$¢PhytoKeys

PhytoKeys 236: 157-178 (2023)
DOI: 10.3897/phytokeys.236.110498

Research Article

Erica L. (Ericaceae): homonyms amongst published names for
African species and proposed replacement names

E. Charles Nelson’®, E. G. H. Oliver2, Michael D. Pirie?®

1 Tippitiwitchet Cottage, 255A Broadgate, Sutton St Edmund PE72 OLT, UK

2 Department of Botany and Zoology, University of Stellenbosch, Private Bag X1, Stellenbosch, 7602 Matieland, South Africa
3 University Museum, University of Bergen, Postboks 7800, NO-5020 Bergen, Norway

Corresponding author: Michael D. Pirie (michael.pirie@uib.no)

OPEN Qaccess

This article is part of:

Systematics, natural history, and
conservation of Erica (Ericaceae)

Edited by Michael Pirie, Félix Forest,

Timo van der Niet, Jaime Fagundez, Seth
Musker, Fernando Ojeda, Anina Coetzee, E.
Charles Nelson

Academic editor: Jaime Fagundez
Received: 3 August 2023

Accepted: 5 November 2023
Published: 20 December 2023

Citation: Nelson EC, Oliver EGH,

Pirie MD (2023) Erica L. (Ericaceae):
homonyms amongst published names
for African species and proposed
replacement names. PhytoKeys 236:
157-178. https://doi.org/10.3897/
phytokeys.236.110498

Copyright: © E. Charles Nelson et al.

This is an open access article distributed under
terms of the Creative Commons Attribution
License (Attribution 4.0 International -

CC BY 4.0).

Abstract

In support of ongoing taxonomic work on the large and complex flowering plant
genus Erica (Ericaceae), we document nineteen pairs of homonyms representing
currently used illegitimate names. We provide replacements for thirteen names and
new typifications for five. We relegate five names to synonymy: Erica aemula Guthrie &
Bolus under Erica distorta Bartl.; Erica armata Klotzsch ex Benth. under Erica umbrosa
H. A. Baker; Erica capensis T.M. Salter under Erica turbiniflora Salisb.; Erica lanata
Andrews under Erica flaccida Link; and Erica tomentosa Salisb. under Erica velutina
Bartl. Finally, we suggest conservation of Erica aristata Andrews. The new names are:
Erica adelopetala E.C. Nelson & E.G.H. Oliv. replacing Erica insignis E.G.H. Oliv.; Erica
bombycina E.C. Nelson & Pirie replacing Erica niveniana E.G.H. Oliv.; Erica concordia E.C.
Nelson & E.G.H. Oliv. replacing Erica constantia Nois. ex Benth.; Erica didymocarpa E.C.
Nelson & E.G.H. Oliv. replacing Erica rugata E.G.H. Oliv.; Erica galantha E.C. Nelson &
E.G.H. Oliv. replacing Erica perlata Benth.; Erica mallotocalyx E.C. Nelson & E.G.H. Oliv.
replacing Erica flocciflora Benth.; Erica notoporina E.C. Nelson & E.G.H. Oliv. replacing
E. autumnalis L.Bolus; Erica oliveranthus E.C. Nelson & Pirie replacing Erica tenuis
Salisb.; Erica oraria E.C. Nelson & E.G.H. Oliv. replacing Erica spectabilis Klotzsch ex
Benth.; Erica oresbia E.C. Nelson & E.G.H. Oliv. replacing Erica demissa Klotzsch ex
Benth.; Erica poculiflora E.C. Nelson & E.G.H. Oliv. replacing Erica stenantha Klotzsch
ex Benth.; Erica rhodella E.C. Nelson & E.G.H. Oliv. replacing Erica rhodantha Guthrie &
Bolus; Erica supranubia E.C. Nelson & Pirie replacing Erica praecox Klotzsch.

Key words: Cape heaths, Erica, Hans Dulfer, International register of heather names,
nineteenth-century English nursery catalogues, nomenclature

Introduction

The nomenclatural history of the genus Erica L. sensu /ato (Oliver 2000, 2012)
is complicated by the extraordinary fashion in western Europe for cultivating
“Cape heaths”, the English name generally given to plants derived from the
Erica species endemic to southern Africa. Erica-mania commenced in the
last decade of the 18" century, burgeoned in the early 1800s and petered out
in the middle of that century (Nelson and Pirie 2022). A consequence of the

57

E. Charles Nelson et al.: Homonyms in Erica

horticultural interest was a proliferation of names, applied often in a haphazard
manner by nurserymen and gardeners, as well as by botanists. Many names
were first published in nurserymen’s and gardeners’ catalogues and in horticul-
tural periodicals, often with accompanying descriptions or diagnoses (Nelson
and Oliver 2004; Nelson and Small 2004-2005; see also Reveal (2012)).

Work carried out between 1995 and 2004 by ECN for The Heather Society
(established in 1963, formally dissolved in 2020) on the second volume of the
International Register of Heather Names (Nelson and Small 2004-2005) nec-
essarily included an extensive survey of nineteenth-century horticultural publi-
cations, resulting in the accumulation of more than 6,000 names, at all ranks
and many times that number of bibliographic citations, for Erica taxa of African
origin. This information was collated in a database, complementing a similar
one for the so-called “hardy heaths” — Andromeda L., Calluna Salisb., Daboecia
D. Don and species of Erica endemic in Europe, Macaronesia and western Asia
(Nelson and Small 2000). We are currently working to make this resource open-
ly accessible, particularly through integration with the World Flora Online (WFO
2023; Elliot et al. in prep.). The database provides an invaluable bibliographic
tool, much more comprehensive and, for a genus of more than 800 species,
more practical and detailed than any existing botanico-bibliographic indexes
including such standard publications as “Index Kewensis” which only provided
publication information for protologues. Inevitably a database of such a com-
prehensive nature revealed a scattering of hitherto unsuspected, or inadver-
tently overlooked, problems with the established and currently accepted names
of African Erica species.

Some of these problems do not arise, in fact, from the unearthing of long-bur-
ied names (most of the binomials discussed here were recorded by Dulfer
(1965)), but rather from a better understanding of the history of certain pub-
lications and more accurate information about dates of publication of, for ex-
ample, George Bentham’s (1839) treatment of Erica and related genera (Nelson
2005) and Henry Cranke Andrews’s multi-volume works, Coloured Engravings
of Heaths (Cleevely and Oliver 2002) and The Heathery (Cleevely et al. 2003).

Although changes in current names are rarely welcomed, particularly outside
the taxonomic community, by following the rules set out in the International
Code of Nomenclature for Algae, Fungi and Plants (Shenzhen Code) 2018 (Tur-
land et al. 2018) (hereafter ICN (Shenzhen Code) 2018), we can reflect the state
of knowledge in the field and maintain overall stability in nomenclature. In this
paper, we provide replacement names for homonyms and clarify typifications
where necessary.

Homonyms in Erica

The Heather Society's database revealed pairs of homonyms within Erica where
the one in current use (compare Oliver and Oliver (2003); Oliver (2012)) is not
the earliest published. The need to replace these later names was signalled in
the four published parts of the International Register of Heather Names, vol-
ume 2, African Species, Hybrids and Cultivars (Nelson and Small 2004-2005).
These names are listed (as summarised in Table 1) and discussed and, where
necessary, replacements are provided when an alternative validly published
name is not available.

PhytoKeys 236: 157-178 (2023), DOI: 10.3897/phytokeys.236.110498 158

E. Charles Nelson et al.: Homonyms in Erica

Table 1. Homonym pairs in Erica in alphabetic order with dates of publication.

Specific epithet

aemula
aristata
armata
autumnalis
Capensis
constantia
demissa
flocciflora
insignis
lanata
niveniana
perlata
praecox
rhodantha
rugata
spectabilis
stenantha
tenuis

tomentosa

Author(s) and date of currently accepted binomial Author(s) and date(s) of earlier binomial
Guthrie and Bolus (1905) Rollisson (1855)
Andrews (1807) Salisbury (1796)
Klotzsch ex Bentham (1839) Sprengel (1825)
L. Bolus (1923) Hort. ex Bentham (1839)
Salter (1935) Regel (1842)
Noisette ex Bentham (1839) Hort. ex Sinclair (1825)
Klotzsch ex Bentham (1839) Hort. ex Sinclair (1825)
Bentham (December 1839) Tausch (October 1839)
Oliver (1981) Hort. (1853)
Andrews (1806) Wendland (1798)
Oliver (2000) Hort. ex Loudon (1830)
Bentham (1839) Sinclair (1825)
Klotzsch (1838) Hort. ex Sinclair (1825)
Guthrie and Bolus (1905) Regel (1842)
Oliver (2000) Hort. ex Sinclair (1825)
Klotzsch ex Bentham (1839) Waitz (1805)
Klotzsch ex Bentham (1839) Sweet (1830)
Salisbury (1802) Moench (1802)
Salisbury (1802) Masson (1776)

It should be noted that nurserymen’s catalogues during the 19" and 20"
centuries were usually annual publications, reissued with minor amendments,
deletions and additions, year after year, decade after decade. The dated cata-
logues noted in this paper are examples only — they must not be assumed to be
the first, the last or the only issue containing a particular name.

1. Erica aemula Rollisson (1855), non Guthrie and Bolus (1905)

This binomial first appeared in print within an advertisement inserted by Messrs
William Rollisson & Sons of Tooting, London, in “The Gardeners’ Chronicle”: 218
(7 April 1855). A brief description, noting the “fine bright crimson” flowers, was
included that validates the name. Subsequently, it was published in several oth-
er English nurserymen’s catalogues (see Nelson and Small (2004: pt 1: 11)): for
example, in William Rollisson and Sons’ catalogue for 1877, which company
claimed it as a hybrid raised and sent out by them; James Fraser, Lea Bridge
Road Nursery, for 1866-1867; E. G. Henderson and Son for Autumn 1871;
James Veitch and Sons for 1873-1874; and B. S. Williams for 1881. Compilers
of gardening dictionaries soon included this heath in their works including Her-
eman (1868: 217) and Wright (ca. 1907: 325).

Guthrie and Bolus (1905), presumably unaware of the previous publication of
the epithet aemula within Erica, used the same epithet for a white-blossomed
species that inhabits rocky, coastal slopes and inland, marshy, high mountain
plateaux in the Western Cape, from Bainskloof to the Steenbras Mountains
(Oliver and Oliver 2000: 429). The species was illustrated by Schumann et al.

PhytoKeys 236: 157-178 (2023), DOI: 10.3897/phytokeys.236.110498 159

E. Charles Nelson et al.: Homonyms in Erica

(1992: 104) and is listed amongst the recognised plant species of southern
Africa by Oliver and Oliver (2003). However, it was subsequently treated as a
local variant from Gordon’s Bay of E. distorta Bartl. (Oliver and Forshaw 2012).
We follow the latter taxonomic opinion and treat E. aemula as a heterotypic
synonym of E. distorta without providing a replacement name.

Erica distorta Bartl., Linnaea 7: 644. 1832
wfo-0000671898

Erica aemula Guthrie & Bolus, Fl. Capensis 4,1: 118. 1905 [wfo-0000671313],
nom. illeg., non Erica aemula Rollisson, Gard. Chron. 1855: 218 (1855) [wfo-
1200040659]. Type. South Africa. “Bains Kloof“ [Bainskloof], Cummings 171
BOL [BOL137142 [https://plants.jstor.org/stable/10.5555/al.ap.specimen.
bol137142], syntype; “Fish Hoek, Gordon’s Bay“, Guthrie 3108 BOL (BOL137141)
[https://plants.jstor.org/stable/10.5555/al.ap.specimen.bol137141], syntype.

Type. South Africa, “Auf Felsenriicken in der Kluft nach der Platte des Tafel-
berges in vierter Hohe” (lectotype tGOET destroyed, fide annotation by N. E.
Brown on K000314148; isolectotype [fragments only], “ad Cap. b. Spei in monte
tabulari leg. Ecklon. Hb. Bg. 1841” K [K0003141 48].

2. Erica aristata Salisb. (1796) [wfo-1000053511], non Andrews
(1807) [wfo-0000671412]

Richard Anthony Salisbury’s (1761-1829) catalogue (Salisbury 1796) of the
plant collection in his own garden at Chapel Allerton, outside Leeds in York-
shire, England, gave the name Erica aristata for a plant from the Cape of Good
Hope, introduced by the nurseryman James Lee (1715-1795). The accompa-
nying diagnosis validates the binomial, but is quite inadequate to identify the
species, which evidently had not bloomed (there is no description of the flow-
ers, only the ternate, linear leaves). Guthrie and Bolus (1905) and Dulfer (1965)
placed Salisbury’s name in synonymy under E. banksii Andrews. No herbarium
specimens, determined by Salisbury, of his E. aristata are known so that equa-
tion is extremely dubious, especially given the brevity of the protologue.

A decade later the same binomial was employed by Henry Cranke Andrews
(fl. 1794-1830) when he illustrated and described the plant that currently bears
this name, an inhabitant of the Kleinrivier Mountains (Andrews 1807: t. 152;
1809a: t. 147). It is an erect, semi-spreading shrublet, to 0.5 m tall, producing
large (to 25 mm long), tubular-inflated flowers that are very sticky and have
spreading lobes; the corolla is longitudinally striped dark and light pink (Oliver
and Oliver 2000).

This species was illustrated by Schumann et al. (1992: 80) and is listed
amongst the currently recognised plant species of southern Africa (Oliver and
Oliver 2003; Oliver 2012: 489). This species is a very well-known plant due to its
striking floral morphology. It is the first Erica species for which pollination by
long-proboscid flies has been demonstrated (Lombardi et al. 2021). Changing
its name would cause considerable confusion, so a proposal will be submitted
for the conservation of E. aristata Andrews over E. aristata Salisb.

PhytoKeys 236: 157-178 (2023), DOI: 10.3897/phytokeys.236.110498 160

E. Charles Nelson et al.: Homonyms in Erica

3. Erica armata Spreng. (1825), non Benth. (1839)

Erica armata was validly published by Sprengel (1825: 2: 184), but Guthrie and
Bolus (1905: 86) and Dulfer (1965: 53) placed it in synonymy under E. sparr-
manii L.f., despite upholding E. armata Klotzsch ex Benth. (1839) as a distinct
species. Thus, Erica armata Klotzsch ex Benth. (1839: 672) is an illegitimate
later homonym. The species to which the binomial is currently applied (see OI-
iver and Oliver (2003: 427); Schumann et al. (1992: 100); Oliver (2012: 489)) is
an erect shrublet, to 0.5 m tall, bearing broadly urn-shaped to tubular, hairy, pink
flowers with exserted anthers; it occurs on rocky slopes. Fortunately, a later
synonym for it is available, Erica umbrosa (Baker 1961).

Erica umbrosa H.A.Baker, J. S. Afr. Bot. 27: 267. 1961
wfo-0000673499

E. armata Klotzsch ex Benth., Prodr. [A. P de Candolle] 7(2): 672. 1839 [wfo-
0000671419], nom. illeg., non Spreng., Syst. Veg. 2: 184. 1825 [wfo-0000671418];
Guthrie and Bolus, Fl. Capensis 4,1: 113. 1905; Dulfer, Ann. Naturhist. Mus, Wein
68: 67. 1965; Oliver, Strelitzia 29: 489. 2012. Type. South Africa. “in herb. reg.
Berol. ...in montibus prov. Worcester et Stellenbosch”, Masson, Niven, Drége (lec-
totypes B destroyed; isolectotype K [Niven 158] (det. E.G.H. Oliver) KO00314208).

Note. Specimens in other herbaria collected by Masson and Drége are variously
labelled as syntypes or isosyntypes, but their identity has not been confirmed:
GDC (Masson, F., s.n., G00494351 https://plants.jstor.org/stable/10.5555/
al.ap.specimen.g00494351); GDC (Drége, J.F., s.n., G00494352 https://plants.
jstor.org/stable/10.5555/al.ap.specimen.g00494352); HBG (Drege, J.F,
s.n., HBG515307 _ https://plants.jstor.org/stable/10.5555/al.ap.specimen.
hbg515307); TUB (Drége, J.F., s.n., TUB003182 left-hand specimen only https://
plants.jstor.org/stable/10.5555/al.ap.specimen.tub003182); S (Drége, J.F., s.n.,
S08-5237 https://plants.jstor.org/stable/10.5555/al.ap.specimen.s08-5237).

Type. South Africa. “Caledon, Elandskloof, Villiersdorp, on a steep S-facing
slope in shade in kloof with a large waterfall, 3 April 1961, E.G.H. Oliver 1423 (ho-
lotype: BOL; isotype: NBG-0199392-1 [https://plants.jstor.org/stable/10.5555/
al.ap.specimen.nbg0199392-1].

4. Erica autumnalis Hort. ex Benth. (1839), non L.Bolus (1923)

This binomial was published under the entry for Erica formosa Thunb. with a
validating diagnostic phrase by Bentham (1839) who attributed the name to
horticulturists. Bentham appended an asterisk indicating that he deemed it to
be a horticultural hybrid. Regel (1842) attributed the same name to English gar-
deners. It was not included in “Index Kewensis”, nor was it taken up or listed by
Guthrie and Bolus (1905). On the other hand, Dulfer (1965) included the name,
for a hybrid, following Regel.

Bolus (1923) probably overlooked Bentham’s use of the binomial and pub-
lished it for a Western Cape species found on moist slopes at middle altitude
distributed from the Hottentots Holland Mountains to Kogelberg (Dulfer 1964:

PhytoKeys 236: 157-178 (2023), DOI: 10.3897/phytokeys.236.110498 161

E. Charles Nelson et al.: Homonyms in Erica

146; 1965: 92; Oliver and Oliver 2000: 430; Sieben et al. 2004; Oliver 2012:
490). This species was illustrated by Schumann et al. (1992: 146) and is listed
amongst the currently recognised plant species of southern Africa (Oliver and
Oliver 2003: 427; Oliver 2012: 490).

A new name is required for the taxon and the new epithet alludes to the
autumn, which is the species’ main, Southern-Hemisphere flowering season.

Erica notoporina E.C.Nelson & E.G.H.Oliv., nom. nov.
urn:lsid:ipni.org:names:77327519-1
wfo-1000053505

pro Erica autumnalis L.Bolus, Ann. Bolus Herb. 3: 178. 1923 [wfo-0000671451],
nom illeg., non E. x autumnalis Hort. ex Benth., Prodr. [A. P. de Candolle] 7(2):
659 (1839) [wfo-1000053512]; Regel, Verh. Vereins Beford. Gartenbaues
Konigl. Preuss. Staaten 16: 307. 1842; Regel, Kult. Aufz. Eriken, 147. 1843;
Dulfer, Ann. Naturhist. Mus, Wein 68: 151 (1965).

Type. South Africa, “Bought in Adderly St., Cape Town’, 2 February 1922,
N.S. Pillans 16784: (holotype: BOL [BOL-137249 (https://plants.jstor.org/sta-
ble/10.5555/al.ap.specimen.bol137249); isotype BOL [BOL-137250]).

5. Erica capensis Regel (1842), non T.M.Salter (1935)

In his monograph on Erica, Eduard August von Regel (1815-1892) described a
plant named E. capensis (Regel 1842: 318, 1843: 158) and the binomial has had
a sporadic existence since the mid-19" century. Guthrie and Bolus (1905) did
not list Regel’s use of the epithet capensis. The binomial reappeared in the early
20" century in, for example, the seed-list of the French nursery Vilmorin Andrieux
& Cie for 1922-1923 and was recorded by Dulfer (1965: 151) who opined that
Regel’s name was synonymous with E. pelviformis Salisb. (= E. mauritanica L.).

Salter (1935) employed the same binomial for a species found in marshes
at low altitude on the Cape Peninsula (Dulfer 1964: 145, 146; 1965: 90; Oliver
and Oliver 2000; Oliver 2012: 493). This species was illustrated by Schumann
et al. (1992: 144), and is listed amongst the currently recognised plant species
of southern Africa by Oliver and Oliver (2003: 429).

Oliver and Oliver (2003: 429, 447) placed Salter’s name in synonymy un-
der Erica turbiniflora Salisb. (wfo-0000673478). As that name was validly pub-
lished and pre-dates Salter’s by more than a century, it is the correct name for
the taxon.

Erica turbiniflora Salisb., Trans Linn. Soc. 6: 377. 1802
wfo-0000673478

Erica capensis T.M.Salter, J. S. Afr. Bot. 1: 34. 1935 [wfo-0000671621], nom.
Illeg., non Regel, Verh. Vereins Beford. Gartenbaues K6nigl. Preuss. Staaten
16: 318 (1842) [wfo-1000053513], 158. 1843. Type: South Africa. “Cape Pen-
insula, marshes on lower Hout and Klaasjager River”, 14 February 1934, TM.

PhytoKeys 236: 157-178 (2023), DOI: 10.3897/phytokeys.236.110498 162

E. Charles Nelson et al.: Homonyms in Erica

Salter 4292 (holotype: BOL-137252 [https://plants.jstor.org/stable/10.5555/
al.ap.specimen.bol137252)).

Type. Without locality, Hibbert ex herb. Salisbury (lectotype, designated here: K
[K-314663]).

6. Erica constantia Hort. Ex G.Sinclair (1825), non Nois. Ex Benth. (1839)

A Cape heath named “Erica Constantia” was included by Messrs Lee & Ken-
nedy in a manuscript list of species that had been introduced into cultivation
by the firm up to 1808 (see Nelson and Oliver (2004: 138)). The binomial has
been traced in print in Conrad Loddiges & Sons’ 1818 catalogue and, three
years later, in Johann Heinrich Friedrich Link’s (1767-1851) list of plants in
Berlin Botanic Garden (Link 1821: 374). Neither of those publications con-
tained a description, but George Sinclair's (1787-1834) catalogue of the
heaths (Sinclair 1825) grown in the Duke of Bedford’s garden at Woburn,
Bedfordshire, England, did include a description (based on at least one living
plant, although Sinclair had not observed this in bloom and a herbarium spec-
imen). Sinclair (1825) attributed the name to “Hortulanis” (gardeners) and
described the heath as having leaves in threes, with bell-shaped flowers in
terminal inflorescences, with awnless anthers. He did not provide (as he usu-
ally did) the colour of the corolla because he had only seen a dried specimen.
The name is also known from at least eight other 19"-century works, includ-
ing those of Bentham (1839: 666), Regel (1842: 300) who remarked “Die als
E. constantia in deutschen Garten gehende Pflanze gehort zur E. trivialis” and
Hereman (1868: 218) whose description indicated that the cultivated plant
bore purple flowers.

However, Bentham (1839: 672) also described a species, stated to have been
named Erica constantia by Noisette — presumably the French horticulturist
Jean Claude Noisette (1772-1849) — on the basis of one of Klotzsch’s spec-
imens, then in the Berlin Herbarium and destroyed during the Second World
War. Guthrie and Bolus (1905: 115) accepted this name, having seen the type
material in Berlin “ex horto [Lee &] Kennedy, 1816”, but overlooked the earlier
usage, whereas Dulfer (1964: 108, 137; 1965: 67) chose to ignore the other use
of the same binomial (including as a synonym for his E. simulans var. trivialis
(Klotsch ex Benth.) Dulfer [wfo-0000673262]) and accepted Bentham’s second
application of the name, citing a specimen collected by Esterhuysen (28788)
from “Oudensberg” [sic. Audebsberg], Worcester District, Western Cape, as
matching Bentham’s protologue.

Frica constantia Nois. ex Benth. is currently applied to a white-flowered
heath from rocky slopes at high altitude, ranging from the Hex River Mountains
to Klein Swartberg (Oliver and Oliver 2000, 2003: 430; Oliver 2012: 492). It is
clearly not the purple-flowered taxon cultivated in European gardens during the
19" century.

Here, we designate the Esterhuysen collection cited by Dulfer (1965) as
neotype to fix the application of the name in its current sense and provide a
replacement name for the species. One meaning of the Latin word constantia
is harmony, although whether that was also the intended meaning of Lee & Ken-
nedy’s epithet cannot be determined. Concordia also means harmony.

PhytoKeys 236: 157-178 (2023), DOI: 10.3897/phytokeys.236.110498 163

E. Charles Nelson et al.: Homonyms in Erica

Erica concordia E.C.Nelson & E.G.H.Oliv., nom. nov.
urn:lsid:ipni.org:names:77327521-1
wfo-1000053500

pro E. constantia Nois. ex Benth., Prodr. [A. P. de Candolle] 7(2): 672. 1839 [wfo-
0000671735], nom. illeg., non Hort. ex G.Sinclair, Hort. eric. woburn.: 6, 32.
1825 [wfo-1000055091]; Guthrie and Bolus, Fl. Capensis 4,1: 115-116. 1905;
Dulfer, Ann. Naturhist. Mus, Wein 68: 67-68. 1965.

Type. Without locality or collector, Herb. Klotzsch (holotype: B, destroyed).
South Africa, Audensberg, S slopes. 15 February 1959, E.E. Esterhuysen 28188
(neotype, designated here, NBG [NBG-0265661-0]).

7. Erica demissa Hort. ex G.Sinclair (1825), non Klotzsch ex Benth. (1839)

“Dwarf green-flowered heath” was the English name used by Sinclair (1825)
when he described a plant cultivated in England in the early 19" century un-
der the name Erica demissa. The name (as “demisa”) was included in Messrs
Lee and Kennedy’s manuscript list, mentioned above (see Nelson and Oliver
(2004: 138)), of Cape heaths that had been introduced into cultivation by the
firm up to 1808. Within a year of this, the name was printed by Donn (1809),
Cushing (1814: 210) and in Conrad Loddiges and Sons’ catalogue for 1818.

Sinclair’s description (1825) is more than adequate to validate the name,
which Dulfer (1965: 29) noted, perhaps correctly, as a synonym of Erica coc-
cinea L., no doubt following Sinclair's grouping of the species (Sinclair 1825:
35). There is a specimen so named in LIV, but we have not examined it.

Meanwhile, Bentham (1839) chose to adopt a name proposed by Klotzsch
and, thus, published the same binomial for an entirely different species with
white to rosy-pink flowers. This is distributed on the lower to middle slopes
of the Swartberg as far east as Grahamstown (Oliver and Oliver 2000).
This species was illustrated by Schumann et al. (1992: 137) and is listed
amongst the currently recognised plant species of southern Africa (Oliver
and Oliver 2003: 432), but clearly is not the “dwarf, green-flowered” heath
known to Sinclair.

A new epithet, derived from the Greek compound opeoBtoc (oresbios)
meaning living on mountains (Stearn 1973), is published here and alludes
to the species’ habitat in “rocky veld on the Swartberg” (Schumann et al.
1992: 137).

Erica oresbia E.C.Nelson & E.G.H.Oliv., nom. nov.
urn:lsid:ipni.org:names:77327522-1
wfo-1000053507

pro E. demissa Klotzsch ex Benth., Prodr. [A. P. de Candolle] 7(2): 666. 1839
[wfo-0000671862], nom illeg., non Hort. ex G. Sinclair Hort. eric. woburn.: 8.
1825 [wfo-0000671861]; Benth., Prodr. [A. P de Candolle] 7(2): 621. 1839;
Guthrie and Bolus, Fl. Capensis 4,1: 47. 1905; Dulfer, Ann. Naturhist. Mus,
Wein 68: 86-87. 1965.

PhytoKeys 236: 157-178 (2023), DOI: 10.3897/phytokeys.236.110498 164

E. Charles Nelson et al.: Homonyms in Erica

Type. South Africa. “Uitenhaag [Uitenhage], in monitbus Vanstadensrivier”, 1000-
AOOO[ft], C.A Ecklon and C.L.P Zeyher s.n. (syntype: not traced); South Africa,
“flum. Camtoo [Gamtoos]”, Masson s.n. (syntype: not traced), Burchell 4709 (syn-
type: HAL [https://plants.jstor.org/stable/10.5555/al.ap.specimen.hal0135770)).

8. Erica flocciflora Tausch (1839), non Benth. (1839)

This is an instance of two names published within weeks of each other. Taus-
ch’s binomial was published on 28 October 1839, more than a month before
Bentham’s and, thus, has priority. Unfortunately, Dulfer (1965) consistently mis-
quoted (as 1838) the publication date of Bentham’s treatment of Erica pub-
lished in the second part of volume 7 of Augustin Pyramus de Candolle’s Pro-
dromus. Late December 1839 is accepted as being the correct publication date
for volume 7 part 2, although standard sources (e.g. Stafleu and Cowan (1976))
contain contradictory dates (for discussion, see Nelson (2005)). The first part
of volume 7 was issued in 1838, but it does not contain the Ericeae.

Given the fact he had an incorrect (earlier) date of publication, Dulfer (1965:
98, 140) maintained Bentham’s binomial as the correct name for the taxon and
disregarded Tausch’s name (which he stated was synonymous with Erica daph-
niflora Salisb.).

Frica flocciflora, as described by Bentham, occurs on the dry, lower slopes
and rocky foothills of the Kouga Mountains and has a cream corolla with a
distinctive woolly calyx (Oliver and Oliver 2000). This species was illustrated
by Schumann et al. (1992: 157) and is listed amongst the currently recognised
plant species of southern Africa (Oliver and Oliver 2003: 434).

Irrespective of synonymy, Bentham’s binomial is illegitimate because it is a
later homonym. The new epithet that we provide below continues the allusion
to floccus (Latin: tuft of woolly hairs) by adopting a Greek equivalent, uaAAWTtoc
(mallotos), fleecy.

Erica mallotocalyx E.C.Nelson & E.G.H.Oliv., nom. nov.
urn:lsid:ipni.org:names:77327523-1
wfo-1000053504

pro E. flocciflora Benth., Prodr. [A. P. de Candolle] 7(2): 660. 1839 (late Decem-
ber) [wfo-0000672068], nom. illeg., non Tausch, Flora Bot. Zeit. Regensb. 22:
629. 1839 (28 October) [wfo-0000672067]; Guthrie and Bolus, FI. Capensis
4,1: 310 (1905); Dulfer, Ann. Naturhist. Mus, Wein 68: 98 (1965).

Type. South Africa. “In colonia capensi” (cit. Bentham 1839) [‘on a rocky hill
near Groot River, Uniondale Div.], 14 March 1814, W.J. Burchell 4992 (lectotype,
here designated, K (K-314571 [https://plants.jstor.org/stable/10.5555/al.ap.
specimen.k000314571]; isolectotype, G)

9. Erica insignis Hort. (1853), non E.G.H. Oliv. (1981)

This binomial, overlooked by the compilers of “Index Kewensis”, has been traced
in no fewer than six publications issued during the latter half of the 19" century

PhytoKeys 236: 157-178 (2023), DOI: 10.3897/phytokeys.236.110498 165

E. Charles Nelson et al.: Homonyms in Erica

including the “Journal of the Royal Horticultural Society” (1853: 8: xl) and “Revue
horticole” (1882: 54: 219-220). Accompanying descriptions indicated it was ap-
plied to a scarlet-blossomed heath. It was also traced in catalogues issued by
the following British and New Zealand nurseries: James Fraser, Lea Bridge Road
Nursery for 1866-1867; William Rollisson & Sons for 1877; B. S. Williams for
1881; James Dickson & Sons, Newton Nurseries, for 1884; Nairn & Sons, Christ-
church (New Zealand) for 1896. It is highly improbable that the plant cultivated at
least until the end of the 19" century in European and New Zealand gardens was
the same species as that first collected by Stokoe in 1935 and described by Oli-
ver (1981). It inhabits rock crevices on upper, north-facing slopes of mountains
including the Anysberg and Groot Swartberg and has remarkable flowers with a
very small corolla (+ 5 mm) concealed within a greatly extended calyx (+ 20 mm).
It was illustrated by Schumann et al. (1992: 215) and is listed amongst the cur-
rently recognised plant species of southern Africa (Oliver and Oliver 2003: 436).
The new epithet alludes to the concealed corolla (from Greek: aSeAoc (ade-
los = unseen), tetadov (petalon = leaf, i.e. petal)) and echoes the name of the
section to which this species and E. nabea Guthrie and Bolus were assigned.

Erica adelopetala E.C.Nelson & E.G.H.Oliv., nom. nov.
urn:lsid:ipni.org:names:77327524-1
wfo-1000053498

pro E. insignis E.G.H. Oliv., Bothalia 13: 446. 1981 [wfo-0000672329], nom. il-
leg., non hort. in J. Roy. Hort. Soc. 8: xl. 1853 [wfo-1000053515]; Rev. hort.
94: 219-220. 1882.

Type. South Africa, “Swartberg, north slopes below Kangoberg”, 1,400 m, 11
December 1979, E.G.H. Oliver 7469 (holotype, NBG [as STE] [https://plants.jstor.
org/stable/10.5555/al.ap.specimen.nbg0133788-0]; isotypes K, PRE).

10. Erica lanata J.C. Wendl. (1798), non Andrews (1806)

Messrs Lee & Kennedy had employed the name “Erica lanata” in the manuscript
list, mentioned previously (see Nelson and Oliver (2004: 138)), of species that
had been introduced into cultivation by the firm before 1808. Andrews (1806:
t. 121) was undoubtedly using this binomial for the same plant. However, the
binomial had been published eight years earlier by Johann Christoph Wendland
(1755-1828) for a different species (Wendland 1798: 45). The persistent use of
Andrews’s binomial, despite the priority of Wendland’s, is inexplicable. As not-
ed by Dulfer (1965: 44), Wendland’s name was a synonym of Erica conspicua
Sol., which Dulfer relegated to a variety of E. curviflora L. (var. splendens (J.C.
Wendl.) Dulfer = E. splendens J.C. Wendl., non Andrews), but is currently re-
garded as a distinct species (Oliver and Oliver 2000, 2003; Oliver 2012). There
is a later name available to replace Erica lanata Andrews and that is E. flaccida
Hort. ex Link; Sinclair (1825: 10) was the first to make this equation in print.
Erica flaccida has been traced in print in several publications (e.g. Anonymous
(1808: 191); Cushing (1812: 224; 1814, 224)) and in Conrad Loddiges & Sons’
catalogue for 1811, before it was taken up by Link (1821: 1: 367), who cited

PhytoKeys 236: 157-178 (2023), DOI: 10.3897/phytokeys.236.110498 166

E. Charles Nelson et al.: Homonyms in Erica

English gardeners as his source. None of the sources published prior to 1821
included a diagnosis or description.

Erica flaccida Hort. ex Link, Enum. hort. berol. alt.: 367. 1821
wfo-0000672048

Erica lanata Andrews, Heathery, 3: t. 121 (1806); Col. engr. heaths, 3, t. 179.
1809b [wfo-0000672411], nom. illeg., non E. lanata J.C. Wendl., Bot. Beo-
bach.: 45. 1798 [wfo-0000672410]; Salisb., Trans. Linn. Soc. 6: 360 (1802);
Guthrie and Bolus, Fl. Capensis 4,1: 73. 1905; Dulfer, Ann. Naturhist. Mus,
Wein 68: 112 (1965); Oliver, Strelitzia 29: 499. 2012. Type. Icontype (illustra-
tion in Andrews, Heathery, 3: t. 121. 1806 [cit. Dulfer (1965): 112]).

Type. South Africa. “Hab. in Pr. b. sp. [Promontorium Bonae Spei = Cape of
Good Hope] ... Hort. ang/.” (B destroyed). Neotype (here designated). South Afri-
ca. Western Cape, George Dist., Outeniqua Pass. June 1960. E.G.H. Oliver 1596
NBG [NBG0112414-0].

11. Erica niveniana Hort. ex Loudon (1830), non E.G.H.Oliv. (2000)

This binomial appeared in print two centuries ago (Donn 1804: 69) and was re-
peated four years later (Anonymous 1808: 193); neither publication contained
a diagnosis. However, Loudon (1830: 147) provided a description and explicitly
attributed it to Andrews’s “heaths 2” (i.e. “Coloured Engravings of Heaths” 2: t.
112. 1802). The name also occurs in at least two mid-19" century publications:
M'Intosh (1855: 709) and Hereman (1868: 219).

Andrews (1802) did not employ the termination —iana (adjectival), but pub-
lished E. nivenia (an incorrect variant of the substantive form nivenii) (see Nel-
son and Oliver (2004: 140)). These epithets are deemed (under ICN (Shenzhen
Code) (2018, Art. 61.1 and 61.2); Turland et al. (2018)) to be simple orthograph-
ical variants, with the same type, of E. nivenii. Regrettably, this means that Oli-
ver's binomial, proposed when Syndesmanthus nivenii N.E. Br. was transferred
into Erica (Oliver 2000: 225), is an illegitimate later homonym.

The new name alludes to the silky hairs that give the inflorescences a fluffy
appearance (Oliver 2012: 486).

E. bombycina E.C.Nelson & Pirie, nom. nov.
urn:lsid:ipni.org:inames:77327525-1
wfo-1000053499

pro FE. niveniana E.G.H.Oliv., Contrib. Bolus Herb. 19: 225. 2000 [wfo-
0000672716], nom. Illeg., non E. nivenii Hort. Ex Loudon (as “nivenia”), Hort.
Brit.: 147. 1830 [wfo-1000053516] (see Nelson and Oliver (2004)).

Type. South Africa. Without locality [“Erica N95 on elevated Situations”], J.
Niven 95 (holotype K (K-000225736 [https://powo.science.kew.org/taxon/
urn:lsid:ipni.org:names:1017329-1]).

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E. Charles Nelson et al.: Homonyms in Erica

12. Erica perlata G.Sinclair (1825), non Benth. (1839)

It appears that no-one has hitherto noticed the inconsistency pertaining to the
application of the binomial Erica perlata (meaning “beset with pearls”) (Baker
and Oliver 1967: 74). Twentieth-century works consistently attribute the name
to Sinclair (1825) and apply it to an erect shrublet, up to 0.6 m tall, with small,
urn- to bell-shaped, hairy, white flowers, possessing partly exserted anthers
(Baker and Oliver 1967: t. 67; Schumann et al. 1992: 115; Oliver and Oliver 2000,
2003; Oliver 2012: 504). However, Sinclair’s protologue (1825: 18) clearly ap-
plies to a different species. He wrote:

Fol. Bractea. Anth. Pistill. Inflorescentia Color. Corol. T. Flor.
4 rem. sub. ex. ex. term. br. umbel.; R.P.1-3. Anthers Spring,
mut. corol. glob. R.R.O.7. Autumn.

In other words: leaves ternate; bracts remote; anthers subulate, exserted,
muticous; pistil exserted, inflorescences terminal, branching, umbellate; corolla
globose, pink-madder; anthers red (between scarlet and Indian red); flowering
[in Northern Hemisphere] spring and autumn.

No herbarium specimens from the Duke of Bedford’s Woburn collection, the
basis of Sinclair's descriptions, are known that could be considered as type
material, so the protologue is paramount. At least the colour of the flower sig-
nals that the protologue does not match the current application of the binomial.
Sinclair's colour codes were very precise, being determined using a specially
constructed “diagram of colours” (a colour-wheel) (Sinclair 1825: [39]—41; see
Nelson (2011: 8), fig. 5).

Dulfer (1965: 74-75, 77, 141, 153) succeeded in making the various 19"-cen-
tury applications of the binomial more confusing, although he correctly accord-
ed priority to Sinclair, albeit quoting an incorrect publication date (“1816?”) for
“Hortus ericaeus Woburnensis” (see Nelson (2003)), an error inadvertently re-
peated in Baker and Oliver (1967: 74).

According to Dulfer (1965: 74), the disparate species that were identi-
fied under this name by various authors included the European Erica /u-
sitanica Rudolph (Regel 1843: 162) and his own E. sphaeroidea Dulfer
(“Lee sec Kl[otzsch] sec Benth. [1839: 672]”) (illustrated by Schumann et
al. (1992: 119)). In synonymy under E. perlata G.Sinclair, Dulfer (1965: 75)
placed E. barbata var. minor Andrews, E. pura Lodd., E. procumbens Lodd.,
E. ephemera Tausch, Gypsocallis procumbens G.Don and Ericodes minus
Kuntze. He concluded (Dulfer 1965: 75): “E. perlata ist eigentlich nur eine in
allen Teilen kleinere Form von E. pannosa und daher kaum eine Art, sondern
eine Var. von E. pannosa.”

Baker and Oliver (1967: 74-75) were not in any doubt that Erica perlata
possessed “pearly-white flowers”, inhabited the Riviersonderend Range and
was allied to E. barbata Benth., but considering Sinclair's protologue (1825:
18), his name cannot apply to that plant. In fact, the white-blossomed Riv-
iersonderend heath appears to have no valid name because Sinclair's is not
applicable and renders Bentham’s illegitimate. Erica pura Loddiges (1817: t.
72) is a nomen nudum, while Loddiges’s E. procumbens (1833: t. 1993) is

PhytoKeys 236: 157-178 (2023), DOI: 10.3897/phytokeys.236.110498 168

E. Charles Nelson et al.: Homonyms in Erica

another illegitimate later homonym (contrary to Taylor (2016: 1127) who was
not aware of the homonymy).

We propose the replacement name Erica galantha for the Riviersonder-
end species, in allusion to the white, pearl-like flowers. Galanthos, from yaAa
(gala = milk) and Gv€oc (anthos = flower), means with a milk-white flower (as in
Galanthus L., Amaryllidaceae, the Eurasian snowdrop).

Erica galantha E.C.Nelson & E.G.H.Oliv., nom. nov.
urn:lsid:ipni.org:names:77327526-1
wfo-1000053503

pro E. perlata Benth., Prodr. [A. P. de Candolle] 7(2): 670. 1839 [wfo-100005351 7],
nom. Illeg., non E. perlata G.Sinclair, Hort. Eric. woburn.: 18 (1825) [wfo-
0000672873].

Type. South Africa, “In colonia capensi”, Drége s.n. (lectotype W, effectively
designated by Dulfer (1965: 75)) [https://plants.jstor.org/stable/10.5555/al.ap.
specimen.w0008571].

13. Erica praecox Hort. Ex G.Sinclair (1825), non Klotzsch (1838)

This binomial was printed, without accompanying descriptions, in Conrad Lod-
diges & Sons’ catalogue for 1807 and in at least two other pre-1825 publica-
tions (Anonymous 1808: 195; Cushing 1814: 210). The next publication, that
of Sinclair (1825: 22), is the significant one. As he often did, Sinclair attributed
this name to horticulturists, in this case specifically to one, “Hortul. Cormack”,
undoubtedly the English nurseryman John Cormack of New Cross, Kent, with
whom Sinclair was to enter partnership in 1827 (Harvey 1973, 1981). Messrs
Lee & Kennedy had employed the name “Erica praecox” in the manuscript list of
species, mentioned previously (Nelson and Oliver 2004: 138), that the firm had
introduced into cultivation.

Sinclair’s (1825: 19) accompanying description is, like the majority of his de-
scriptions, precise and accurate, sufficient to validate the binomial and render
Klotzsch’'s one illegitimate.

Klotzsch (1838) described a quite different species under this same
binomial. His name is currently applied to a spreading, compact shrublet, to
0.2 m tall, bearing urn-shaped to tubular urn-shaped, pink flowers between
December and February (Oliver and Oliver 2000, 2003: 443; Oliver 2012:
505). It inhabits mountain summits and ridges between the Du Toitskloof
Mountains and Villiersdorp and was illustrated by Schumann et al. (1992:
88, 89).

Dulfer (1965: 60) placed Erica praecox Klotzsch in synonym under E. ven-
tricosa var. meyeriana Dulfer and noted E. behen E.Mey. ex Klotzsch as a syn-
onym. However, Klotzsch (1838), noting E. behen as a name used in Drége’s
Herbarium, gave it as synonymous with E. savileae Andrews.

Our replacement epithet refers to the species’ occurrence high in the moun-
tains — above the clouds.

PhytoKeys 236: 157-178 (2023), DOI: 10.3897/phytokeys.236.110498 169

E. Charles Nelson et al.: Homonyms in Erica

Erica supranubia E.C.Nelson & Pirie, nom. nov.
urn:lsid:ipni.org:names:77327527-1
wfo-1000053510

pro E. praecox Klotzsch, Linnaea 12: 517. 1838 [wfo-0000672980], nom. illeg.,
non Hort. ex G.Sinclair, Hort. eric. woburn.: 19, 32. 1825 [wfo-1200010024];
Benth., Prodr. [A. PR. de Candolle] 7(2): 678. 1839.

Type. South Africa. “Dutoitskloof”, Drege 1147 (lectotype P (P-00110863), des-
ignated here (det. E.G.H. Oliver) [https://plants.jstor.org/stable/10.5555/al.ap.
specimen.p00110863]; isolectotype W [https://plants.jstor.org/stable/10.5555/
al.ap.specimen.w18890186298, https://plants.jstor.org/stable/10.5555/al.ap.
specimen.w18890158225].

14. Erica rhodantha Regel (1842), non Guthrie and Bolus (1905)

Regel (1842, 1843) published this name for a plant which Dulfer (1965: 154)
equated with Erica pelviformis (= E. mauritanica) (see above no. 4). The later
publication of the same binomial by Guthrie and Bolus (1905) is illegitimate.

The plant described and named by Guthrie and Bolus (1905) is poorly repre-
sented in herbaria, but is listed amongst the currently recognised plant species
of southern Africa (Oliver and Oliver 2003: 444). The type locality was Garcia's
Pass and the taxon has recently also been collected on the lower, northern
slopes of the Langeberg (Oliver and Oliver 2003: 444). Erica rhodantha Guthrie
and Bolus resembles E. gillii Benth., but can be distinguished from that species
by its unridged, brown anthers (ridged and black in E. gillii) (Oliver and Oliver
1994: 27). It is an erect shrublet with small, subcalycine, cup-shaped, rose-co-
loured flowers (Oliver and Oliver 2000).

The new epithet continues the allusion to the rose-red (rhodellus) flowers.

Erica rhodella E.C.Nelson & E.G.H.Oliv., nom. nov.
urn:lsid:ipni.org:names:77327528-1
wfo-1000053509

pro FE. rhodantha Guthrie and Bolus, Fl. Capensis 4,1: 288. 1905 [wfo-
0000673111], nom. illeg., non Regel, Verh. Vereins Beférd. Gartenbaues
K6nigl. Preuss. Staaten 16: 318. 1842 [wfo-1000053518], Regel, Kult. Aufz.
Eriken, 158 (1843); Dulfer, Ann. Naturhist. Mus, Wein 68: 154. 1965.

Type. South Africa. Riversdale Div.; Garcias Pass, 1200 ft, Galpin 3706 (lecto-
type BOL, effectively designated by Dulfer (1965: 131) [https://plants.jstor.org/
stable/10.5555/al.ap.specimen.bol137442]; isolectotypes K [https://plants.
jstor.org/stable/10.5555/al.ap.specimen.k000314992], NBG _ [https://plants.
jstor.org/stable/10.5555/al.ap.specimen.nbg0199737-0], PRE [https://plants.
jstor.org/stable/10.5555/al.ap.specimen.pre0309255-0], SAM [https://plants.
jstor.org/stable/10.5555/al.ap.specimen.sam0010474-0], W _ [https://plants.
jstor.org/stable/10.5555/al.ap.specimen.w19610016719].

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E. Charles Nelson et al.: Homonyms in Erica

15. Erica rugata Hort. ex G.Sinclair (1825), non E.G.H.Oliv. (2000)

This binomial was printed, without accompanying descriptions, in Conrad
Loddiges & Sons’ catalogue for 1811 and in at least two other pre-1825
publications (Cushing 1814: 227; Link 1821: 374). The next publication, that
of Sinclair (1825: 22), is, as before, the significant one. Sinclair attributed
this name to gardeners (“Hortulanis”), noting it in synonymy under the
entry for Erica rugosa Andrews; there is a cryptic (cf. ICN (Shenzhen Code)
(2018, Art. 38.14); Turland et al. (2018)) reference to Andrews'’s publication
“Heaths, vol. iv.” (i.e. “Coloured Engravings of Heaths” 4: t. 267 (post 1809;
see Cleevely and Oliver (2002))). Andrews’s E. rugosa is regarded as a
horticultural hybrid.

Unaware of the previous use of this binomial — it is not listed in botanical
indexes such as “Index Kewensis”, nor was it noted by Dulfer (1965) — Oliver
(2000: 368) chose and published the same binomial when transferring
Coccosperma rugosum Klotzsch into Erica. The new epithet alludes to
the rugose ovary of this species which is always bi-locular (fide Oliver
(2000: 369)).

Erica didymocarpa E.C.Nelson & E.G.H.Oliv., nom. nov.
urn:lsid:ipni.org:names:77327529-1
wfo-1000053501

pro E. rugata E.G.H.Oliv., Contrib. Bolus Herb. 19: 368 (2000) [wfo-0000673157],
nom. illeg., non Hort. ex G.Sinclair, Hort. eric. woburn.: 22 (1825) [wfo-
1000053519] (= E. x rugosa Andrews [wfo-0000673159)).

Type. South Africa. “Cap, im Gebirge bei der Kapstadt” [mountains near Cape
Town] [loc. 84], Zeyher s.n. (lectotype K, designated by Oliver 2000); isolectotype
MEL [https://plants.jstor.org/stable/10.5555/al.ap.specimen.mel623226)).

16. Erica spectabilis C.F. Waitz (1805), non Klotzsch ex Benth. (1839)

The name Erica spectabilis appeared in print more than thirty years before its
publication by Bentham (1839). Under his E. spectabilis, Waitz (1805: 220) quot-
ed Andrews’s description making E. formosa Andrews a synonym, an equation
he reinforced in the “Alphabetisches VerzeichnifS§ der Heidenarten” (Waitz 1805:
324). Andrews’s name was illegitimate, because of the prior publication of E.
formosa Thunb. and the plant concerned, which possessed vermilion-coloured
flowers, is regarded as being a horticultural hybrid (it was claimed by Messrs
Rollisson of Tooting).

As the variant “spectabilia’, the binomial has been traced in the 1804 cata-
logue issued by Conrad Loddiges & Sons who corrected the spelling to “spect-
abilis” in 1818.

The handsome, variable, white-, cream- to green-flowered species for
which Bentham (1839: 659) published the same binomial is restricted to
the limestone hills near the coast from Bredasdorp to Gouritsmond, whilst

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E. Charles Nelson et al.: Homonyms in Erica

a similar species, E. syngenesia Compton, with larger cream-white flowers
occurs inland, from the Witteberg to Swartberg (Oliver 2012) and Kam-
manassie Mountains (Oliver et al., in prep.). Both species were illustrated
by Schumann et al. (1992: 156), E. syngenesia from the Klein Swartberg
(Schumann et al. 1992: 156, figs 12, 13 and 14) and E. oraria (as E. spectabi-
lis) from coastal habitats near Still Bay (Schumann et al. 1992: 156, figs 10
and 11). E. syngenesia and E. oraria (as E. spectabilis) are included amongst
the currently recognised plant species of southern Africa (Oliver and Oliver
2003: 445; Oliver 2012).

The new epithet, from the Latin ora (edge or sea coast), reflects the coastal
distribution of the species.

Erica oraria E.C.Nelson & E.G.H.Oliv., nom. nov.
urn:lsid:ipni.org:names:77330003-1
wfo-1000056203

pro E. spectabilis Klotzsch ex Benth., Prodr. [A. P. de Candolle] 7(2): 659. 1839
[wfo-0000673283], nom. illeg., non Waitz, Beschreibung der Gattung und Ar-
ten der Heiden: 220. 1805 [wfo-0000673282]: Guthrie and Bolus, Fl. Capensis
4,1: 57. 1905.

Type. South Africa. “in Strandweld [Strandveld]”, Drége s.n. (syntypes: +B,
GDC [https://plants.jstor.org/stable/10.5555/al.ap.specimen.g004651 65],
HBG [https://plants.jstor.org/stable/10.5555/al.ap.specimen.hbg507932], K,
W _ [https://plants.jstor.org/stable/10.5555/al.ap.specimen.w18890186305,
https://plants.jstor.org/stable/10.5555/al.ap.specimen.w18890321973,
https://plants.jstor.org/stable/10.5555/al.ap.specimen.w0005951)).

17. Erica stenantha Sweet (1830), non Klotzsch ex Benth. (1839)

Sweet (1830: 340) published this binomial with a reference to the fourth vol-
ume of Andrews’s “heath.” and also the synonym “tenuiflora y carnea. A. H. v.
4.” (i.e. “Coloured Engravings of Heaths” 4: t. 281 (post 1824)); this indirect
reference to a previously published description validates Sweet's binomial
(ICN (Shenzhen Code) (2018, Art. 38.13); Turland et al. (2018)). Thus, Sweet
was raising Andrews’s Erica tenuiflora var. carnea to the rank of a species with
this binomial.

As noted by Dulfer (1965: 61), following Andrews’s Erica tenuiflora, E.
stenantha Sweet is a synonym of E. cylindrica Thunb. Dulfer (1965) did not
recognise that Bentham’s binomial (1839: 685) was a later, illegitimate hom-
onym. The species named E. stenantha by Bentham (1839) inhabits the up-
per slopes of the Langeberg; it is an erect shrub, with small, calycine, cup-
shaped, dark pink flowers (Oliver and Oliver 2000) and is listed amongst
the currently recognised plant species of southern Africa (Oliver and Oliver
2003: 446).

The new epithet is derived from Latin poculus (cup) and alludes to the cup-
shaped flowers.

PhytoKeys 236: 157-178 (2023), DOI: 10.3897/phytokeys.236.110498 172

E. Charles Nelson et al.: Homonyms in Erica

Erica poculiflora E.C.Nelson & E.G.H.Oliv., nom. nov.
urn:lsid:ipni.org:names:77327531-1
wfo-1000053508

pro E. stenantha Benth., Prodr. [A. P. de Candolle] 7(2): 685. 1839 [wfo-
0000673321], nom. illeg., non Sweet, Hort. Brit.: 340. 1830 [wfo-0000673320]
(= E. cylindrica Thunb.).

Type. South Africa. “Berge bei Zwellendam", C.F.Ecklon and C.L.PR Zeyher 221
(lectotype W, effectively designated by Dulfer 1965: 130) [https://plants.jstor.
org/stable/10.5555/al.ap.specimen.w0005950]; isolectotypes MEL [https://
plants.jstor.org/stable/10.5555/al.ap.specimen.mel2384382]; S__ [https://
plants.jstor.org/stable/10.5555/al.ap.specimen.s08-6141)).

18. Erica tenuis Moench (1802), non Salisb. (1802)

According to Stafleu and Cowan (1981), the supplementary volume to Moench’s
“Methodus plantas Horti Botanici et Agri Marburgensis” was issued on 2 May
1802, a little more than three weeks before Salisbury’s paper was published
in “Transactions of the Linnean Society” between 24 and 27 May 1802. Thus,
Moench’s binomial renders Salisbury’s illegitimate.

Mysteriously, Dulfer (1965: 141) stated that Moench’s name was a synonym
of “Ceramia tenuis G. Don sec. Benth., Pr. 693 (1838) [sic]”. However, Don’s pub-
lication contains no such name, nor is there any reference on p. 693 in Bentham
(1839) to this synonymy. The identity of the plant described by Moench, there-
fore, remains unknown.

Figure 1. Erica oliveranthus E.C. Nelson and Pirie, renamed in honour of E.G.H. (Ted) Oliver
and Inge M. Oliver (photo: MDP; https://www.inaturalist.org/observations/19098927).

PhytoKeys 236: 157-178 (2023), DOI: 10.3897/phytokeys.236.110498 173

E. Charles Nelson et al.: Homonyms in Erica

Erica tenuis Salisb. is the current name for a white-flowered heath (Fig. 1)
that occurs in the Western Cape from Clanwilliam to Humansdorp (Schumann
et al. 1992: 172), but it must now be replaced.

The new epithet published here is a tribute to our co-author, Dr E.G.H. (‘Ted’)
Oliver, pre-eminent authority on the taxonomy of Erica, and to his late wife
and collaborator, Inge Magdalene Oliver (1947-2003), who was an authority
on Erica in her own right. They have previously been honoured separately in
E. ingeana E.G.H. Oliv. and E. oliveri H.A.Baker (Schumann et al. 1992: 131).

Erica oliveranthus E.C.Nelson & Pirie, nom. nov.
urn:lsid:ipni.org:names:77327532-1
wfo-1000053506

pro E. tenuis Salisb., Trans. linn. Soc 6: 329. 1802 [wfo-0000673389], nom. illeg.,
non Moench, Methodus: 17. 1802 [wfo-0000673388].

Type. Without locality or collector, Ex herb. R. A. Salisbury (lectotype
K [KO00314799] [https://powo.science.kew.org/taxon/urn:|sid:ipni.
org:names:329729-1)).

19. E. tomentosa Masson (1776), non Salisb. (1802)

Masson only published one account of his botanical explorations at the Cape
of Good Hope (Masson 1776) and, in this, he recorded that on 30 December
1773, during his second journey (Masson 1776: 298-299; see Bradlow (1994:
124)), he reached:

... the hot bath, which is situated at the foot of a ridge of dry mountains: ...
Next morning, we went up to the top of this ridge of mountains ... We found
here a species of heath remarkable for having its branches and leaves all
covered with a fine hoary down or nap, which we thought singular in that
genus: we Called it Erica tomentosa.

Bradlow (1994: 157 n. 212) identified the “hot bath” as the spring situated
4 km east of the southern entrance to Toorwater Poort, in the Groot Swartberg
Range. Thunberg was with Masson on this occasion and his corresponding
specimen became the type of Erica passerina Montin (1775) (fide J.P. Rourke,
in Bradlow (1994: 157 n. 212)).

It is often difficult to decide whether a sentence such as Masson's consti-
tutes a diagnosis as defined in the International Code of Nomenclature: ‘... a
statement of that which, in the opinion of its author, distinguishes the taxon
from others’ (ICN (Shenzhen Code) 2018, Art. 38.2; Turland et al. 2018). Given
that, at this time, the early 1770s, only about fifty Erica species from the Cape
Region had been described (Oliver 2000: 4, figure 1; Nelson and Oliver 2004)
and that Masson was, by then, familiar with many more undescribed species
in their wild habitats, the clause “which we thought singular in that genus” sug-
gests that this is precisely what Masson wrote. Thus, his binomial was validly
published. However, it is a junior synonym of E. passerina.

PhytoKeys 236: 157-178 (2023), DOI: 10.3897/phytokeys.236.110498 174

E. Charles Nelson et al.: Homonyms in Erica

It follows that Salisbury’s binomial is illegitimate and has to be replaced.
The heterotypic synonym, Erica velutina Bartl. (fide Dulfer (1965: 67)), may be
employed for this taxon. It is found on the rocky, lower, southern slopes of the
Riviersonderend Mountains (Oliver and Oliver 2000, 2003: 446). An erect shru-
blet, to 0.5 m tall, E. velutina bears small, urn-shaped, finely hairy, lilac or dark
pink flowers.

Erica velutina Bartl., Linnaea 7: 645. 1832.

Erica tomentosa Salisb., Trans. Linn. Soc. 6: 327. 1802 [wfo-0000673425], nom.
illeg., non Masson, Phil. Trans. 66: 299. 1776 [wfo-1000053520] (= E. passer-
ina Montin). Type. South Africa. “Hottentots Holland”, |. Mulder s.n. ex herb.
Salisbury (not located).

Note. A label identifying the specimen labelled “C.B.S. Niven 16” (K-
000314197) [https://plants.jstor.org/stable/10.5555/al.ap.specimen.
k000314197] as a lectotype is incorrect as the protologue cited only a collec-
tion from Hottentots Holland by |. Mulder and the Niven collection is, there-
fore, not original material.

Type. South Africa. “Am Fusse des Babylonschenthurmbergen [Babilon-
storing]”, Ecklon s.n. (holotype GOET-003270 [https://plants.jstor.org/sta-
ble/10.5555/al.ap.specimen.goet003270)).

Acknowledgements

The authors thank Dr Alan Elliot (Royal Botanic Garden Edinburgh) for his in-
valuable input relating to WFO and rediscovery of outstanding nomenclatural
issues in Erica addressed here. Thanks are also due to Dr Larry Dorr and Dr
John Manning for their careful reviews of the paper and for their many pertinent
comments that have improved the paper. We acknowledge the support of The
Heather Society (now disbanded) in the development of the International Reg-
ister of Heather Names which originally brought these issues to light.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding
The Heather Society.

Author contributions

ECN: Conceptualisation, Investigation, Writing - Original draft, Writing - Review and Ed-
iting. EGHO: Investigation, Writing - Review and Editing (prior to 2023). MDP: Investiga-
tion, Writing - Review and Editing (from 2023).

PhytoKeys 236: 157-178 (2023), DOI: 10.3897/phytokeys.236.110498 175

E. Charles Nelson et al.: Homonyms in Erica

Author ORCIDs

E. Charles Nelson © https://orcid.org/0000-0002-3909-4388
Michael D. Pirie © https://orcid.org/0000-0003-0403-4470

Data availability

All of the data that support the findings of this study are available in the main text.

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