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Volume 63, 1934
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– 529 –

Series II—Two-needle Pines (leaf section semi-circular).

P. austriaca (P. nigra var. austriaca). (Rotorua.) (Fig. 5.)

Comparative with: P. laricio, which cannot be distinguished by any fundamental differences; but has fewer resin ducts, sometimes internal, and few stomatic lines. P. austriaca has an uneven, ridged upper surface. P. densiflora, P. sylvestris, which have a similar number of peripheral resin ducts.

Leaf section: Semi-circular.

Epidermis: Regular, oblong, much thickened cells with dot lumen.

Stomata: 8 on upper face, 12 on lower face. Guard cells sunk below level of the hypoderm, thickened on inner face; opening urn-shaped, inner air space bounded by U-cell.

Hypodermis: 1 row of regular cubical cells, with thicker walls than in a water layer, and 1 row of sclerosed cells increased at angles and in groups.

Mesophyll: 3 rows of plicate, 5 lobed cells.

Resin ducts: 14 median, arranged around the stele, small, of about 8 slightly thickened outer cells, lined with epithelial cells.

Endodermis: Small, rounded, slightly thickened cells.

Vascular strand: Kidney shaped; vascular bundles 2, separated by thin-walled parenchyma and a narrow band of thickened cells; immediately bounding the phloem is a band, 1 cell thick, of sclerenchyma cells with contents.

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Fig. 5

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P. Banksiana. (Fig. 6.)

  • Comparative with: P. luchuensis, which differs definitely in no diagnostic characters, and from which P. Banksiana can only be distinguished by its widely separated vascular bundles, and the presence of an indefinite “water layer” of hypodermis.

    • P. caribaea, which has a greater number of stomatic lines.

    • P. Thungergii and P. muricata, with usually 3 resin ducts.

  • Leaf section: Semi-circular and elongated.

  • Epidermis: Regular, cubical, much thickened cells.

  • Stomata: 6 on upper face, 9 on lower face. Guard cells slightly sunk below hypodermis, sclerosed on both walls; opening bounded by crescent-shaped epidermal cells; inner space bounded by U-cell.

  • Hypodermis: A pseudo water layer and one row of sclerosed cells much flattened in places, increased at the angles and in scattered groups.

  • Mesophyll: Two rows increased at corners, of much plicated cells, with distinct palisade infoldings.

  • Resin ducts: 2 median; large, of 1 row of sclerosed cells.

  • Endodermis: Elongated, elliptical cells, thickened on outer and lateral walls.

  • Vascular strand: A flattened ellipse, vascular bundles 2, widely separated by thin-walled parenchyma; on flanks of the phloem are scattered storage cells, and a few pitted transfusion cells are visible; distinct storage “rays” traversing xylem and phloem. N.B.—In many sections examined there were no resin ducts.

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Fig. 6

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P. densiflora. (Fig. 7.)

  • Comparative with: P. resinosa, which has only 2 or 4 resin ducts peripheral, and some median.

    • P. luchuensis, austriaca, muricata, Thunbergii, all of which have median resin ducts.

    • P. Massoniana, which also has one water layer only.

  • Leaf section: Semi-circular.

  • Epidermis: Cubical cells with rounded outer wall and distinct cuticle layer.

  • Stomata: 7–8 on upper face, 10–14 on lower face; often in pairs close together; not sunken; guard cells sunk below hypoderm; inner chamber bounded by U-cell.

  • Hypodermis: 1 row of water layer cells.

  • Mesophyll: 3 rows of plicate, roughly 6 lobed circular cells, with palisade infoldings.

  • Resin ducts: 10–12 peripheral, 3 on the upper plain face; fairly small, of 7–9 thickened cells, with slight epithelial layer.

  • Endodermis: Distinct, elliptical cells thickened on outer wall.

  • Vascular Strand: Elliptical, vascular bundles 2; surrounded by thick-walled parenchyma; on the outer flank of the phloem is a mass of “albuminous” cells with storage cell contents, and outside these a group of transfusion cells.

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Fig. 7

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P. echinata (Rotorua). (Fig. 8.)

  • Comparative with: P. Murrayana, with fewer stomatic lines and vascular bundles widely separated; P. Banksiana, with indefinite water layer, fewer stomata, and bundles very widely separated; P. luchuensis, with no water layer.

  • Section: Semi-circular.

  • Epidermis: Regular, small cells with even outer surface.

  • Stomata: 11 on upper surface, 15 on lower surface, guard cells only slightly sunk.

  • Hypodermis: 1 row of regular, thin-walled water cells and an inner, slightly thickened scleroderm layer.

  • Mesophyll: A distinct palisade layer together with an inner row of plicate parenchyma, increased at angles.

  • Resin ducts: 2 median, of about 12 thickened cells with an inner epithelial layer of cells; sometimes 1–3 median or internal additional small ducts in the lower mesophyll.

  • Endodermis: Elliptical, regular, of large cells thickened on the outer and lateral walls.

  • Vascular strand: Of two vascular bundles, closely placed, with transfusion tissue on the flanks.

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Fig. 8

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P. halepensis (Rotorua). (Fig. 9.)

  • Comparative with: P. Massoniana, which has one hypoderm layer only.

    • P. sylvestris, which has one hypoderm water layer and no sclerosed layer, and with outline much broken by stomata.

    • P. densiflora, which has usually more resin ducts and has a water layer only.

  • Leaf section: Semi-circular.

  • Epidermis: Regular, cubical cells with thickened walls and reduced lumen; much thickened cuticle.

  • Stomata: 3 on upper face, 6 on lower face. Guard cells much sunk below level of hypodermis, thickened on both faces; opening flanked by the projecting tips of epidermal cells, and bounded inside by U-shaped mesophyll cell.

  • Hypodermis: 1 row of thin-walled water layer; with a further sclerosed layer along the lower surface.

  • Mesophyll: Markedly plicate circular cells, with palisade infoldings.

  • Resin ducts: 6 peripheral, sometimes 1 or 2 median; 2 only on upper face; of about 9 thickened cells, around an epithelial layer.

  • Endodermis: Round cells, unthickened.

  • Vascular strand: Elliptical, 2 V-B connected by thickened parenchyma cells; a mass of these also face the protoxylem; a few transfusion cells are scattered on the outer flanks of the bundles; large single thickened parenchyma cells scattered below endodermis. Some “rays” apparent in the phloem.

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Fig. 9

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P. laricio (P. nigra, var. calabrica). (Fig. 10.)

  • Comparative with: P. pinaster, which has a distinct water layer.

    • P. caribaea, which has some resin ducts internal, and up to 15 stomata on the lower face.

    • P. Banksiana, which has 2 resin ducts.

    • P. Thunbergii, which has 2–3 resin ducts.

    • P. muricata, which has a water layer, 3 resin ducts, and deeply sunken stomata.

    • P. Murrayana, which has no resin ducts (in samples examined).

  • Leaf section: Semi-circular.

  • Epidermis: Regular, large, narrowly oblong cells, much thickened.

  • Stomata: 9 on upper surface, 8 on lower surface. The cup-shaped narrow openings bounded by the elongated epidermal cells; guard cells sunk below hypodermis, fully sclerosed.

  • Hypodermis: 1–2 rows of thickened cells, a single row only on the lower surface.

  • Resin ducts: 8–10 median, sometimes 1 or 2 internal; usually of 2 rows of thickened cells, not sclerosed.

  • Endodermis: Very regular, elliptical cells, thickened on lateral walls.

  • Vascular strand: Flatly elliptical, vascular bundles 2, surrounded by large-celled parenchyma. Subtending the phloem is a band of collenchymatous cells. No “rays” visible.

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Fig. 10

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P. luchuensis. (Fig. 11.)

  • Comparative with: P. Banksiana, with widely separated vascular bundles and an indefinite water layer.

    • P. caribaea (2-needle), with many (15–11) lines of stomata.

    • P. Thunbergii and P. muricata, with usually 3 resin ducts and the latter with deeply divided outline.

  • Leaf section: Semi-circular.

  • Epidermis: Regular, much thickened cubic cells, with lumen reduced to a dot; outer cuticle distinct.

  • Stomata: 6 on upper face, 9 on lower face. Deeply sunk, opening bounded by roughly triangular epidermal cells, inner opening bounded by U-cell.

  • Hypodermis: 2 rows of thickened, irregular cells.

  • Mesophyll: 2 rows of plicate cells, showing palisade infoldings.

  • Resin ducts: 2 median, of 1 or 2 rows of slightly thickened cells.

  • Endodermis: Large celled, thickened on lateral walls.

  • Vascular strand: Elliptical, vascular bundles 2, spreading; large cells of sclerenchyma scattered opposite the phloem.

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Fig. 11

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P. Massoniana. (Fig. 12.)

  • Comparative with: P. halepensis and P. densiflora, but differs from the former in having 1 row of sclerosed hypoderm cells on all faces.

  • Leaf section: Semi-circular.

  • Epidermis: Regular, cubical cells; all walls much thickened, giving a dot lumen; outline even.

  • Stomata: 7–8 on upper face, 8–10 on lower face.

  • Hypodermis: 1 or 2 rows of sclerosed hypoderm cells increased at angles.

  • Mesophyll: Much plicated, hexagonal cells with palisade formation on all surfaces.

  • Resin ducts: 6–8 peripheral, on both surfaces, each duct of 1 row of thickened cells surrounding a large cavity approximately 1–16th of the plain face length.

  • Endodermis: Oval cells with thickened outer and adjacent walls, not very distinct from inner parenchyma.

  • Vascular strand: Elliptical; V-B 2, a few transfusion rays through xylem and phloem, scattered thickened parenchyma cells around the bundles.

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Fig. 12

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P. muricata (Rotorua). (Fig. 13.)

  • Comparative with: P. laricio, which has no water layer.

  • P. pinaster, with 9 resin ducts.

    • P. Murrayana, which has no resin ducts and the two vascular bundles widely separated in the stele.

    • P. luchuensis, which has no distinct water layer.

    • P. austriaca, which has 14 resin ducts.

    • P. pinea, with no resin ducts and no water layer.

  • Leaf section: Semi-circular.

  • Epidermis: Deeply broken by stomata; regular, almost cubical cells rounded on outer edge and much thickened.

  • Stomata: 8 on upper face, 11–12 on lower face. Large, sunk below hypodermis; the wide opening bounded by narrow epidermal cells; guard cells completely thickened, inner air space bounded by a U-cell.

  • Hypodermis: 1 “water layer” and one sclerosed layer increased at angles and sometimes in groups.

  • Mesophyll: 2 rows of deeply plicate circular cells, with palisade infoldings on either face.

  • Resin ducts: 2 median; of 10 sclerosed cells.

  • Endodermis: Large oval cells.

  • Vascular strand: Elliptical, vascular bundles 2, surrounded by thin-walled parenchyma; a little transfusion tissue on flanks; distinct “rays” visible.

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Fig. 13

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P. Murrayana (P. contorta var. latifolia). (Fig. 14.)

  • Comparative with: P. laricio, which differs in having many (8–10) resin ducts.

    • P. Banksiana, which has a water layer.

    • P. luchuensis, with V-B not widely apart.

    • P. austriaca, with many (14) resin ducts.

    • P. pinea, of greater size of leaf section.

  • Leaf section: Semi-circular.

  • Epidermis: Regular, almost cubical, thickened cells, rounded on outer edge.

  • Stomata: 7 on upper surface, 11 on lower surface. Guard cells slightly sunk.

  • Hypodermis: 1 row of water layer cells, and 1 row, increased in places, of rounded sclerenchyma.

  • Mesophyll: 2 rows of shortly plicate circular cells.

  • Resin ducts: Nil.

  • Endodermis: Regular, elliptical cells, thickened on outer and lateral walls.

  • Vascular strand: Flattened elliptical, vascular bundles 2, small, surrounded by and widely separated by large, thin-walled parenchyma.

  • N.B.—Dallimore and Jackson, Elwes and Henry, and Mathews give this species as possessing median resin ducts.

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Fig. 14

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P. pinaster (Rot. 2). (Fig. 15.)

  • Comparative with: P. laricio, with no water layer.

    • P. austriaca, with no water layer.

    • P. pinea, with few or no resin ducts, and no parenchyma “rays.”

  • Section: Semi-circular.

  • Epidermis: Regular, cubical, thickened cells with dot lumen.

  • Stomata: 7 on upper face, 15 on lower face. Deeply sunk; guard cells below level of the hypodermis. Tubular opening is bounded by an elongated epidermal cell which is “beaked” at the inner margin, restricting the stomatic opening; inner air space bounded by a U-cell.

  • Hypodermis: 1 “water layer” of large, somewhat irregularly shaped cells, with one or two rows of large, irregularly shaped sclerenchyma cells interrupted at the stomata, increased at angles and with cell walls much thickened.

  • Mesophyll: 2 or 3 rows of deeply plicate, roughly 4–6 lobed, square parenchyma cells.

  • Resin ducts: 9 median, placed around are on the lower face only; of 1 row of 9 sclerosed cells.

  • Endodermis: Large, almost circular cells, scarcely thickened.

  • Vascular strand: Flattened elliptical; vascular bundles 2, large; a row of thickened parenchyma bounds the phloem masses and extends between the bundles. A mass of distinct albuminous cells flanks the phloem masses adjoined by the transfusion tissue. “Rays” distinct in xylem phloem.

Note.—This specimen differs from other types examined, which have (i) only 2 resin ducts; (ii) 20 ducts; the leaves of these types have no water layer.

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Fig. 15

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P. pinea (Rotoura). (Fig. 16.)

  • Comparative with: P. pinaster, which has resin ducts and a distinct water layer, with hypodermal sclerenchyma projecting into the mesophyll between deeply sunk stomata.

    • P. Murrayana, with a distinct water layer and greater number of stomata.

  • Leaf section: Semi-circular.

  • Epidermis: Even, large, much thickened cells; cuticle thin.

  • Stomata: 6 on upper face, 9 on lower face. Guard cells sunk below hypodermis, partly thickened.

  • Hypodermis: 2 rows of thickened cells, the outer layer having the appearance of a water layer, but with thickened walls, the inner row sclerosed.

  • Mesophyll: 3 rows of plicate, 6 lobed cells, with outer deep palisade infoldings.

  • Resin ducts: Nil.

  • Endodermis: Circular cells not distinct from inner parenchyma, slightly thickened on outer walls.

  • Vascular strand: Elongated plano-convex, 2, small vascular bundles, surrounded by small celled parenchyma; xylem very regular, flanked on outer edges by transfusion cells; no rays visible.

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Fig. 16

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P. resinosa. (Fig. 17.)

  • Comparative with: P. luchuensis, which has only median resin ducts, and no water layer.

    • P. halepensis and P. Massoniana, in both of which the resin ducts are bounded by 1 row of cells only.

  • Leaf section: Semi-circular.

  • Epidermis: Cubical, even cells with a dot lumen.

  • Stomata: 7 on upper face, 10 on lower face. Slightly sunk inner chamber bounded by U-cell.

  • Hypodermis: One row of pseudo water layer cells.

  • Mesophyll: 3–4 rows of roughly 6 lobed plicate cells, wth slight palisade infoldings.

  • Resin ducts: 2 median, 4 peripheral, the latter fairly large of 12 thickened cells in a double row; those median are smaller and apparently rudimentary.

  • Endodermis: Of flattened, elliptical cells, thickened on outer walls.

  • Vascular strand: Elliptical, vascular bundles 2, surrounded by thin-walled parenchyma; a little transfusion tissue on flanks; no visible rays.

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Fig. 17

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P. sylvestris (Rotorua). (Fig. 18.)

  • Comparative with: P. halepensis, which has 2 hypoderm layers.

    • P. resinosa, which has fewer resin canals—2–4 peripheral and sometimes 2 median.

  • Leaf section: Semi-circular.

  • Epidermis: Outline broadly crenate, of very regular, narrowly oblong cells with reduced lumen.

  • Stomata: 9–12 on lower face, 8 on upper face. The urn-shaped opening bounded by projecting lips of epidermal cells; guard cells sunk below the hypodermal layer.

  • Hypodermis: A water layer of flattened, thin-walled cells and one row sclerosed cells.

  • Mesophyll: Shortly plicate, hexagonal cells, with granular cell contents.

  • Resin ducts: 8 peripheral, of 1 row of 8–9 irregular spherical, much thickened cells with lumen much reduced.

  • Endodermis: Regular, elliptical, small cells, laterally thickened.

  • Vascular strand: Elliptical flattened; vascular bundles 2, small, separated by a mass of thickened parenchyma cells; a band of collenchymatous cells crosses the stele facing the phloem.

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Fig. 18

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P. Thunbergii. (Fig. 19.)

  • Comparative with: P. Massoniana, with 6–8 peripheral ducts.

    • P. halepensis, with 6–7 ducts, and stomata in lines of 6 + 3.

    • P. muricata, with deeply sunken stomata and hypodermis in groups.

    • P. luchuensis, with 2 layers of scleroderm hypodermis.

    • P. densiflora, with peripheral resin ducts.

  • Leaf section: Semi-circular.

  • Epidermis: Regular, thickened cells with dot lumen.

  • Stomata: 8 on upper surface, 12 on lower surface. Guard cells sunk below hypodermis; crescent-shaped epidermal cells bounding the wide opening.

  • Hypodermis: 1 row of small thin-walled cells next to epidermis, together with 1 row, doubled in places, of larger irregularly shaped sclerosed cells.

  • Mesophyll: Slightly plicate 5–6 lobed cells, with palisade infoldings on both faces.

  • Resin canals: 2 median placed at the angles; of 1 row of sclerosed cells, lined with epithelium.

  • Endodermis: Regular, almost spherical cells.

  • Vascular strand: Elliptical, vascular bundles 2, surrounded by thin-walled parenchyma.

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Fig. 19