WOOD ANATOMY OF SOLANACEAE

Transcription

Allertonia, 6(4), April 1992, pp. 279—326
WOOD ANATOMY OF SOLANACEAE: A SURVEY
SHERwIN CLQu1sT’
ABSTRACT
Quantitative and qualitative wood anatomy data are presented for 106 collections of Solanaceae,
representing 21 genera and 82 species. Wood expressions are diverse in Solanaceae: wood is ring
porous to diffuse porous; vessels have simple perforation plates (rarely vestiges of bars); lateral wall
pitting is alternate; grooves andlor helical thickenings are present in vessels ofsome species; imperforate
tracheary elements range from true tracheids with pits 5—7 m in diameter (Brunfelsia, Fabiana) to
fiber-tracheids with pit cavities I m in diameter; vasicentric tracheids and vascular tracheids occur
in a scattering of species; axial parenchyma is diffuse, diffuse-in-aggregates, narrow banded, rayadjacent, and vasicentric scanty; rays are basically Heterogeneous Type IIB, but in particular species
they approach or attain Heterogeneous III, Homogeneous I, Homogeneous III, Paedomorphic I, and
Paedomorphic III; crystal sand and rhomboidal crystals are present in fIbriform thin-walled idioblasts,
fiber-tracheids, axial parenchyma, and ray cells in a scattering of species. Solanaceae range into many
habitats, and degree of wood xeromorphy or mesomorphy sensitively correlates these with respect to
vessel diameter, vessel density, vessel element length, presence of vasicentric tracheids, and presence
of helical sculpture on vessels. There is in Solanaceae as well as in other families a continuum,
demonstrable with SEM, between the phenomena of grooves interconnecting pit apertures (confluent
pit apertures), grooves accompanied by thickening bands, and presence of helical thickenings (without
grooves) in vessels of Solanaceae; these conditions are included under the inclusive heading of helical
sculpture. Degree of grouping of vessels is proportional to xeromorphy except in Brunfelsia and
Fabiana, which have tracheids that deter vessel grouping as in other dicotyledonous woods. Wood of
scandent Solanaceae is distinctive and similar to that of other climbing dicotyledons. A predominance
of upright ray cells, tallness of multiseriate rays, and other features indicate paedomorphosis clearly
in wood of some Solanaceae (e.g., Datura meteloides, Solanum xantii), but the family appears to have
had a woody ancestry. Imperforate tracheary elements with living contents show minimal diameter
of pit cavities. A feature such as crystal sand presence links Solanaceae closely to the satellite families
Duckeodendraceae, Goetzeaceae, and Nolanaceae. Newly reported for the family are silica bodies in
rays (Acnistus) and bordered pits in sclerosed tyloses (Solanum gayanum).
INTRODUCTION
The family Solanaceae contains about 85 genera and 3600 species (D’Arcy
1979, 1986). The present study describes wood features of 21 genera and 82
species, which may seem like a small proportion of the family. However, several
reasons may be given for this representation. Many Solanaceae are herbaceous
and offer too little wood for study of wood anatomy; wood of herbs and nearherbs is essentially primary xylem or like primary xylem, and is not really com
parable with data obtained from woody cylinders of appreciable size (a few some
what woody herbs have nevertheless been included in this study for purposes of
‘Rancho Santa Ana Botanic Garden and Department of Biology, Pomona College, Claremont,
California 91711.
279
280
ALLERTONIA
6.4
comparison). Most xylaria contain material primarily of trees; Solanaceae are not
typically trees, and thus xylaria offer only a very small fraction of the family.
Furthermore, Solanaceae are most abundant in tropical areas and wood oftropical
genera has not been sampled so extensively (based on species number) as has that
of temperate genera. More significantly, identification of tropical Solanaceae—
particularly in large genera such as Solanum—is often difficult, so that not only
are wood collectors tempted to avoid some large genera, but specimens when
taken are sometimes not reliably identified to species, if they are identified to
species at all. If these various circumstances are considered, the proportion of
species included in the present study does not seem so scanty. In fact, samples
determined only to genus have not been included.
Despite the number of woody Solanaceae, there has been no previous survey
of wood anatomy in the family, other than the familial summaries of Solereder
(1908) and Metcalfe & Chalk (1950). Details on wood anatomy have been offered
for a few species (a single species in some papers listed) by Ahmad (1964), Baas
& Schweingruber(1987), Bonnemain(1970), Cariquist & Hoekman (1985), Cozzo
(1946), D’Arcy (1970), Descole & O’Donell (1937), Fahn et al. (1986), Gottwald
& Parameswaran (1964), Greguss (1959), Melville (1949), Norverto (1989), Tor
torelli (1940), and Williams (1936). Inamdar & Murthy (1977) and Murthy et al.
(1980) surveyed vessel elements in selected Solanaceae, but these studies are
mainly concerned with primary xylem, and their results are not really applicable
here. The present study is thus the first attempt to survey wood anatomy for the
family. The present effort must be termed a survey rather than a monograph, and
hopefully the inherent interest of solanaceous wood as revealed here will encourage
further collection of woods that can lead to monographs of wood of particular
solanaceous genera.
The inherent interest in wood anatomy of Solanaceae is considerable because
the family shows a wide range of features. Although perforation plates are char
acteristically simple in the family, a very few vestigial scalariform or aberrant
plates may be observed. Imperforate tracheary elements range from cells that
must be termed tracheids (according to the scheme of Bailey, 1936, or the IAWA
Committee on Nomenclature, 1964) to fiber-tracheids with vestigial borders on
pits; some species have vasicentric tracheids. Axial parenchyma is exclusively
diffuse in some species, exclusively vasicentric in others. Rays are predominantly
multiseriate in some species, predominantly uniseriate in others, and in histology,
ray cells vary from exclusively procumbent to exclusively upright. Crystal types
include rhomboidal crystals and crystal sand in axial parenchyma, ray cells, and
in fibriform idioblasts.
This wide range of wood features invites comparison to taxonomic systems.
Systems for subfamilial and tribal groupings reviewed by D’Arcy (1979) and
Hunziker (1979) offer frameworks for comparison to wood data.
The wide range of wood features in Solanaceae represents more numerous
instances of ecological patterning of wood than character state distributions that
primarily relate to taxonomic groupings. Certainly Solanaceae occupy a wide range
of habitats. The following classification of the species in the present study can be
offered, together with subsequent modifying comments:
1992
CARLQUIST: SOLANACEAE
281
Tropical trees or large shrubs: Brugmansia sanguinea, B. suaveolens, Cypho
mandra hartwegii, Solanum auriculatum, S. australe, S. erianthum, S. gran
diflorum, S. hayesii, S. hirtum, S. hispidum, S. leucocarpon, S. nigricans, S.
nudum, S. paludosum, S. rugosum, S. saponaceum, and S. triste.
Subtropical trees or large shrubs: Acnistus arborescens, A. grandiflorus, Nico
tiana otophora, N. raimondii, N. setchellii, N. tomentosa, Nothocestrum (all
species), Solanum acropterum, S. albidum, S. bahamense, S. kauaiense, S.
oblongifolium, S. sandwicense, and S. trichoneuron.
Temperate tree or large shrubs: Duboisia myoporoides.
Subtropical shrubs: Acnistus parviflorus, Brunfelsia calycina, B. nitida, Cestrum
(all species), Dunalia (both species), Iochroma tubulosa, Lycium sandwicense,
Nicotiana cordifolia, N. glauca, Solanum crispum, S. gayanum, and S. nel
sonii.
Temperate shrubs: Anthocercis littorea, Fabiana (all species), Grabowskya (all
species), Lycianthes lycioides, Lycium (all species except L. sandwicense),
Solanum nitidum, and S. simile.
Temperate subshrubs: Solanum douglasii, S. xantii.
Temperate herbs: Datura meteloides, Lycopersicon esculentum.
Subtropical climbers: Solandra guttata, Solanum appendiculatum, S. jasmi
noides, S. sodiroi, S. tetrapetalum, and Streptosolen jamesonii.
Within the above categories, different climatic regimes are represented. For
example, “subtropical” is applied to Andean shrubs that experience minimal frost,
despite latitude near the equator, as well as to shrubs from higher latitude but
lower elevation (e.g., Solanum bahamense). Some of the shrubs termed temperate
here are from tropical latitudes but at high elevation where frost is prevalent
(Solanum nitidum), whereas others are from extreme habitats such as the Atacama
Desert of Chile (Fabiana bryoides) or areas of Patagonia that are both dry and
cold (Fabiana viscosa, Grabowskya ameghinoi). Some of the species of Lycium
are from areas that are moderately dry (L. europaeum, shores ofthe Mediterranean
Sea), whereas others are from desert areas (L. brevipes, L. fremontii). Although
at first glance these seem like quite different kinds of habitats in terms of total
rainfall, the adaptations of wood in Mediterranean-type climates and desert areas
are very similar (Cariquist & Hoekman, 1985), a fact that reflects the necessity
of wood to maintain an intact water conducting system during a prolonged dry
season.
MATERIALS AND METHODS
TLE 1 indicates the sources of materials. Numerous samples were provided
by xylaria. These are supplemented by specimens I have obtained during field
work in California, Chile, Peru, and the Hawaiian Islands, and from botanic
gardens or other sites of cultivation. Herbarium vouchers for my collections are
located in the herbarium of the Rancho Santa Ana Botanic Garden (RsA). Her
barium voucher specimens for the other collections are specified in TABLE 1.
Where herbarium specimens of small shrubs provided woody stems sufficiently
large for study and comparable in diameter to those one might collect in the wild
282
ALLERTONIA
T.aLE 1.
WOOD CHARACTERISTICS OF SoLAcEAE.
TAx0N
Acnistus arborescens Dunal
A. grandiflorus Miers
A. parvifiorus Griseb.
Anthocercis littorea Ruiz & Pay.
Brugmansia sanguinea Ruiz & Pay.
B. suaveolens Humb. & Bonpl.
Brunfelsia calycina Benth.
B. nitida Benth.
Capsicum ciliatum (H.B.K.) Kuntze
Cestrum conglomeratum Ruiz & Pay.
C. diurnum L.
C. hirtum Sieber
C. mcscrophyllum Salzm. ex Dunal
C. nocturnum L.
C. parqui L’Héritier
C. pubens Griseb.
Cyphomandra hart wegii (Miers) Dunal
C. pendula Sendtn.
Datura mete/aides DC.
Duboisia myoporoides R. Br.
Dunalia arborescens (L.) Sleum.
D. obovata Dammer
Fabiana bryoides Phil.
F. imbricata Ruiz & Pay.
F. vjscosa Hook. & Arnott
Grabowskya ameghinoi Speg.
G. duplicatum Arnott
Iochroma tubu/osa Benth.
Lycianthes lycioides (L.) Hassi.
Lycium brevipes Benth.
L. carolinianum Walter
L. cestroides Schlechtend.
L. elongatum Miers
L. europaeum L.
L. fremontii A. Gray
L. sandwicense A. Gray
Lycopersicon esculentum Mill.
Nicotiana cordifolia Phil. (ROOT)
N. cordifolia (STEM)
6.4
COLLECrION & LOCALITY
SJRw-32009, Dominica
USw-36913, Dominica
PRFw-10526, Venezuela
MADw-10535, Argentina
MADw-20535, Argentina
Carlquist 968, Albany, Australia
cult. UCBBG
USw-6030, cult. Puerto Rico
cult. Vavra Estate, UCLA
Cariquist 15911, cult. Claremont, CA
Cariquist 15875, cult. Claremont, CA
Carlquist 7108, Carpish, Peru
Cariquist 7347a, Tiltil, Chile
USw-6000, Jamaica
USw-6035, Puerto Rico
PRFw-12830
USw-12818
cult. UCBBG
PRFw-1051 9. Argentina
Nee & Mon 3564 (i.iy), Costa Rica
Caniquist 15948, Claremont, CA
PRFw-24269, N.S.W., Australia
SFCw-R-950, N.S.W., Australia
SFCw-R-4182, N.S.W., Australia
Mexia 4001 (uC), Peru
Cariquist 7155, Tarma, Peru
Werdermann 352 (uC), Copiapo, Chile
Caniquist 7180, central Chile
Mexia 7839 (uC), Chubut, Argentina
Donat 37 (ucl, S. Argentina
PRFw-10560, Argentina
Car/quist 7347, Urubamba R., Peru
RSABG 14429, Imperial Co., CA
PRFw-24080, Florida
PRFw-10582, Argentina
PRFw-10520, Chile
PRFw-24676
Carlquist 7801, cult. RSABG
Carlquist 2376, Makapuu, Hawaii
cult. Lawai Valley, Hawaii
Skottsberg 18, Juan Fernandez Is.
Skottsberg 18 (ItsAw)
cult. UCBBG 60.095
Soibrig 3691 (oH), Juan Fernandez
N. glauca R. Graham
N.
N.
N.
N.
otophora Griseb.
raimondii Macbr.
setchellii Goodspeed
tomentosa Ruiz & Pay.
Cariquist s.n., Claremont, CA
Car/quist sn., Claremont, CA
Cariquist 7345, Pisac, Peru
cult. UCBBG 51.001
Cariquist 7345, Calca, Peru
Carlquist 15667, cult. UCBBG
Canlquist 7043, Tarma, Peru
UCBBG 36.001
Carlquist 7347, Calca, Peru
1992
Tii 1
1
2
3
4
5
VD
54
45
76
40
67
112
99
121
81
32
32
44
60
66
67
58
60
57
29
53
109
75
66
78
63
65
36
49
16
25
11
15
24
33
31
16
30
63
58
46
29
28
50
32
28
95
38
53
65
36
95
38
113
52
71
85
VM
56
36
59
73
59
38
30
13
20
147
185
64
35
60
43
74
37
59
110
100
5
9
52
56
53
65
102
68
433
200
367
252
650
57
105
331
110
52
121
54
620
104
82
81
44
39
43
77
81
130
23
63
21
40
28
22
VL
365
358
223
245
227
350
369
321
389
415
359
414
441
416
539
605
331
425
341
293
564
498
340
690
539
589
454
348
192
322
175
164
189
321
149
218
208
208
123
208
179
123
301
217
220
284
227
397
407
398
482
356
482
506
449
435
VG
1.8
2.3
2.3
2.3
2.9
1.9
1.8
1.6
1.5
1.1
1.1
2.3
1.7
1.6
1.8
1.4
1.5
2.1
1.2
2.0
1.5
1.4
1.6
7.6
8.1
5.3
1.1
2.1
1.1
1.1
1.1
VT
2.8
3.0
2.5
3.0
2.5
2.0
2.0
1.3
2.9
2.5
2.3
2.8
2.6
3.1
2.8
2.5
2.0
2.5
2.0
1.0
2.2
2.0
2.4
2.0
2.0
2.2
3.2
3.2
1.3
1.7
2.1
2.0
3.5
3.8
2.5
2.3
2.4
2.5
2.5
2.3
2.3
2.4
2.3
2.3
2.0
1.5
2.3
1.0
1.0
2.3
2.8
2.3
2.5
3.6
2.9
2.0
1.4
1.9
x
2.0
3.1
3.4
2.2
1.9
2.1
2.2
2.4
2.8
2.3
4.7
2.4
5.1
3.6
3.1
6
7
PD
5
5
7
5
5
5
12
12
12
5
4
8
8
5
10
6
8
5
6
8
10
8
7
7
6
6
4
5
5
7
5
4
5
5
5
6
5
5
5
6
5
5
6
5
5
8
5
6
6
6
5
7
7
6
7
7
TD
15
13
11
18
11
20
13
27
30
25
27
28
32
22
20
20
23
26
18
15
25
30
28
28
20
29
24
31
12
14
13
13
13
18
13
14
14
18
11
10
12
20
20
26
26
18
18
28
20
23
22
25
28
24
31
17
283
(Cor.rrIr.uED)
8
TL
704
890
572
518
643
889
724
351
403
976
727
724
976
737
936
1120
758
785
730
633
1234
123
605
1200
1000
1280
662
646
261
553
271
346
520
656
470
578
350
529
350
511
535
350
590
610
532
662
558
737
804
739
917
630
927
912
900
737
9
10
11
12
13
14
TF
2.5
2.4
2.5
2.0
2.2
2.5
2.5
1.5
2.6
3.9
4.1
3.1
3.2
2.9
2.8
4.1
2.1
2.1
2.2
1.5
1.8
2.1
2.6
2.5
2.5
2.5
3.2
3.3
2.0
3.1
2.6
2.1
2.0
2.5
3.1
2.4
1.5
2.5
2.3
2.3
2.5
1.0
2.0
2.4
2.7
2.3
2.1
1.0
1.0
2.5
2.0
2.1
2.3
2.1
2.2
1.5
TP
2
2
2
2
2
5
5
5
5
5
4
3
3
3
3
3
3
3
3
2
2
4
2
6
5
5
3
3
5
7
5
2
4
4
3
3
3
3
3
2
2
2
4
4
4
4
4
5
5
5
3
3
4
4
4
4
MR
362
280
614
UR
128
106
114
119
113
104
170
132
283
340
289
368
250
265
227
255
113
227
161
151
434
402
568
142
189
135
132
250
48
62
75
112
95
113
79
151
236
123
95
123
121
236
266
105
127
132
132
170
142
168
217
190
138
384
322
142
RH
usP
usP
usP
P
USP
USP
usP
USP
USp
usP
usP
USp
uSP
USP
USP
USp
usP
USp
USP
usP
Usp
USP
Us
sP
usP
sP
Usp
usP
Us
Usp
USp
sP
sP
usP
usP
USp
USp
USP
uSP
sP
sP
USP
USp
USP
usP
USp
USP
Usp
Usp
USp
USp
USp
USP
USp
USP
USP
MR
351
447
584
134
258
1032
1218
3238
1575
90
62
285
756
458
840
474
538
411
90
155
11599
4150
432
961
641
589
123
251
7
40
5
10
7
186
44
11
57
252
59
177
8
33
184
86
140
692
201
273
327
110
1990
215
2594
658
1138
1681
—
—
274
917
936
882
415
295
903
481
539
558
548
435
671
520
671
1526
608
1060
331
378
365
488
581
—
141
—
131
123
246
247
239
—
170
123
217
369
236
893
265
193
246
236
643
463
406
378
292
—
483
410
255
284
ALLERTONIA
TABLE 1.
TAx0N
Nothocestrum breviflorum A. Gray
N. tat folium A. Gray
N. longif’olium A. Gray
Solandra guttata D. Don
Solanum accrescens Standi. & C. Morton
S. acropterum Griseb.
S. albidum Dunal
S. appendiculatum Humb. & Bonpi.
S. auriculatum Aiton
S. australe C. Morton
S. bahamense L.
S. chrysotrichum Schiectend.
S. crispum Ruiz & Pay.
S. douglasii Dunal in DC.
S. erianthum D. Don
S. gayanum Phil.
S. grandiflorum Ruiz & Pay.
S. hayesii Fernald
S. hirtum Vahi
S. hispidum Pers.
S. jasminoides Paxt.
S. kauaiense Hillebr.
S. leucocarpon Dunal
S. nelsonii Dunal
S.
S.
S.
S.
S.
S.
nigricans M. Martens & Galeotti
nitidum Ruiz & Pay. (TRUNK)
nitidum (BRANCH)
nudum Dunal
oblongifolium Humb. & Bonpi.
paludosuni Dunal
S. rugosum Dunal
S. sandwicense Duanl
S. saponaceum Dunal
S. simile F. v. Muell.
S. sodiroi Bitter
S. tetrapetalum Rusby
S. torvum Sw.
S. trichoneuron Lillo
S. triste Jacq.
S. xantii A. Gray
Streptosolen jamesonii Miers
6.4
(Cor.mNuED)
C0LLECrI0N &
LOCALITY
Cariquist 2087, Puuwaawaa, Hawaii
Cariquist 1944, E. Maui, Hawaii
USw-26014, Kokee, Kauai, Hawaii
Nee & Mon 4040 (I.4Y), Venezuela
USw-6002, Jamaica
Carlquist 7066, San Ramon, Peru
Anderson 479 (us), Mexico
PRFw-12277
SFCw-R-4313
PRFw-24224, Bahamas
Nee & Illis 16695 (MA.r), Mexico
Carlquist 7160, Tiltil, Chile
Carlquist 7271, Cutipay, Chile
Canlquist 15907, S. California
PRFw-12792, India
SFCw-.R-166-9, India
SFCw-R-977-1 75, India
USw-8466, Chile
Carlquist 7143, Tingo Maria, Peru
Canlquist 7388, Iquitos, Peru
Nee & Mon 3667 (NY), Colon, Panama
Nee & Mon 4131 (Ny), Mérida, Venezuela
Carlquist 15867, cult. Pomona, CA
USw-15288, Kokee, Kauai, Hawaii
PRFw-16 150
Cariquist 2333, Pearl & Hermes Reef Hawaii
SFCw-R.-988-11, Hawaiian Islands
Nee & Taylor 28900 (Ny), Mexico
Cariquist 7079, C. de Pasco, Peru
Caniquist 7079, C. de Pasco, Peru
Nee & Mon 4154 (Ny), Mérida, Venezuela
cult. UCBBG
PRFw-1 7925, Guayana
PRFw-16104
USw-1840, Dominican Republic
USw-1844, Dominican Republic
USw-6032, Puerto Rico
PRFw-17376, Hawaiian Islands
Carlquist 7045, Acobamba, Peru
SFCw-R-936-324, Australia
Anderson 715 (us), Colombia
Anderson 749 (us), Colombia
USw-6033, Indonesia
PRFw- 10508, Argentina
PRFw-10558
Wo(f 3496 (RSA), S. California
Key to columns: 1, VD, mean vessel diameter, tim; 2, VM, mean number vessels per mm
; 3, VL,
2
mean vessel element length, m; 4, VG, mean number of vessels per group; 5, VT, mean vessel wall
thickness, sm; 6, PD, mean diameter of lateral wall pit of vessel (if circular), jsm; 7, TD, mean
imperforate tracheary element diameter at widest point, tim; 8, TL, mean imperforate tracheary element
length, tim; 9, TT, mean imperforate tracheary element wall thickness, tim; 10, TP, mean diameter
of pit cavity on imperforate tracheary elements, tim; 11, MR mean multiseriate ray height, m; 12,
UR, mean umseriate ray height, m; 13, RH, ray histology (U = upright; S = square; P = procumbent;
upper case indicates predominant or common types); 14, MR, Mesomorphy ratio (vessel diameter
times vessel element length divided by vessels per mm
).
2
1992
TLE 1
1
80
73
87
120
63
38
100
71
107
106
64
38
25
49
103
87
110
35
81
177
161
99
71
81
29
59
103
83
71
80
25
20
63
79
123
101
114
94
80
86
79
109
44
31
45
123
84
95
35
41
2
30
22
9
12
61
40
17
115
14
16
56
14
304
37
21
26
23
220
80
11
8
14
33
18
206
30
35
59
73
34
436
795
19
65
17
20
19
17
14
13
11
11
69
468
155
17
28
22
236
80
3
274
284
303
264
320
236
408
246
427
340
284
431
222
361
321
298
392
280
321
477
495
344
304
461
224
270
501
208
231
364
268
251
479
350
444
402
397
416
375
397
227
359
260
274
289
388
303
340
210
408
4
3.5
3.5
1.9
4.6
2.8
2.1
1.8
4.9
1.9
1.3
2.1
1.6
6.0
1.9
1.8
2.2
1.6
2.5
1.9
1.4
1.2
2.5
1.6
1.5
3.2
3.2
2.2
2.3
2.2
1.7
7.9
18.6
1.7
3.3
1.9
1.5
1.9
1.6
1.5
2.2
1.0
1.3
3.6
::
8.9
1.6
2.2
1.5
1.4
5
5.5
5.1
5.5
2.5
3.7
2.0
2.6
3.4
3.9
2.2
2.3
2.4
1.8
2.2
3.0
2.6
3.2
3.0
1.8
4.3
2.8
3.2
2.8
2.8
2.9
2.5
3.5
3.1
3.4
3.1
2.0
1.8
3.2
3.2
3.0
2.8
2.1
2.1
2.5
3.1
2.7
2.2
2.9
2.4
2.9
3.0
2.8
3.5
2.4
2.4
6
8
7
9
12
9
5
10
7
10
10
7
9
8
7
5
6
8
5
12
12
12
12
6
10
7
7
3
5
6
4
8
8
5
10
6
6
8
8
7
8
4
10
6
7
10
10
3
7
5
7
7
14
18
16
15
14
14
21
30
32
16
17
26
18
28
13
23
24
19
18
28
30
25
20
28
22
25
22
12
17
22
16
16
24
14
18
17
17
17
14
12
23
25
20
15
23
18
18
17
22
30
285
(CoNTu’uED)
8
624
615
770
606
706
624
779
556
983
794
643
813
567
672
737
651
828
480
700
1210
1130
733
536
857
461
502
1010
539
559
779
493
600
891
766
813
799
756
832
804
823
766
759
695
505
491
737
709
737
489
758
9
2.5
2.5
3.0
2.9
1.8
2.3
2.0
2.9
2.2
2.3
3.8
2.1
2.2
0.8
1.8
2.0
2.1
2.3
2.5
2.0
1.8
2.2
2.2
2.2
2.5
3.8
2.0
2.3
2.3
2.4
1.7
1.7
2.1
2.3
0.7
0.8
0.6
0.7
1.2
0.8
2.9
0.8
2.3
2.3
2.2
2.3
2.5
2.3
2.2
2.7
10
11
5
4
5
4
2
3
2
2
2
3
3
3
2
3
3
3
3
2
5
3
3
2
2
3
2
5
3
3
2
3
3
3
4
4
4
4
3
3
2
2
5
4
2
2
3
4
3
4
2
5
265
227
274
463
155
284
820
1000
248
227
444
339
620
498
189
167
312
399
662
312
248
224
261
350
3140
236
407
199
345
258
389
342
363
374
170
199
274
180
189
217
378
259
339
1500
12
142
142
189
255
99
170
448
215
91
76
161
164
168
236
68
95
90
238
132
119
68
144
178
171
140
72
199
113
168
125
110
130
233
198
142
132
113
104
90
123
79
123
103
120
331
208 180
284 161
227 142
408 109
737 208
—
13
14
sP
sP
sP
USp
usP
Us
sP
USP
sP
usP
usP
usP
usP
usP
usP
usP
usP
Usp
usP
P
P
usP
USP
usP
USP
USP
USP
USP
USP
USP
sP
USP
USP
usP
sP
sP
sP
usP
usP
usP
USP
usP
usP
Us
Us
usP
USp
USP
731
942
2929
2640
330
224
2561
152
3263
2075
325
1170
18
478
1574
997
1875
45
325
7675
9962
2433
654
2075
32
531
1474
293
225
856
15
6
1588
425
3212
2030
2382
3008
1765
2626
1630
3557
166
18
84
2807
909
1468
31
209
Us
USp
(e.g., Fabiana bryoides, F. viscosa, and Grabowskya ameghinoi), portions removed
from herbarium specimens were used. Wood samples were mostly available in
dried form. Liquid-preserved specimens were studied for some species (Datura
meteloides, Lycium brevipes, L. fremontii, Nicotiana glauca, Solanum douglasii,
S. jasminoides, and S. xantii). Liquid preservation of woods is desirable because
286
ALLERTONIA
6.4
living (nucleated) fibers are probably characteristic of many species in the family,
judging from the sampling listed by Wolkinger (1970).
Most woods were sectioned on a sliding microtome. As some Solanaceae have
rather soft woods, vessels in these often break during sectioning. Vessels also tend
to break in species with numerous large vessels, such as the climbing species
Solanum appendiculatum, S. sodiroi, and S. tetrapetalum. For woods that pro
vided these problems, an alternative technique involving softening in ethylene
diamine, embedding in paraffin, and sectioning on a rotary microtome (Cariquist,
1982) was employed. In most instances, counterstaining with fast green was em
ployed in order to clarify pit diameter in fiber-tracheids and cell contents for those
species for which liquid-preserved material was available. Starch was readily
discernible in liquid-preserved material. In other specimens, starch remnants
could be identified.
TABLE 1 lists quantitative characters (as well as ray histology, column 13) of
the species studied. Most figures in TABLE 1 represent means derived from 25
measurements. For vessel wall thickness, diameter of pits on lateral walls of
vessels, imperforate tracheary element diameter, imperforate tracheary element
wall thickness, and pit cavity diameter on imperforate tracheary elements, means
could not be readily determined because of difficulty in sampling when a feature
is highly variable; for these features, a few representative measurements were
averaged. Vessel diameter was measured as lumen diameter at widest point.
Number of vessels per group was calculated on the basis that a solitary vessel =
1, a pair of vessels = 2, etc. (and a mean was derived from 25 measurements).
Terms follow the recommendations of the IAWA Committee on Nomenclature
(1964) except for the usages ofthe terms vasicentric tracheid and vascular tracheid,
which follow usages presented in my review of these topics (Carlquist, 1 985a).
The usage of the term “vasicentric” follows that of Kribs (1935). The term “pae
domorphosis” follows my use of the term in an earlier paper (Carlquist, 1962),
and is not to be confused with ‘juvenile”, which may refer to earlier-formed
wood in stems of dicotyledons at large, including those that do not show pae
domorphic wood characters. In TAJ3LE 1, vessel diameter of earlywood has not
been calculated separately from latewood. In most species, the difference between
the two is not extreme; although in strongly ring-porous species earlywood vessels
may embolize while latewood vessels do not (suggesting a reason for separate
calculation), the less extreme differences between earlywood and latewood in
Solanaceae probably do not show differential embolisms, so a mean based on all
vessels is functionally more accurate. Range in vessel diameter is not functionally
meaningful, and has not been given. The narrowest vessels cannot be distinguished
from vasicentric tracheids (or tracheids) with certainty, and an unusually large
vessel in a particular section is not significant. The term “angular” in connection
with vessel outline follows the usage of Frost (1930), which has phylogenetic
significance; the somewhat angular vessels of some Solanaceae do not qualify as
angular under his definition. Borders on ray cells have been overlooked by various
workers seeking to find them in face view; sectional views of the ray cell wall are
more reliable and have been utilized here to decide if ray cell wall pits are bordered
or not. Grooves interconnecting pit apertures, paired bands beside pit apertures,
1992
287
and helical thickenings represent three types of helical sculpture but with inter
mediate expressions; studies with SEM (e.g. Cariquist, 1992) in Lamiaceae and
Asteraceae have demonstrated this.
ANATOMICAL DESCRIPTIONS
GROWTH RINGs
Temperate to subtropical Solanaceae have growth rings demarcated by appre
(FIGuns 1, 23, 32, 36, 38). Growth rings of
this sort qualify as ring porous and were observed in Acnistus grandijiorus (FIGuRE
1), Fabiana bryoides, F. viscosa (FIGURE 23), Grabowskya duplicatum, Lycianthes
lcioides (FIGu 32), Lycium cestroides, L. elongatum (FIGUIu 36), L. europaeum
(FIGURE 38), and L. fremontii. In the remaining Solanaceae, growth rings are
demarcated by either slightly wider earlywood vessels or radially narrower fiber
tracheids in latewood or both (FIGUREs 9, 50, 66, 73). Demarcation of earlywood
from latewood is so gradual in some of these instances that one may equally well
choose to call them diffuse porous or semi-ring porous. Even in humid tropical
areas, a slight degree of seasonality in rainfall can correspond to formation of a
weakly demarcated growth ring. Transections in which no growth rings are ap
parent in portions illustrated (and in which little growth ring demarcation occurs
elsewhere in the species listed) include Anthocercis littorea (FIGuRE 4), Acnistus
parvflorus (FIGu 5), Cestrum diurnum (FIGU1 23), Nicotiana cordifolia (FIGU
41), N. raimondii (FIGURE 46), Solanum acropterum (FIGURE 56), S. gayanum
(FIGURE 61), and S. oblongifolium (FIGuRE 70).
ciably larger vessels in earlywood
VEssEL ELEMENTS
Solanaceae characteristically have simple perforation plates. In Cestrum diur
num, a few perforation plates with irregular thread-like bars traversing the plate
can be seen (FIGURES 15, 16). These can be compared more to a pit-like than to
a scalariform pattern on account of the reticulate conformation of the wall strands.
In Solanum sandwicense a very few scalariform perforation plates were observed;
the remainder of perforation plates were simple. Aberrant perforation plates were
reported for Solanaceae by Gottwald and Parameswaran (1964). Occasionally two
perforation plates on an end wall of a vessel element could be observed in a few
of the Solanaceae studied. This condition has been reported for a few Solanaceae
by Inamdar & Murthy (1977) and Murthy et al. (1980). These authors also
report occasional instances of three or even four perforation plates on an end wall.
Occurrence of pairs of perforation plates on the end walls of a vessel is to be
expected occasionally with relation to forking of a vessel, but the occurrence of
three or four perforation plates on an end is unlikely to have this explanation.
Vessel elements of Solanaceae are uniformly rounded (rather than angular) in
transectional view, even when narrow (FIGU1s 1, 5, 9, 13, 23, 32, 34, 35, 36,
38, 41, 46, 47, 50, 56, 58, 61, 66, 70, 73, 78). The term “angular” here conforms
with the usage of Frost (1930), and refers to vessels like those of Cercidiphyllum,
which are much more angular in transectional view than those of Solanaceae.
288
ALLERTONIA
6.4
Vessels are wider radially than tangentially in Cestrum diurnum (FIGURE 13) and
other species.
Mean vessel diameter is given in TABLE 1, column 1. One can readily see that
in some species, mean diameter exceeds 100 m: Anthocercis littorea (FIGu 4),
Solandra guttata (FIGURE 28), and Solanum paludosum (FIGURE 72), for example.
One must remember that in some ring porous Solanaceae, earlywood vessels
exceeding 100 m in diameter may be common (FIGuREs 1, 32, 38), but narrow
latewood vessels are much more numerous in these species, so that mean diameter
well below 100 m is registered. Likewise, scandent species of Solanaceac, such
as a S. appendiculatum (FIGURE 58), commonly have vessels that exceed 100 tm
in diameter, but in these species, narrow vessels (many of which are little if at all
wider than imperforate tracheary elements) are quite common and are intermixed
with the wide vessels. This accounts for relatively low mean vessel diameter in
Solanum appendiculatum, S. jasminoides, S. sodiroi, S. tetrapetalum, and Strep
tosolen jamesonii.
Vessel density is commonly thought to be approximately inversely proportional
to vessel diameter in dicotyledonous woods, but this is not true because few
dicotyledonous woods closely approach packing limits for vessels of a given mean
diameter. Thus, in the figures for vessels per mm
2 of transection (TABLE 1, column
2), one notices figures that vary inversely with vessel diameter (column 1), but
not at all closely. Lack of correlation is most notable in the scandent species listed
at the end of the preceding paragraph; this is also true in dicotyledons in general
(Cariquist, 1 985b). One can see in the photomicrographs that the vessels in the
scandent Solanum appendiculatum (FIGURE 58) are much more dense than those
in the arborescent S. paludosum (FIGURE 73). The presence of narrow vessels, not
so conspicuous as large vessels, in the wood of scandent species such as S. ap
pendiculatum makes the vessel density much greater than that of S. paludosum,
in which narrow vessels are scarce. In fact, the contrast between such a pair of
species may be more marked than the figures in TABLE 1 or the photomicrographs
indicate, because very narrow vessels as seen in transection are often disregarded
because they are not readily distinguished from imperforate tracheary elements
of the same diameter. Solandra guttata is not genuinely a vine: it can be described,
rather, as a sprawling or ascending shrub.
Very low vessel densities (fewer than 20 per mm
) characterize the tropical or
2
subtropical wet forest trees Brugmansia suaveolens, Cyphomandra hartwegii, C.
pendula, Nothocestrum longifolium, Solanum grandiflorum, S. paludosum (FIGUlu
73), S. rugosum, S. sandwicense, S. saponaceum, and S. torvum. Notably high
vessel densities (more than 200 per mm
) occur in species belonging to more than
2
one ecological or habit category. One finds high vessel densities in woody herbs
(Solanum douglasii, S. xantii), shrubs ofvery dry or desert areas (Fabiana bryoides,
Grabowskya ameghinoi [FIou 28], Lycium brevipes, and L. fremontii), shrubs
of cold alpine areas (Solanum nitidum), and shrubs of seasonally dry areas such
as Fabiana imbricata, F. viscosa (FIGURE 23), and Grabowskya duplicatum. Two
climbing species (S. jasminoides, S. sodiroi) are reported here to exceed 200 vessels
per mm
. Very likely, if one could accurately count very narrow vessels in scandent
2
species, one could add other scandent Solanaceae to this list.
1992
289
Vessel element length is shown for TABLE 1, column 3. Metcalfe & Chalk (1950,
p. 1360) claim that the mean vessel element length for dicotyledons at large is
649 tm. Only one collection of one species in the family, Duboisia myoporoides,
rises to that level. One must keep in mind that the Metcalfe & Chalk sample is
undoubtedly biased in favor of wet forest trees, which tend to bulk larger in wood
sample and wood slide collections. When data are drawn from areas where dry
habitats predominate, results are quite different. For example, in the southern
California flora, the mean vessel element length for the 207 woody species studied
is 233 im; projected to all 512 woody species in the flora the mean would be 225
m (Carlquist & Hoekman, 1985). By this standard, the mean vessel element
length for all Solanaceae studied here, 348 tim, is relatively long. If we use this
figure by which to compare vessel element lengths within the family, we find
notably short vessel elements in Acnistus grandflorus, Cestrum pubens, Fabiana
bryoides, F. viscosa, Grabowskya ameghinoi, G. duplicatum, Lycium (all species),
Lycopersicon esculentum, Nicotiana cordifolia, Nothocestrum (all species), Solan
dra guttata, Solanum acropterum, S. appendiculatum, S. chrysotrichum, S. cris
pum, S. jasminoides, S. nelsonii, S. simile, S. sodiroi, S. tetrapetalum, and S.
xantii. Nearly all of these could be described as shrubs (particularly ofdry climates)
or climbers.
Vessel grouping in Solanaceae shows an extraordinary range: from species with
vessels solitary or nearly so to species in which vessels are aggregated into groups
of indefinite extent. Predominantly solitary vessels are illustrated here for Fabiana
imbricata (FIGuRE 20), F. viscosa (FIGURE 23), and Solanum paludosum (FIGuRE
73). Vessel grouping of 1.2 vessels per group or less characterizes the genera
Brunfelsia and Fabiana, as well as a scattering of other species: Cestrum noc
turnum, Dunalia arborescens, Solanum grandiflorum, and Solanum sandwicense.
The significance of this will be examined at the end of the Ecological Correlations
section of this paper.
Larger vessel groupings may be defined as more than 2.5 vessels per group.
Using this criterion, all species of Grabowskya, Lycium (except L. sandwicense),
and Nothocestrum, as well as Acnistus parviflorus, Lycopersicon esculentum, Ni
cotiana cordifolia (root), N. otophora, N. tomentosa, and 11 species of Solanum
have larger vessel groupings. However, the vessel groupings in Grabowskya (FIGuu
28) and Lycium (FIGURES 34, 36, 38) are more extensive. Such vessel groupings,
which extend across rays frequently, as seen in transection, were designated “vessel
aggregations” in an earlier paper (Carlquist, 19 87a). Vessel aggregations are present
also in latewood of Acnistus australis (FIGU 1, center). Vessel groupings in
Solanaceae usually take the form of radial multiples, as in Anthocercis littorea
(FIGURE 4), A. parviflorus (FIGuRE 5), Nicotiana cordifolia (FIGURE 41), N. raE
mondii (FIGURES 46, 47), Nothocestrum latifolium (FIGURE 50), Solanum acrop
terum (FIGURE 56), S. oblongifolium (FIGURE 70), and Solandra guttata (FIGURE
78).
Vessel wall thickness within Solanaceae, as shown in TABLE 1, column 5, ranges
from 1.0 m (Nicotiana glauca) to 5.5 tm (Nothocestrum spp.); the latter genus
has vessel walls characteristically thicker than those seen in other genera. The
range just indicated cannot express the variations seen within a single wood
290
ALLERTONIA
6.4
section. Thinner-walled vessels are mostly latewood vessels; the large earlywood
vessels in Solanaceae characteristically have thick walls. The tendency for wider
vessels to be thicker-walled is clearly shown throughout the genus Solanum, in
which most species are diffuse porous, and in which the correlation can be shown
on the basis of means, as indicated by comparing the figures for columns 1 and
5 in TABLE 1 for Solanum species. Estimating vessel wall thickness in a wood is
very difficult because within a single vessel, wall thickness varies so greatly. For
example, if one sees a pair of adjacent vessels in a transection, the walls where
the vessels are in contact are appreciably thicker than the wall portions not in
contact.
Mean diameter of vessel-to-vessel pits (where isodiametric; elliptical pits are
excluded intentionally) is shown in TABLE 1, column 6. This range is unexpectedly
wide. The median vessel-to-vessel pit cavity diameter is about 6—7 m in Sola
naceae; the genera Brunfelsia, Dunalia, Fabiana, Grabowskya, and Lycium (FIGu1u
40) have pits smaller than that. Larger pits are evident in Solandra and in various
species of Solanum, such as S. hayesii (FIGuRE 59), S. gayanum (FIGuRE 65), and
S. kauaiense (FIGURE 68). However, certain species of Solanum have notably
small vessel-to-vessel pits: for example, pits are about 3 m in diameter in S.
leucocarpon and S. sandwicense. Vessel-to-vessel pits are typically round in outline
in Solanaceae (FIGu1us 65, 75, 76), but pits polygonal in outline are occasionally
encountered, as in S. kauaiense (FIGURE 68). Vessel-to-vessel pits are typically a
little wider laterally than vertically. For example, if pits when circular are specified
as 6 tm in diameter, pits in that species will mostly be about 6 m vertically by
7 tm horizontally, i.e., elliptic.
Vessel-to-ray and vessel-to-axial parenchyma pits tend to be similar in mor
phology but slightly larger in pit cavity diameter in comparison to vessel-to-vessel
pits. In some species, vessel-to-ray pits are characteristically elongate; Nicotiana
tomentosa, for example, has vessel-to-ray pits that should be termed pseudosca
lariform (like scalariform pits in general appearance, but derived from an alternate
pattern by lateral widening). Vestigial borders are present on the ray side of vesselray pit pairs. Apertures of pits are narrowly elliptical on vessel-vessel pits, and
somewhat wider in vessel-ray pits (on the vessel side).
The vessels of Solanaceae illustrate why helical “sculpture” must include both
“grooves” (“coalescent pit apertures” of some authors) and thickenings, as recently
advocated (Carlquist, 1 988a). In Solanaceae, a continuum between these phe
nomena exists; such a continuum can be confirmed with SEM, as in the vessels
I have studied of such Lamiaceae as Poliomintha (Carlquist, 1992). Workers who
have not worked with the tubifiorous families of dicotyledons, which show this
continuum clearly when one looks for it, may be unfamiliar with the structures
discussed here. Grooves interconnecting pit apertures, shown here for Nothoces
trum latifolium (FIGURE 52), were observed in Acnistus grandorus, Anthocercis
littorea, Cestrum nocturnum, Cyphomandra pendula, Lycium sandwicense, Noth-.
ocestrum (all species), Solanum auriculatum, S. chrysotrichum, S. crispum, S.
douglasii, S. erianthum, S. grandiflorum (short grooves), S. hayesii, S. kauaiense,
S. leucocarpon (short grooves), S. nigricans, S. nudum, S. oblongifolium, S. sand
wicense, S. saponaceum, S. sodiroi (short grooves), and S. triste. Grooves accom
panied on their margins by inconspicuous bands (a thickening on either side of
1992
291
a groove) were observed in Anthocercis littorea, Cestrum pubens (FIGURE 17),
Solanum rugosum (FIGUREs 75, 76), and S. xantii. Grooves plus pronounced
helical thickenings paired along the grooves were observed in Grabowskya (both
species), Lycium brevipes, L. carolinianum, L. cestroides, L. elongaturn, L. eu
ropaeum (FIGURE 40), Solanum crispum, S. hayesii (FIGuRE 59), and S. jasmi
noides (narrow vessels). Prominent helical thickenings, without any evident grooves,
were observed in Fabiana bryoides, F. imbricata, F. viscosa (FIGURES 26, 27),
Lyciumfremontii, Solanum crispum (Cariquist 7160), 5. gayanum (FIGuRE 65),
and S. nitidum.
VAsIcENTRIc T1&cHEIDs
Vasicentric tracheids were reported earlier in Grabowskya duplicatum, Lycium
brevipes, L. cestroides, L. elongatum, L. europaeum, and L. fremontii (Cariquist,
1985a) and for three species of Lycium by Baas and Schweingruber (1987). At
the time of my (1 985a) review, I had not yet examined material of scandent
Solanaceae. Vasicentric tracheids occur in the climbing species Solanum appen
diculatum, S. jasminoides, S. sodiroi, and S. tetrapetalum, intermixed with narrow
vessel elements. Presence of vasicentric tracheids in a greater proportion of climb
ing woody dicotyledons than had hitherto been appreciated was noted in a review
of wood of scandent dicotyledons (Carlquist, 1 985b). Vasicentric tracheids are
also newly reported here for nonclimbing Solanaceae not examined at the time
of the 198 5a survey: Grabowskya ameghinoi, Solanum douglasii, S. nitidum, and
S. xantii. Vasicentric tracheids are scarce in the wood of S. douglasii. Vasicentric
tracheids have helical thickenings in those species in which vessels bear such
thickenings: Grabowskya (both species), Lycium (all species except L. sandwi
cense), and Solanum nitidum.
VAscuL TItcHEIDs
Vascular tracheids may be defined as tracheids occurring at the end of a growth
ring in a wood that otherwise contains fiber-tracheids or libriform fibers; the
vascular tracheids may be considered the last-formed vessel elements in the latewood, so narrow that they lack perforation plates (Carlquist, 1 988a). Vascular
tracheids thus do not sheathe vessels, as vasicentric tracheids do. Vascular tra
cheids present as a layer one or two tracheids thick at the end of latewood were
observed in the wood of Solanum erianthum (PRFw- 12792).
F1BER-TicHjDs AND TRUE TIcHEir.s
Following the definition of Bailey (1936) and the IAWA Committee on No
menclature (1964), Solanaceae must be claimed to have fiber-tracheids with the
exception ofBrunfelsia and Fabiana. These two genera must be regarded as having
tracheids on the basis of several considerations mentioned below. I prefer to call
these tracheids true tracheids, a term required where a contrast with vasicentric
tracheids is required (Cariquist, 1 988a).
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Mean diameter of fiber-tracheids (or tracheids) at widest point is given in TABLE
1, column 7. Interestingly, there is not a high degree of correlation between vessel
diameter and the diameter of the imperforate tracheary elements in any given
wood. Notably wide fiber-tracheids (28 tm or wider mean diameter at widest
point) characterize Brugmansia suaveolens, Capsicum ciliatum, Cestrum con
glomeratum, Duboisia myoporoides, Nicotiana glauca, N. setchellii, Solanum ap
pendiculatum, S. auriculatum, S. grandiflorum, S. hispidum, and Streptosolen
jamesonii. Most of these have vessel elements less than 80 tm in diameter.
Nicotiana cordifolia (FIGuRE 41) has fiber-tracheids the mean diameter of which
is nearly as great as that of its vessel elements. Although little attention has been
paid to diameter of imperforate tracheary elements in dicotyledons, instances like
the above deserve investigation.
Mean length of imperforate tracheary elements of Solanaceae is given in TABLE
1, column 8. In most dicotyledonous woods, imperforate tracheary element length
parallels vessel element length, and Solanaceae also show this trend. Metcalfe &
Chalk (1950, p. 1361) compute the mean imperforate tracheary element length
of dicotyledons as a whole as 1317 m. All of the Solanaceae studied here have
mean imperforate tracheary element lengths shorter than that figure. If one divides
mean imperforate tracheary element length in Solanaceae (724 tm) by mean vessel
element length in the family (348 Lm), one obtains a ratio of 2.08. Metcalfe &
Chalk (1950, p. 1301) report mean “fibre length” (imperforate tracheary element
length) in dicotyledons as a whole to be 1317 m and mean vessel element length
to be 649 tim. Thus the ratio between lengths of the two cell types (termed “F/V
ratio” by some authors) is 2.03, so Solanaceae are close to the ratio figure for
dicotyledons as a whole, despite having much shorter vessel elements and im
perforate tracheary elements. The “F/V ratio” was considered to be an index of
phyletic advancement within certain limits (Carlquist, 1975).
Mean wall thickness of imperforate tracheary elements is given in TABLE 1,
column 9. This figure as obtained here does not represent the thickest point in
the wall (e.g., the angles of a cell), but rather a point midway between angles, and
thus a rather minimal thickness of the wall. The range of figures obtained for
Solanaceae in TABLE 1 is from 0.6 tm in Solanum rugosum to 4.1 m in Brunfelsia
nitida and Cestrum macrophyllum. A high proportion of Solanaceae have im
perforate tracheary element wall thickness less than 2.5 m; this accounts for the
softness of many solanaceous woods. The fact that woods of Brunfelsia and Duna
ha are harder in texture can be attributed to the greater wall thickness of imper
forate tracheary elements in these two genera.
Pit diameter in imperforate tracheary elements is given for the species of So
lanaceae in TABLE 1, column 10. Pit diameter is given to the closest integer because
often there is a range ofvalues (e.g., 1.5 tm to 2.5 nm), often in a single imperforate
tracheary element. Pit cavity diameter is notably small in Lycium (2.5—3 am); it
is also small in many species of Solanum, such as S. nigricans. Even in imperforate
tracheary elements with small pits, vestigial borders can be observed.
Solanaceae were singled out as a family in which some genera have tracheids,
whereas others have fiber-tracheids (Cariquist, 1 988a). Fabiana was cited as hav
ing tracheids, but Brunfelsia can be added also. Both Fabiana and Brunfelsia have
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293
bordered pits maximal in diameter for the family on imperforate tracheary ele
ments. More significantly, the pits are much more densely placed than in some
other species of Solanaceae in which pit diameter is as great as it is in Brunfelsia
or Fabiana (FIGURE 25). Another interesting test of whether tracheids are present
or not is whether grouping of vessels is deterred (Cariquist, 1984). This phenom
enon is represented in Solanaceae: only in Brunfelsia and Fabiana does the number
of vessels per group approach 1.0 (TABLE 1, column 4), despite the fact that the
species of Fabiana grow in dry situations where vessel grouping would be expected
(in woods in which tracheids are not present). One species of Solanum, S. sand
wicense, also meets pit diameter, pit density, and vessel grouping criteria for
possession of tracheids. Pit diameter greater that 5 tm is present in imperforate
tracheary elements of Brugmansia, but pits are not as densely placed as in the
tracheids of the examples cited above, and Brugmansia must be said to have
fiber-tracheids. Although Baas (1986) has been skeptical of this distinction, it
continues to prove feasible and of significance with respect to wood physiology
in my opinion (for a discussion of this question, see Carlquist, 1988a, pp. 104—
107). Helical thickenings are characteristic of conducting cells in wood, and in
this connection, it is interesting that they occur in vasicentric tracheids of some
Solanaceae (see above) and in true tracheids of Fabiana viscosa (FIGu1u 25; FIGURE
26, far right), but never in fiber-tracheids in Solanaceae.
Fiber-tracheids of Solanaceae typically have more pits on radial than on tan
gential walls, as tends to be true in dicotyledons at large. A radial wall is shown
for Solanum nigricans in FIGURE 77. The radial walls of solanaceous fiber-tra
cheids that face ray cells are more densely pitted than walls that face other fiber
tracheids. Although this phenomenon may be common in dicotyledons, it is not
often reported in literature on wood anatomy
Living fibers have been reported in Solanaceae by Wolkinger (1970) in Cestrum
elegans Schlectendahl, C. nocturnum, C. purpureum (Lindl.) Standl., Solanum
convolvulus Sendtner, S. dulcamara L., and Streptosolen jamesonii. Liquid-pre
served material in the present study revealed nuclei in fiber-tracheids of Datura
meteloides, Lycium brevipes, L. fremontii, Nicotiana glauca, Solanum douglasii,
S. jasminoides, and S. xantii. Starch, indicative of the living nature of wood cells,
was observed in fiber-tracheids of species made from dried wood samples in the
case of Acnistus grandiflorus, Brugmansia sanguinea, Capsicum ciliatum, Cestrum
conglomeratum, and C. pubens. Undoubtedly, starch is more common in fiber
tracheids of Solanaceae than these scattered observations based on dried material
(which is less favorable than liquid-preserved material) would indicate. Another
indicator of the presence of protoplasts of protracted longevity in fiber-tracheids
is the occurrence of septa. Septate fibers were recorded in Acnistus arborescens,
A. grandiflorus (FIGURE 3), A. parvijiorus (fiber-tracheids more than once septate),
Dunalia obovata, Solanum hayesii, and S. jasminoides.
FIBRw0RM CRYsTAL-BEAJUNG IDIOBLASTS
Septate fibriform cells that contain crystals were observed in Solanaceae. These
septate idioblasts have thin primary walls, but are of the same length as fiber-
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ALLERTONIA
6.4
tracheids. These are illustrated here for Solanum hayesii (FIGURE 60). These
idioblasts are definitely fibriform (prosenchymatous) in shape and are nonseptate
(unlike axial parenchyma, which occurs in strands). Nonseptate crystal sand id
ioblasts of the same sort are illustrated here for Grabowskya ameghinoi (FIGUREs
30, 31), and were also observed in Brugmansia sanguinea, B. suaveolens, Lycium
europaeum, L. fremontij (crystal sand intermixed with rhomboidal crystals), L.
sandwicense, Solandra guttata, Solanum albidum, S. grandiflorum (FIGURE 81),
and S. torvum (FIGURE 82). Fibriform crystal sand idioblasts have been reported
for Lycium elongatum Miers (Norverto, 1989), and crystal sand was reported for
three species of Lycium by Fahn et al. (1986). Thin-walled septate fibriform
idioblasts containing numerous septa and one crystal per subdivision (chambered
crystals) were observed in material ofAcnistusparvzflorus (FIGURE 7). Small rhom
boidal crystals in septate fibriform idioblasts were observed in Acnistus arbores
cens. In Grabowskya duplicatum (FIGURE 22), fibriform crystal-bearing idioblasts,
each of which bears one large crystal, are conspicuous (FIGURE 22). Axial paren
chyma has morphology and distributions in Solanaceae that differ from the dis
tributions of the fibriform crystal-bearing idioblasts.
AxiAl, PARENcHYii
Axial parenchyma is diverse in Solanaceae: it shows a greater range than that
in many families of dicotyledons. Diffuse axial parenchyma was observed in
Brunfelsia (both species), Cyphomandra pendula (diffuse mainly, with a little
vasicentric scanty and some ray-adjacent parenchyma), Duboisia myoporoides
(diffuse plus diffuse-in-aggregates plus diffuse-in-clusters in PRFw-24269; diffuse
plus ray-adjacent in SFCw-R-950; diffuse only, moderately scarce, in SFCw-R
4182), Fabiana bryoides (sparse), F. imbricata (moderately sparse), F. viscosa
(moderately common), Grabowskya ameghinoi (FIGURE 30), G. duplicatum (abun
dant), Lycianthes lycioides (diffuse plus diffuse-in-aggregates), Lycium brevipes
(diffuse, diffuse-in-aggregates, wide-banded), L. cestroides (abundant diffuse plus
diffuse-in-aggregates), L. elongatum (diffuse plus diffuse-in-aggregates), L. fre
montii (diffuse, a little diffuse-in-aggregates, initial), L. sandwicense (abundant
diffuse plus diffuse-in-aggregates), Nicotiana cordifolia, N. glauca, N. otophora,
N. raimondii (FIGURE 47), N. setchellii (abundant), N. tomentosa, Solandra guttata
(diffuse plus wide bands), and Solanum acropterum (scarce). In Nicotiana cordi
folia and N. tomentosa (Cariquist 7043) there are what appear to be wide bands
of parenchyma; in longitudinal section these cells do prove to have apparently
simple pits and are wide and thin-walled like axial parenchyma cells, but they
are not subdivided as is axial parenchyma elsewhere in Solanaceae.
Axial parenchyma so scarce as to be effectively absent was recorded for Cap
sicum ciliatum, Cestrum conglomeratum, C. diurnum, C. nocturnum, C. parqui,
and Lycopersicon esculentum. The other species of Cestrum (C. macrophyllum,
C. pubens) proved to have very small amounts of vasicentric scanty parenchyma.
Vasicentric scanty parenchyma was observed in Acnistus (all collections), Brug
mansia (all collections), Cyphomandra hartwegii (FIGURE 19), Datura meteloides,
Dunalia (both species), Iochroma tubulosa, all species of Solanum except S. ac
ropterum, and Streptosolen jamesonii. In Nothocestrum, axial parenchyma is
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295
scanty vasicentric plus banded (FIGu1s 50, 53). The bands are 1—5 cells wide
(FIGURE 50), most commonly 2—3 cells wide (FIGuRE 53). Scanty vasicentric
parenchyma in Solanaceae may be only a single cell adjacent to a vessel, as in
Cestrum pubens. More commonly, 1—2 layers of cells surrounding half to threequarters of a vessel’s circumference are present, as in Solanum kauaiense (FIGURE
66). Complete sheaths are common in Solanum triste. As viewed in longisection,
the axial parenchyma of Solanaceae occurs in strands of two to five cells, most
commonly two to three.
Crystal sand was observed in axial parenchyma of Duboisia myoporoides,
Nothocestrum latifolium (FIGuRE 54), and Solandra guttata. Small rhomboidal
crystals were observed in axial parenchyma of Nicotiana otophora (FIGuRE 44).
Starch was noted in axial parenchyma of Grabowskya ameghinoi (FIGURE 30),
Lycianthes lycioides, Lycium brevipes, L. elongatum, L. europaeum, Nicotiana
cordifolia, N. raimondii, Solanum albidum, and S. nudum.
RAYS
Solanaceae have great diversity in ray structure. Both multiseriate and uniseriate
rays are present. Even within a small genus such as Acnistus, extremes are evident.
In Acnistus grandiflorus, multiseriate rays predominate, although uniseriate rays
are also present (FIGuRE 2). In A. parviflorus, rays are almost exclusively uniseriate
(FIGuRE 6). Within the genus Solanum, multiseriate rays predominate in S. cris
pum, S. erianthum, S. grandijiorum, S. saponaceum, S. simile, and S. xantii.
Uniseriate rays predominate in S. acropterum (FIGuRE 57), 5. hirtum, S. nudum,
S. paludosum (FIGuRE 74), S. rugosum, S. sodiroi, S. torvum, S. trichoneuron,
and S. triste. Multiseriate rays are about as frequent as uniseriate rays in the
remaining species, such as S. kauaiense (FIGURE 67). In Fabiana, uniseriate rays
are present almost exclusively (FIGu1 24), as they are also in Grabowskya (FIGu1s
21, 29). Uniseriate rays predominate in Lycium (FIGURES 33, 37, and 39). In
Nicotiana, both types may be equally abundant, as in N. cordifolia (FIGURE 42),
but in most species of the genus, uniseriate rays predominate.
Mean height of multiseriate rays is given for Solanaceae in TABLE 1, column
11. Multiseriate rays average from one to two times the length of vessel elements
in most collections. Species in which multiseriate rays are more than twice the
length of the vessel elements include Acnistus grandiflorus, Brugmansia (all col
lections), Capsicum ciliatum, Cestrum pubens, Datura meteloides, Lycopersicon
esculentum, Solanum appendiculatum, S. jasminoides, S. sodiroi, and S. tetra
petalum. Multisenate rays that average appreciably less than the length of vessel
elements can be cited for Anthocercis littorea, Brunfelsia nitida, Cestrum noc
turnum, Duboisia myoporoides, Fabiana viscosa (multiseriate rays are virtually
absent in other species of Fabiana), Grabowskya (both species), Iochroma tubu
losa, Lycicum cestroides, L. elongatum, Nicotiana otophora, Nothocestrum (all
species), Solanum accrescens, S. auriculatum, S. australe, S. chrysotrichum, S.
erianthum, S. grandiflorum, S. hayesii, S. hirtum, S. nigricans, S. nudum, S.
paludosum, S. rugosum, S. saponaceum, S. torvum, S. trichoneuron, and S. triste.
The two groups of species, based on whether multiseriate rays exceed vessel
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ALLERTONIA
6.4
elements in length or are shorter than vessel elements, prove to be significant in
that they mostly differ with respect to ray histology, as indicated later in this
section.
Mean height of uniseriate rays is shown in TABLE 1, column 12. The mean
height of umseriate rays is less than half the height of multiseriate rays in most
species of Solanaceae. Exceptions occur in species in which multiseriate rays are
few and narrow (often biseriate), such as Grabowskya duplicatum (FIGURE 21),
Lycium elongatum (FIGui 37), L. europaeum (FIGu 39), Nothocestrum lati
folium (FIGURE 51), Solanum acropterum (FIGuRE 57), and S. paludosum (FIGuRE
74). Thus the more closely multiseriate rays tend to resemble uniseriate rays, the
more similar they are to each other in dimensions.
Ray histology is summarized with a few symbols in TABLE 1, column 13. The
rays in Solanaceae are most commonly Heterogeneous Type IIB of Kribs (1935);
in this type, upright cells occur in some (or all) uniseriate rays, in tip (“marginal”)
cells of multiseriate rays (cells at upper and lower edges of multiseriate rays), and
as occasional sheathing cells (cells along the sides of multiseriate rays); uniseriate
wings are absent on multiseriate rays. Such rays would be summarized with the
formula usP in TABLE 1, column 13, because the majority of ray cells (most cells
in the multiseriate portions of multiserate rays) would be procumbent. Rays of
this type are illustrated for Lycianthes lycioides (FIGURE 33), Nicotiana cordifolia
(FIGURE 42), and Solanum kauaiense (FIGURE 67).
Rays nearly all uniseriate, but heterocellular, occur in some species of Lycium,
such as L. elongatum (FIGURE 37). These fall into Heterogeneous Type III of Kribs
(1935), or are transitional between Heterogeneous Type IIB and Heterogeneous
Type III.
Rays in which upright cells are about as abundant as procumbent cells probably
still qualify as Heterogeneous Type IIB, although they may be said to indicate a
degree of paedomorphosis in the presence of more than a minimum of upright
cells. Examples of these rays, indicated by the formula USP in TABLE 1, column
13, are illustrated here by Cestrum diurnum (FIGURE 14).
Rays both multiseriate and uniseriate with a high proportion of upright cells
and in which procumbent cells are scarce or lacking (e.g., the formula Us or U
in TABLE 1, column 13) are illustrated here by Datura meteloides (FIGURES 11,
12), Nicotiana raimondii (FIGURE 48), and Solanum acropterum (FIGURE 57).
Other species in which this type was recorded include Nicotiana cordifolia (So/brig
3691), N. g/auca, Solanum douglasii, S. simile, S. sodiroi, S. xantii, and St rep
tosolen jamesonii. These correspond to Paedomorphic Type I (Carlquist, 1 988a),
or are very close to that type, transitional from Heterogeneous Type IIB. Note
should be taken that these species are mostly the same as those listed above as
having muhiseriate rays twice as tall or taller than the uniseriate rays. One could
ascribe the paedomorphic condition of rays in the species listed above to im
maturity of the stems. However, small stem sizes were studied only for Datura
meteloides, Solanum douglasii, and S. xantii, which never become large plants.
The wood samples for the other species were at least as great in woody cylinder
thickness as the species for which rays showed no juvenile features. Thus, I believe
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297
the species listed above have genuinely paedomorphic woods, with the possible
exception of the three cited for smaller stem sizes.
Some species have rays all uniseriate (or with few multiseriate rays) and ray
cells upright or mostly so. Included in this category are Fabiana bryoides, F.
imbricata, F. viscosa (FIGURE 24), Lycium brevipes, L. carolinianum, and L. ces
troides. These rays correspond to Paedomorphic Type III (Carlquist, 1988a).
The reverse tendency, rays with cells almost exclusively procumbent, occurs in
a scattering of Solanaceae. Those with this tendency, and with rays both multi
seriate and uniseriate, include Acnistus arborescens, A. grandijiorus (FIGURE 2),
A. parviflorus, Brunfelsia (both species), Duboisia myoporoides, Solanum nitidum
(trunk), Solanum oblongifolium (FIGURES 71, 72), S. paludosum (FIGURE 74), and
S. rugosum. Acnistus grandiflorus is close to Homogeneous Type II (cells all
procumbent, uniseriate rays lacking), whereas the remainder in the preceding
sentence are transitional between Heterogeneous Type IIB and Homogeneous
Type I. Predominance of procumbent cells in species with few multiseriate rays
is shown by Grabowskya ameghinoi (FIGu 29), G. duplicatum (FIGURE 21), and
Lycium europaeum (FIGu 39). Species in this category are closer to Homoge
neous Type III than to any other type; the few square (or even upright) cells in
rays of these species indicate only a small degree of intermediacy in the direction
of Heterogeneous Type JIB. The list of species with rays containing a predomi
nance of procumbent cells proves to be nearly the same as the list of species in
which multiseriate rays are less than twice as tall as uniseriate rays.
Ray cells contain crystals in a few Solanaceae. Crystal sand was observed in
ray cells with thin primary walls, intermixed as idioblasts among ordinary ray
cells with lignified secondary walls, in Duboisia myoporoides, Nicotiana cordifolia
(FIGu1 43), N. tomentosa, Nothocestrum latifolium (FIGu1u 55), N. longifolium,
Solanum bahamense, S. leucocarpon, S. nelsonii (both collections), and S. sodiroi.
Crystal sand plus rhomboidal crystals was observed in ray cells of Solanum tn
choneuron (FIGURE 80). Some ray cells of S. trichoneuron contain only rhomboidal
crystals (FIGuRE 79). Rhomboidal crystals were also observed in ray cells of S.
sandwicense. Small rhomboidal crystals occur in ray cells of Cestrum pubens
(FIGURE 18), Nicotiana otophora (FIGuiE 45), Solanum jasminoides, and S. ob
longifolium.
Silica bodies were observed in ray cells ofAcnistus arborescens and A. parviflorus
(FIGURE 8). The identity of these bodies is based on their lack of birefrigence with
polarized light, their shape, and their pale violet color when slightly out of focus.
Silica bodies have not hitherto been reported for Solanaceae (Carlquist, 1 988a).
Starch was observed in ray cells of Capsicum ciliatum, Cestrum pubens, Cy
phomandra pendula, Lycianthes lycioides, Lycium elongatum, L. europaeum, L.
sandwicense, Nicotiana cordifolia, N. raimondii, Solanum albidum, S. chrysotri
chum, S. erianthum, S. hayesii, S. oblongifolium, and Streptosolen jamesonii.
Undoubtedly more species than listed here contain starch in ray cells; starch is
degraded in drying and processing of wood for sectioning, and in some of the
specimens listed above, starch grains were present as partially dissolved or de
graded masses.
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ALLERTONIA
6.4
Trabeculae in ray cells were observed in Nicotiana raimondii (FIGURE 49).
Ray cells have moderately thin but lignified walls in Solanaceae (FIGuREs 8,
12, 18, 31, 45, 48, 49, 55, 59, 69, 79, 80, and 82). Pits on the tangentially oriented
walls of ray cells of Solanaceae are denser than those on the horizontally and
radially oriented walls. The pits on the tangential walls are very frequently bor
dered throughout Solanaceae. In fact, most of the pits on horizontally and radially
oriented walls of Solanaceae are inconspicuously bordered also. In only a few
species with relatively thin ray cell walls, such as Nicotiana glauca and Solanum
auriculatum, were nonbordered ray cell pits characteristically present; borders
increase in their overarching of the pit cavity with increasing wall thickness, so
this is understandable. Bordered pits on ray cells are illustrated here in FIGuREs
18, 45, 49, 69, 72, and 79.
AMORPHOUS DEPosrrs IN WOOD
Dark-staining amorphous deposits in parenchyma cells are common in woods
of some Solanaceae, although notably absent in some genera (e.g., Cestrum). Darkstaining deposits are illustrated here for Acnistus parvifiorus (FIGu1s 5, 6, and
8), Grabowskya duplicatum (FIGuREs 21,22), Lycium brevipes (FIGURE 35), Notho
cestrum latifolium (FIGuRES 50, 51, and 53), and Solanum kauaiense (FIGulus
67, 68, and 69). In the transection of Acnistus parviflorus wood (FIGu 5), the
dark cells are not all parenchyma cells; the deposits have spread into fiber-tracheids
as well. In the section of Lycium brevipes (FIGURE 35), the dark-staining com
pounds are in the form of small droplets, individually inconspicuous, that make
the parenchyma cells that contain them slightly darker gray than neighboring cells
in the FIGURE 35 photomicrograph.
CRYsTAIs: SUMMARY
Crystals occur in thin-walled fibriform idioblasts, axial parenchyma, and ray
cells in wood of Solanaceae. Crystals are more abundant in plant portions other
than wood in Solanaceae. As an example of this, I observed crystal sand frequently
in phloem parenchyma of Solanaceae (in species in which bark portions were
adherent to wood sections), although crystal sand was observed in wood cells of
only a small proportion of the Solanaceae studied.
Rhomboidal crystals were observed in septate fibriform idioblasts in Acnistus
arboreus (small crystals) and A. parviflorus (large crystals, chambered by numerous
septa, FIGuRE 7). Exceptionally large single rhomboidal crystals occur in fibriform
idioblasts of Grabowskya duplicatum (FIGURE 22). Rhomboidal crystals (mixed
with crystal sand) occur in thin-walled fibriform idioblasts of Lycium fremontii.
Rhomboidal crystals occur in axial parenchyma in Nicotiana otophora (FIGuRE
44). Rhomboidal crystals were observed in ray cells of Cestrum pubens (FIGURE
18), Nicotiana otophora (FIGURE 45), Solanum oblongifolium, S. sandwicense, and
S. trichoneuron (FIGus 79, 80).
Crystal sand in thin-walled fibriform idioblasts occurs in Brugmansia sangui
flea, B. suaveolens, Grabowskya ameghinoi (FIGus 30, 31), G. duplicatum, Lyci
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urn europaeurn, L. frernontii (mixed with rhomboidal crystals), L. sandwicense,
Solanurn albidurn, S. grandiflorurn, S. hayesii (septate idioblasts, FIGu 60), and
S. torvurn (FIGURE 82). Crystal sand in axial parenchyma occurs in Duboisia
myoporoides, Nothocestrurn latifoliurn (FIGuRE 54), and Solandra guttata. Crystal
sand in thin-walled ray cells was observed in Duboisia myoporoides, Nicotiana
cordifolia, N. tornentosa, Nothocestrurn breviflorurn, N. latifolium (FIGuRE 55),
N. longifoliurn, Solanurn baharnense, S. grandiflorurn (FIGuRE 81), S. leucocarpon,
S. nelsonii, S. sodiroi, S. torvurn (FIGuRE 82), and S. trichoneuron (FIGuRE 80:
rhomboidal crystals also present).
TYLOsEs
Thin-walled tyloses were observed in vessels in Acnistus parviflorus, Cyphornan
dra hartwegii (FIGURE 19), Lycopersicon esculenturn, Nicotiana otophora, Notho
cestrurn latifoliurn, Solanum gayanurn, S. nitidurn (trunk), and S. sodiroi. Thinwalled tyloses were reported by Ahmad (1964) in Solanurn indicurn. Sclerosed
tyloses were observed in Cestrurn macrophyllum and in Solanurn gayanurn
(FIGURES 62, 63, and 64). The scierosed tyloses of S. gayanurn have only mod
erately thick walls (FIGuRES 62, 64), but the pits are uniformly bordered (FIGURES
63, 64). The occurrence of bordered pits in scierosed tyloses elsewhere in dicot
yledons must be very rare; I am not acquainted with any prior reports.
CAMmAL, VARIANTS
Only normal cambial activity was observed in the species studied here. The
report of Bonnemain (1970) on “included phloem” in woods of Solanaceae could
not be confirmed on the basis of my material.
CONCLUSIONS
EcoLocucAL CONCLUSIONS
In dealing with such a large group as Solanaceae, ecological generalizations are
difficult to present, especially when a large proportion of the samples have been
obtained from xylarium collections, for which locality data and ecological infor
mation are often minimal. More significantly, Solanaceae often grow in somewhat
disturbed habitats, such as road cuts or cultivated fields, so that a particular plant
may be growing in a habitat appreciably drier or wetter than would be expected
in a particular locality. If we use the Mesomorphy ratio (TABLE 1, column 14),
an arbitrary index devised earlier (Carlquist, 1977), ecological status of habitats
occupied by particular woods can be estimated. Mesomorphy values lower than
50 generally indicate dry habitats (rainfall 50 cm or less per year), whereas mod
erately mesic habitats would tend to have woods with values above 800. These
levels are suggested on the basis of two earlier studies (Carlquist, 1981; Carlquist
& Hoekman, 1985), in which wide ranges of ecological sites were represented. If
we look at the Mesomorphy values for the Solanaceae studied, we find values
below 50 in Fabiana bryoides (M = 7), which grows in the Atacama Desert of
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ALLERTONIA
6.4
Chile; F. viscosa (M = 5), in dry and cold areas of Patagonia; Grabowskya amegh
inoi (M = 10), from Patagonia; and G. duplicatum (M = 7), from cool dry areas
of central Argentina. Lycium is interesting, because values parallel habitat closely:
L. brevipes (M = 11) is from the Colorado (Sonora) Desert of California and L.
fremontii (M = 7) is a Mojave Desert species, whereas L. carolinianum (M = 57)
is from dry but humid areas of Florida and L. sandwicense (M = 33) is from
similarly exposed (but humid) dry coastal lowlands of easternmost Oahu. In
Nothocestrum, the wet forest species N. longifolium (M = 2929) has wood much
more mesomorphic than wood ofthe species from dry (but humid) lowland forests,
N. breviflorum (M = 731) and N. latifolium (M = 942). Dryland species of Notho
cestrum are at least partially drought deciduous, moderating the selective effect
of environment on wood anatomy. Notably low Mesomorphy values in the genus
Solanum may be found in S. crispum (M = 18), from the Chilean matorral, as
well as Californian species from similar chaparral habitats: S. douglasii (M = 45)
and S. xantii (M = 31). The high montane Peruvian shrub Solanum nitidum
reflects the dryness and coldness of its alpine habitat in its wood Mesomorphy
values (trunk, M = 15; branch, M = 6).
One might not have expected that scandent species of Solanum would also
have relatively low Mesomorphy values: S. appendiculatum (M = 152), S. jas
minoides (M = 32), S. sodiroi (M = 18), and S. tetrapetalum (M = 84). Scandent
plants in general tend to have wide vessels (see Carlquist, 198 Sb), which woods
of plants in dry habitats usually do not, but scandent plants also tend to have
numerous narrow vessels intermixed with the large vessels (Carlquist, 198 Sb),
accounting for the low Mesomorphy values of the abovementioned species. So
landra guttata can be called a climbing or sprawling subtropical shrub rather than
a true liana, so its relatively high Mesomorphy value (2640) is not unexpected;
Solandra species are generally intolerant of drought.
All of the species in the present study with Mesomorphy values in excess of
2000 qualify as rain forest or cloud forest trees or else plants provided with
abundant water from cultivation. Tropical rain forest habitats are occupied by
species with notably high values: Cyphomandra hartwegii, from Puntarenas, Costa
Rica, and Solanum grandj1orum, from Iquitos in the Amazonian rain forest of
Peru.
The presence of true tracheids is clearly correlated with ability to withstand
drought in Fabiana. The presence of true tracheids, sometimes but certainly not
always indicative of xeromorphy in dicotyledons, may seem surprising in Brun
felsia in view of the chiefly tropical and subtropical distribution of this genus
(mostly Antilles and northern South America: Plowman, 1979). Some of the areas
inhabited by Brunfelsia are less than mesic, however.
Vasicentric tracheids are a clear indicator of drought resistance. The two genera
all species of which have vasicentric tracheids, Grabowskya and Lycium, are from
areas that are seasonally very dry (e.g., southern Europe) or dry for prolonged
periods, such as the deserts of the southwestern U.S. The species of Solanum from
the most extreme habitat represented by materials of the genus here, S. nitidum
(Cerro de Pasco, ca. 4800 m elevation), has vasicentric tracheids. To be sure,
vasicentric tracheids are present in moderate numbers in scandent species of
1992
301
Solanum, but woody vines and lianas tend frequently to have moderate numbers
of vasicentric tracheids, perhaps as a safety device (Cariquist, 1 985b).
Grouping of vessels is a feature that is indicative of wood xeromorphy (Carl
quist, 1984). However, presence of tracheids has been shown to deter grouping
of vessels (Carlquist, 1984), and this accounts for the very low number of vessels
per group in Brunfelsia and Fabiana. Extremely large numbers of vessels per
group, indicated by the co sign in TABLE 1, column 4, characterize the genera
(Grabowskya, Lycium) and species (Solanum xantii) from rather extreme habitats;
the high alpine Solanum nitidum also has very high numbers of vessels per group.
HABIT AND
WooD
ANATOMY
In the above discussion, the relationship between ecology and vessel features
(in the Mesomorphy ratio) implies that ecology is of overriding importance. That
does appear to be true, although the vining habit is an exception. If quantitative
data for Solanaceae (other than Solandra guttata) are compared, one sees that the
scandent species provide a distinctive pattern. The means for the five scandent
species as a group are as follows (with means for Solanaceae as a whole in paren
theses): vessel diameter, 41 m (43 Lm), vessels per mm
, 204 (80), vessel element
2
length, 288 m (408 am), and Mesomorphy ratio, 99 (1080). Interestingly, the
mean vessel diameter for the vines is close to that of Solanaceae as a whole; vessel
diameter of the scandent species may in fact be even somewhat lower, because
very narrow vessels as seen in transection are often counted as imperforate tra
cheary elements. Number of vessels per mm
2 is notably high; vessel density in
the scandent Solanaceae is probably not comparable to that of woody lianas,
which have few but very wide vessels, analyzed earlier (Carlquist, 1975, p. 206).
Indeed, the climbing Solanaceae are not strongly woody. Vessel element length
is appreciably less for the vines than for Solanaceae as a whole, confirming the
trend from the liana sample of dicotyledons (Carlquist, 1975, p. 206). By having
vessel diameter similar to that for Solanaceae as a whole, but a much greater
vessel density, the climbing Solanaceae have a conductive area per unit area of
transection about 2.5 times that of Solanaceae as a whole.
Vessel features of shrubby or arboreal Solanaceae could be calculated separately
from those of the climbing species, but have not been, because the range from
small shrub to tree is so continuous. However, selection of species of shrubs shows
that they have narrower vessels, more numerous per mm
, than do trees. This is
2
not surprising, because one finds this in shrubs and trees in other samplings
(Carlquist, 1975, p. 206; Baas et al., 1983; Cariquist & Hoekman, 1985). One
can also say that wood of shrubs in general tends to be more xeromorphic than
that of trees.
Paedomorphosis in dicotyledonous woods can be recognized in terms of a
predominance ofupright cells in rays (Carlquist, 1962). On the basis ofthis feature,
the following woods show paedomorphosis: Brugmansia suaveolens, Capsicum
ciliatum, Cyphomandra hartwegii, Datura meteloides, Dunalia arborescens, Fa
biana bi’yoides, F. imbricata, F. viscosa, Lycium brevipes, L. carolinianum, Lyco
persicon esculentum, Nicotiana glauca, N. otophora, Solandra guttata, Solanum
302
ALLERTONIA
6.4
accrescens, S. crispum, S. simile, S. sodiroi, S. xantii, and Streptosolen jamesonii.
The species of Capsicum, Cyphomandra, and Nicotiana in this list may represent
instances of secondary woodiness, despite the apparent woodiness of the family
as a whole, because their woods are paedomorphic although woody cylinder
development is considerable (ca. 10 cm in diam.). Datura meteloides, Lycopersicon
esculentum, and Solanum xantii qualify as woody “herbs” and are very likely
woodier than their ancestors; however, these are relatively small plants, and more
paedomorphic characters are to be expected, whereas in the other species cited
above, wood sample size is just as large as for woods that showed no paedomorphic
features. The species cited above as having paedomorphic rays also have multi
senate rays twice as long as vessel elements in those respective species, except for
Fabiana and Lycium. Paedomorphic woods tend to have tall multiseniate rays
(Carlquist, 1 988a). Fabiana and Lycium cannot be expected to have multiseniate
rays twice as tall as vessel elements because in those genera, multiseriate rays are
scarce or absent (where present, they are usually biseriate and otherwise like
uniseriate rays).
Fabiana and Lycium may represent a special category with respect to paedo
morphosis. Certain small, woody shrubs that tend to have a finite size and that
have rays uniseriate (or nearly so) constitute a category of plants that demonstrate
paedomorphosis in wood, as defined by a decrease in vessel element length as the
stem increases in size (Carlquist, 1989). Fabiana and Lycium agree with the criteria
cited for Bruniaceae, Empetraceae, Myrothamnaceae, and Tremandraceae.
SYsTE&TIc CoNcLusIoNs
Most of the genera studied here are represented by only a fraction of their
species. Therefore, this study must be considered a survey of woods in the family,
not a monograph. A few genera are represented here by species so distinctive in
their wood features as to merit discussion with respect to specific criteria. Thus
the comments on systematic implications of wood anatomy must be limited to
a limited number of situations that merit comment in view of the limitations of
the present study. To go beyond the implications of the data at hand would be
unwarranted. For this reason, a data summary arranged according to systematic
groupings would be premature.
The three species ofAcnistus are remarkably diverse. Acnistus grandiflorus wood
is ring-porous (FIGURE 1), but wood is diffuse-porous or nearly so in A. arborescens
and A. parvWorus (FIGuRE 5). Fiber-tracheids are septate in Acnistus, but with
various contents: small rhomboidal crystals in A. arborescens, starch but no crys
tals in A. grandiflorus, and large crystals separated by septa (chambered crystals)
in A. parvorus (FIGuRE 7). Vasicentnic tracheids are present only in A. parviflorus.
Rays are predominantly multiseriate in A. arborescens and A. grandiflorus but
almost exclusively unisenate in A. parviflorus (FIGURE 6). Silica bodies are present
in A. arborescens and A. parvflorus (FIGURE 8). Doubtless as any genus of Sola
naceae becomes better known with respect to its wood anatomy, distinctive wood
features at the species level will emerge.
Solanaceae have been subdivided into two subfamilies, Solanoideae and Ces
1992
303
troideae, by several authors (see D’Arcy, 1979 for a history of these concepts).
The feature in the present study that corresponds most closely to this division is
parenchyma type. The genera employed in the present study fall into the two
subfamilies as listed below. Parenchyma distributions are given in terms of these
subfamilies.
Subfamily Solanoideae: Acnistus, Brugmansia, Capsicum, Cyphomandra, Da
tura, Dunalia, Grabowskya, Iochroma, Lycianthes, Lycium, Lycopersicon,
Nothocestrum, Solandra, and Solanum. All of these have scanty vasicentric
axial parenchyma except for Capsicum (little or no axial parenchyma), Gra
bowskya (diffuse), Lycianthes (diffuse, diffuse-in-aggregates), Lycium (diffuse,
diffuse-in-aggregates), Lycopersicon (diffuse, diffuse-in-aggregates), and So
landra (diffuse plus wide bands).
Subfamily Cestroideae: Anthocercis, Brunfelsia, Cestrum, Duboisia, Fabiana,
Nicotiana, and Streptosolen. All ofthese have diffuse axial parenchyma except
for Cestrum (axial parenchyma absent or very sparse vasicentric) and Strep
tosolen (scanty vasicentric).
The parenchyma distributions tend to reinforce the division into two subfamilies.
The exceptions to the predominant parenchyma type in the two subfamilies are
not necessarily to be regarded as indicative that the subfamilies should be recon
structed. Rather, the exceptions may represent parallel evolution (or possibly
reversals). Diffuse parenchyma is the more primitive type in dicotyledons, ac
cording to the data of Kribs (1937). Indeed, the two species of Cyphomandra
differ with respect to axial parenchyma (scanty vasicentric in C. hart wegii, diffuse
plus scanty vasicentric and ray-adjacent in C. pendula). The specimens on which
these studies were done seem likely to be correctly determined. All species of
Solanum were recorded as having vasicentric axial parenchyma except for S.
acropterum; this instance should be re-examined on the basis of authenticated
material. Interesting with regard to the phylogenetic position of the two subfam
ilies is that although diffuse parenchyma would mark Cestroideae as more prim
itive, evidence from floral anatomy (Armstrong, 1986) is cited in support of the
sequence of D’Arcy (1979), in which Solanoideae are placed in a basal position
in the family.
If there is difficulty in using wood features to aid generic or subfamilial concepts,
support of the tribes with wood data is even more tenuous at present. If one looks
at the assignment of the genera in the present study to tribes, such as those
recognized by Hunziker (1979), one notes that only one or two genera per tribe
have been studied here, and thus any basis for tribal distinctions by means of
wood anatomy is insufficient. An exception is Lycieae: Grabowskya and Lycium
are well represented here, although the third genus (Phrodus, with two species) is
not. Grabowskya and Lycium have well marked growth rings, vessels intermixed
with vascular tracheids in large aggregations, diffuse axial parenchyma, and crystal
sand in thin-walled fibriform idioblasts (some of these features in only some of
the species of both genera, but all features are present in both genera). Armstrong
(1986), using data from floral anatomy, also stresses the distinctiveness of the
Lycieae. In the tribe Nicotianeae, Fabiana with its distinctive tracheids and uni
304
ALLERTONIA
6.4
•‘‘i 11%
: ‘!k .J4
:
.
0
tji
FIGuRES 1—4. Wood sections ofAcnistus andAnthocercis. 1—3. Acnistus grandiflorus (PRFw-10525).
1. Transection; wide earlywood vessels, above. 2. Tangential section; wide multiseriate rays predom
inate. 3. Fiber-tracheids from radial section, showing septa. 4. Anthocercis littorea (Carlquist 968),
transection; vessels in radial multiples. FIGuREs 1, 2, and 4, scale above FIGuRE 1 (divisions = 10
gm); FiGuna 3, scale above FIGuaa 3 (divisions = 10 tim).
1992
305
FIGuRES 5—8. Wood sections of Acnislus parvorus (USw-4193). 5. Transection; dark-staining
deposits visible in many cells. 6. Tangential section; rays are uniseriate. 7. Portion of radial section
to show chambered crystals (left) and septate fibers. 8. Portion of radial section showing ray cells (long
axis of ray oriented vertically) containing silica bodies. FlouRas 5, 6, scale above FIGuRE 1; FiGuRas
7, 8, scale above FIGuRE 3.
306
ALLERTONIA
6.4
FIGURES 9—12. Wood sections of Capsicum and Datura. 9—10. Capsicum ciliatum (Carlquist 7108).
9. Transection; growth rings are indistinct. 10. Tangential section; most ray cells are upright. 11—12.
Datura meteloides (Cariquist 15848). 11. Tangential section; tall ray cells are upright. 12. Radial
section; ray cells are markedly upright. FIGuREs 9—11, scale above FIGu1a 1; FIGuaa 12, scale above
FIGURE 12 (divisions = 10 nm).
1992
a;
307
F?
FIGu1s 13—18. Wood sections of Cestrum. 13—16. C. diurnum (cult. Vavra estate, UCLA) 13.
Transection, showing thin-walled nature of fiber-tracheids. 14. Tangential section; rays are uniseriate
or biseriate. 15—16. Perforation plates from radial section, showing strands of wall material traversing
the plates. 15. Arcuate strands. 16. Slender strands forming network around polygonal area. 17—18.
C. pubens (FPRw-10519). 17. Vessel from tangential section, showing inconspicuous thickenings. 18.
Ray cells from radial section, showing sparse small rhomboidal crystals. FIGuIs 13, 14, scale above
FIGuRE 1; Fiour.as 15—18, scale above FIGuRE 3.
308
ALLERTONIA
6.4
FIGuiis 19—22. Wood sections of Cyphomandra, Fabiana, and Grabowskya. 19. Cyphomandra
hartwegii (Nee & Mon 3564); transection, showing tyloses in vessel and thin-walled liber-tracheids.
20. Fabiana imbnicata (Cariquist 7180), transection; narrower vessels and narrower tracheids delimit
growth rings; vessels are solitary. 21—22. Grabowskya duplicatum (PRFw-10560). 21. Tangential sec
tion; rays are uniseriate. 22. Large rhomboidal crystals in fibriform idioblasts from tangential section.
FIGuRE 19, scale above FIGuRE 12; FIGuREs 20, 21, scale above FIGuRE 1; FIGURE 22, scale above
FIGURE 3.
1992
309
FIGuRES 23—27. Wood sections of Fabiana viscosa (Mexia 7838). 23. Transection; numerous
growth rings evident; vessels are mostly solitary. 24. Tangential section; rays are uniseriate composed
of upright cells. 25. Portion of radial section showing tracheids containing helical thickenings; walls
between tracheids show bordered nature of pits. 26—27. SEM photomicrographs of radial section to
show helical thickenings. 26. Earlywood vessel (left) plus tracheids (extreme right). 27. Latewood
vessels. FIGus.a 23, 24, scale above FIGURE 1; FIGuRE 25, scale above FIGUa 3; FIGUREs 26, 27, scales
indicated at upper right, respectively (bracket in each
10 am).
310
ALLERTONIA
6.4
31
FiGuans 28—31. Wood sections of Grabowskya ameghinoi (Donat 36). 28. Transection; vessels are
very narrow, and are in large diagonal aggregations. 29. Tangential section; rays are uniseriate or
biseriate. 30. Transection, to show libriform crystal sand idioblasts (above) and diffuse axial paren
chyma cells containing starch. 31. Thin-walled fibriform crystal sand idioblasts from tangential section.
FiGuans 28, 29, scale above FIGuI 1; Fiouias 30, 31, scale above FIGuIa 3.
1992
311
FiGuiras 32—35. Wood sections of Lycianthes and Lycium. 32—33. Lycianthes lycioides (Cariquist
7347). 32. Transection; earlywood vessels are much wider than latewood vessels. 33. Tangential
section; multiseriate rays and uniseriate rays are about equally abundant. 34—35. Lycium brevipes
(RSABG-14429). 34. Transection, showing extensive diagonal vessel aggregations. 35. Transection
portion, to show vessel group containing vasicentric tracheids and diffuse axial parenchyma cells
(identifiable by dark contents). FIGuREs 32—34, scale above FIGuRE [;F1GuIE 35, scale above FIG
URE 12.
312
ALLERTONIA
6.4
FIGuRES 36—40. Wood sections of Lycium. 36—37. L. elongatum (PRFw-10520). 36. Transection;
large vessels are intermixed with narrow vessels and vasicentric tracheids in aggregations. 37. Tan
gential section; rays are predominantly uniseriate, ray cells are upright. 38—40. L. sandwicense (PRFw
246 76). 38. Transection; large earlywood vessels contrast with narrow latewood vessels in large ag
gregations. 39. Tangential section; rays are uniseriate and biseriate. 40. Vessels from tangential section,
showing helical thickenings. FIGuREs 36—39, scale above FIGuRE 1; FIGuRE 40, scale above FIG
URE 3.
1992
313
I
FIGuREs 41—45. Wood sections of Nicotiana. 41—42. N. cordifolia (Skottsberg 18, Stem). 41. Tran
section; vessels are in radial multiples. 42. Tangential section; multisenate rays are wide, short. 43—
45. N. otophora (UCBBG-51.OO1). 43. Crystal sand idioblast in ray from tangential section. 44. Small
rhomboidal crystals in axial parenchyma, from tangential section. 45. Small rhomboidal crystals in
ray cells from radial section. Fiouis 41, 42, scale above FIGuRE 1; FIGuRES 43—45, scale above FIG
URE 3.
314
ALLERTONIA
6.4
.1
‘49
FIGuas 46—49. Wood sections of Nicotiana raimondii (Cariquist 7345). 46. Transection; vessels
are in long radial multiples. 47. Transection to show diffuse parenchyma cells (rounded, with gray
contents, mixed among angular fiber-tracheids). 48. Radial section; almost all ray cells are upright,
few are square. 49. Ray cells containing trabecula (horizontally across center), from radial section.
Ficjupz 46, scale above FIGuRE 1; FIGuIas 47, 48, scale above FIGua.a 12; FIGuRE 49, scale above
FIGuI 3.
1992
315
FIGuREs 50—55. Wood sections of Nothocestrum latifolium (Cariquist 2087). 50. Transection; axial
parenchyma is scanty vasicentric plus banded apotracheal. 51. Tangential section; rays are uniseriate
plus biseriate. 52. Grooves interconnecting pit apertures, from vessel wall of tangential section. 53.
Portion of transection, showing thick vessel walls and dark-staining compounds in parenchyma cells.
54. Crystal sand in axial parenchyma cells from radial section. 55. Crystal sand idioblast in ray, from
tangential section. Fiouis 50, 51, scale above FIGuRE 1; FIGuREs 52, 54, 55, scale above FiGuan 3;
Fiousn 53, scale above Frouan 12.
316
ALLERTONIA
Fiousas 56—60. Wood sections of Solanum. 56—57. S. acropterum (USw-6002). 56. Transection,
illustrative of low vessel density despite narrowness of vessels. 57. Tangential section; rays are uni
senate, composed of upright ray cells. 58. S. appendiculatum (Anderson 479); transection, showing
dense placement of wide vessels. 59—60. S. hayesii (Nee & Mon 3667). 59. Helical thickenings in
vessel (right) and vasicentric tracheid (left), from tangential section. 60. Septate thin-walled fibriform
crystal sand idioblast, from radial section. FIGURaS 56—58, scale above FIGuRE 1; FIGuREs 59, 60,
scale above FIGuRE 3.
1992
317
FIGuIas 61—65. Wood sections of Solanum gayanum (USw-8466). 61. Transection, showing rel
atively great density of vessels. 62—64. Sclerosed tyloses in vessel from radial section. 62. Vessel
showing density and size of tyloses. 63. Wall between tyloses in sectioning view, showing bordered
nature of pits. 64. Wall portions of tyloses in face view; wall thickness evident. 65. Vessel from
tangential section, to show helical thickenings and lateral wall pitting. FIGuRE 61, scale above FIGuIa
1; FiGuan 62, scale above FIGu 12; FIGuREs 63—65, scale above Fiouaa 3.
318
ALLERTONIA
6.4
FIGURES 66—69. Wood sections of Solanum kauaiense (USw-15288). 66. Transection; wood is
semi ring porous. 67. Tangential section; multiseriate rays are notably wide 68. Vessel wall from
tangential section, to show angular outline of pits. 69. Ray cells from radial section (horizontal ray
axils arranged vertically) to show dark-staining contents, borders on pits interconnecting ray cells.
FIGuRES 66, 67, scale above FIGURE 1; FIGuREs 68, 69, scale above FIGuRE 3.
1992
319
FIGuIas 70—74. Wood sections of Solanum. 70—71. S. ob1ongfolium (cult. UCBBG). 70. Tran
section; wood is diffuse porous, vessels are in radial multiples. 71. Tangential section; multiseriate
and uniseriate rays are about equally abundant. 72—74. S. paludosum (PRFw-17926). 72. Ray cells
from radial section to show procumbent nature of cells (horizontal axis of ray oriented vertically). 73.
Transection; growth rings are inconspicuously demarcated; vessels are wide. 74. Tangential section;
rays are sparse, mostly uniseriate. FIGUREs 70, 71, 73, 74, scale above FIGURE 1; FIGURE 72, scale
above FIGuRE 12.
6.4
ALLERTONIA
320
!4
L4
0
.—.
—I1
—.
—
I
-
—
4_
-
,
FIGuRES 75—78. Wood sections of Solanum and Solandra. 75—76. Solanum rugosum (PRFw
16104), portions of tangential sections. 75. Vessel-vessel pitting, showing grooves interconnecting pit
apertures. 76. Vessel to axial parenchyma pitting, showing grooves interconnecting pit apertures; pairs
of inconspicuous thickenings accompany the grooves. 77. Solanum nigricans (Nee & Taylor 28900);
fiber-tracheids from radial section, showing very small borders on pits. 78. Solandra guttata (cult.
Vavra estate, UCLA); transection to show wide vessels, banded parenchyma. Fiouis 7 5—77, scale
above FIGus 3; Fious 78, scale above FIGuRE 12.
1992
321
FiGuans 79—82. Wood sections of Solanum to show crystals. 79—80. S. trichoneuron (PRFw
10508), ray cells from radial section. 79. L.arge rhomboidal crystals. 80. Rhomboidal crystals, inter
mixed with crystal sand in some ray cells. 81. S. grandflorum (Carlquist 7388), portion of thin-walled
fibriform crystal sand idioblast from radial section. 82. S. torvum (USw-6033), lIbriform crystal sand
idioblast from tangential section. FIGuREs 79—82, scale above Fiouan 3.
322
ALLERTONIA
6.4
seriate rays seems quite different from Nicotiana, but it would be a distinctive
genus in any assemblage of genera. In the Salpiglossideae, the genus Streptosolen
shows no appreciable differences from Solanum in wood anatomy, but Brunfelsia,
with its distinctive tracheids and diffuse parenchyma, differs from Streptosolen.
Although evolution from tracheids to fiber-tracheids may be reversible within
relatively narrow limits, as a first interpretation one tends to consider that tra
cheids are more primitive than fiber-tracheids (data that tend to show that tra
cheids are primitive are presented in Metcalfe & Chalk, 1950, p. xlv, TABLE 3).
Relationship between Solanaceae and other families will be reviewed when a
study of wood anatomy of Convolvulaceae, a project now in progress, is com
pleted. Solanaceae and Convolvulaceae are often regarded as closely related. The
data from the present study support the interpretation of Hunziker (1979) that
Duckeodendraceae and Goetzeaceae are satellite families of Solanaceae; these
latter two families have no wood features not also seen in Solanaceae (Carlquist,
1 988b). Nolanaceae also prove to be close to Solanaceae in wood characteristics
(Cariquist, I 987b), which is in agreement with Hunziker (1979). Both Goetzeaceae
and Nolanaceae possess crystal sand, a feature not common in woods of dicot
yledons: it is found in only seven families other than Goetzeaceae, Nolanaceae,
and Solanaceae (Carlquist, 1 988a, p. 234). Other wood features by means of which
recognition of an order, Solanales (e.g., Dahlgren, 1975; Thorne, 1976), is sup
ported will be reviewed in a forthcoming paper. That paper summarizing wood
anatomy of tubiflorous families of dicotyledons is planned as a conclusion to the
series of papers of which the present study is a portion.
ACKNOWLEDGMENTS
Wood samples have kindly been provided from the Samuel G. Record Collec
tion (SJRw) of the Forest Products Laboratory, Madison, Wisconsin, and from
the wood collection of the Princes Risborough Laboratory (PRFw). Appreciation
is expressed to Dr. Regis B. Miller, and Dr. J. D. Brazier for woods from these
two xylaria, respectively. The samples cited with USw numbers in TABLE 1 are
actually located in the RSAw wood collection, but represent part of a very large
group of wood specimens sent to me by the Smithsonian Institution through the
courtesy of Dr. William L. Stern. Dr. Gregory Anderson provided samples of
Solanum sect. Basarthrum, which are notable for their climbing habit (Anderson
et al., 1987). The wood samples collected by Dr. Michael Nec and coworkers
proved very helpful, and his courtesy, as well as that of the New York Botanical
Garden, in providing me with these deserves special thanks. Wood samples from
the University of California Botanic Garden, Berkeley (UCBBG in TABLE 1) were
collected with the permission of Dr. Robert Ornduff. Specimens from the Vavra
Estate, a garden that belonged to UCLA for several years, were made available
by Dr. Mildred A. Mathias. I was able to collect woods of Solanaceae in South
America by means of a grant from the National Science Foundation, DEB 8108810. Several former students aided in the sectioning of wood samples and
collection of data; for their help, I would like to express appreciation to David
Barnard, Vincent M. Eckhart, David A. Hoekman, Charles F. Quibell, and Scott
1992
323
Zona. Sandra Knapp kindly identified herbarium specimens documenting my
South American wood collections. Dr. David F. Cutler, Dr. David H. Lorence,
and an anonymous reviewer read the manuscript and offered helpful suggestions.
LITERATURE CITED
Ai-IMA.u, K. J. 1964. Anatomy of plants—Il. Solanum indicum Linn. Bull. Nat. Bot. Gard. (Lucknow,
India) 108: 1—14.
ANDERsON, G. J., T. P. STEINHARTER, & G. CoopER-DRIvER. 1987. Foliar flavonoids and the sys
tematics of Solanum sect. Basarthrum. Syst. Bot. 12: 534—540.
A111sr1oNG, J. E. 1986. Comparative floral anatomy of Solanaceae: a preliminary survey. In: D’Arcy,
W. G. (ed.). Solanaceae. Biology and Systematics, pp. 101—113. Columbia University Press, New
York.
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DESCOLE,
1992
325
iNDEX
An asterisk (*) after a page number indicates a figure .N ames occurring in abstract, acknowledgments,
and literature cited are not indexed.
Acnistus, 294, 295, 302, 303, 304*
—arborescens, 281, 282, 293, 294, 297, 298, 302
—australis, 289
—grandiflorus, 281, 282, 287. 289, 290, 293,
295, 297, 302, 304*
—parviflorus, 283, 282, 287, 289, 293, 294, 295,
297, 298, 299, 302, 305*
Anthocercis, 303, 304*
—littorea, 281, 282, 287, 288, 289, 290, 291,
295, 304*
Asteraceae, 287
Brugmansia, 293, 294, 295, 303
—sanguinea, 281, 282, 293, 294, 298
—suaveolens, 281, 282, 288, 292, 294, 298, 301
Brunfelsia, 289, 290, 291, 292, 293, 294, 297,
300. 301, 303, 322
—calycina, 281, 282
—nitida, 281, 282, 292, 295
Bruniaceae, 302
Capsicum, 302, 303, 306*
—ciliatum, 292, 293, 294, 295, 297, 301, 306*
Ccrcidiphyllum, 287
Cestrum, 281, 294, 298, 303, 307*
—conglornerasum, 282, 292, 293, 294
—diumum, 282, 287, 288, 294, 296, 307*
—elegans, 293
—hirtum, 282
—macrophyllum, 282, 292, 294, 299
—nocturnum, 282, 289, 290, 293, 294, 295
—parqui, 282, 294
—puhens, 282, 289, 291, 293, 294, 295, 297,
298, 307*
—purpureum, 293
Convolvulaceae, 322
Cyphomandra, 302, 303, 308*
—hartwcgii, 281, 282, 288, 294, 299, 300, 301,
303, 308
—pendula, 288, 290, 294, 297, 303
Datura, 303, 306*
—mcteloides, 281, 282, 285, 293, 294, 295, 296,
301, 302, 306*
Duhoisia, 303
—myoporoides, 281, 282, 289, 292, 294, 295,
297, 299
Duckeodendraccac, 322
Dunalia, 281, 290, 292, 294, 303
—arborescens, 282, 289, 301
—obovata, 282, 293
Empetraceac, 302
Fahiana, 281, 289, 290, 291, 292, 293, 295, 300,
301, 302, 303, 308*
—bryoides, 281, 282, 285, 287, 288, 289, 294,
297, 299, 301
—imbricata, 282, 288, 289, 294, 297, 301, 308*
—viscosa, 281, 282, 285, 287, 288, 289, 293, 294,
295,297,300,301,309*
Goetieaceae, 322
Grabowskya, 281, 289, 290, 291, 295, 300, 301,
303, 308*
—ameghinoi, 281, 282, 285, 288, 289, 291, 294,
295, 297, 298, 300, 310*
—duplicatum, 282, 287, 288, 289, 291, 294, 296,
297, 298, 300, 308*
Iochroma, 303
—tubulosa, 281, 282, 294, 295
Lamiaccae, 287, 290
Lycianthes, 303, 311*
—lycioides, 281, 282. 287, 294, 295, 296, 297,
311*
Lycium, 281, 289, 290, 291, 292, 294, 295. 296,
300, 301, 302, 303, 311*, 312*
—brevipes, 281, 282, 285, 288, 291, 293, 294,
295, 297, 298, 300, 301, 311*
—carolinianum, 282, 291, 297, 300, 301
—cestroides, 282, 287, 291, 294, 295, 297
—elongatum, 282, 287, 291, 294, 295, 296, 297,
312*
—europaeum, 281, 282, 287, 291, 294, 295, 296,
297, 298
—fremontii, 281, 282, 285. 287, 288, 293, 294,
298, 299, 300
—sandwicense, 281, 282, 289, 290, 291, 294, 297,
299, 300, 312*
Lycopersicon, 303
—esculentum, 281, 282, 289, 294, 295, 299, 301,
302
Myrothamnaceae, 302
Nicotiana, 295, 302, 303, 313*, 322
—cordifolia, 281, 282, 287, 289, 292, 294, 295,
296, 297, 299. 313*
—glauca, 281, 282, 285, 289, 292, 293, 294, 296,
298, 301
—otophora, 281, 282, 289, 294, 295, 297, 298,
299, 301, 313*
—raiinondii, 281, 282, 287, 289, 294, 295, 296,
297, 298, 314*
—setchellii, 281, 282, 292, 294
—tomentosa, 281, 282, 289, 290, 294, 297, 299
Nolanaceae, 322
Nothocestrum, 281, 289, 290, 294, 295, 300, 303
—breviflorum, 284. 299, 300
—latifolium, 284, 289, 290, 295, 296, 297, 298,
299, 300, 315*
—longifolium, 282, 288, 297, 299, 300
Polioinintha, 290
Solanaceae subfam. Cestroideae, 302, 303
trib. Nicotianeae, 303
trib. Salpiglossideae, 322
—
—
——
326
ALLERTONIA
—subfam. Solanoideae, 302, 303
trib. Lycicac, 303
Solanales, 322
Solandra, 290, 300, 303, 320*
—guttata, 281, 284. 288, 289, 294, 295, 299,
300, 301, 320*
Solanum, 280, 289, 290, 292, 293, 294, 295, 300,
301, 303, 3 16*, 319* 320*, 321*, 322
—accrescens, 284, 295, 301
—acropterum, 281, 284, 287, 289, 294, 295, 296,
303, 316*
—aihidum, 281, 284, 294, 295, 297, 299
—appendiculatum, 281, 284, 286, 288, 289, 291,
292, 295, 300, 31*
—auriculatum, 281, 284, 290, 292, 295, 298
—australe, 281, 284, 295
—bahamense, 281, 284, 297, 299
—chrysotrichum, 284, 289, 290, 295, 297
—convolvulus, 293
—crispum, 281,284,289, 290, 291, 295, 300,
302
—douglasii, 281, 284, 285, 288, 290, 291, 293,
296, 300
—dulcamara, 293
—erianthum, 281, 284, 290, 291, 295, 297
—gayanum, 281, 284, 287, 290, 291, 299, 317*
—grandiflorum, 281, 284, 288, 289, 290, 292,
294, 295, 299, 300, 321*
—hayesii, 281, 284, 291), 293, 294, 295, 297,
299, 316*
—hirwm, 281, 284, 295
—hispidum, 281, 284, 292
6.4
—indicum, 299
—jasmrnoides, 281, 284, 285, 288, 289, 291, 293,
295, 297, 300
—kauaiense, 281,284, 290, 295, 296, 298, 318*
—leucocarpon, 281, 284, 290, 297, 299
—nclsonii, 281,284, 289, 297, 299
—nigricans, 281, 284, 290, 292, 293, 295, 320*
—nitidum, 281, 284, 288, 291, 297, 299, 300, 301
—nudum, 281, 284, 290, 295
—oblongifolium, 281, 284, 287, 289, 290, 297,
298, 319*
—paludosum, 281, 284, 288, 289, 295, 296, 297,
319*
—rugosum, 281,284,288,291,292,295,297,
320*
—sandwicense, 281, 284, 287, 288, 289, 290, 293,
297, 298
—saponaceum, 281, 284, 288, 290, 295
—simile, 281, 284, 289, 295, 296, 302
—sodiroi, 281,284,286,288,289,290, 291,295,
296, 297, 299, 300, 302
—tetrapetaium, 281, 284, 286, 288, 289, 291, 295,
300
—torvum, 284, 288, 294, 295, 299, 321 *
—trichoneuron, 281, 284, 295, 297, 298, 299, 321*
—triste, 281, 284, 290, 295
—xantii, 281, 284, 285, 288, 289, 291, 293, 295,
296, 300,301, 302
Strepiosolen, 303, 322
—jamesonhi, 281, 284, 288, 292, 293, 294, 296,
297, 302
lrcmandraceac, 302

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