ARTIFICIAL KEYS TO THE FRULLANIA SPECIES OF THE NEW ZEALAND BOTANICAL REGION

1. Leaf lobes delicately crenulate-denticulate or dentate, the teeth often formed by paired projecting cells, teeth hyaline, to ca. 5 µm long.
• F. chevalierii [Auckland, Northland]
1. Leaf lobes edentate, smooth-margined or at most crenulated.
• 2

   

2. First branch leaf of the lateral branch (BL1) with TWO saccate segments.
• 3
2. First branch leaf of the lateral branch (BL1) either a) ± characteristic of normal stem leaves (dorsal lobe may be reduced and strap-shaped) with only ONE saccate segment or lobule, or, b) all BL1 segments or lobes explanate (or at mostchannel shaped) and leaves characteristic of the main stem developing at BL2 or rarely at BL3.
• 5

   

3. Pseudo-dimorphic branching, i.e., the secondary and tertiary branches have markedly smaller leaf-lobes and underleaves (compared to those of the main stem), branches often becoming reduced in length; lobuli clavate, small (in comparison to the dorsal lobe), occupying < 0.2_ the exposed area of the lobe, and only 1-1.5(2)_ long as wide, usually spreading at angles of 45-90 degrees from the stem; 4-6 oilbodies per cell; plants medium to large.
• F. ptychantha [Throughout: particularly in moist/wet habitats]
3. Pseudo-dimorphic branching absent to inconspicuous; lobuli typically cylindrically pitcher-shaped, large in proportion to the lobes, its area obscuring 0.25-0.4_ the exposed area of the dorsal lobe, lobuli usually spreading at angles greater than or equal to 45; (1)2-3 (4) oilbodies per cell; plants small to medium.
• 4

   

4. Perianth plicate, ± 5-keeled, rim of beak with single-celled protuberances; leaf lobes often with weakly incurved distal margins, when flat with widely rounded apices; lobe median cells with small trigones, intervening walls ± leptodermous; oilbodies collectively occupying ca. 1/3 of cell lumen, surface appearing coarsely granular to weakly botryoidal, spherical, 3-5 µm in diam., to ovoid or ellipsoidal, 4.0-10.0 µm long _ 2.0-4.0 µm wide.
• F. congesta [Throughout]
4. Perianth 3-keeled, rim of beak smooth, single-celled protuberances absent; leaf lobe with appendiculate to acute apices lobe median cells with large and concave trigones; oilbodies collectively occupying ca. 3/4 of cell lumen, oilbody surface appearing strongly botryoidal, oil bodies ovoid or ellipsoidal, 6.0-12.0 µm long _ 3.0-5.0 µm wide.
• F. bartlettii [Northland, Radar Bush]

   

5. Lobules asymmetrical, cucullate (hood-shaped) to broadly galeate (hollow and vaulted; like a helmet), upper portion inflated (then with mouth and/or sector adjacent to stem appearing "pinched in"), lobules wider than long (high) or nearly as wide as long, free outer margin typically rostrate with well-developed beak.
• 6
5. Lobules ± symmetrical, either cylindrical-campanulate (bell-shaped) and ± constricted above the mouth, or cylindrically pitcher-shaped, lobules ± inflated throughout or nearly so, mouth not strongly compressed, lobules as wide as long or 1-3.5 (4.5)X long as wide, outer margin never developing into well-developed beak.
• 19

   

6. Perianth surface smooth, lacking any form of ornamentation, e.g., uni- or multicellular projections or outgrowths, papillae, or tubercles.
• 7
6. Perianth surface with some form of ornamentation.
• 14

   

7. Perinath appear dorsi-ventrally appressed, with only lateral keels (ventral keels indistinct or absent); perinath to half- exserted, subspherical; or exserted, inflated throughout cylindrical to sac-like; no evidence of plicae.
• 8
7. Perianths 3-keeled or plicate with multiple, accessory keels.
   • 11

      

8. Perianth dorsi-ventrally appressed or almost so; plants (small) medium to large.
• 9
8. Perianth globose or subspeherical, or inflated throughout and cylindrical or sac-like; plants large or at least often forming large patches.
• 10

   

9. Lobules typically large, obscuring 50-75% of the surface area of the exposed dorsal lobe; stem underleaves to c. 0.2 bifid, suborbicular in outline; lobe median cells with subtriangular trigones and intermediate thickenings rare or absent; plants medium to large.
• F. scandens [Central & southern North Island, South Island, incl. Stewart Is., Auckland Is.]
9. Lobules typically small to medium, only obscuring to 25% of the surface area of the exposed dorsal lobe; stem underleaves to c. 0.3-.4 bifid, subobovate in outline; lobe median cells with subtriangular to weakly nodulose trigones and intermediate thickenings often present; plants small to medium.
• F. deplanata [Throughout]

   

10. Lobules typically small, obscuring > 20% of the surface area of the exposed dorsal lobe, lobules almost totally obscured by the underleaves; perianth large, emergent and conspicuous, inflated, cylindrical to sac-like; lobe median cells with ± small to medium, subtringular to subnodulose trigones, occasionally intermediate thickening; (4) 5­9 (10) oil-bodies/cell, lacking any significant ornamentation; plants often pale green.
• F. spinifera [Throughout]
10. Lobules typically large, obscuring < 50 % of the surface area of the exposed dorsal lobe, underleaves not obscuring lobules (occasionally some phenotypes with both small and large lobules); perianth inconspicuous, only c. half-exserted, immersed by bracts and bracteoles, subspherical; lobe median cells with ± large distinctive subtringular to subnodulose trigones; (2,3) 47 (8) oil-bodies/cell, coarsely granular or coarsely segmented; plants olive green to almost reddish brown.
• F. patula [Throughout]

   

11. Perianth strongly 3-keeled, i.e., 2 dorsal keels + 1 ventral keel - keels usually occurring from perianth base to apex (occasionally maybe plicate towards beak giving the appearance of 1-2 extra minor keels on the ventral face).
• 12
11. Perianth with multiple, accessory keels on the dorsal and/or ventral surface, i.e., 5-12 (14) keeled in transverse profile.
• 13

   

12. Leaf lobes strongly squarrose when wet, standing almost vertically away from the substrate, base cordate, usually both dorsal and ventral bases auriculate-appendiculate; apex of underleaf bilobed to 0.20-0.3 its length, underleaf ± orbicular in outline.
• F. squarrosula [Throughout]
12. Leaf lobes not strongly squarrose when wet, base cordate with only dorsal base auriculate-appendiculate; shortly bifid, apex of underleaf bilobed to 0.10-0.20 its length, lower lateral margins ± straight.
• F. falsiloba [Throughout]

   

13. * 2 large lateral keels + 1 dorsal keel + 2 ventral keels; thus 5-keeled in transverse profile; stylus minute to small, filiform to narrowly triangular, (3) 4-6 (7) cells in total; leaf lobe median cells with usually semi-straight to occasionally sinuose cell walls; leaf lobes, lobules, stem may develop gemmae and propagula.
• F. pentapleura [Throughout]
13. 2 +/- large lateral keels + 2-3 dorsal keels + 3-4 (5) ventral keels; thus 7-10 keeled in transverse profile; stylus medium to large in size, foliose, lanceolate-triangular, (6)8-26(30) cells in total; leaf lobe median with strongly sinuose cell walls; gemmae and propagula absent.
• F. probosciphora [Central North Island, South Island]
* See also key for F. probosciphora and F. pentapleura

   

14. Perianths with projections, outgrowths, papillae, or tubercles DENSLY distributed over the surface of the perianth; lobule often with obtuse or obliquely truncate at beak.
• 15
14. Perianths with projections or outgrowths, papillae, or tubercles only SPARESLY over the surface of the perianth; lobule often with a long-pilose beak.
• 18

   

15. Perianth inflated, obpyriform, or subspherical, without distinctive keels, and without accessory keels.
• 16
15. Perianth 3-keeled or plicate with accessory keels.
• 17

   

16. Perianth densely covered by irregular-sized, subcircular, scalelike, foliar or plate-like outgrowths (mimicking a pine cone); perianth exserted; gemmae absent.
• F. nicholsonii
16. Perianth densely covered by irregular-sized, spinose-like outgrowths, ranging from unicellular outgrowths of 3-5 cells to larger outgrowths of 2-4 (5) cells wide and 4-8 (10-12) long; perianth half-exserted or half- immersed by bracts and bracteoles; gemmae occasionally present.
• F. rostellata

   

17. Leaf lobes squarrose when wet, lobes semi-convolute when dry, so that shoots becoming tubular; projections often densely distributed throughout surface from base to almost the apex.
• F. pcynantha var. typica [Throughout]
17. Not squarrose, lobes ± flat when wet or dry; projections often becoming sparse towards apex.
• F. pcynantha var. setchellii [North Island]

   

18. Underleaves toothed, with 1-3 pairs of acute to spinose teeth produced on lateral margins; perianth with multicellular, tuberculate-spinose, outgrowths, partuclarly on the keels.
• F. monocera
18. Underleaves entire, or a blunt angulation or an indistinct, obtuse tooth on lateral margins; perianth without spinose outgrowths, but margins of keels (occ. between keels) appearing angular-crispate, with low tubercles
• F. allanii

   

19. Perianth smooth, lacking multicellular projections or outgrowths, or at most buliform, with the surface bearing numerous, very low, broad and rounded elevations; only a single elevation per cell.
• 20
19. Perianth surface with irregular-sized, subtringular to oblong, multicellular tubercles or outgrowths.
• F. anomala

   

20. First branch underleaf with only 2 distinct and explanate appendages: 1 large unlobed ventral segment + 1 slightly smaller dorsal segment; median cells with thin semi-stright cell walls or heavily thickened and nodulose; oil-bodies with ± distinct segmentation appearing finely to coarsely granular or segmented.
• 21
20. First branch underleaf with 3 distinct divisions: 1 ventral, explanate, bilobed segment + 1 dorsal saccate segment; or 3 explanate segments; median cells with subequally thickened walls; oil bodies lacking any significant ornamentation and frequently appearing as almost homogeneous oil droplets (except ocelli or enlarged oilbodies, if present).
• 25

   

21. Lobules incumbent, with the apices inclined towards the stem, dense and partially imbricate to each other; lobes densely imbricate; plants medium in size; perianth ± 3-keeled.
• F. incumbens
21. Lobules ± parallel to the stem; lobes loosely imbricate, contiguous, to distant; plants small to medium; perianth 3-keeled or multi-keeled.
• 22

   

22. Lobules campanulate, constricted immediately above the mouth; the lobule usually with a conspicuously large globose, thick-walled cell occupying the base of the lobule near point of attachment; lobe median cells with sinuous or nodulose cell walls.
• 23
22. Lobules helmet-shaped and NOT constricted above the mouth; large globose, thick-walled cell occupying the base of the lobule absent or inconspicuous; lobe median cells semi-straight or nodulose.
• 24

   

23. Plants small (shoots c. 500-750 µm wide); gemmae and propagula often present and caducous; often sterile; perianth when present, small, to 800 µm long; surface smooth;
• F. fugax
23. Plants medium (shoots 750-1000 µm wide); gemmae and propagula absent and non-caducous; perianths often present; perianth with very low lying irregular outgrowths or tubercles.
• F. media

   

24. * 2 large lateral keels + 1 dorsal keel + 2 ventral keels; thus 5-keeled in transverse profile; stylus minute to small, filiform to narrowly triangular, (3) 4-6 (7) cells in total; leaf lobe median cells with usually semi-straight to occasionally sinuose cell walls; leaf lobes, lobules, stem may develop gemmae and propagula.
• F. pentapleura [Throughout]
24. 2 ± large lateral keels + 2-3 dorsal keels + 3-4 (5) ventral keels; thus 7-10 keeled in transverse profile; stylus medium to large in size, foliose, lanceolate-triangular, (6) 8-26 (30) cells in total; leaf lobe median with strongly sinuose cell walls; gemmae and propagula absent.
• F. probosciphora [Central North Island, South Island]
* See also key for F. probosciphora and F. pentapleura

   

25. Lobuli typically parallel to subparallel, often bicoloured with the basal 2-5 cells towards the mouth (up to 0.25 of the lobule) hyaline to subhyaline - in contrast to the olive-green to brown pigmentation elsewhere of lobule; oil-bodies of the lobe median cells dimorphic in form; Type One: 1-2 per cell, very large, finely granular, and occur from the basal cells through to the central region of the lobe, occupying 0.75 to almost the entire cell lumen; Type Two: 2-3 (4) oilbodies per cell, typically small, spherical (1) 2-4 (5) µm in diam. to ovoid or ellipsoidal (2) 3-5 (6) µm _ (1) 2-3 (4) µm, subhyaline, without any significant, visible, internal structure i.e giving the appearance of being almost homogeneous; styli ± triangular.
• F. maritimus [Stewart Island, Auckland Islands]
25. Lobuli at angles of (25 degrees) to 45 degrees (80 degrees) from the stem, so that the lobuli are tilted outwards; lobuli not bicoloured; Type One oil-body (described above) absent, ocelli if present, confined to basal cells, otherwise Type Two only; styli small, forming an arched filament, to large and subtriangular or oblong, of up to 15 (20) cells.
• 26

   

26. Underleaves entire, or at most lateral margins rarely with a blunt angulation or an indistinct, obtuse tooth; basal ocelli absent; styli various, +/- large in comparison to leaf lobe, either subtriangular or a broadly arched filament or subrectangular.
• 27
26. Underleves toothed, with 1-2 (3) distinct teeth on the lateral margins; basal ocelli present; styli typically small, usually an arched filament, terminating in a unicellular row of 2-3 cells.
• 30

   

27. Stylus a distinct, subrectangular shape with a truncate apex; a distinctive angular projection on the lobule immediately above the slit; plants small (c. 250-500 µm) µm; branching sparingly.
• F. truncata [Coromandel (Table Mt.), Westland, Stewart Island]
27. Stylus subtriangular or a broadly arched filament, acute to subacute apex; distinctive angular projection absent (not to be confused with the +/- discoloured, gibbous, cell above mouth); plants small to medium.
• 28

   

28. Plants monoicous; leaves characteristic of the main stem not developing until either BL2, BL3 or BL4; cells of the perianth surface often each elevated as small, to ca. 4µm high tubercles; elaters (12) 14-20 (22) in total.
• F. pauciexiliens [Coromandel (Table Mountain), South Island].
28. Plants dioicous; leaves characteristic of the main stem developing at BL1; perianth surface smooth; elaters (24) 26-34 (42) in total.
• 29

   

29. Perianth 3-keeled, caducous leaves.
• F. rostrata var. typica [Throughout]
29. Perianth plicate, 2 dorsal keels + 1 ventral keel + multiple accessory keels.
• F. rostrata var. plicata [Tablelands, NW Nelson]

   

30. Plants dioicous; first branch leaf with a saccate appendage, and similar to those of the main stem.
• F. atterima [Throughout]
30. Plants monoicous; first branch leaf (BL1) without any saccate appendages, dorsal and ventral lobes often conspicuously reduced in size, and frequently strap-shaped, leaves characteristic of the main stem developing at BL2 or BL3.
• 31

   

31. Plants typically small (300-500µm wide); perianth, smooth, with a conspicuous groove on the dorsal face; underleaves of primary branches typically with 1, rarely 2, inconspicuous lateral teeth (to 4 cells in length); underleaves of secondary branches less prominently toothed than those of the main stem.
• F. subrostrata [Throughout]
31. Plants small to medium in size (to 1000 µm wide); perianth, plicate with conspicuous elevated hill-like elevations; underleaves of primary branches with 1-3 conspicous lateral teeth (to 6 cells in length); underleaves of secondary branches similar in form to those of the main stem.
• F. colliculosa [Subalpine: NW Nelson, Mt Arthur; Arthurs Pass]