segments margin with catenate trichomes ........................................................................................................................... 15

14. Abaxial laminar surface between veins glabrous ( Fig. 21A ) ............................................................................. C. laetevirens

–. Abaxial laminar surface between veins with sparse bacilliform trichomes ( Fig. 22C ) ...................................... C. nigrovenia

15. Rachis scales dark brown to blackish (Fig. 06C) ............................................................................................. C. paranaensis

–. Rachis scales castaneous (Fig. 06B) or light castaneous (Fig. 06A) ................................................................................... 16

16. Costal scales lanceolate with cordate base ( Figs. 26A–B ) ............................................. C. submarginalis var. submarginalis

–. Costal scales linear or filiform with truncate or rounded base ( Figs. 2C–D ) ........................ C. submarginalis var. tenuifolia

17. Sori inframedial; costule abaxially with scales like the ones on costa, but shorter, at least basally (but with or without proscales).............................................................................................................................................................................. 18

–. Sori medial or supramedial; costule abaxially without scales like the ones on costa (but with or without proscales)........ 22

18. Stem short-creeping (Fig. 01C); rachis and costal scales abaxially with many short and long fimbriae along all margin, the ones on rachis dark brown with pale edges and the ones on costa uniformly light castaneous ( Figs. 13G–H ) ....... C. deflexa

–. Stem erect (Fig. 01A) or ascending (Fig. 01B); rachis and costal scales abaxially with or without some short fimbriae at base and laterally, the ones on rachis uniformly light castaneous or uniformly dark brown to blackish, and the ones on costa uniformly light castaneous, or uniformly dark brown to blackish or mostly castaneous but dark brown to blackish towards apex ...................................................................................................................................................................................... 19

19. Rachis scales light castaneous, entire (e.g. Figs. 17G ; 21D ) or slightly denticulate (e.g. Fig19B ) ..................................... 20

–. Rachis scales dark brown to blackish, dentate, the teeth claw-shaped (e.g. Figs. 15B, C, E ) ............................................. 21

20. Rachis and costa abaxially with scales lanceolate with cordate base, slightly denticulate ( Fig. 12I ) .................... C. anniesii

–. Rachis and costa abaxially with scales subulate, vaulted at base, entire ( Fig. 17G ) ........................................... C. eriocaulis

21. Costa abaxially with scales mostly castaneous, clathrate, but dark brown to blackish towards apex; abaxial laminar surface between veins glabrous or with sparse bacilliform trichomes ( Figs. 15A–B ) ...................................... C. distans var. distans

–. Costa abaxially with scales dark brown to blackish, mostly subclathrate, dark brown to blackish; abaxial laminar surface between veins with sparse catenate trichomes ( Figs 15C–D ) ........................................................... C. distans var. isabellina

22. Segments margin glabrous .................................................................................................................................. C. nigrovenia

–. Segments margins with catenate trichomes ......................................................................................................................... 23

20 • Phytotaxa 335 (1) © 2018 Magnolia Press

VIVEROS ET AL.

23. Rachis and costa abaxially with scales with many short and long fimbriae along all margin, the ones on rachis dark brown with pale edges and the ones on costa uniformly light castaneous (Figs. G–H) ...................................................... C. deflexa

–. Rachis and costal scales abaxially with or without some short fimbriae at base and laterally, the ones on rachis uniformly castaneous or uniformly dark brown to blackish, and the ones on costa uniformly castaneous, or uniformly dark brown to blackish................................................................................................................................................................................. 24

24. Adaxial laminar surface between veins with dense filiform trichomes (illustrated in Schwartsburd et al. 2007 and Viveros & Salino 2015) ..................................................................................................................................................... C. bigarellae

–. Adaxial laminar surface between veins glabrous or with sparse to dense catenate trichomes, sometimes also with sparse bacilliform and glandular trichomes..................................................................................................................................... 25

25. Segment apex apiculate........................................................................................................................................................ 26

–. Segment apex acute, rounded, or obtuse .............................................................................................................................. 27

26. Costa abaxially with scales lanceolate with cordate base ( Figs. 26A–B ) ...................... C. submarginalis var. submarginalis

–. Costa abaxially with scales filiform with truncate or rounded base ( Figs. 26 C–D ) ............. C. submarginalis var. tenuifolia

27. Petiole, rachis and costa abaxially with glandular trichomes; abaxial laminar surface between veins with glandular and bacilliform trichomes; indusia with glandular trichomes (Fig. 07D; illustrated in Viveros & Salino 2015) ..... C. glandulosa

–. Petiole, rachis and costa abaxially without or rarely with glandular trichomes; abaxial laminar surface between veins glabrous or with catenate and bacilliform trichomes, or filiform trichomes and bacilliform trichomes, rarely glandular trichomes; indusia with bacilliform or catenate trichomes .................................................................................................. 28

28. Rachis scales dark brown to blackish (Fig. 06C); scales on costa abaxially lanceolate ...................................................... 29

–. Rachis scales castaneous (Fig. 06B); scales on costa abaxially linear-lanceolate ............................................................... 30

29. Rachis scales 1–2 mm long; costa and laminar surface between veins with dense catenate trichomes, sometimes also with bacilliform and glandular trichomes (adaxially and abaxially; illustrated in Viveros & Salino 2015) ............... C. falciculata

–. Rachis scales 3–6 mm long; costa and laminar surface between veins with sparse catenate trichomes (adaxially and abaxially), sometimes also with bacilliform trichomes (abaxially), rarely glabrous (adaxially and abaxially; illustrated in Viveros & Salino 2015) ........................................................................................................................................... C. paranaensis

30. (4) 7–11 pinnae pairs; adaxial laminar surface glabrous; plants from southeastern Brazil (illustrated in Viveros & Salino 2015) ............................................................................................................................... ................................. C. christensenii

–. 19–27 pinnae pairs; adaxial laminar surface with sparse to dense catenate trichomes; plants from Venezuela, Colombia, Ecuador, Peru, Bolivia and Brazilian Amazon ............................................................................................... C. microchlaena

Figures

FIGURE 2. Transversal sections of petiole bases of four Ctenitis species from South America. A: C. nigrovenia, showing three vascular bundles (Salino 15212, BHCB). B: C. christensenii, showing four vascular bundles (Salino 13408, BHCB). C: C. laetevirens, showing five vascular bundles (Schimitt 561, BHCB). D: C. nervata, showing six vascular bundles (Almeida 1682, BHCB). Scale bars = 1 mm.

FIGURE 12. Some morphological characters of Ctenitis from South America. A–B: C. abyssi (Sehnem 6315, PACA). A: detail of abaxial surface of a medial pinna, showing indument B: abaxial costa scale (scale bar = 1 mm). C–D: C. ampla (Salino 3257, BHCB). C: abaxial surface of a portion of pinna rachis and two pinnules of medial pinnae. D: abaxial costa scale. E–J: C. anniesii (Gasper 851, BHCB). E: abaxial surface of a portion of rachis and a medial pinna. F: sorus with indusium. G: detail of abaxial surface of a medial pinna, showing indument. H: petiole scale. I: abaxial costa scale. J: adaxial costa scale. Scale bars in A, B, D, G–J = 1 mm; C, E, = 1 cm; F = 0.5 mm.

FIGURE 13. Some morphological characters of Ctenitis from South America. A–E: Ctenitis aspidioides (Salino 14908, BHCB). A: abaxial surface of a portion of rachis and two basal segments of a medial pinna. B: sorus with indusium. C: detail of abaxial surface of a medial pinna showing indument. D: petiole scale. E: abaxial costa scale. F–H: C. deflexa (Salino 2383, BHCB). F: abaxial surface of a portion of rachis and two basal segments of a medial pinna. G: rachis scale. H: abaxial costa scale. Scale bars in A = 1 cm; B = 0.5 mm, C–H = 1 mm.

FIGURE 15. Some morphological characters of Ctenitis from South America. A–C: Ctenitis distans var. distans (Fontana 4996, BHCB). A: detail of abaxial surface of a medial pinna showing indument. B: abaxial costa scale. C: adaxial costa scale. D–E: C. distans var. isabellina (Lombardi 6830, BHCB). D: detail of abaxial surface of a medial pinna showing indument. E: abaxial costa scale. Scale bars = 1 mm.

FIGURE 17. Some morphological characters of Ctenitis from South America. A–B: Ctenitis equestris (Breedlove 33659, CAS). A: abaxial surface of portion of pinnules and pinna rachis (scale bar = 1 cm). B: abaxial costa scale. C–G: C. eriocaulis (Pietrobom 5007, BHCB). C: abaxial surface of a portion of rachis and two basal segments of a medial pinna. D: adaxial surface of a portion of a segment. E: abaxial surface of a portion of a pinna. F: sorus with indusium. G: abaxial costa scale. Scale bars in A and C = 1cm; B, D, E, G = 1mm; F = 0.5 mm.

FIGURE 19. Some morphological characters of Ctenitis from South America. A–C: Ctenitis fenestralis (Salino 7285, BHCB). A: abaxial surface of a portion of a pinna. B: abaxial costa scale. C: adaxial costa scale. D–E: C. flexuosa (Luederwaldt 1890, UC). D: abaxial surface of a portion of pinnule. E: abaxial costa scale. F–G: C. grisebachii (Grayum 3707, MO). F: abaxial surface of a portion of pinnule. Scale bars = 1 mm.

FIGURE 21. Some morphological characters of Ctenitis from South America. A–B: Ctenitis laetevirens (Schimitt 561, BHCB). A: abaxial surface of portion of pinna. B: abaxial costa scale. C–F: Ctenitis microchlaena (unknown s.n., UC). C: abaxial surface of a portion of rachis and two basal basiscopic segments of a medial pinna. D: abaxial costa scale. E: abaxial surface of a pinna, showing indument. F: sorus with indusium. Scale bars A, B, D e E = 1 mm; C = 1 cm; F = 0.5 mm

FIGURE 22. Some morphological characters of Ctenitis from South America. A–B: C. nervata (Almeida 1682, BHCB). A: abaxial surface of a portion of rachis and two basal segments of a medial pinna. B: abaxial costa scale. C–D: C. nigrovenia (Salino 15212, BHCB). C: abaxial surface of a pinna, showing indument. D: abaxial costa scale. Scale bars in A = 1 cm; B = 0.5 mm; C, D = 1 mm.

FIGURE 24. Some morphological characters of Ctenitis from South America. A–C: Ctenitis refulgens var. refulgens (Salino 15242, BHCB). A: abaxial surface of a portion of rachis and two basal segments of a medial pinna. B: abaxial surface of rachis, costa and segment, showing indument. C: abaxial costa scale. D–E: C. refulgens var. peruviana (Tuomisto, 13392 UC). D: abaxial surface of a portion of medial pinna, showing indument. E: abaxial costa scale. Scale bars in A = 1 cm, B–E = 1mm.

FIGURE 26. Some morphological characters of Ctenitis from South America. A–B: Ctenitis submarginalis var. submarginalis (Verdi 1843, BHCB). A: abaxial surface of a portion of rachis and two basal segments of a medial pinna. B: abaxial costa scale. C–D: C. submarginalis var. tenuifolia (Pietrobom 5015, BHCB). C: abaxial surface of a portion of medial pinna, showing indument. D: abaxial costa scale. Scale bars in A = 1 cm; B–D: = 1 mm.