BotanicalJournal ofthe Lannean Sockp (1997), 125: 273-293
Typification of nineteen names of K c a n
Solanurn species described by A. Richard
and others, including S. campylacanthum
and S. panduriforme
RICHARD N. LESTER
School of Biological Sciences, Universip of Birmingham, Birmingham B15 2TT
Receiued October 1996; accepted for publication June 1997
The Paris Herbarium (P) contains many important but neglected specimens, collected from
Abyssinia (now Eritrea and Ethiopia) by Quartin-Dillon and Petit, as well as by Schimper,
and first described as new species by Achille Richard. A brief account is presented here of
the circumstancesof the collection, description and publication of these. Definitive typification
and nomenclatural details are provided for twelve names of Abyssinian Solanurn species
described by A. Richard in 1850 in Paris, and of five other African species described by
Dunal(l852) and of two others. These are S.adoense Hochst. ex A. Rich., S. b@rcaturn Hochst.
ex A. Rich. (=S.b$murn Hochst. ex Dunal), S. carnpylacanthurn Hochst. ex A. Rich., S.
grossidentaturn A. Rich., S.hirtulurn Steud. ex A. Rich., S. milenturn A. Rich., S. mamuanthum
A. Rich., S. p$mymrn A. Rich., S. plebeiurn A. Rich., S. pobanthmurn Hochst. ex A. Rich., S.
schirnperianurn Hochst. ex A. Rich. and S.unguiculaturn A. Rich., and also S. albicauk Kotschy
ex Dunal, S. a'elagome Dunal, S. hudelotii Dunal, S. pandurifom E. Mey. ex Dunal and S.
senegarnbicurn Dunal, and also S. kotscbii Steud. (ined.) and S. senegahe Perr. (ined.). Some of
these are synonyms of S. anguivi Lam., Sforskalii Dunal, S.incanurn L., S. teninale Forssk. and
S.uillosurn Mill. Hitherto, places of publication of several of these names, and the authorship
of many of them have been cited incorrectly.
0 1997 The Linnean Society of London
ADDITIONAL KEY WORDS:-Abyssinia
taxonomy - typification.
- Ethiopia
-
Solanaceae - Solanurn incanurn -
CONTENTS
Introduction . . . . . . . . . . . . . . . . . . .
Taxonomic history of some African Solanurn species . . . . .
Typification and nomenclature of Solanurn species . . . . . .
Solanurn adome Hochst. ex A. Rich. . . . . . . . . .
Solanurn bajkrcaturn Hochst. ex A. Rich. . . . . . . . .
Solanurn carnpylacanthurn Hochst. ex A. Rich. . . . . . .
Solanurn pandunyome E. Mey. ex D u d ( = S. delagoense Dunal)
Solanurn gmssidentatum A. Rich. . . . . . . . . . .
Solanurn hirtulurn Steud. ex A. Rich. . . . . . . . . .
Solanurn rnacilentum A. Rich. . . . . . . . . . . .
Solanurn rnacracanthurn A. Rich. . . . . . . . . . . .
00244074/97/120273
+ 21 $25,00/0/bt970126
273
. . . .
. . . . .
. . . . .
. . . . .
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
. . . . .
274
274
280
280
281
282
283
285
285
286
288
0 1997 The Linnean Society of London
274
R. N. LESTER
Solanuin p$enjruni A. Rich.
. . . . . . .
tSolanutn pkbeium A. Rich. . . . . . . . .
Solanum pobanthnnum Hochst. ex A. Rich. . . .
Solununz .srhii?@erianum Hochst. ex ‘1.Rich. . . .
Solunutn unguiculahcrn A. Rich. . . . . . . .
Other Solanurn species . . . . . . . . .
Acknowledgemcnts . . . . . . . . . . . .
Referenccs . . . . . . . . . . . . . . .
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
. . . . . . . .
. . . . . . . .
.
.
.
.
.
.
288
289
290
290
291
292
292
292
IKTRODUCTIOK
The genus Solanum is pantropical and very large, with over 1000 species and
about 5000 names. Solanum is ridden with taxonomic conlusion and we are still far
from a comprehensive and conclusive taxonomic treatment of the whole genus.
Even in Africa, where there are probably fewer than 150 species, there is serious
disagreement between different authors. Although Jaeger (1985) worked hard and
long to produce a synoptic kiew of Solanum throughout Africa, his treatment does
not resolve several important taxonomic problems: it is uncertain of the priority of
several commonly used names, and it lacks proper typification of many of these
names, let alone many others which are probably synonyms. Ideally typification
should be included within a taxonomic revision or monograph, but since a complete
revision of Solanum in Africa is unlikely in the forseeable future, 19 species names
arc here typified, many of which have been neglected or misinterpreted until now.
The main result of the present work is to show that several species originally
named by Hochstetter or Steudel, and commonly considered to have been first
described properly by Dunal ( 1 852), had in fact already been described by Richard
(1850). Names of most of these have not hitherto been typified properly, even
following Dunal, so this is done here appropriately according to Richard, almost
entirely involving lectotypification using specimens in the Paris herbarium (Table
1). Besides these, names of several species both named and described by Richard
are typified, some of which have been neglected, although others have already been
correctly reduced to synonymy. At the same time typification is here effected for
several other species names represented at Paris, most of which are also synonyms.
The particular cases of Solanum campylacanthum, S. delagoense and S. pandurZfomze are
dealt with properly here for the first time. Although they have been mentioned in
some previous publications, consideration of their type specimens has now resolved
important taxonomic and nomenclatural problems that were hitherto confusing or
at least uncertain.
This paper also draws attention to the excellent Solanum collections of QuartinDillon and Petit and others in Paris, and the neglected work of Achille Richard,
which is an important foundation for African plant taxonomy.
.lSONOJIIC HISTORY OF SOME AFRICAV SOMVLLi1 SPECIES
Intensive exploration of Africa, for both scientific and political purposes, was
undertaken in the 19th century by expeditions from Britain, France, Germany and
other nations, many of which collected vast numbers of botanical specimens which
are the basis of African plant nomenclature today.
TYPIFICATION OF A. RICHARD'S AFRICAN SOLRn/iM NAMES
275
TABLE
1. Typification of some names of African Solanun species described by Richard (1850) and
Dunal (1852)
Species describcd by A. Richard
Modern taxonomic designation
Page references from Richard A (1850). Tentamen Florae Abyssinicae. Vol. 2. Arthus Bertrand, Pans.
S. adoense Hochst. ex A. Rich. @. 105)
Schimper 147, Abyssinia, in valleys, near Adoa, 26 May 1837
lectotype P!, isolectotpes P!, P!, K!, G-DC (IDC Microfiche 2083/4!)
[S. adoense Hochst. ex A. Rich.]
S b@rcatum Hochst. ex A. Rich. @. 98)
[S.terminale Forssk. subsp. terminale]
Schimuer 201., Abyssinia.
near Adoa
,
lectotype P!, isolectotypes P!, P!, P!. P!, P!, K!, BM!, G-DC (IDC Microfiche
2067/3!)
S. campylacanthum Hochst. ex A. Rich. @. 102)
[S. campylacanthum Hochst. ex A. Rich.]
Schimper 1082, Abyssinia, Adoa, 25 Nov. 1838
lectotype P!, isolectotypes P!, K!, K!, BM!, G-DC (IDC Microfiche 2088/
7!\
S. gmssidentatum A. Rich. @. 101)
Quartin-Dillon and Petit, Abyssinia, TchClikote province.
lectotype F!, isolectotype P!.
[?? S. uillosum Mill.
??I
S. hirtulum Steud. ex A. Rich. @. 101)
[S. hirtulum Steud. ex A. Rich.]
Schimper '977, Abyssinia, Semiene province, in fields near Enschadcap,
29Jan. 1838
lectotype P!, isolectotpes P!, P!, K!, BM!, BM!, G-DC (IDC Microfiche
2061 /5!), G-BOIS, MPU.
S. man'lentum A. Rich. @. 105)
Quartin-Dillon and Petit, Abyssinia, Choho province.
lectotype P!, isolectotype P!.
[S. fonkalzi Dunal]
S. macracanthum A. Rich. @. 106)
Quartin-Dillon and Petit, Abyssinia, Choa kingdom.
lectotype P!, isolectotypes P!, P!, P!.
[S. macracanthum A. Rich.]
S. pipenj%mmA. Rich. @. 106)
Quartin-Dillon and Petit, Abyssinia, Tcheliiote province.
lectotype P!, isolectotypes P!, P!, P!, P!, P!, K!.
[S. p;Pmjmm A. Rich.]
S.plebeium A. Rich. @. 100)
[?? S.uillosum Mill.
subsp. miniatum (Bernh. ex Willd.)
Edmonds ??I
Quartin-Dillon and Petit, Abyssinia, Chire province.
lectotype P!, isolectotype P!.
S. po(yanthemum Hochst. ex A. Rich. (p. 99)
[S. schimperianum Hochst. ex A. Rich.
Schimper 1526, Abyssinia, near h u m .
var.polyanthemum(Hochst. {exA. Rich.}
lectotype P!, isolectotypes P!, K!. BM!, G-DC (IDC Microfiche 2068/24!) ) Bitter]
S. xhimperianum Hochst. ex A. Rich. @. 98)
Schimper 202, Abyssinia, in mountains near Adoa, 5 June 1837
lectotype P!, isolectotypes P!, BM!, G-DC (IDC Microfiche 2068/23!)
[S. schimperianum Hochst. ex A. Rich.]
S. unguiculatum A. Rich. @. 102)
Quartin-Dillon and Petit, Abyssinia, Chire province.
lectotype P!, isolectotpes P!, P!.
[S.incunum L. var. unguiculatum (A. Rich.)
Bitter]
R.N.LESTER
276
TABLE
1. Continued
Species described by Dunal or others
~
~
p o d e r n taxonomic designation]
~~~
Page and number references from Dunal MF (1852). Solanaceae. In: de Candolle AP (ed) Aodmmus Systemutzs
nuturultr Regni uegetu6ilu 13 (1): 1 4 9 0 . Pans.
[S. pundun@mt E. Mey. ex Dunal]
S. pundunifonne E. Mey. ex Dunal @. 370, no. 851)
Drege s.n., South Africa, Transkei, banks of the River Mbasche, below
IOOOft., in scrub. p o t Cape of Good Hope]. January 1832.
holotype G-DC (IDC Microfiche 2092/13!), isotypes P!, K!, K!.
S. delugome Dunal (p. 349, no. 802)
Forbes, Mozambique, Delagoa Bay (=B. de Maputo) 1822.
holotype G-DC (IDC Microfiche 2089/12!), isotypes K!, K!, K!, K!, K!.
[S. pandunfonne E. Mey. ex. Dunal]
S. ul6traub Kotschy ex Dunal @. 204, no. 500)
[S.forshlii Dunal]
Kotschy 309, Nubia, Cordofan, Chuni, in savanna. 29 Dec. 1839.
lectotype (G-DC, Microfiche 2078/8!), isolectotypes MPC, P!, P!, P!, K!.
S.heudelotii Dunal (p. 205, no. 501)
[S.forskalii Dunal]
Heudelot 417, Senegambia, near Gabor, 1837.
lectotype P!, isolectotypes P!, P!, K!, OXF!.
.S. kotrchyzi Steud (ined.)
[S.forskalii Dunal]
Kotschy s.n., Xubia.
holotype (?) P!
S. senegalenre Perr. (iied.)
Perrottet 340, Senegal, 10 Sept. 1824.
holotype (?) P!
[S.forskalii Dunal]
S. saegam6uum D u d @ . 194, no. 473)
Heudelot 7 13, Senegambia
holotype P!
[S.unguiui Lam.]
Apart from several minor earlier works (see Stearn, 1982),the first comprehensive
botanical treatment of Ethiopia and Eritrea (then called Abyssinia) was Achille
Richard’s Titamen Florae Abyssinicae (‘An attempt at a Flora of Abyssinia’) published
between 1847 and 1851, consisting of two volumes of text and one volume of plates.
Here, many species were described properly for the first time. Although Richard
deprecatingly referred to his work as merely an essay or an imperfect sketch of the
botanical riches of Abyssinia, it was in fact an extensive and very valuable work,
especially when used in conjunction with his herbarium in Paris. Sadly, both of
these have been overlooked, although the recent facsimile reproduction of the former
(Richard, 1982) should go some way towards redressing this neglect.
Achille Richard (17961852) and his father L.-C.-M. Richard (1 754-1821) were
eminent plant taxonomists (Boiteau, 1976), who together created one of the most
important herbaria in Paris in their time. The Richard Herbarium was bought after
Achille’s death by Franqueville in 1856; it was then incorporated into the herbarium
of Drake del Castillo in 1891, and subsequently into the Herbarium of Paris in
1905 (Bureau, 1904;Jolinon, pers. comm.). It is now part of the Museum d’Histoire
naturelle. The Herbarium received specimens from many sources, but a particularly
rich supply of Afiican material came from an expedition to Abyssinia during
1838-43. In the preface to the Tintamen Richard included a sensitive and moving
tribute to his two young friends and colleagues, Richard Quartin-Dillon and Antoine
TYPIFICATION OF A. RICHARD’S AFRICAN SOLGNLiM NAMES
277
Petit, who died whilst collecting the specimens which were later described in the
book.
Naturalists who had initially trained in medicine, Richard Quartin-Dillon, a
student of Achille Richard, was the expedition’s botanist and Antoine Petit its
zoologist. The team also included ThCophile Lefebvre, a lieutenant in the Royal
Marines, responsible for surveying physical geography, ethnology and history. The
expedition members set out from Paris at the end of 1838, arriving in Abyssinia in
June 1839. They settled first at Adoua, capital of TigrC, which they explored for 6
months, extending their excursions into the valley of TaccazC and the province of
ChirC. From Adoua they sent back the first consignment of plants and birds (Lasegue,
1845; Richard, 1840, 1847). During 1840 Quartin-Dillon visited the province of
Samen, the kingdom of Amhara, and SirC. In 1841 he returned to Adoua to put
his new collections in order, and to prepare to explore other parts of Abyssinia with
Petit. In October 1841, he and Petit went down into the valley of the Mareb, one
of the principal rivers of Abyssinia, against the advice of local people who knew
dangerous fevers to be endemic in that hot and humid region. After only a few days
both explorers were laid low by fever, to which Quartin-Dillon succumbed on 22
October 1841. Petit was more fortunate, but recovered his health only after a
lengthy convalescence (Lasegue, 1845).(Note: Richard, 1847 and Stearn, 1982 state
that Quartin-Dillon died in 1840, but Lasegue (1845) and Stafleu & Cowan (1983)
specifSI 1841, which accords better with other information in Richard (1847)).
During a stay of about 2 years, Quartin-Dillon had collected more than 1000
plant species; these were obtained principally in TigrC, in the vicinity of Adoua, and
in the valley of TaccazC, especially on the lofty slopes, around the village of
TchClatchtkannt (one of the richest localities in Abyssinia). He also found specimens
around Gondar, capital of Amhara, in the mountainous province of Semiene, and
finally and fatally in the valley of the Mareb.
Following the tragic demise of Quartin-Dillon, Petit took over the botanical as
well as the zoological research. During 1842 he visted the provinces of Lasta, Schoa
(Choa) and parts of the kingdom of Amhara; he also made valuable plant collections
in the provinces of Ouodgerate, Enderta, TchClenkote and Yedjou. Sadly, he did
not long survive his partner. On 3 June 1843, when finally returning to Gondar,
he was seized by a crocodile while attempting to swim across the Nile. Subsequently,
Thtophile Lefebvre managed to collect some additional specimens before returning
alone to Europe in early 1844 (Ladgue, 1845; Richard, 1847). The total number
of plant species collected was about 1500, of which about 75% were new to science.
Most of their specimens went to the Herbarium either directly or sequentially.
However, some duplicates were sent to the herbaria of both Cosson and Franqueville
at Paris, as well as to Kew, and also Delessert, now at Geneva (Vallot, 1882: 187).
Richard received the first consignment of about 600 species from Quartin-Dillon
in May 1840. He promptly set about studying and determining them and published
an initial selection of 20 species the following November, naming three of the new
genera in honour of the three young naturalists (Lasegue, 1845; Richard, 1840). He
continued to work on the collections over the next few years; recognizing that almost
all of the species were new to science, he gave them adequate descriptions and new
names. It was difficult work, because the collectors’ notes accompanying the
specimens were minimal. Quartin-Dillon and Petit could not have anticipated their
untimely deaths and had doubtless committed much valuable information to memory:
most specimens lack collectors’ numbers or even dates. However, by scrutinizing
278
R. N. LESTER
the journals of their travels, Richard was able to fill in much useful information.
He further supplemented his species descriptions and notes from other sources such
as Vignaud’s plant portraits from Abyssinia, as well as several other sources such as
the Prodrornus of De Candolle, Flore de S@gambie and various collections from
Abyssinia. He also referred to reports on these by Bruce, Salt, Brown, Ruppell,
Fresenius, d’Hericourt, Feret, Galinier and Delile (Richard, 1847; Stearn, 1982).
A major additional source of information, but one which also presented Richard
with a dilemma, was the three series of specimens collected by the prodigious
German explorer and botanist Lt‘ilhelm Georg Schimper from 1837 onwards for
the Botanical Society of Esslingen. These were carefully named and published (but
not described),by C. F. Hochstetter and E. G. Steudel, and distributed with printed
labels to many subscribers, including most of the major European herbaria. ‘The
first three series comprised many thousands of specimens, numbering around
1900 collections; occasionally, a single number included specimens from different
populations or even from different species, and sometimes specimens of different
collections were interchanged (e.g. between Schimper 20 1 and 97 7 at Paris).
Subsequently, Schimper made 15 more series of collections in Abyssinia, but these
Lvere destroyed when his house was sacked (note by Cufodontis in the Kew copy of
Richard [ 18471). Altogether, Schimper’s first three series of collections comprised
about 1600 species.
Richard had already recognized, described and named about 1500 species which
he generally attributed to both Quartin-Dillon and Petit, when he received the
materials collected by Schimper. By careful comparison he found that most species
were represented by both sets of collections, although about 200 species had been
discovered by Quartin-Dillon and Petit but not by Schimper, and a similar number
vice versa. Since the Schimper collections were merely named but not yet described,
the names were not validly published according to our present Code, and Richard
could have described all these new species with the names he had already given
them. However, before printing his flora, he ma<gnanimouslyexpunged his own
names in favour of all the names applied to Schimper’s specimens by Hochstetter
and Steudel, except for those species not found by Schimper. The published
descriptions were therefore based mostly on the excellent and abundant specimens
collected by Quartin-Dillon and Petit, with additional information about localities
and flowering times obtained from Schimper. Richard cited both sets of specimens,
but in those cases where he adopted the Hochstetter or Steudel name he attributed
it to that respective author and gave pre-eminent citation to the Schimper specimens.
However, it should be borne in mind that it was Richard who first described these
species, so the attribution should be, for example, Solanurn campylacanthurn A. Rich.,
or at most Solanum campylacanthuni Hochst. ex A. Rich. Since in most cases Richard
cited several specimens,lectotypificationis necessary. His descriptions being primarily
based on the collections of Quartin-Dillon and Petit, it is generally appropriate to
designate one of their specimens as lectotype, with any duplicates as isolectotypes,
and any conspecific collections also cited by Richard, whether from Quartin-Dillon
and Petit or from Schimper, as lectoparatypes. However, further complications were
caused by Cufodontis (1963) who listed all of the species described by Richard,
frequently noting that the type specimens were at Pans, and even citing some
collectors and numbers. However, it seems that he never examined the Solanum
specimens at Paris, and certainly did not annotate them authoritatively.For example,
for Solanuni adoense Hochst. ex A. Rich. Cufodontis cited only Schimper 147 as the
TYPIFICATION OF A. RICHARD’S AFRICAN SOZANUM NAMES
2i‘J
type; this may be considered as a partial typification and I therefore here complete
the typification by choosing a lectotype from the three specimens of Schimper 147
at Paris, even though I would have preferred to have used one of the QuartinDillon and Petit specimens. Although Richard (1847) replaced his own names with
those given by Hochstetter and Steudel to Schimper’s first three series of specimens,
he warned that any names given to future Schimper series would be considered
void. In the event, all subsequent collections by Schimper were destroyed and never
reached Europe.
Richard (1850) listed 18 species of Solanum from Abyssinia, with descriptions in
Latin and additional notes in French. Six of these were already well known (e.g. S.
tuberosum), but the other 12 were new, six of which were definitely his own new
species names (designated by the sufKx ‘Nob.’=nobis =ours). For the remaining six
he used the available names provided by Hochstetter (5) or Steudel(1) for Schimper’s
specimens, and acknowledged them. However, strictly speaking, because he published
the first validating descriptions these names must now be attributed to him as
‘Hochst. ex A. Rich.’ or ‘Steud. ex. A. Rich.’ or merely ‘A. Rich.’. Since these
names were validated in 1850 (Stafleu & Cowan, 1983), they take precedence over
the subsequent use of these same names by Dunal (1852).
A further consideration is that whereas the descriptions of Dunal were based on
the relatively few Schimper specimens available to him, the descriptions of Richard,
in contrast, were derived from examination of many excellent specimens collected
by Quartin-Dillon and Petit as well as those of Schimper. The availability of more
collections and specimens of each species not only allowed Richard a better
understanding of the species, but also provided us with a wider choice of syntypes.
This is particularly important in the case of Solanum campylacanthum, as discussed
below.
Unfortunately, when Dunal’s work was published in May 1852, he, and thus
many subsequent taxonomists, ignored the earlier and highly relevant publication
of Richard (1850). It is a pity that Dunal and Richard did not confer before their
respective publications, which could have prevented the problems now addressed
in this paper. Amongst all the specimens at Paris of the species discussed here, there
is only one annotated by Dunal; it is not significant and was not collected by
Quartin-Dillon and Petit.
None of the specimens cited here were annotated by Bitter (1913, 1917, 1921,
1923), although a few other Paris specimens of the same species were seen by him,
possibly on loan to Berlin.
The nomenclatural significance of these specimens, and also of Richard’s (1850)
publication, has been ignored generally by Solanum taxonomists, eclipsed by the
magnum opus of Dunal (1852). It is probable that the type specimens of many names
of species of other plant families, first effectively published by Richard, still await
discovery in Paris.
Cufodontis (1963) recognized the priority of Richard’s publication (which he
incorrectly dated 1851), notably for S. campylacanthum Hochst. ex A. Rich. His
synopsis gave basic nomenclatural details and suggestions for type collections as
published by Richard (1 850), but almost certainly he did not see the type specimens
at Paris (W. Stearn, pers. comm.). He treated 84 species of Solanum from Ethiopia,
including almost all those discussed here, but made few taxonomic judgements, and
stated the need for a complete revision of the whole genus.
Complete taxonomic revision of Solanurn in all parts of Africa is still required.
280
R. N. LESTER
Meanwhile the extensive but tentative conspectus ofJaeger (1985) requires piecemeal
additions such as this present contribution. This paper is based upon examination
of all the African Solanum specimens at Paris, especially those from Abyssinia. Many
type specimens have been recognized, as detailed below, some of which had been
vaguely recognized previously, but not sufficiently to count as effective typification
(W. Stearn, pers. comm.). All type specimens cited here have now been labelled
and rearranged appropriately, and photocopies, kindly donated by Paris, have been
deposited at K and ETH.
All the Latin species descriptions of Richard referred to here have been examined
by the present author and found to be satisfactory. Additional notes presented here
as quotations were translated from the original Latin and French (or German).
However, any verbal descriptions of Solanum species, such as those of Dunal, or even
Bitter, are inadequate without reference to type specimens.
While the main objective of this paper is the typification of names of species
described by Richard (1850), this opportunity has also been used to discuss their
present nomenclatural position. Jaeger’s comprehensive conspectus (1985), which
covers virtually all species of Solanum from the whole of Africa and adjacent areas,
has proved to be of immense value in this task, even though few names are properly
typified. It encompasses all relevant information from the major works of Dunal
(1813, 1852) and Bitter (1913, 1917, 1921, 1923), but also, and in some ways more
importantly, it reviews all the minor treatments of Solanum species in different parts
of Africa, which, whilst often based on adequate field observations, and therefore
describing real species, frequently generated synonymy and other nomenclatural
problems. By treating all species from all of Africa, Jaeger (1985),Jaeger & Hepper
(1 986) and Jaeger & Lester (unpubl.) have produced a relatively comprehensive
and well balanced taxonomic treatment which provides a sound basis for future
investigations of various species in various parts of Africa or in various parts of the
genus Solanum. However, many still require typification, and this is attempted here
for a few of these species (Table 1). While this paper is concerned chiefly with
typification of names of Solanum species first described by Richard, some interlinked
species described by Dunal and others are also considered.
TYPIFICATION AND NOMENCLATURE OF SOLRlwM SPECIES
Solanum adoense Hochst. ex A. Rich.
7yptjication
Besides the formal Latin description of “S. adoense Hochst.” (citing Schimper 147
and 1865), Richard stated:
It grows in hills and waste places near Adoua, flowering and fruiting in the month
of September (Quartin Dillon).
Observation.-The specimens which I have under my eyes, and which were
collected in the neighbourhood of Adoua, by Dr Quartin Dillon and by Mr
Schimper, appear to me to have much resemblance with the species which Jacquin
has described and figured in his Hortur Schoenbmnnensk, under the name of Solanum
r@escm. That species was first observed at the Cape of Good Hope.
TYPIFICATION OF A. RICHARD’S AFRICAN SOLeNuM NAMES
28 1
Richard referred principally to the three specimens collected by Quartin-Dillon (and
Petit), Abyssinia, Adowa, 17 September 1839. All three are from the Herbarium
Richard, and were previously named “S. rigescens Jacq.” On one “h. Schon t. 42”
was added to this name, but later both were deleted and replaced by “S. adoense
Hochst. I. 147., 111. 1865.” This main specimen has both flowers and fruits. One
of the other specimens, which appears to have been broken from the main plant
and is definitely a second sheet of the same collection, has the number ‘361’, which
may be the collection number, on the label. The third sheet might have been taken
from the same plant.
Since Richard paid more attention to the Quartin-Dillon and Petit collection,
one of these specimens might have been chosen as lectotype. However, Cufodontis
(1963), without visiting Paris, cited Schimper 147 as type. This may be considered
as a partial lectotypification, and so I have chosen one of the Schimper 147 specimens
at Paris, with the best leaves, flowers and fmits, as lectotype, and have labelled it
appropriately: the other four specimens of this collection are thereby isolectotypes,
as also are those at K (!), G-DC (IDC Microfiche 2083/4!) and elsewhere. The
specimens of Schimper 1865 at Paris (!), Kew (!), G-DC (IDC Microfiche 2083/5!),
as well as those collected by Quartin-Dillon and Petit from Adowa, 17 September
1839 at Paris (!), are lectoparatypes or isolectoparatypes.
Nomenclature
Based on Schimper 147 and 1865 (at MPU, G-DC and Delile), Dunal (1852)
described this taxon as S. adoense Hochst. Bitter relegated it to S. indicum L. subsp.
adoense (Hochst.) Bitter (citing Dunal’s description, and specimens of Schimper 147
and 1865, probably at B).Jaeger called it S. adoense Hochst. ex Dunal (citing Schimper
147 and 1865 at G-DC). However, Cufodontis (1963) realised that it should correctly
be called S. adoense Hochst. ex A. Rich. S. adoense is distinct from S. anguivi Lam.,
but the same as S. uollense (Chiov.) Pic. Serm. (basionym S. indicum auct. non L.
subsp. mesodolichum Bitter var. uollenris Chiov. 1939; holotype FI!, isotype K!).
Solanum bajircatum Hochst. ex A. Rich.
Gpijication
Besides the formal description of “S. bajircatum Hochst.” (citing Schimper 20 l),
Richard stated:
It grows in rugged places and valleys around Adoua, flowering and fruiting in
June (Schimper) and in Ouodgerate province (Ant. Petit).
Observation.-Here again [is] a neighbouring species of the bitter-sweet [i.e. S.
dulcamara], but that is easy to distinguish. First its leaves are all entire and not
lobed; its flowers are much more numerous, with the corolla tomentose on the
external surface, and its fruits globular and not elongated. It offers also a certain
resemblance with Solanum schimpmanum,but its leaves are more elongated, attenuated
at their base, less hairy; the flowers are smaller, less numerous, and the lobes of
the corolla deeper and narrower.
The lectotype chosen here (Table 1) has two pieces with flowers and fruits respectively,
282
R. N. LESTER
which were collected by Schimper No. 201 from Abyssinia, near Adoa. On the
label is also written “W. Schimper pl. Abyssin. Ed. 11. Hohenack. 1852” and “201
Solanum bfurcum Hochst.” This sheet was successively in the herbaria of Steudel,
Richard, Drake and Paris. There are isolectotypes on five other sheets (but also an
admixture from another collector; there is also another sheet of a different species,
S.hirtulum, which is labelled Schimper 201, but is probably Schimper 977). There
are two lectoparatypes collected by Quartin-Dillon and Petit, from Oudgerate.
There are isolectotypes at G-DC (IDC Microfiche 2067/3!), K (!), and probably
elsewhere.
.,Vomenclature
Based on Schimper 201 (at G-IX, and probably at B, respectively) both Dunal
and Bitter called this taxon S. bzjircum Hochst., but Bitter also gave S. bgurcatum A.
Rich. as a synonym.Jaeger gave both S, b@rcum Hochst. and S. b@rcatum A. Rich.,
without typification, as synonyms of S. terminale Forssk. subsp. terminale, a name
recocgnized by Heine (1960), but not by Bitter or Dunal. This appears to be the
correct name for this taxon.
The two apparently different names S. bfurcum and S. bfurcatum are in fact merely
orthographic variants. Inspection of the specimens in Paris (placed under S. terminale
by Heine, but now relocated under S. bzjircatum) suggests that Richard saw specimens
of Schimper 201 labelled S. bijiurcum Hochst., and thereby identified the specimens
of Quartin-Dillon and Petit as the same species. However, when published by
Richard, the name was printed as S. b@rcatum Hochst. Further inspection of
specimens and documents at Paris might reveal when Richard changed the spelling
of this name, which should be cited as S. bijiurcatum Hochst. ex A. Rich. (fide
Brummitt), despite the note to the contrary by Milne-Redhead (1949).
Although Heine (1960) included too many taxa within S. terminale, including very
distinct species such as S. welwikchii C. H. Wright (Jaeger, 1985), S. bfurcatum
Hochst. ex A. Rich. (and S. bzjiurcum Hochst. ex Dunal) are definitely synonyms of
S. terminale Forssk. subsp. terminale, as noted by Cufodontis (1963).
Solanum campylacanthum Hochst. ex A. Rich.
@$cation
Besides the formal description of “S. campylacanthum Hochst.” (citing Schimper
123 & 1082), Richard stated that:
It grows in Chirt. province (Quartin Dillon) and near Adoua (Quartin Dillon and
Schimper) flowering in May and fruiting in November.
Vernacular names: in Tigre language: Anggouldi; and in Amhara language: Amboie.
Observation.--The species which M. Hochstetter has named S. campylacanthum
does not differ much from Solanurn sanctum L. or S. incanum Forsk. Only the colour
of the tomentum, which covers almost all the parts of the plant, is rusty-green and
not white; its flowers [are] bigger and its berries larger.
Although Richard made ample reference to the Quartin-Dillon (and Petit) collections,
and some of these at Paris are very good specimens, it may be preferable to choose
TYPIFICATION O F A. RICHARD’S AFRICAN SOLANL!V NAMES
283
a Schimper collection as type, because these are represented more widely at other
herbaria, where they have been seen by Dunal, Bitter and others. Cufodontis cited
both Schimper 123 and 1082 as syntypes, but he probably did not see the Paris
specimens, and in any case this does not even constitute a partial lectotypification,
and can be ignored.
One specimen of Schimper 1082 at Paris, which has good flowers and fruits and
best fits Richard’s description and the Quartin-Dillon and Petit lectoparatypes, has
now been designated and labelled as the lectotype for S. campylacanthum Hochst. ex
A. Rich., and another is an isolectotype, but a third sheet of that collection is
excluded, because it seems to be a different species, namely S.incanum L. The two
isolectotypes at Kew (!) are like the lectotype, but that at G-DC (IDC Microfiche
2088/7!) has very narrow leaves. Thus Schimper 1082 was a mixed collection, from
various plants in the general region of Adoa, but from it can be selected the
lectotypes and some isolectotypes which accord with Richard’s concept of S.
campylacanthum, and which are very similar to the excellent Quartin-Dillon and Petit
lectoparatypes.
The lectoparatype collections of Schimper 123 at P (!), K (!) and G-DC (IDC
Microfiche 2088/6!) are uniform, but atypical in having slightly narrower leaves
and very prickly stems.
Of the Quartin-Dillon and Petit specimens at Paris, there are four which qualify
as lectoparatypes, and all four are very similar to the lectotype. One is an excellent
specimen with both flowers and fruit, collected in the valley of Adowa [sic!] 25
August 1839; two were from ChirC, four from Adoa, and also one specimen with
merely Abyssinia as locality but with a small note “4e. Envoi No. 94. anggoullai
tig., amboie amh.” (i.e. “fourth sending, number 94, local names ‘anggoullai”
(Tigre), ‘amboie’ (Amhara)”). There are some other specimens that may also be
lectoparatypes.
Nomenclature
Dunal described S.campylacanthum Hochst. ex Dunal from specimens of Schimper
123 and 1082 in G-DC, and Bitter referred to these and other collections of
Schimper and others, probably mostly in Berlin.
Realizing the vast range of variation in Africa, Bitter recognized 58 infraspecific
taxa, in 22 species, which he placed together in subseries Campylacantha Bitter, most
of which I call S.campylacanthum Hochst. ex A. Rich., although some are S. pandurzforme
E. Mey. ex Dunal. However, a few distinctive local taxa may also be recognized,
such as S. stellativillosum Bitter.
Jaeger (1 985) provided an objective, but tentative and cautious, taxonomic
treatment of this and other species in the S. incanum aggregate. Results from large
scale biosystematic investigations of the whole range of African and Asian relatives
of S. incanum L. and S.melongena L. (Lester & Hasan, 1991, and subsequent studies),
have proved the distinctness of at least four species corresponding to S. lichtensteinii
Willd., S.incanum L., S.pandurzfomze E. Mey. ex Dunal and S.campylacanthum Hochst.
ex A. Rich., and necessitated the typification of these last two species. Almost a
thousand specimens at Kew have now been determined to these four species.
Solanum pandurifomze E. Mey. ex Dunal ( = S. delagoense Dunal)
Although this species was not considered by A. Richard, it must be considered
284
R. N. LESTER
here alongside its close relative S. camfilacanthum A. Rich. Furthermore, there is also
an important specimen at Paris.
Specimens collected by Dr2ge in January 1832 from scrub below 1000 feet alt.
on the banks of the River Mbasche (or Bashee), Transkei, South Africa (Drkge,
1843) (not from the Cape of Good Hope, as generally stated on the labels), were
distributed as “Solanurn panduraefome E.M.”, which was published without diagnosis
by E. Meyer in Drege (1843: 147).
The first valid publication with diagnosis was by Dunal (1852) as S. pandura&me,
but this should be corrected to S. pandunturn E. Mey. ex Dunal (fide Brummitt).
Dunal cited Drkge s.n. (Cape of Good Hope . . . 1000 feet) G-DC (IDC Microfiche
2092/ 13!),which is therefore the holotype: isotypes are at Paris!, Kew! (one designated
‘ISOTYPE’ by Jaeger) and probably elsewhere. Although these specimens are
labelled ‘Cape of Good Hope’, the holotype also bears a handwritten note referring
to “Basche. lOOOft altitude”, which agrees with the locality published by Drkge
(1843).
Both Bitter (1923) and Jaeger (1985) used the name S. pandunzmze E. Meyer,
although the correct citation should be S. pandunyome E. Mey. ex Dunal, which may
be shortened to S. pandunyome Dunal. (This name must not be confused with S.
panduriznne Dunal “mss in h. Boiss.)’,which is now S. anoplocadum Dunal, as detailed
in Index Kmensis).
For the allied taxon, Sohnum delagoense Dunal (1852) there are no problems. The
holotype is Forbes, Delagoa Bay [i.e. B. de Maputo, Mozambique] 1822, from
“Herb. SOC.Hort. Lond.”, G-DC (IDC Microfiche 2089/ 12!). There are five isotypes
on four sheets at Kew!, with several notes identifying them as S. panduriznne. Bitter
(1923) did not cite the type collection, but saw a wide range of material which he
classified into 14 subspecies and varieties, under S. delagoense.
Having established the typification and nomenclature, the taxonomic position of
S. panduriiome and other taxa must be considered. Although Bitter distinguished
many species and even more infraspecific taxa, detailed biosystematic studies by
Lester & Hasan (1991) and others have shown that the S. campylacanthum-S. bojerl
S. delagoense-S. pandu?zfonne complex is best considered as two species, referred to as
groups A and B by Lester and Hasan. Taxon A, S. campylacanthum, comprises
relatively broad-leaved shrubs, common in East Africa in tropical savanna-woodland,
etc. There is great genetic diversity both within and between populations, and
several distinct species, as well as many varieties, may deserve recognition. Taxon
B comprises narrower leaved, less robust shrubs or herbs, which are more uniform
genetically, but present very different phenotypes depending on the age of growth
and the environmental conditions. These have probably evolved from taxon A and
spread south into drier and/or colder areas. Both S. delaguense Dunal and S.
pandunyome E. Mey. ex Dunal belong to this taxon B: the two holotypes are virtually
indistinguishable, and the five excellent isotypes of S. delagoense at Kew, comprising
both lush sucker shoots and meagre twigs from old bushes, encompass the morphology
of the two isotypes of S. pandun$nne at Kew. Bitter attempted to distinguish these
as two distinct species, but encountered great difficulties, and admitted that there
were transition forms such as S. delagoense var. transvaalense Bitter.
S. pandul?’fome E. Mey. ex Dunal, and S. delagoense Dunal were both validly
published by Dunal (1852), but in my opinion both names definitely refer to the
same species, and therefore S. delagoense should be treated as a synonym of S.
pandurifonne, as first stated by Wright (1904).However, Rossler (in Merxmuller, 1969)
TYPIFICATION OF A. RICHARD’S AFRICAN SOLAJfUbf NAMES
285
used the name S. delugome Dunal, giving S. pandurdorme auct. non E. Meyer as
listed by Dinter (1927) as a synonym. The reasons for this decision are not known,
but may reflect Heine’s notes on Munich specimens, concerning Bitter’s concepts
of these two species.
In southwest Africa, including the Cape, occur very prickly xerophytic plants with
deeply lobed leaves, called S.pandunzrme by Bitter, but these may deserve recognition
as a separate species, distinct from S. pandunzrme E. Mey ex Dunal, and even more
distinct from S. campylacanthum Hochst. ex A. Rich.
Solanum grossidentatum A. Rich.
iryp$cation
Besides the formal description of “S. grossedentatum Nob.”, Richard stated:
It grows in the province of TchClikote (Ant. Petit).
Observation.-The species which we are describing belongs to the tribe Maurella
of M. le prof. Dunal; but it seems to me to be perfectly distinct from all the species
arranged in this group. It is placed at the side of Solanum villosum Lamkc. [sic!],
but it presents a rusty hue due to the hairs which cover almost all its parts. Its
leaves are very deeply incised, in large unequal and pointed teeth; its cymes, long
pedunculate, are composed of only two to four flowers, of which the corolla is
twice as large as that of Solanum villosum; its calyces are excessively hairy, with
elongated divisions, narrow and obtuse.
The lectotype designated here was collected by Quartin-Dillon and Petit from
“Abyssinia, Ichtlikote”, was in the Herbarium Richard, and was named “Solanum
grossedentatum Nob.” (On the Herb. Mus. Paris label is lightly written ‘?T’.)The
isolectotype,which appears to be the lower parts of the same shoots as the Iectotype,
was named “Solanum grossedentatum Rich.”.
Nomenclature
Although originally published as S. grossgdentatum, this should be corrected to
Solanurn gross9mtatum A. Rich. (fide Brummitt). Neither Dunal nor Bitter treated this
species.Jaeger maintained it, based on Richard‘s description, but suggested it might
be included in S. villosum Mill., if it proved to be a tetraploid with yellow to red
fruits. Cufodontis (1963)gave S. grossidentatumA. Rich. and S.plebeium var. pssidentatum
(A. Rich.) Chiov. as synonyms of S. memphiticumJ. F. Gmel. var. abyssinicum (Dunal)
Cufod., citing Schimper 46 (Adoa) as the type of that species.
Solanum hirtulum Steud. ex A. Rich.
irypzjication
Besides the formal description of “S. hirtulum Steudel” (citing Schimper 977),
Richard stated:
It grows in fields near Entchetkab, in the Semiene province, flowering in January
(Schimper).
R. N.LESTER
28ti
Ohsenation.-Small species of the tribe of the Afaurella, having resemblance to
Solanunz uzlluwn Lamkc. [sic!], which also grows in Egypt. But it is smaller, the
angles of the stem are less conspicuous and its flowers are less numerous and
larger.
The lectotype was collected by Schimper No. 977, 29 January 1838, Abyssinia,
Semiene pro\.ince, in fields near Enschadcap. It was distributed in 1842 under the
name Solanum hirtulum Steud., and was first in Herbarium Steudel, then Herbier E.
Drake, then Herb. hlus Paris. A ‘TYPE’ label was stuck on this sheet some years
ago. and at the same time ‘ISOTYPE’ labels were stuck on three other sheets of
Schimper 977. Two of these are true isolectotypes (but one of these is not S. hirtulum,
probably by reciprocal confusion with the collection Schimper 20 1 of S. bfurcatum
Hochst. ex ’4. Rich.). Edmonds (pers. comm.) agrees with my choice of lectotype.
Other isolectoty-pes are at K (!), G-DC (IDC hlicrofiche 2061/5!), G-BOIS and
AWE.
%,IOntenclature
The specimens collected by Schimper 977, and named by Steudel ined., were
seen separately by Richard (P!), Dunal (G-DC, G-BOIS), Bitter (B, MPU), Jaeger
(K), who a<greedin recognizing this distinct species. Bitter also cited several other
collections as S. hirtulum Steud. ex Dunal (1852), but Cufodontis, and also Jaeger,
recognized it as S. hirtulum Steud. ex A. Rich. (1850).
Solanum macihitum A. Rich.
(‘This category includes other synonyms of S. forskulii Dunal, etc.)
qp$ication
Besides the formal description of “S. macilentum Noh.”, Richard stated:
It is said that it grows in the sandy region of Choho (Quartin Dillon).
Observation.--It is to the section Melongena that this species belongs, by its
polygamous flowers, of which a very small number are fertile. Its fertile flowers
occupy the same position as in the other species of this group. They are longer
pedunculate and part from the base of the cyme, but their calyx is, like that of
the other flowers, equally delvoid ofprickles; a few of these prickles show themselves
on the peduncles and on the pedicels.
This species has a lank and completely peculiar habit; by its rather small, oval,
cordiform, obtuse leaves; by its flowers forming a long peduncled and bifurcate
cyme, by its fertile flomers devoid of prickles, one can easily distinLguishit from
the other species of the same group.
The lectotj-pe, with the most abundant material, and the isolectotype wert collected
by Quartin-Dillon and Petit from Abyssinia, Choho, and were in Richard’s herbarium
and were named by him as S. marzbztum Nob. and S. macilentum Rich. respectively.
A third sheet, which does not state Choho and was not in Richard’s herbarium,
cannot be designated with certainty as a type specimen: it was determined as S.
alhzcaule Kotschy by Bitter in 1925.
TYPIFICATION OF A. RICHARD’S AFRICAN SOLANUM NAMES
287
Nomenclature
The three specimens named S. macilentum A. Rich. had been placed under S.
forskalii Dunal at Paris, together with type specimens of S. albicaule Kotschy and S.
heudelotii Dunal, and also specimens of “S. senegalense Perrott. (non Dunal)” and “S.
kotschyii Stead.”, but all these type specimens have now been separated. Bitter
annotated several of these specimens. Someone else annotated the lectotype of S.
nzacilenturn A. Rich. with “S. albicaule (Kotsch) Dun = S. Heudelotii Dun” [sic!].
Dunal (1852) did not consider S. macilentum A. Rich. or S. senegahse Perr., but he
did describe S. senegambicum Dunal (citing Heudelot 713 at Paris, which is now
labelled ‘isotype’, and is S. anguiui Lam. P!). Dunal treated S. albicaub Kotschy
immediately before describing S. heudelotii Dunal (citing Heudelot 417 at Paris), but
S. forskalii Dunal, which he had described in 1813, he only mentioned briefly in his
appendix of insufficiently known Solanum species.
Bitter (1923: 297), working only from Richard’s description of S. macilentum A&
Rich., agreed with Wright (1906) that it was an imperfectly known species. He did
not mention S. forskalii Dunal, but gave full treatments of S. heudelotii Dunal (in
Section Oliganthes, series 1. Ajoindica), and S. albicaule Kotschy (in Section Oliganthes,
series 6. Albicaule). Subsequently, in 1925, he determined one S. macilentum specimen,
now at Paris, as S. albicaule Kotschy.
Cufodontis (1963) cited the type collection of S. macilentum from Richard (1850,
as 1851), and stated that without doubt the holotype would be preserved at Paris,
and should be inspected.
Jaeger (1985, citing Hepper & Wood, 1983) stated that S. albicaule Kotschy ex
Dunal and S. heudelotii Dunal (and also S. villosum Forssk. non Mill. and S. scindicum
Prain) are synonyms of S.forskalii Dunal. He did not mention S. macilentum A. Rich.
(nor S. senegalense Perr., nor S.kotschyii Steud.). D’Arcy & Rakotozafy (1994)described
S. batoides from Madagascar, which is very similar to S. forskalii.
Types of several of the other species mentioned above are also at Paris:
S. albicaule Kotschy ex Dunal (1852)
Dunal cited Kotschy 309 (Nubia, Cordofan, Chursi) at both G-DC and MPU
(“herb. Dunal”). The G-DC specimen (IDC Microfiche 2078/8!) is hereby designated
lectotype, and the MPU specimen is an isolectotype: there are more isolectotypes
on two sheets at Paris!, and one at Kew (!).
S. heudelotii Dunal (1852)
Dunal cited Heudelot 417 (Senegambia, near Gabor) at Paris, but there are three
such specimens there. The lectotype has been selected because it has good flowers
and fruits, and has the fullest collection notes. (‘Type’had been written informally
on this sheet.) The other two specimens are isolectotypes, one of which had been
determined as S. albicaule Kotschy by Bitter in 1925. Other isolectotypes are at K!
and OXF!.
S. kotschyii Steud. (ined.?)
At Paris, a collection by “Kotschy s.n., Nubia”, from Herbarium Steudel, is
labelled “418 S.Kotschyii Steud”, mentioning its affinity to S. hegazeme Dunal (XIII,
1.292). It is a good specimen of S.forskalii Dunal.
288
R.N. LESTER
S. senegulense Perr. (ined.?)
The specimen Perrottet 340 (Senegal, 10 Sept. 1824) (P!), which may be the
holotype if this species has been published, was determined as S. ulbicaule Kotschy
by Bitter in 1925. S. s m g a h e Perr. was probably intended for publication in Flora
Senegambiae Tentamen, of which only volume 1 was published by Guillemin,
Perrottet and Richard in 1830-33 (Vallot, 1882: 186), and it was not mentioned by
Dunal or Bitter, and is not in Index Kewensis. It should not be confused with S.
senegambicum Dunal.
S. senegambicum Dunal (1 852)
The holotype, Heudelot 7 13 (Senegambia), Cp!), has been incorrectly labelled
“Isotype” previously. It has also been identified as Solanum indicum auct. non L.
subsp. dbtichum (Thonn.) Bitter var. distichurn, and is indeed a cultivated form of S.
anguiui Lam.
Solunum macracanthum A. Rich.
7ypZfication
Besides the formal description of “S. macracanthum Nob.”, Richard stated:
It grows in the mountainous kingdom of Choa, where it was discovered by the
blessed doctor A. Petit. [Note: this species is dangerously armed!]
Observation.-Species remarkable for the length of its prickles which are straight
and not recurved, by its small leaves generally having an inch of length, entire or
panduriform, by its cymes containing only a small number of flowers of which the
calyces are bristling with prickles, and the fruits the size of a cherry, of a beautiful
yellow orange colour.
The lectotype was collected by Quartin-Dillon and Petit in Abyssinia, Choa. In the
Herbarium Richard, it was first named Solunum sp. nova, then Solunum macracanthum
Nob. There are three sheets of isolectotypes at Paris.
Jllomencluture
This striking species has been neglected, although Bitter (1923) recognized and
described it in detail, based on Schimper 141 (possibly at B) and even created a
distinct series for it. Cufodontis listed S. macracanthum, but both Dunal and Jaeger
ignored it, although Jaeger treated S. cymbalarifolium Chiov. (1929), which is a
somewhat similar species.
Solanurn p + m ? t m A. Rich.
Gp$ication
Besides the formal description of “S.p+m@mrn Nob.”, Richard stated:
It grows in the mountainous province of Tchelikote, where friend doctor Ant.
Petit discovered it.
TYPIFICATION OF A. RICHARD’S AFRICAN SOZAAZW NAMES
289
Observation.-It is a little bushy shrub having a slight resemblance with Solanum
macracunthum Nob.; but its shorter prickles, its excessively small leaves, which are
very variable in their shape, its globular fruits scarcely exceeding the size of a
pepper grain, form a combination of characters which together distinguish this
species from the former.
The lectotype was collected by Quartin-Dillon and Petit in EtchClikote (Abyssinia).
It was named Solanum p$eriferurn Nob. in Herbarium Richard. More recently ‘type’
was written informally on it. There are five isolectotypes at Paris (with the locality
spelled Etchelicote), three from Herbarium Richard, and two from Herb. A. de
Franqueville with the number 226.
JVomencluture
There is an isolectotype at Kew (Petit 226!), seen by Jaeger. Bitter did not see
type material, but used Richard’s description to identlf). Hildebrandt 479 as this
species, and then elaborated a full description. S. p$m@rum was not known to Dunal,
but Cufodontis listed it.
Sohnum plebeium A. Rich.
5pijication
Besides the formal description of “S. plebeium Nob.”, Richard stated:
It grows on rubbish dumps and cultivated places of the province Chire‘ (Quartin
Dillon).
Observation.-Is this a new species? I believe so, without however having complete
certainty of it, because, in the group Maurella, the forms are very variable. However,
our plant from Chirt appears to constitute a distinct type. It differs from Sohnum
n i p m and S.pterocaulon, which it resembles most, by its pale yellow fruits, by its
elliptic-lanceolate leaves, equally attenuated to a point at their two extremities,
entire, ciliate, sessile, studded with several stiff short hairs. In one word, when one
examines our plant and if one compares it to specimens of the two other species,
one is hit by the difference which it presents in its habit and the shape of its leaves.
The lectotype was collected by Quartin-Dillon and Petit from Abyssinia, Chirt:
province. It was named Solanum plebeium Nob. in Herbarium Richard. A ‘TYPE’
label was correctly attached to this sheet some years ago (although Jaeger (1985)
stated ‘(P?)’indicating doubt of the whereabouts of this type). At the same time the
isolectotype was labelled ‘ISOTYPE’. However, there are three other sheets which
were also labelled ‘ISOTYPE’ at this same time, but since their localities do not
state ‘ChirC’, (and since they were never in Herbarium Richard), their status is
doubtful and they should not be considered as types. Edmonds (pers. comm.) agrees
with my choice of lectotype.
JVomenclature
Several subsequent authors have misspelled the name S.plebgum, with a 3’. Bitter
did not see Richard’s specimens, but used his description to identlf). other collections,
and thence produced more detailed descriptions and distinguished two varieties of
R. N. LESTER
2’90
this species. Although Cufodontis (1 963) treated it as a distinct species,Jaeger (1985),
after consultation with Edmonds (1 979, 1984), considered that this species was
probably a synonym of S. uillosum Mill. subsp. miniatum (Bernh. ex Willd.) Edmonds.
Solanum polyanthmum Hochst. ex A. Rich.
Gp$ication
Besides the formal description of “S. pobanthmum Hochst.” (citing Schimper 1526),
Richard stated:
It tgro\\.snear Awrn, flowering and fruiting in Nolxmber (Schimper).
Observation.- Solanurn pofyanthemum Hochst. has some resemblance to Solanum
rchinipenanum, but its leaves, which are tomentose and of a rusty colour on the
underside: its numerous flowers, forming a terminal cyme, distinguish it easily
from this other species.
T h e lectotype was collected by Schimper No. 1526, 24 November 1842, near Axum,
in Abyssinia. Attached is a handwritten note in German, possibly by Schimper
himself “. . . in the . . . at 6000- 8000 feet above sea level. Axum. 24 November
1842. Rare. 1526. Solanum”. This specimen was first in Herbarium Steudel, then
Herbier E. Drake, then together \\-ith the isolectotype, in Herb. Mus. Paris, where
both had been placed in a type folder, but without any annotation. There are other
isolectotypes at G-DC (IDC Microfiche 2068/24!), K (!), and BM.
Zomenclature
This taxon was called S.pobanthmum Hochst. by Dunal, based on Schimper 1526
(G-DC). Bitter called it S. schimpm’anum Hochst. var. polyanthemum (Hochst. [ex A.
Rich]j Bitter, again based on Schimper 1526 and other specimens, and also described
subvar. cordz$lium Bitter, based on a Quartin-Dillon and Petit collection (last sending
1844) he sakv at Paris. Jaeger, who saw an isotype at BM, recognized it as 5’.
polyanthemum Hochst. ex A. Rich., but then treated it as a synonym of S. schimperianum
Hochst. ex A. Rich., as also did Cufodontis.
~
Solanum schimperianum Hochst. ex A. Rich.
Typijcation
Besides the formal description of “S. schimperianum Hochst. (citing Schimper 202),
Richard stated:
It grows in mountainous places around Adoua, flowering and fruiting in June
(Quartin Dillon and Schimper).
Abyssinian name: Agol.
Obsenation-The number of species referred to the first part of section I1 [ix.
Dukarnara], is of little importance. Solanum schirnperianum, which belongs to this
,group, is strongly distinct from the other species. It has some analogy with Solanurn
TYPIFICATION OF A. RICHARD’S AFRICAN SOLA,VUM NAMES
29 I
dulcamara L., but its leaves, which are always entire and not lobed, and are covered
with a green and ashy tomentose down, and its globular fruits, distinguish it from
the rest.
The lectotype was collected by Schimper No. 202, 5 June 1837, in mountains near
Adoa, Abyssinia. Abyssinian name ‘Agol’. This specimen, distributed in 1840, which
was first in Herbarium Steudel, then in Herbier E. Drake, and then in Herb. Mus.
Paris, was almost certainly seen by Richard: another sheet of Schimper 202, is an
isolectotype. (Both specimens were previously labelled ‘ISOTYPE’,probably referring
to S. schimperianum Hochst. ex Dunal, of which the TYPE is at G-DC). There are
more isolectotypes at G-DC (IDC Microfiche 2068/23!) and BM. Four more sheets,
all from Herbarium Richard, and collected by Quartin-Dillon and Petit from ‘Adoua’
(no date) and from “Vallee d’Adowa, 25 August 1839” are lectoparatypes.
Nomenclature
Based on Schimper 202 (at G-DC and probably B, respectively), th‘
is taxon was
called S. schimperianum Hochst. by Dunal and by Bitter, but was correctly recognized
as S. schimperianum Hochst. ex A. Rich. by Cufodontis and also by Jaeger, who also
saw an isolectotype specimen of Schimper 202 at BM.
Solanum unguiculatum A. Rich.
3p$ication
Besides the formal description of “S. unguiculatum Nob.”, Richard stated:
It grows in Chirt! province (Quartin Dillon and Ant. Petit).
Observation.-I believe this species [is] very close to Solanurn indicum L. [sic!]. It
differs from it nevertheless by its leaves, which are almost entire and scarcely lobed
in their outline, which are tomentose and of a greenish yellow colour on both
surfaces, and not almost white beneath.
The lectotype was collected by Quartin-Dillon and Petit from Abyssinia, Chirt. On
the label is written “Solanum tomentosum ? ? L.”, but also “Solanum unguiculatum Nob.”.
The two isolectotypes are named Solanum unguiculatum Rich. All three sheets appear
to be from a single collection, and have been kept together in Herbarium Richard
and subsequently. (I have determined these specimens as S. incanum L. sens. str., i.e.
group C, Lester & Hasan, 1991).
Nomenclature
Although Bitter did not see the type material, he saw other similar specimens
from Abyssinia (Schimper 56, 181, 383, 527, etc.), and correctly relegated this taxon
to varietal rank as S. incanum L. var. unguiculatum (A. Rich.) Bitter, which was followed
by Cufodontis (1963).
R. N.LESTER
292
Other Solanum species
Six other species of Solanum, namely S. tuberosum L., S. n i p m L., S. villosum Lam.,
S. malginatum L.f., S. sanctum L. [ = S. incanum L.] and S. dubium Fresen. [ = S. coagulans
Forssk., non S. coagulansauct.], were also described by Richard (1850)from Abyssinia.
ACKNO\\’LEDGESf EKTS
The research presented here was undertaken in Paris after a meeting of the
European Solanaceae Information Network at INRA, Montfavet, funded by the
European Union Biotechnology Programme (Grant BI02-CT93-0397). Subsistence
costs were provided by the British Council Anglo-French cooperation ALLIANCE
programme (Grant PN 91.065). I am deeply indebted to M. J.-C. Jolinon (P), who
helped greatly with finding and photocopying obscure specimens and literature in
Paris. Professor P. Morat (P) welcomed me there. Dr Henk Beentje (K) and Dr
Dick Brummitt (K) assisted in finer points of interpretation of Latin diagnoses and
nomenclatural procedures. I thank them all for their kind help. The thesis of Jaeger
(1985) provided an invaluable initiation to this taxonomic minefield.
REFERENCES
Bitter G. 1913. Solana africana I. (Aforellae novae vel minus cognitae) Botanische Jahrbucher 49:
560-569.
Bitter G. 1917. Solana africana 11. Botnnische3ahrbiichtr 54: 416-506.
Bitter G. 1921. Solana africana 111. Botanische Jahrbucher 57: 248-286.
Bitter G. 1923. Solana africana W. Repertonurn specierum muarum regni vegetabilis 16: 1-320.
Boiteau P. 1976. La dynastie des Richard, jardiniers-botanistes. Comptes Rendus du 11 Oe Congris National
des Soci&s Savantes, Paris, 1975 3 13-29.
Brongniart A. 1854. Notice historique sur M. Achille Richard. Bulletin de la Sociiti botanique de France
1: 373-386.
Bureau E. 1904. Notice sur Emmanuel Drake del Castillo. Bulletin de la SocGtl botanique de France 51:
cxvii, cxxk-cxxxiii.
Chiovenda E. 1929. Flora somala. Sindicato Italian0 h i Grafiche, Rome.
Cufodontis G. 1963. Solanum in Enumeratio Plantarum Aethiopiae Spermatophyta. Bulletin du Jardin
Botunique de I’Etat Bmrelles 33 (Suppl.) 861-880.
D’Arcy WG,Rakotozsfy A. 1994. Flore de .kladagascar et des Comores. Famille 176 - Solanaceae. Museum
national d’Histoire naturelle, Paris.
Dinter K. 1927. Index der aus Deutsch - Siidwestafrika bis zum Jahre 19 17 bekannt gewordenen
Manzenarten. XXII. Repertonurn specierum novarum regni vegetabilis 23: 363.
D+ge JF. 1843. Zwei pflanzengeografischeDokumente. Flora 2: (Suppl.) 147.
Dunal MF. 1813. Histoire ,Vatuzlle Medirinale ef Economique des Solanum. Paris, Strasbourg, Montpellier.
Dunal MJ?. 1852. Solanaceae. In: de Candolle rip, ed. Pmdmmus Systematis naturalis Regni vegetabilis 13
(1): 1690. Paris.
Edmonds JM. 1979. Nomenclatural notes on some species of Solanum L. found in Europe. Botanical
Journal ofthe Linnean Sock9 78: 213-233.
EdmondsJM. 1984. Solanum L. section Solanum - a name change in S. villosum Miller. BotanicalJoumal
of the Linnean Sock& 8 9 165-170.
Heine HH. 1960. Notes on Solanum. Kpu! Bulletin 14: 245-249.
Hepper FN, Wood FRI. 1983. New combinations and notes based on Forsskal’s Arabian collection.
K m Bulletin 38(1): 83-86.
TYPIFICATION OF A. RICHARDS AFRICAN SOIANUM NAMES
293
Jaeger P-ML. 1985. Systmtic studies in the genus Solanum in A&ca. Unpublished Ph.D. Thesis,
University of Birmingham, UK. 272 pp.
Jaeger P-ML, Hepper FN. 1986. A review of the genus Solanum in Africa. In: D’Arcy WG, ed.
Solanaceae: biology and systematics. New York Columbia University Press.
Lashgue A. 1845. Mush botanique de M. Benjamin Delessert. Notices sur les collections deplantes et la bibliothkque
p i le composent; contenant en outre des documents sur les principaux herbiers d’Europe et l’exposd des vvages
entrepris dans l‘inhi2 de la botanique. Librairie de Fortin, Masson et Cie., Paris. 588 pp.
Lester RN, Hasan SMZ. 1991. Origin and domestication of the brinjal egg-plant, Solanum melongena,
from S.incanum, in Africa and Asia. In: Hawkes JG, Lester RN, Nee M, Estrada RN, eds. Solanaceae
III - Tmonomy, Chmisb, Evolution. Kew: Royal Botanic Gardens.
Merxmuller H. 1969. (ed.) Pmdrmus einer Flora von Sudwestu$&. 124. Solanaceae.
Milne-Redhead E. 1949. Solanum b@rcatum A. Rich. Kew Bulletin 1948: 467.
Richard A. 1840. Plantes nouvelles d’Abyssinie, recueillies dans la province du TigrC: par M. le Dr
Richard Quartin-Dillon, voyageur naturaliste du Musbum d’Histoire naturelle de Paris. Annales des
Sciences naturelles, Botanique series 2 14: 257-276.
Richard A. 1847. Zntamen Florae Abyssinicae. Vol. 1. Arthus Bertrand, Paris.
Richard A. 1850. Zntumen Florae Abyssinicae. Vol. 2. Arthus Bertrand, Paris.
Richard A. 1982. Zntamen Florae Abyssinicae. li0l.s 1, 2 & 3. Facsimile. Institute of Systematic Botany,
Uppsala.
Stafleu FA, Cowan RS. 1983. Taxonomic literature: Volume I E P-Sak. Regnum vegetabile. The Hague: I.t:
Junk. p. 452.
Stearn WT. 1982. Preface to facsimile edition of Achille Richard’s Entamen Florae Abyssinicae. Vol. 1.
Institute of Systematic Botany, Uppsala.
Vallot J. 1882. Notice sur les voyages botaniques accomplis dans 1’Afrique tropicale. Bulletin de la
Socikti botanique de France 2 9 171-205.
Wright CH. 1904. Solanaceae. In: Thistleton-DyerWT, ed. Flora Capensis 4(2).London: Lovell Reeve,
87-121.
Wright CH. 1906. Solanaceae. In: Thistleton-Dyer WT, ed. Flora of Tropical A&ca 4(2). London:
Lovell Reeve, 207-26 1.