Bark anatomy of lianescent Bignoniaceae: a generic synopsis
Author
Pace, Marcelo R.
Author
Marcati, Carmen R.
Author
Lohmann, Lúcia G.
Author
Angyalossy, Veronica
text
Adansonia
2023
3
2023-05-30
45
12
167
210
journal article
10.5252/adansonia2023v45a12
1639-4798
8006364
XIII.
Amphilophium
clade
TAXONOMIC
INFORMATION
. — This clade contains a single genus,
Amphilophium
, with multiples of four phloem wedges in transversal section.
Amphilophium
currently includes species from six previously recognized genera (see
Fischer
et al.
2004
;
Table 1
),
Amphilophium
,
Distictella
Kuntze
,
Distictis
Mart. ex Meisn
,
Glaziova
Bureau
,
Haplolophium
Cham.
, and
Pithecoctenium
Mart. ex Meisn.
TOTAL
NUMBER
OF
SPECIES
IN
THIS
CLADE
. — 47 species (
Lohmann & Taylor 2014
).
STUDIED
SPECIES
. — Seven species,
Amphilophium bracteatum
(Cham.) L.G.Lohmann
,
A. crucigerum
(
L
.)
L
.
G
.Lohmann,
A. dolichoides
(Cham.) L.G. Lohmann
,
A. elongatum
(Vahl.) L.G. Lohmann
,
A. magnoliifolium
(Kunth) L.G. Lohmann
,
A. paniculatum
(
L
.) Kunth, and
A. pulverulentum
(Sandwith) L.G. Lohmann.
Regular phloem
Thin to thick fiber bands (
Fig. 15D
), assemblages absent.
Variant phloem
General configuration.
Non-fibrous(
Fig.15C
),with 20-30 rows of sieve tubes and phloem parenchyma cells between fiber bands (
Fig. 15C
). The axial elements tend to be diffuse, sometimes forming short radial and tangential groups (
Fig.15C, F
).As seen in tangential section, the axial parenchyma is storied (
Fig.15G
).
Sieve-tube elements.
As seen in transverse section, each sieve element is associated with one companion cell (
Fig. 15F
). The sieve tubes are generally solitary or in multiples of two, showing no distinct arrangement (
Fig. 15F
), often scattered among the phloem parenchyma cells or forming radial or tangential rows of two or three cells (
Fig. 15F
). In
A. crucigerum
and
A. elongatum
sieve tubes of two distinct diameters are present (
Fig.15F
). In
A. crucigerum
the large and narrow sieve elements occur adjacent to one another in the phloem (
Fig.15F
), while in
A. elongatum
they are organized radially from wide (70 µm in average) to narrow (35 µm in average), gradually decreasing in diameter. As seen in longitudinal section, the sieve elements are rather short (<
1 mm
) and their end walls are transverse to inclined, with most sieve elements bearing simple sieve plates(
Fig.15E
). Compound sieve plates with 2-3 sieve areas are sometimes encountered.
Axial parenchyma.
The phloem parenchyma tends to intermingle all the cells (
Fig. 15F
), although at some portions they can be recognized as forming radial rows of 3-4 cells (
Fig.15F
).Crystalliferous parenchyma surrounds the fiber bands (
Fig.15E
), except for
A. magnoliifolium
and
A. pulverulentum
where parenchyma cells surround the fiber bands, but crystals are lacking.
Fibers.
Fiber bands occur at considerable distances (more than 20 cells) from one another in
Amphilophium
(
Fig. 15C
), the
Bignonieae
genus with the most distant fiber bands. The fiber bands possess 2-3 rows of cells and are discontinuous between two limiting rays, i.e., they never occupy the entire space between two rays (
Fig. 15C
).
Rays.
The limiting rays are only lignified to the xylem face (
Fig. 15C
). The wedge rays are non-lignified, not even when crossing the fiber bands.
Crystals.
The crystalliferous parenchyma bears styloid crystals in
Amphilophium elongatum
, prismatic crystals in
A. crucigerum
(
Fig. 15E
), and
A. paniculatum
; crystals are lacking in
A. magnoliifolium
. Most, if not all of the axial and ray parenchyma cells contain acicular and navicular crystals.
FIG
. 15. — Secondary phloem of
Amphilophium
:
A -G
,
Amphilophium crucigerum
;
A -E
, transverse sections;
A
, periderm with a stratified phellem, with an alternation of thick-walled lignified cells, and thin-walled, non-lignified cells. Phelloderm thick;
B
, prismatic crystals abundant in the phellem (
arrowheads
);
C
, included phloem wedge forms interxylary phloem in the genus.The cambium remains active and produce phloem in the enclosed wedgde,causing a conspicuous collapse of the nonconducting phloem;
D
, regular phloem with thin to thick fiber bands. A large portion of the nonconducting undergoes extensive collapse. Some of the rays greatly dilate by division of the ray margins, forming wedge shaped rays. Some ray cells undergo belated sclerosis forming sclereids (
arrows
);
E
, detail of the secondary phloem,showing turgid sieve tubes (
arrows
) with one companion cell lying on its corner. Loss of conductivity happens a few layers away from the cambium, with collapse of the sieve tubes (
arrowheads
);
F
, longitudinal radial section. Wide sieve tubes with simple sieve plates. Crystalliferous parenchyma bearing prismatic crystals around fiber band;
G
, longitudinal tangential section, regular phloem. Axial parenchyma storied, rays wide, with over 5 cells in width. Abbreviations:
cz
, cambial zone. Scale bars: A, C, G, 200 μm; B, F, 100 μm; D, 600 μm; E, 50 μm.
FIG
. 16. — Secondary phloem of
Dolichandra
:
A -D
, transverse sections;
B
,
Dolichandra unguiculata
. All other photos are from
Dolichandra unguiscati
;
A
, limiting rays non-lignified, tend to expand and divide (
arrows
), non-fibrous phloem. Thin fiber bands;
B
, non-fibrous phloem, sieve tubes in short radial multiples. Limiting rays non-lignified;
C
, pith also undergo divisions, uniting to divisions at the limiting rays (
arrowheads
). Nonconducting phloem collapsed;
D
, detail of the conducting variant phloem. Sieve tubes (
arrows
) with one or two companion cells.Acicular crystals abundant;
E
, tangential section. Storied structure in both axial and ray cells;
F
, radial section. Sieve tube elements with compound sieve plates (
arrow
). Abbreviations:
cz
, cambial zone;
pi
, pith. Scale bars: A, 200 μm; B, E, 150 μm; C, 500 μm; D, 50 μm; F, 100 μm.
Periderm
A
single periderm is formed (
Fig. 15A
). The phellem is non-stratified, with evenly thin-walled cells in
Amphilophium elongatum
,
A. magnoliifolium
, and
A. paniculatum
; stratified, with alternating thin and thick-walled cells in
A. crucigerum
(
Fig. 15A, B
). Prismatic crystals are found in the phellem of
Amphilophium crucigerum
(
Fig. 15B
). The phelloderm is thin in
Amphilophium elongatum
and thick in
A. crucigerum
and
A. magnoliifolium
, with more than three cell layers (
Table 1
). Stratified lenticels are found in
Amphilophium crucigerum
.