Nord. J. Bot.

- Section of tropical taxonomy

The genus Theophrasta (Theophrastaceae). Foliar structures,

floral biology and taxonomy
Bertil StPhl

Stihl, B. 1987. The genus Theophrasta (Theophrastaceae). Foliar structures, floral

biology and taxonomy. - Nord. J. Bot. 7: 52%538. Copenhagen. ISSN 0107-055X.

The genus Theophrasta is endemic to the island of Hispaniola. Two species are
recognized, viz. T. jussieui and T. americana. T. jmsieui is restricted to the central
part of northern Haiti, whereas the latter is distributed over most of Hispaniola. Only
floral characters have been practically useful to distinguish the species. T. jussieui
differs from T. americana primarily by its larger flowers and inflorescences. The two
species are considered closely related. An investigation of leaf surface structures and
foliar anatomy, particulary the sclerenchyma, is presented. Floral biology, including
adaptions to sapromyophilous pollination, is described and discussed, and notes on
habit, ecology, and pollen morphology are given.

B. Stihl, Dept. of Systematic Botany, Univ. of Goteborg, Carl Skottsbergs Gata 22,
S-413 19 Goteborg, Sweden.

Introduction Material and methods

The small Neotropical family Theophrastaceae includes This investigation is based primarily on herbarium spec-
five genera of shrubs and small trees. They are dis- imens. About 65 collections from the following herbaria
tinguished, in the first place, by reproductive charac- have been studied: A, B, BM, BP, BR, C, CGE, F, G,
ters, but differ to some extent also in habit and in size GB, GH, GOET, HBG, IJ, JBSD, K, L, LE, M, MO,
and shape of the leaves. Four of the genera, Deherainia MPU, NY, P, PH, S, U, US (abbreviations according to
Decne., Jacquinia L., Neomezia Votsch, and Theo- Holmgren et al. 1981). During a stay in the Dominican
phrasta L., have an essentially Antillean distribution, Republic in April 1985 the author had the opportunity
whilst Clavija R. & P. is centered in continental South to make field observations on several populations of T.
America. americana.
The last revision of Theophrasta, endemic to the is- Measurements of vegetative parts and fruits were
land of Hispaniola, was made by Mez (1901, 1903). He made on dry specimens, those of flowers on rehydrated
recognized two species, T. americana L. and T. jussieui material. A small number of flowers pickled in FAA
Lindl., based on differences in floral size, shape of the were used for complementary studies and for the prep-
stamina1 appendages, and in leaf serration. This study aration of illustrations.
supports Mez’s classification and no nomenclatural For the investigation of microscopic foliar structures,
changes have been found necessary. However, the cir- small leaf fragments, consistently taken from the broad-
cumscription of the two species is here revised. Only the est part of the lamina midway between the main vein
difference in floral characters has shown practically use- and the margin, were soaked over night in a weak
ful to distinguish between the two species. ammonia solution. Leaf sections were cut by hand with
a razor blade and embedded in glycerine without stain-
ing. Macerates of sclerenchymatous tissue were pre-

0 NORDIC JOURNAL OF BOTANY

NORD. J. BOT. 7: 529-538, TROP 156

N o d J. Bot. 7 ( 5 ) 1987 529

Fig. 1. Leaf epidermis. - A, D, F-G: Theophrasra arnericana; B-C,E, H:T. jussieui. - A-B (X35), D-E ( ~ 1 6 0 ) :Peels of upper
epidermis. - C (X15): Upper epidermis under dissection microscope: note sunken glands and the fine striation caused by
subepidermal fibre bundles. - F (X370): SEM of sunken glandular hair on the upper side; the epicuticular structures are probably
wax secretions. - G ( x 150), H (x535): SEM of lower epidermis; note sunken gland (G), cuticular ledges above the stomata pores,
and the irregularly ridged surface. - A, D: Stah1 & Lindstrom 238; B-C, E, H:Ekman H4425; F-G: Stihl & Lindstrom 178.

pared with chlorine water (Eau d e Javelle, Kisser 1931). and 24.5 mm’ respectively. Complementary frequency
This method also yielded good epidermal peels. On counts of glands on the adaxial leaf surface were also
each leaf examined, frequencies and measurements of made directly o n dried leaves under a dissecting micro-
stomata and epidermal glands were obtained from scope; the values were in these cases obtained from
counts made on leaf surface areas comprising 0.6 mm2 counts made on areas comprising 18 mm’.

530 N o d 1. Bot. 7 ( 5 ) 1987

 A
Within the family, the same fundamental habit is
z
.’
1 2 3 shown by Clavija and the very short-stemmed, mono-
typic genus Neomezia (Mez 1903, Hall6 & Oldeman
5
z
1 * 1970). However, both genera differ from Theophrasta
c
4 10- in being more apt to branching when damaged. In addi-
m . tion, these genera do not have the strictly suprafoliar

7.5-
.... . a.
* 2

*’ 2
.
4
placement of the flowers.

The leaf
In Clavija and Jacquinia, both genera of about 30 spe-
cies, the leaves exhibit a considerable variation in
5-

e 50 100 glands/crn2

Fig. 2. Correlation between leaf width and frequency of glands

shape, size, texture and anatomy (Votsch 1904, Stihl,
unpubl.). Mostly, this variation provides useful charac-
ters for delimiting taxa. It is therefore surprising that
this study fails to demonstrate any significant foliar
on the adaxial leaf surface in Theophrastuumericunu (0)and characters to distinguish between the two species of
T. jussieui (+). Labelled arrows designate measurements from
different leaves of the same collection number. Theophrasta, which differ so prominently in floral char-
acters. Morphological features such as leaf serration,
which Mez (1903) considered decisive, and leaf width,
are highly variable and overlap too much to be of any
For the measurements of pollen grains, anthers from taxonomic significance. Furthermore, leaf length seems
herbarium specimens were soaked in a 0.25% NH, solu- to be positively correlated to plant size.
tion for an hour before being crushed on microscope
slides. Twenty pollen grains from each of the examined Epidermal structures. The adaxial leaf surface has a
collections were measured. thick cuticula; it is smooth with regard to primary sculp-
For supplementary SEM study of pollen and leaf ture, but epicuticular secretions, here interpreted as
structures, dry samples were taken from herbarium wax flakes (Figs 1A-B,F), were seen in some of the
specimens and coated with a 50A layer of gold-palla- specimens examined in SEM. The thinner cuticula of
dium. the abaxial side is furnished with ridges, which were
present but variously well pronounced in all specimens
examined (Fig. 1G).
The stomata are anomocytic and confined to the
Morphology abaxial leaf side, where they are distributed in rows,
being absent from areas below the subepidermal fibre
Habit
bundles. Due to overarching of cuticular ledges, the
Both species of Theophrasta coincide with Corner’s stomatal pores are provided with small antechambers
Model (Hall6 et al. 1978). They are monocaulous (Fig. 1H). No significant differences in stomatal size
shrubs and treelets, generally less than 2 m high. (29.5-32.5 pm long) and frequency (100-200/mm2) were
Branching is extremely rare, and if the apical meristem found between T. americana (N=12) and T. jussieui
is damaged, growth continues from a single dormant (N=3).
bud at the very distal part of the stem. Sometimes, and The foliar glands, which are similar in construction
presumably as a result of disturbances, plants may pro- throughout the Theophrastaceae, are composed of a
duce a * few additional subterraneously originating short stalk cell and a broad, multicellular head. They
shoots. are sunken into the epidermis (Figs 1A-B, F) and occur
The leaves are clustered in 3 or 4 pseudo-whorls along on both sides of the leaf. Under a dissecting microscope
the upper part of the stem. Each pseudo-whorl is com- they appear as small, evenly distributed dots (Fig. 1C).
posed of up to ca. 20 leaves arranged in a condensed The glands are most abundant on the abaxial leaf side,
spiral. The apical meristem is protected by narrow bud viz. 159-384/cm2 (%= 288; N = 10) in T. americana and
scales, persisting as spines on the older parts of the 275-280/cm2 (R =277; N = 3) in T. jussieui. The figures
stem. Periodic growth, imposed by the seasonally dry for the adaxial leaf side are 60-200/cm2 (% = 108; N = 30)
climate, is thus reflected by alternating whorls of spines in T. americana and 155-196/cm2 (%=176; N=7) in T.
and leavedeaf scars. jussieui.
The racemose inflorescences appear above the up- The adaxial gland frequency, which seemed to be a
permost leaf whorl, but originate laterally, just below possible character after a preliminary study, is very
the apical bud (Fig. 5H). The fruits are always found at variable, also within the same population. Furthermore,
the top of the stem and mature before the flush of new the frequency seems to be partly negatively correlated
leaves. to the leaf width (Fig. 2). It is noteworthy, however,

Nord. J. Bot. 7 ( 5 ) 1987 531

Fig. 3. Foliar sclerenchyma in Theophrasra arnericana. - A (X40): Fibre bundles of the lower side. - B (X295): TS of leaf. - C
(x45): Fibre cells of macerated fibre bundles. - A-C: Ekman H8121.

that the values for T. jussieui, the collections of which Sclerenchyrna. A characteristic feature of the Theo-
are from a geographically limited area, vary relatively phrastaceae is the presence of extra-xylary foliar scle-
little. In contrast, though, similar geographical correla- renchyma developed as subepidermal layers or bundles
tion for the collections of T. americana is not discerni- of tightly packed fibre cells. In Theophrasta this scleren-
ble. chyma consists of very coarse fibre bundles forming an
The anticlinal cell walls of the adaxial epidermis are anastomosing net-like pattern on both sides of the leaf
slightly undulated to almost straight (Fig. 1D-E). The (Fig. 3A-B). On dried leaves these bundles are visible
almost straight condition prevailed in T. jussieui, to the naked eye as fine striations running almost paral-
whereas the examined material of T. americana exhib- lel to the lateral veins (Fig. 1C). They are less easily
ited the whole series of this variation. observed on fresh leaves. The number of fibre cells in a
bundle vanes considerably, from 40 to 150 in an ar-

532 Nard. I. Bot. 7 ( 5 ) 1987

bitrary cross-section. Generally, the adaxial leaf side yellowish pollen grains below. Calcium oxalate crystals,
possesses coarser and more closely placed bundles than arranged in the same manner, have been described in
does the abaxial side. Deherainia smarugdina (Pohl 1931) and are also spar-
The individual fibre cells always have a narrow lumen sely present in the anthers of Clavija and Jacquinia
and sometimes sinuous outer walls. They are mostly (StBhl, unpubl.).
straight with pointed tips (Fig. 3C), but curved cells, The colour of the corolla changes during the course of
sometimes with a tendency to branch, are often met anthesis from pale buff to reddish-brown and finally to
with at points where the bundles interconnect. The brownish-black. This change is expedited by desicca-
straight fibre cells are 0.50-1.10 mm long on the adaxial tion, so that herbarium specimens always have brow-
side and 0.354.85 mm on the abaxial side. nish-black flowers. The matt appearance of the corolla
No difference was found in foliar sclerenchyma be- is caused by a dense striation of the epidermal surfaces.
tween T. americana and T. jussieui. The statement by The surfaces of the staminodes are smooth, however.
Votsch (1904: 505) that T. jussieui differs from T. amer- During field work in the Dominican Republic, flo-
icana by the presence of a hypodermal layer on the wering plants of T. americana were studied in two
adaxial leaf side, was not verified. No hypodermal lay- widely separated populations (vouchers: Stdhl & Lind-
ers were seen in any of the collections examined. Only strom 178, 262). In both populations the flowers were
in two collections, one of each species, I found single or found to have a fungal scent. The only earlier record of
small groups of large, empty cells in sub-epidermal posi- flower odour in Theophrasta is that of a cultivated plant
tions. These cells never formed a continuous hypoderm, of T. jussieui: “. .. nicht von dem angenehmsten Ge-
however. ruch.” (Otto 1837).
The flowers are protandrous and during the staminate
phase the ovary is enclosed by the loosely coherent,
Floral biology and pollination
erect stamens. At the onset of the pistilate phase, corre-
The floral biology of Theophrasta has not been de- sponding fairly well to the colour shift of the corolla to
scribed before. Many of the features, morphological as reddish-brown, the stamens bend outwards.
well as ecological, agree with the “sapromyophilous Although the inflorescences are located at the top of
pollination syndrome” as defined by Faegri and van der the stem, well above the forest floor, a simulated close-
Pijl (1979) and thoroughly studied by Vogel (1978). In to-the-ground condition arises as the flowers are cov-
short, pollination is affected by Diptera. Features of the ered by litter from the forest canopy (Fig. 5F). In the
sapromyophilous blossoms attract the insects and acti- field one usually has to remove a considerable amount
vaies the; instinct to feed or deposit eggs. Since the of dead leaves to see whether the plant is flowering or
resulting larvae perish the system is parasitic on behalf not.
of the plant. Eggs and larvae of the suborder Cyclorrhapha (pos-
The corolla is bell-shaped in Theophrasta. Within the sibly fam. Syrphidae; H . Anderson, pers. corn.) were
tube there is a thick ridge composed of staminodes and found in some pickled T. americana flowers of the red-
emergences from the corolla wall (Fig. 5C-D). In T. dish-brown colour. Eggs were found exclusively on the
jussieui, the emergences seem to be less well developed lower parts of the puckered corolla lobes, whereas the
and the ridge is dominated by the staminodes. The larvae were found mainly in the chamber below the
surface of the ridge is fairly smooth in T. americana but staminodal ridge.
papillose in T. jussieui. In contrast, the inner sides of Since no real attempts to undertake pollination stud-
the corolla lobes are irregularly puckered in T. amer- ies were made while in field, conclusions about the
icana and smooth in T. jussieui. At least in T. amer- pollination biology of Theophrasta must be drawn from
icana, the staminodal ridge becomes more irregular in indirect evidence. Indications of sapromyophily in T.
shape during anthesis. Below the ridge, where a small americana, and pollination by insects that normally
chamber is formed, lamellae from the staminodes and breed in mushrooms, are the fungal scent and the *
the emergences continue down to the bottom of the hidden position of the blossoms, the colour, shape and
corolla. Both calyx and corolla are furnished with sunk- k irregular structures of the corolla, and, finally, the
en glandular hairs of the same construction as those on presence of Diptera eggs and larvae in the flowers.
the leaves. The glands are most abundant within the Also, the flowering period (April-July; recorded from
corolla and are especially plentiful on and in close prox- herbarium material) coincides with the humid period of
imity of the staminodes. It is not known if the activity of the year (Moscoso 1943). The role of other k peculiar
these glands compensate for the absence of true nectar- floral features, i.e. color shift, protandry, calcium ox-
ies. The secretory products are in any case very sparse. alate crystals in the anthers, and appendaged stamens,
The stamens are apically appendaged and, as in all remain obscure. There is indirect evidence suggesting a
members of the family, the anthers are extrorse. In- slightly deviant pollination biology in T. jussieui.
terestingly, the upper third of the anthers are filled up Firstly, compared to those of T.americana, the flowers
with small calcium oxalate crystals. This mass of crystals are much larger and lack irregular (“fungoid’) struc-
is brilliantly white and contrasts noticeably with the tures above the staminodal ridge. Secondly, as seen in

34 Nord. J . Bot. 7 ( 5 ) 1987 533

 in the lower part of the inflorescence narrow, lanceolate
and adnate to the base of the pedicel, further up nar-
rowly ovate to ovate, with sparsely serrulate margins,
gradually fused higher up on the pedicel; calyx yellow-
ish-brown, persisting in fruit, sometimes puberulous in-
side at base, glandular-punctate, lobes connate at base,
margins membranaceous and gnawed; corolla pale buff,
turning brownish-black with age, glandular-punctate
(especially within), campanulate, the lobes broadly
ovate, eventually spreading; staminodes gibbous, trans-
Fig. 4. SEM of dried and partly collapsed pollen of Theo- versely oblong, densely glandular-punctate, fused with
phrasta americana (A) and T. jussieui (B). - A: StHhl & Lind-
strom 178; B: Ekman H8309. - Scales: 5 pm. emergences from the corolla, forming a thick ridge
within the tube; stamens loosely coherent, forming a
tube around the gynoecium, eventually spreading, fil-
early drawings (Lindley 1821, Vnese 1839), the longer aments free, flattened, broadening towards base, con-
inflorescences of T . jussieui may eventually protrude nate at base and fused with the corolla tube, anthers
among the surrounding leaves and are thus less easily narrowly oblong with white crystals in the upper part of
covered by plant litter. Third and lastly, the sole record the thecae, connective apically produced to form a tri-
of flower odour in T . jussieui (Otto 1837, see above) angular appendage; ovary broadly ovoid, glandular-
does not explicitely state a fungal scent. Indeed, the punctate, gradually narrowing into a style of equal
differentiation of the two species is probably best under- length, ovules numerous, spirally inserted in 5-7 rows
stood as adaptions to different pollination strategies. on a basal column, stigma capitate, orange and resin-
ous. Fruits yellow, subglobose; seeds pale brown,
smooth, variable in size and shape, often within the
Pollen
same fruit, ovoid to irregularly obtuse-angled, some-
The pollen of Theophrasta is 3-colporate, subspheroi- times compressed, immersed in a juicy, yellowish-white
dal, and 22.5-26.5 pm long (polar axis; %=24.0, N=7). pulp, the endosperm copious and hard.
No difference was found in size and shape of the pollen
grains between the two species. However, the exine
Key to the species
pattern of T . americana appears coarser than that of T .
jussieui in the two specimens examined in SEM (Fig. 1. Corolla %15 mm long, calyx lobes 2 . 5 4 . 0 mm long. . . . . .
4A-B). Pollen with similar morphology has previously ....................................... T. americana
1. Corolla 2C30 mm long, calyx lobes 6-10 mm long. . . . . . .
been reported in Clavija and Jacquinia (Erdtman 1952). ......................................... T. jussieui

Taxonomy Theophrasta americana L.

Theophrasta L. LinnC 1753: 149. - Type: The copy of a Plumier drawing in the
Lime 1737: 45; 1753: 149. - Type: Theophrasta americana L. “Codex Boerhaavianus”, now in the Library of the Rijksuni-
versiteit, Groningen (not seen by me; see nomenclatural note).
Theophrastafusca Decaisne 1876: 143. - Type: “Hort. Bot.
Evergreen, unbranched shrubs or small trees; stems Parisiensis. Anno 1859.” (P lectotype, selected here).
spiny, puberulous. Leaves arranged in 3 or 4 closely
placed pseudo-whorls along the upper part of the stem, Nomenclatural note: When describing T. americana,
the leaves of the uppermost pseudo-whorl ascending, Linne (1753) cited Plumier (1703), where the species
those of the lower whorls becoming deflexed; petioles occurs under a phrase name. Both the description and
short and broad; blades oblanceolate or narrowly oblan- the accompanying plate in Plumier’s work refer entirely
ceolate, coriaceous, attenuate at base, usually shortly or to fruits and flowers, while the description of Linne
very shortly so, obtuse or acute at apex, margins slightly covers vegetative characteristics as well. A large part of
revolute and strongly serrate, rarely serrulate, the teeth Plumier’s original drawings were compiled and copied
pungent and unequal in length; primary vein somewhat for H. Boerhaave in 1733 (Codex Boerhaavianus; Ur-
impressed above, raised beneath, lateral veins not very ban 1920). These copies, which LinnC saw during his
conspicuous; surface glandular-punctate and very stay in Holland in 1737-38 (Polhill & Stearn 1976),
faintly striated laterally. Inflorescences racemose, ap- formed the basis for later copies published by J. Bur-
pearing above the uppermost pseudo-whorl of leaves man. However, from letters cited by Gillis and Stearn
from just below the apical bud; rachis sparsely to den- (1974) it is clear that Lime not could have seen the
sely puberulous, the indumentum brownish-red, irreg- drawing of T . americana in Burman’s edition (Plumier
ularly distributed, most copious at the nodes and in 1757) before the publication of the name in Species
decurrent lines from the pedicels. Flowers 5-merous, plantarum (Linne 1753). The copy of T . americana in
*
bisexual, each subtended by a sepaloid bract: bracts the “Codex Boerhaavianus” is obviously the sole ele-

534 Nord. J . Bot. 7 ( 5 ) 1987

Fig. 5. A : Theophrasta
jussieui; B-I: T. americana.
- A-B: Inflorescences. - C:
Flower. - D: Flower in
longitudinal section. - E:
Habit. - F: Flowering shoot
tip from above. - G :
Leaves. - H: Uppermost
part of stem with the
inflorescences indicated; the
leaves cut. - I: Fruits. - A:
Photograph in herb. S;
B-D, F, H: Stahl &
Lindstrom 262; E, I: Stahl
& Lindstrom 238; G : Stahl
& Lindstrom 237 and
Ekman H8121 (above). - A ,
E-F drawn from
photographs; B-D drawn
from pickled material; G-I
drawn from herbarium
material. - Scales: A-B 2
cm; C 5 mm; D 2 mm; G 10
cm; H 15 mm; I 3 cm.

ment used by LinnC and can thus be regarded as the irregularly bullate above; staminodes k smooth or
holotype. faintly pappilose, eventually becoming more irregular in
shape; filaments 2-3 mm, anthers 2-3 mm, the append-
Illustrations: Figs lA,D,F,G; 3; 4A; 5B-I. ages 1 mm long; ovary with 5C90 ovules, style ca. 2 mm

Shrub or treelet to 2 m, occasionally higher; stem den-

sely t o sparsely puberulous, t o ca. 5 cm diam at base,
spines subulate to cuspidate, to 11 mm long. Leaves
+
Theophrarfa americana
T IYSSD~YI
I
with the petioles 3-7 mm long and 3-9 mm broad;
blades oblanceolate to narrowly oblanceolate, 13.5-
55.0 X 3.5-10.0 cm, the teeth (38-)50-120(-138) per
leaf, to 6(-11) mm long. Inflorescences 3-7 cm long
with 15-35 flowers, erect and tightly crowded together.
Flowers with the pedicels 2.0-8.5 mm long; bracts 2 4
mm long; calyx lobes very broadly ovate to ovate, I

2.5-4.0 X 3.0-5.5 mm; corolla tube 4.5-7.0 mm long,

the lobes very broadly ovate, 3.5-7.0 X 3.5-6.0 mm, Fig. 6 . Distribution of Theophrasta americana and T. jussieui.

34' Nord. J . Bot. 7 ( 5 ) 1987 535

long. Fruits 20-45 mm diam, sometimes with slightly 1985: 8-9). Thus, Theophrasta jussieui seems best typ-
tappering ends, pericarp thin and brittle, smooth or ified on the plate in Lindley (1821).
often faintly pitted; seeds 5-40(-63) per fruit, 6-11(-14) When describing T. jussieui, Lindley (1821) adopted
mm long. the idea earlier proposed by Smith (1817), that T. amer-
icana of LinnC (1753) was to be considered a species of
Distribution and ecology: T. americana is distributed all Clavija R. & P. (Ruiz & Pavon 1794). But since Lindley
over Hispaniola (Fig. 6). It grows as an understory chose to use the generic name Theophrasta for his new
plant in seasonal, semi-evergreen to almost evergreen species, no nomenclatural complications result.
forests, from near sea level up to altitudes of 800, rarely T . henrici, T. spinosa, and T. densiflora are super-
1000 m. fluous names. Both Hamilton (1825) and Koch and
Fintelmann (1859b) cite T. jussieui as a synonym and
Decaisne (1876) include T. henrici in the circumscrip-
Remarks: One collection, Eggers 1560, here included in tion of T. densiflora.
T. americana, was identified as T. jussieui by Mez
(1903). The size of the flowers in this collection falls Illustrations: Figs lB-C, E, H; 4B; 5A.
within the upper part of the variation range of T. amer-
icana, a feature which it shares with two later collections Shrub or treelet to 3 m high; stem puberulous, spines
(Ekman 13232, 15089). All three collections have flow- cuspidate, to 16 mm long. Leaves with the petioles 6 1 0
ers in which the inner sides of the corolla lobes are mm long and 5-9 mm broad; blades oblanceolate, 28.5-
irregularly puckered, typical of T. americana. 60.5 x 5.CL11.0 cm, the teeth YCL130(-150) per leaf, to
A collection from the Gonave Island (Eyerdam 93) lo(-12) mm long. Inflorescences 6.5-16.0 cm long with
deviates by its narrow leaves and by having conspic- 13-20 flowers, erect at first later becoming deflexed.
uously high frequency values of stomata (ca. 270 mm’) Flowers with the pedicels 15-32 mm long; bracts 4-8
and foliar glands (205-245 cm2adaxially). However, the mm long; calyx lobes narrowly ovate, sometimes ovate,
microscopical investigation revealed a fungal infection, 6.5-9.5 x 3.0-4.5 mm; corolla tube 14-20 mm long, the
located to the adaxial epidermis. I have excluded the lobes very broadly ovate, 6 1 0 X 6-8 mm, k smooth
measurements obtained from this collection from the above; staminodes papillose, obtusely apiculated above
circumscription of T. americana, although it is unknown in the centre; filaments 2-3 mm, anthers 2-3 mm, the
to what degree this infection has influenced on the appendages 1.0-1.5 mm long; ovary with 60-80 ovules,
appearance of the plant, the leaves of which are super- style ca. 2 mm long. Fruits: only incomplete herbarium
ficially intact. material known.

Distribution and ecology: Restricted to the central parts

of northern Haiti (Dept. du Nord and Dept. du Nord-
Theophrasta jussieui Lindl. Ouest) (Fig. 6). The sparse information available in-
Lindley 1821: Tab. 26. - Theophrustu henrici Hamilton 1825: dicates an ecology much the same as for T. americana.
27-28. - Theophrustu spinosu Willd. ex Koch & Fintelmann Altitudinal records range from 400 to 850 m.
1859b: 100. - Theophrustu densifloru Decaisne 1876: 142. -
Type: Lindley 1821: Tab. 26 (lectotype, selected here).
Remarks: Besides having larger floral parts than T.
Nomenclatural notes: One specimen present in Lind- americana, T. jussieui is also characterized by having
ley’s herbarium (CGE) is annotated “Theophrasta jus- papillose staminodes and smooth inner corolla walls.
sieui” in Lindley’s hand, but lacks further information. T. jussieui and T. americana are obviously closely
It resembles the Poiteau collection in Jussieu’s herbari- related. The vegetative similarities are great and floral
um (P-JUSS), from which Lindley states that he has differences are mainly in size. The differences in size of
seen material of Theophrasta. However, the figure in corolla and calyx between the two species are hardly at
Lindley (1821) is also said to have been drawn from all met by a corresponding differentiation in the size of
material “communicated to Mr. Lambert by Mr. Hamil- stamens and ovary.
ton”. It is known (Urban 1902) that Hamilton, who It is not known if Hamilton was the sole visitor to
visited Haiti in 1814, brought home material for culti- northern Haiti to introduce T. jussieui as an ornamental
vation in European gardens. Although not explicitly plant in heated glass-houses in Europe. Still, the species
stated, it seems obvious that the vivid illustrations, as became widely spread during the 19th century, and until
well as a part of the description, must have been made the first decades of this century it was or had been
from a living plant. There are several specimens at K cultivated in at least nine of the larger botanic gardens
that could have been prepared from the same plant as in Europe. A recent inquiry for surviving plants has, so
Lindley’s illustration. However, none of these can be far, been negative.
considered original material, since they all were pre- T. jussieui is probably a threatened species, the last
pared after the publication of the name (cf. Andersson record being an Ekman collection from 1927.

536 Nord. I. Bot. 7 ( 5 ) 19x7

 M; Holdndge 499, NY; Liogier 11203, NY (ep, Is, !); Liogier
Excluded names (nomina nuda omitted) 16509, NY; Proctor 39290, JBSD(!); Raunkiaer 744, C (ep). -
Samana: Abbot 254, GH, US(!); Abbot 384, US(!); Ekman
T. cochinchinensis (Lour.) Sprengel 1825: 671 = Melo- H15089, K, NY, S (ep, p, sc, !), US; Stihl & Lindstrom 261,
dinus cochinchinensis (Lour.) Merrill 1935: 310 (Apoc- GB; Stihl & Lindstrom 262, G B (ep, p, !), JBSD; Zanoni &
ynaceae) . Mejia 17643, JBSD(!); Zanoni & Mejia 17786, JBSD. Haiti. -
T. cubensis Radlkofer 1890: 188-189 = Neomezia Dept. du Sud: Eyerdam 412, NY, US (ep). - Dept. de I’Ouest:
Buch 2159, GH, IJ; Buch 3531, G , NY, S, US(!); Ekman
cubensis (Radlk.) Votsch 1904: 541. H2039, IJ, S (ep, !); Ekman H2341, S(!), U; Ekman H8121,
T. glycycarpa (R. & P.) Sprengel 1825: 671 = Leonia GB (ep, Is, m, !), K; Eyerdam 93, GH, US (ep, p); Jacque-
glycycarpa Ruiz & Pavcin 1799: 69 (Violaceae). mont s.n., K, P(!). - Dept. du Nord: Leonard 7257, NY, US. -
T. imperialis Linden ex Koch & Fintelmann 1859a: 91 Dept. du Nord-Ouste: Ekman H4440, F, GH, S (ep, Is, !);
Leonard & Leonard 13499, NY, US; Leonard & Leonard
(Regel 1864: 323, Tab. 453; Andre 1874: 152, Tab. 184) 14837, US(!). Without locality: Bertero s.n., G-DC (microf-
= Planchonella imperialis (Linden ex Koch & Fintel- iche), M, MO. Cultivated: BR, G, P.
mann) Baehni 1965: 67 (Sapotaceae).
T. madagascariensis Willd. ex Roemer & Schultes Theophrasta jussieui
Haiti. - Dept. du Nord: Ekman H2642, S(!); Ekman H8309, IJ
1819: 787 = Brexia rnadagascariensis (Lam.) Thou. (ep, Is), S (m, p, sc, !); Nash & Taylor 1247, NY. - Dept. du
(Saxifragaceae) . Nord-Ouest: Ekman H4425, A (!), F, G, IJ, S (ep, sc, !), U ,
T. pinnata Jacquin 1809: 49, Tab. 64-66 = Talisia US(!). Without locality: Poiteau s.n., LE, P (ep), P-JU, PH (p,
!). Cultivated: B (Is), BP, GOET, K, L, LE, MPU, NY.
princeps Oliver 1888: Tab. 1769 (Sapindaceae; Radl-
kofer 1933: 854). Acknowledgements - I wish to thank Dr Lennart Anderson
The following combinations in Theophrasta refer to and Dr Ulf Molau for critical reading of the manuscript. I am
species of Clavija, which I am presently revising: also indebted to Miss Mane Lindstrom and Dr Thomas Zanoni
and his staff at the Jardin Botanic0 Nacional, Santo Domingo,
T. biborrana (Oerst.) Kuntze 1891: 404. for valuable help during my field work in the Dominican Re-
T. cauloneura (Mart. & Miq.) Kuntze 1891: 404. public, to Dr Sy Sohmer for checking the English, and to Dr
T. fulgens (Hook. f.) Kuntze 1891: 404. Hugo Anderson for identifying the Diptera larvae. Technical
T. grandis (Decne.) Kuntze 1891: 404. assistance with SEM was given by Mrs Tudlik Bergqvist and
Fig. 5 was skillfully drawn by Mrs Liselotte Molau. Financial
T. kalbreyeri Kuntze 1891: 403. support was given by The Royal Swedish Academy of Sciences,
T. lancifolia (Desf.) Kuntze 1891: 404. Stiftelsen Lars Hiertas Minne, and Anna Ahrenbergs Fond.
T. latifolia Willd. ex Roemer & Schultes 1819: 787.
T. longifolia Jacquin 1790: 136.
T. macrocarpa (R. & P.) Kuntze 1891: 404.
T. macrophylla Link ex Roemer & Schultes 1819: References
187. Anderson, L. 1985. Revision of Heliconia subgen. Stenochla-
T. pendula (R. & P.) Kuntze 1891: 404. mys (Musaceae-Heliconioideae). - Opera Bot. 82.
T. pungens Willd. ex Roemer & Schultes 1819: 787. Andre, E. 1874. Theophrasta (?) imperialis. - L’Illustration
T. riedeliana (Regel) Kuntze 1891: 404. horticole 21: 151, Tab. 184.
Baehni, C. 1965. Memoire sur les Sapotacees 111. Inventaire
T. ruiziana Kuntze 1891: 404. des genres. - Boissiera 11: 1-262.
T. serrata Hoffmansegg 1826: 218. Decaisne, M. J. 1876. Note sur quelques plantes du groupe des
T. sparsifolia (Miq.) Kuntze 1891: 404. Theophrastees. - Ann. Sci. Nat. Bot., Ser. 6, 3: 138-145.
T. tetramera Martius 1841: 27. Erdtman, G . 1952. Pollen morphology and plant taxonomy.
Angiosperms. - Almqvist & Wiksell, Stockholm.
T. undulata (D. Don) Kuntze 1891: 404. Fiegri, K. & Pijl, van der L. 1979. The principles of pollination
ecology, ed. 3. - Pergamon press, Oxford.
Specimens examined Gillis, W. T. & Stearn, W. T. 1974. Typification of the names
of the species of Leucaema and Lysiloma in the Bahamas.
Vouchers are indicated as follows: ep = epidermal peel, Is = - Taxon 23: 185-191.
leaf section, m = macerate of foliar sclerenchyma, p = mea- Hall&,F. & Oldeman, R. A. A. 1970. Essai sur I’architecture et
surements of pollen, sc = SEM of leaf surface. Specimens used la dynamique de croissance des arbres tropicaux. - Mason
for Fig. 2 are indicated by an “!”. et Cie, Paris.
- , Oldeman, R. A. A . & Tomlinson, P. B. 1978. Tropical
Theophrasta americana trees and forests. An architectural analysis. - Springer,
Dominican Republic. - Distr. Nacional: Stdhl & Lindstrom Berlin.
178, GB (ep, p, sc, !); Pelaez Feucht 1104, JBSD. - San Hamilton, G . 1825. Prodromus plantarum Indiae occidentalis.
Cristobal: Liogier & Liogier 21591, F, JBSD (ep, m, !), NY. - - London.
San Pedro de Macoris: Taylor 141, NY. - Altagracia: Mejia & Hoffmannsegg, J. C. 1826. Zweiter und Dritter Nachtrag zu
Zanoni 5079, JBSD (Is, p, !). - Pedernales: Howard & Howard dem Verzeichniss der Pflanzenkultur. - Dresden.
8202, BM, GH(!), NY, S, US; Zanoni et al. 27078, JBSD. - Holmgren, P. K., Keuken, W. & Schofield, E. 1981. Index
Barahona: Fuertes 1097, A(!), NY. - Independencia: Stihl & Herbariorum 1. The herbaria of the world, ed. 7. - Reg-
Lindstrom 237, GB; Stdhl & Lindstrom 238, GB (ep, !), JBSD. num Veg. 106.
- Elias PiAa: Stdhl & Lindstrom 205, GB; Zanoni 32948, Jacquin, N. J. 1790. Collectanea, etc. 4. - Wien.
JBSD. - San Juan: Liogier 14696, NY(!); Miller 1239, US (ep, - 1809. Fragmenta botanica, etc. - Wien.
!). -La Vega: Liogier 11327, NY(!). -Duarte: Abbot 2109, US Kisser, J. 1931. Die Venvendung von Eau de Javelle und
(ep, !). - Santiago: Ekman H13232, IJ, S ( ! ) ; Jimenez 2348, Wasserstoffsuperoxyd als Mazerationsmittel fiir Pflanzen-
US(!). - Puerto Plata: Eggers 1560, BM, G , HBG (Is, !), K, L, gewebe. - Cytologia 2: 56-66.

Nord. J. Bot. 7 ( 5 ) 1987 537

Koch, C. & Fintelmann, G. A. 1859a. Neue Linden’sche Pflan- ren von Deherarnia smaragdina. - Jahrb. Wiss. Bot. 75:
Zen. - Wochenschr. Gartnerei Pflanzenk. 12: 90-94. 481493.
- 1859b. Theophrasta und Clavija. Unverhtelte Baume mit Polhill, R. M. & Steam, W. T. 1976. Linnaeus’s notes on
grossen Blattern, besonders Thephrasta spinosa Willd. und Plumier drawings with special reference to Mimosa latisili-
Clavija latifolia C. Koch. - Wochenschr. Gartnerei Pnan- qua. - Taxon 25: 323-325.
zenk. 13: 98-100, 14: 108-110. Radlkofer, L. 1890. Ueber die Gliederung der Familie der
Kuntze, 0. 1891. Revisio generum plantarum etc. 2. - Leipzig. Sapindaceen. - Sitzungsber. Math.-Naturwiss. KI. Bayer.
Lindley, M. 1821. Collectanea botanica, etc. - London. Acad. Wiss. Miinchen 20: 105-379.
Linnt, C. von 1737. Genera plantarum, ed. 1. - Leiden. - 1933. Sapindaceae 1. - In: Engler, A. (Diels, L.), Das
- 1753. SDecies Dlantarum 1. ed. 1. - Stockholm. Pflanzenreich 98 (IV 165). DD. 1-1018.
Martius, M. C. F.’P. von 1841’. Theophrasta tetramera Mart. - Regel, E. 1864. Theophrastari&ierialis Linden. - Gartenflora
Flora 24 (2; Beibl.): 27. 13: 323, Tab. 453.
Merrill, E. D‘. 1935. A commentary on Loureiro’s “Flora Co- Roemer, J. J. & Schultes, J. A. 1819. Systema vegetabilium
chinchinensis”. - Trans. Amer. Philos. SOC., new ser., etc. 4. - Stuttgardt.
24(2): 1445. Ruiz, H. & Pavon, J. 1794. Flora peruvianae, et chilensis
Mez, C. 1901. Theophrastaceae. - In: Urban, I., Symbolae prodromus, etc. - Madrid.
antillanae 2, pp. 434451. - 1799. Flora peruviana, et chilensis, etc. 2. - Madrid.
- 1903. Theophrastaceae. - In: Engler, A., Das Pflanzen- Smith, J. 1817. Theophrasta. - In: Rees, A., The Cyclopaedia
reich 15 (IV 236a), pp. 1 4 8 . 35(2).
Moscoso, R. M. 1943. Catalogus florae domingensis. Parte I. Sprengel, K. 1825. Systema vegetabilium 1. - Gottingen.
Spermatophyta. - New York. Urban, I. 1902. Notae biographicae peregrinatorum Indiae
Oliver, D. 1888. Talisia princeps, O h . - Hooker’s Icones occidentalis botanicorum. - In: Urban, I., Symbolae antil-
plantarum 8: Tab. 1769. lanae 3, pp. 14-158.
Otto, F. 1837. Beitrag zur Cultur der Clavija ornata Don (The- - 1920. Plumiers Leben und Schriften. - Fedde, Repert. sp.
ophrasta Iongifolia Jacq.), Theophrasta Jussieui Lindl. und nov. reg. veg., Beih. 5, pp. 1-196.
Brexia madagascariensis Pers. - Allg. Gartenzeitung 5: Vogel, S. 1978. Pilzmiickenblumen als Pilzmimeten. - Flora
281-282. 167: 325L398.
Plumier, C. 1703. Nova plantarum Americanum Genera. - Votsch, W. 1904. Neue Systematische-anatomische Untersu-
Paris. chungen von Blatt und Achse der Theophrastacken. - Bot.
- 1757. Plantarum Americanarum etc. (ed. J. Burmannus). - Jahrb. Syst. 33: 502-546.
Leiden. Vriese, W. H. de 1839. Hortus Spaarn-bergensis. - Amster-
Pohl, F. 1931. Uber sich offnende Kristallraume in den Anthe- dam.

538 Nord. 1. Bot. 7 ( 5 ) 1987