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New combinations in South American Myricaceae
CARLOS PARRA-O.
Parra-O., C. (Instituto de Ciencias Naturales--Universidad Nacional de Colom-
bia, Apartado Aereo 7495, Bogota, Colombia; email: carlospa@ciencias.unal.
edu.co). New combinations in South American Myricaceae. Brittonia 54: 322-
326. 2002.--New combinations and lectotypification are made of the South Amer-
ican species formerly of
Myrica
in the genus
Morella. A
key and a list of syn-
onyms are presented.
Key words: Myricaceae,
Morella, Myrica,
South America.
Parra-O., C. (Instituto de Ciencias Naturales---Universidad Nacional de Colom-
bia, Apartado Aereo 7495, Bogot~i, Colombia; email: carlospa@ciencias.unal.
edu.co). New combinations in South American Myricaceae. Brittonia 54: 322-
326. 2002.--Se presentan las nuevas combinaciones y las lectotipificaciones ne-
cesarias para transferir a
Morella
las especies suramericanas anteriormente asig-
nadas a
Myrica.
Adem~is se incluyen una clave y una lista de sinrnimos.
Various problems of nomenclature and
typification of the Myricaceae have arisen
in recent years. In this article, I present a
synthesis of the nomenclature that includes
the core genus of Myricaceae,
Myrica L.
(sensu Chevalier, 1901). The problem be-
gan when the strongly different morpholo-
gy led to the genus
Myrica
being separated
into three major groups, which are well dif-
ferentiated from each other and are sup-
ported by different kinds of evidence
(Baird, 1968; Macdonald, 1989; Wilbur,
1994). As Wilbur pointed out, the level as-
signed to these groups (i.e., whether three
genera or one genus with three subgroups)
is a decision that depends on "... both tra-
dition and the prevailing philosophy of the
period." Traditionally, one of these groups
(represented by
M. asplenifolia
L.) was sep-
arated long ago as the genus
Comptonia
L'Hrr. ex Aiton because it had pinnatifid
leaves, stipules, and a burrlike fruit with 6-
8 persistent bracteoles (Bornstein, 1997;
Chevalier, 1901; Elias, 1971; Wilbur, 1994).
The other groups have been treated as
distinct taxonomic categories throughout
their history. Authors such as Elias (1971)
proposed two subgenera: subgen.
Myrica
(with two species,
M. gale
and
M. hartwegii
S. Watson) and subgen.
Morella
(with ca.
40 species).
Early authors such as Spach (1841) and
Chevalier (1901) recognized three genera:
Myrica
L. (40-50 species),
Gale
Duhamel
(two to three species), and
Comptonia
L'Hrr. ex Aiton (one species). After Che-
valier (1901) published his "Monographie
des Myricacres," other authors (Greene,
1910a, 1910b; Nieuwland, 1910) tried to re-
solve the ranking of these taxa, but some
changes were not formalized or the propos-
als were not adopted.
Recent authors (Baird, 1968; Wilbur,
1994) listed several characters that support
Spach's and Chevalier's criteria for consid-
ering the three groups as different genera.
The characters that separate
Myrica
and
Gale are
principally the terminal buds (pres-
ent vs. lacking), the vessel element distri-
bution (diffuse porous vs. ring porous), the
insertion of aments (on old wood mainly
below the leaves vs. at the summit of the
branchelets of the preceding year), the sta-
men number (highly variable, to 22 per
flower vs. consistently reduced, mostly 5 or
less), the adnation of bracteoles to the fruit
wall (none vs. strongly adnate), the fruit
(drupelike with hard wall and covered with
Brittonia,
54(4), 2002, pp. 322-326. ISSUED: 16 April 2003
9 2002, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.
2002] PARRA-O.: MYRICACEAE 323
papillae vs. nutlike with thin wall and with-
out papillae), the wax production of the
fruit (present vs. absent), and the chromo-
some number (n = 8 vs. n = 24, 48) (Baird,
1968; Wilbur, 1994).
The crux of the nomenclatural problem
(which originated with the acceptance of
Myrica
subgen.
Myrica
and
Myrica
subgen.
Morella
as two separate genera) has been
the lectotypification of the genus
Myrica.
If
M. cerifera
were designated as the lectotype
of
Myrica,
then
Myrica
could be treated in
the sense of Chevalier (1901), and
M. gale
and
M. hartwegii
should be segregated into
Gale.
However, if
M. gale
were designated
as the lectotype of
Myrica,
then all the spe-
cies belonging to subgen.
Morella
(most of
the species occurring in the Americas, Af-
rica, eastern Asia, the Philippines, and Ma-
laysia) would be included in
Morella
Lour.
Small (1903) adopted
Morella
Lout. as the
correct name for all the sSutheastern United
States species of
Myrica.
Later criticism by
Greene (1910b) discouraged the use of this
genus. Baird (1968) analyzed the problem
and proposed the suitable nomenclatural
combinations with typification of each one,
supporting the use of
Morella
(to accom-
modate all the species of
Myrica
sensu Che-
valier), but he never published these new
combinations. Wilbur (1994) independently
came to the same conclusion as Baird and
published the nomenclatural transferrals of
four species of
Myrica
from North America
and Canada to
Morella.
He proposed new
names and combinations to define subgen-
era and series within
Morella.
This proposal
was not followed by Bornstein (1997) in his
Flora of North America. Verdcourt and Pol-
hill (1997), unaware of Wilbur's paper,
summarized the most important points of
the problem and proposed the conservation
of
Myrica
and
Gale
following the criteria
of Hylander (1945) and Rehder (1949),
who considered
Myrica cerifera
to be the
lectotype of
Myrica
and made the transfer
of
Myrica gale
and
Myrica hartwegii
to
Gale.
This proposal required that
Myrica
gale
become
Gale belgica
Dumort. to avoid
tautonomy. Although some recent papers
(e.g., Ferguson, 1998) supported the lec-
totypification of
Myrica
by
M. cerifera,
the
proposal was rejected by the Committee for
Spermatophyta. The only species growing
in Sweden
(M. gale)
was well known to
Linnaeus and it seemed only right that that
species should serve as the generic type
(Killick et al., 1998; Brummitt, 1999). After
this rejection, Killick et al. (1998) trans-
ferred the African species previously ac-
cepted by K_illick (1969) and White (1993)
to
Morella,
making 13 new specific com-
binations and five infraspecific ones.
It is evident that the remaining species of
Myrica
should be transferred to
Morella,
which I do in this publication. A key to the
species of
Morella
from South America and
their synonymy are included. At present
many publications continue using names
that were already in synonomy since Che-
valier (1901) [e.g.,
M. arguta, M. caracas-
sana, M. polycarpa (M. pubescens)]. I
agree that all the South American species
of
Morella
belong to subgen.
Cerothamnus
(Tidestr.) Wilbur ser.
Cerothanmus
(Ti-
destr.) Wilbur (Wilbur, 1994).
Morella rotundata
(Steyermark & Ma-
guire) C. Parra-O. is excluded from this key
because the species' description was based
only on sterile specimens and I have seen
no fertile specimens. This species has been
poorly collected in the Chimant~i Massif in
Venezuelan Guayana. Nevertheless, vege-
tative characteristics of the isotype and pa-
ratype indicate that the species is distinct
within
Morella.
Morella Lour., F1. Cochinch. 458. 1790.
TYPE:
Morella rubra
Lour.
Cerophora
Raf., Alsog. Americana 11. 1838 (p:p.).
Myrica
L. subgen.
Morella
(Lour.) Engler, Nat.
Pflanzenfam. III 1: 27. 1893.
Myrica
sensu Chevalier, non L., Mem. Soc. Sci. Nat.
Cherbourg 32: 286. 1901.
Cerothamnus
Tidestr., Elys. Marian., Ferns. 41. 1910.
1.
Key to the Species of Morella of South America
Plants monoecious; leaf blades lanceolate, pubescent beneath (generally), the base attenuate, the main,
secondary, and/or tertiary veins beneath densely or sparsely pubescent; aments with staminate
324 BRITTONIA [VOL. 54
flowers on the lowest part of the rachis, pistillate flowers on the upper part; main bract of the
pistillate flowers lanceolate or short-lanceolate, densely pubescent at apex, margin, and dorsal
portion; fruits pubescent ................................................................... Morella pubescens
Plants dioecious (rarely monoecious); leaf blades variously shaped, glabrous or minutely and sparsely
pubescent beneath, the base acute or cuneate, the main vein (and sometimes secondary veins
beneath) densely or sparsely pubescent; aments with only staminate flowers or pistillate flowers;
main bract of pistillate flowers rhomboid, sometimes lanceolate, sparsely pubescent at apex and
margin (with shorter hairs sometimes present on dorsal portion); fruits usually glabrous.
2. Leaves lanceolate-linear, 5-7 times longer than wide; staminate inflorescence with compound
male flowers; in the Andes of southern Peru and northern Chile .......................... Morella pavonis
2. Leaves of various shapes, 2-4 times longer than wide; staminate inflorescence with single male
flowers; elsewhere in Andean South America.
3. Leaves 3-4 times longer than wide, elliptic-oblong or oblong; fruits ovoid ...........
............................................................................................................... Morella funckii
3. Leaves in a different proportion and shape; fruits spherical.
4. Young twigs usually pubescent; leaf blades obovate or elliptic-obovate, 3.1-9.1 • 1.3-
3.8 cm, the apex obtuse or subobtuse, the margin dentate or dentate-serrate; secondary
bracts in the pistillate flowers 4-6; staminal column 0.5-2.5 mm long ............
.............................................................................. Morella singularis
4. Young twigs pubescent or glabrous; leaf blades of various shapes, 1.1-6.4 • 0.4-2.2
cm, the apex acute or subacute, the margin entire or serrate; secondary bracts in the
pistillate flowers 2-4; staminal column 0.2-1.2 mm long
5. Leaf blades elliptic or narrowly elliptic, the margin entire or sometimes serrate and
then only in the distal 1/3, glabrous; staminate flowers with 2 ovoid or rhomboid
secondary bracts; secondary bracts in the pistillate flowers 3-4; Andean Vene-
zuela, Colombia, Ecuador, and Peru, 2000-4000 m ........................... Morella parvifolia
5. Leaf blades obovate-lanceolate or obovate-oblong, the margin serrate (at least in the
distal 2/3), sometimes pubemlent in and/or between teeth; staminate flowers with
or without secondary bracts, if present the bracts linear; pistillate flowers with 2-
3(4) secondary bracts; Andean Bolivia and Argentina, 1400-2700 m ............
.................................................................................. Morella chevalieri
Morella
funckii (A. Chev.) C.
Parra-O.,
comb. nov. Myrica funckii A. Chev.,
Mem. Soc. Sci. Nat. Cherbourg 32: 286.
1901.
TYPE: VENEZUELA. M6rida: Culata,
8000 ft, Apr 1847, Funck & Schlim 1541
(LECTOTYPEI
here designated, P;
ISOLEC-
TOTYPE" G).
Morella parvifolia (Benth.) C. Parra-O.,
comb. nov.
Myrica parvifolia Benth., P1. Hartweg. 251. 1846.
TYPE: ECUADOR. Monte de Tiopullo dicto prope
Quito, Hartweg 1378
[HOLOTYPE: K;
photos:
COL,' MO, US (F negative no. 11552)].
Myrica parvifolia var. obtusa Benth., Pl. Hartweg
25l. 1846. TYPE: COLOMBIA: prope Bogota,
Hartweg 1379
(HOLOTYPE: K;
photo: COL).
Myrica parvifolia var. longipedunculata A. Chev.,
Mem. Soc. Sci. Nat. Cherbourg 32: 284. 1901.
TYPE: COLOMBIA. Boyac~i: "entre Tunja et Pay-
pa, ~ 1400 poises d'alt," Mar 1843, Linden 1305
[LECTOTVPE: here designated,
P; photo: MO (F
negative no. 11581)].
In the original description of Myrica par-
vifolia var. longipedunculata, Chevalier
(1901) cited two specimens of Linden 1305,
one of them deposited in "H. Mus." (P),
the other in "H. Bruxelles" (BR). The du-
plicate at BR has not been found there (R
Stoffelen, pers. comm.).
Myrica parvifolia vat. lucens A. Chev., Mem. Soc.
Sci. Nat. Cherbourg 32: 285. 1901. TYPE: CO-
LOMBIA. Prov. Bogot~i, 1844, Goudot s.n.
(LEC-
TOTYPE: here designated, P).
Myrica parvifolia vat. macrostachya A. Chev.,
Mem. Soc. Sci. Nat. Cherbourg 32: 284. 1901.
TYPE: COLOMBIA. "Prov. Bogot~i: dans les An-
des, ~t 2900 m d'alt," Triana 825 p.p. (LECTO-
TYeE:
here designated,
P; ISOLECTOTYPES:
MO,
NY, US).
Myrica parvifolia vat. major A. Chev., Mem. Soc. Sci.
Nat. Cherbourg 32: 286. 1901. TYPE: ECUADOR.
"Andes," 1861, Spruce 5135 (rlOLOTYPE: P).
Morella
pavonis (C. DC.) C.
Parra-O.,
comb. nov.
Myrica pavonis C. DC., Prodr. 26 (2): 151. 1868.
TYPE: PERU: prope Huayaquil, Ruiz & Pavdn s.n.
[LECTOTYPE:
here designated,
G; photos: COL,
MO, US (F negative no. 8538)].
Myrica pavonis var. glandulosa A. Chev., Mem.
Soc. Sci. Nat. Cherbourg. 32: 288. 1901. TYPE:
2002] PARRA-O.: MYRICACEAE 325
PERU: Ruiz & Pavdn s.n. [LECTOTYPE:
here des-
ignated,
P; ISOLECTOTYPE: BM (not found)].
In the original description of Myrica pa-
vonis, de Candolle (1864) cited two speci-
mens in the protologue: "In Peruvi~ prope
Huayaquil (herb. Pavon in h. Boiss.)" and
"Santa prope Lima (herb. Kew. Sim n.
88)." I designate the specimen at G as the
lectotype of M. pavonis for a few reasons:
the specimen "Sim n. 88" was not found
at K (S. Owens, pers. comm.); the sheet de-
posited in G is an excellent sample showing
all the diagnostic characters of the species;
and G is the depository of de Candolle's
herbarium.
The label of the lectotype evidently con-
tains a geographical error. The original la-
bel says: "Myrica salicifolia de Huayaquil.
1785. Peru. Flor Masculina." However, it is
likely that the label incorrectly identifies
"Huayaquil" as the collection locale and
that the lectotype comes from Peru and not
from Ecuador. This is supported by the fol-
lowing facts. Though Ruiz and Pavrn had
wanted to collect in Guayaquil and nearby
provinces in Ecuador, various factors pre-
vented them from visiting these localities
(Steele, 1964). Furthermore, at the time this
specimen was collected, Ruiz and Pavrn
were in Peru, near Hminuco and adjacent
areas (Steele, 1964). Based on available ev-
idence, there are no herbarium specimens
of Morella from Guayaquil, Ecuador (in-
cluding nearby areas), or from zones near
sea level (Laegaard, 1999). Morella pavonis
is known to grow only in central and south-
ern Peru and in northern Chile, between
1000 and 3200 m.
Morella pubescens (Humb. &
Bonpl. ex
Willd.) Wilbur, Rhodora 103: 121. 2001.
Myrica pubescens Humboldt & Bonpland ex Will-
denow, Sp. P1. 4: 746. 1806. TYPE: "Habitat in
Nova Granada," Humboldt & Bonpland s.n. [HO-
LOTYPE: herb. Willdenow in B; photos: NY, US
(F negative no. 11553)].
Myrica arguta Kunth, Nov. Gen. Sp., 2: 17, t. 98.
1817. TYPE: COLOMBIA. "Crescit in regno No-
vogranatensi in Paramo de San Fortunato, juxta
viam Fusagasugensem, alt. 1480 hexap. Floret
Septembri," Humboldt & Bonpland s.n. [HOLO-
TYPE: P (n.v.); microfiche: NY; photo: NY (F neg-
ative no. 39603)].
Myrica caracasana Kunth, Nov. Gen. Sp., 2: 18.
1817. TYPE: VENEZUELA. "Crescit in alsis Pro-
vinciae Venezuelae, in monte Silla de Caracas,
alt. 1200 hex., Floret Januario," Humboldt &
Bonpland s.n. [HOLOTYPE: P (n.v.); photo: US (F
negative no. 39605); microfiche: NY]. Myrica pu-
bescens var. caracasana (Kunth) A. Chev., Mem.
Soc. Sci. Nat. Cherbourg 32: 292. 1901.
Myrica macrocarpa Kunth, Nov. Gen. Sp. 2: 16. 1817.
TYPE: PERU. "Crescit in temperatis regni Peruviani
prope Ayavaca, alt. 1370 hexap." Humboldt & Bon-
pland s.n. (HOLOTYPE: P; microfiche: NY).
Myrica polycarpa Kunth, Nov. Gen. Sp. 2:18.1817.
TYPE: "Crescit cum Myrica arguta in regno No-
vogranatensi in Paramo de San Fortunato, juxta
viam Fusagasugensem, alt. 1480 hex, floret Sep-
tembri," Humboldt & Bonpland s.n. [HOLOTYPE:
P (n.v.); microfiche: NY].
Myrica interrupta Benth., P1. Hartweg. 251. 1846.
TYPE: COLOMBIA: "Juxta cataratam Tequinda-
ma, prov. Bogota," Hartweg 1380 (HOLOTYPE: K;
photos: COL, NY, US (F negative no. 11554)].
Myrica pubescens var. interrupta (Benth.) A.
Chev., Mem. Soc. Sci. Nat. Cherbourg 32: 294.
1901.
Myrica arguta var. peruviana C. DC., Prodr. 16(2):
153. 1868. TYPE: PERU. Pavon s.n. (HOLOTYPE:
G; photo: COL).
Myrica arguta var. tinctoria C. DC., Prodr. 16(2):
153. 1868. TYPE: PERU. Pavon s.n. (LECTOTYPE:
here designated,
G; photo: COL; ISOLECTOTYPE:
P).
Myrica pubescens vat. glandulosa A. Chev., Mem.
Soc. Sci. Nat. Cherbourg. 32: 293. 1901. TYPE:
PERU: Hartweg 832
(LECTOTYPE: here desig-
nated,
P; ISOLECTOTYPE: NY).
Myrica pubescens var. tomentosa A. Chev., Mem.
Soc. Sci. Nat. Cherbourg 32: 294. 1901. TYPE:
PERU: "Rivrro-Canta" (LECTOTVPE:
here des-
ignated, P).
Myrica costata Rusby, Descr. S. Amer. P1.8. 1920.
TYPE: BOLIVIA. La Paz. Prov. Noryungas: Un-
duavi, Buchtien 2810 (HOLOTYPE: NY; ISOTYPES:
NY, US).
Morella rotundata
(Steyermark & Maguire)
C. Parra-O., comb. nov.
Myrica rotundata Steyermark & Maguire, Mem.
New York Bot. Gard. 17 (1): 441. 1967. TYPE:
VENEZUELA. Bolfvar: Apacar~i-tepui, Chimantzl
Massif, Steyermark 75084 [HOLOTYPE: VEN
(n.v.); tSOTYPE: NY].
Morella singularis
(C. Parra-O.) C. Parra-
O., comb. nov.
Alchornea cerifera Croizat, Caldasia 2: 128. 1943.
TYPE: COLOMBIA. Putumayo: Lado sur de la
Laguna de la Cocha, Cuatrecasas 11824 (HOLO-
TYPE: US; ISOTYPE: COL). Myrica singularis C.
Parra-O., Caldasia 21: 5. 1999. nom. nov. for A1-
chornea cerifera Croizat, non Myrica cerifera L.,
Sp. P1. 2: 1024. 1753.
326 BRITTONIA [VOL. 54
Morella chevalieri C. Parra-O., Brittonia
52: 321. 2000. TYPE: BOLIVIA. Santa
Cruz: Prov. Florida, 4 km W of center of
Samaipata, 18~ 63~ 2000-
2100 m, 31 Dec 1992 (fr), M. Nee & L
Vargas 43436 [HOLOTYPE: NY; ISOTYPES:
AD (n.v.), MO, LPB (n.v.), SI (n.v,),
USZ (n.v.)].
Myrica pubescens var. glabra A. Chev., Mere. Soc.
Sci. Nat. Cherbourg 32: 293. 1901. TYPE: PERU:
"Andes de l'interior," 1836-1837, Mathews 939
(HOLOTYPE: P).
Acknowledgments
I thank the fellowship program of the
Andrew W. Mellon Foundation for financ-
ing my research as a visiting student at The
New York Botanical Garden from July
1999 to January 2000. I thank the staff at
The New York Botanical Garden for allow-
ing access to the herbarium and the library,
and for providing the necessary facilities
for completing this investigation. Thanks to
D. Lentz (NY) and M. Nee (NY)
for their
continuous support in all aspects related to
this study. Thanks to S. Owens and C.
Barker (K), P. Morat and M. Pignal (P), M.
Baessler (B), E Jacquemoud (G), P. Stof-
felen and E. Robbrecht (BR), and C. Jarvis
and R. Vickery (BM) for loans, photo-
graphs, and information concerning the
types. Thanks to the curators of the herbaria
at COL, MO, and US for providing access
to their collections, and to Instituto de Cien-
cias Naturales of the Universidad Nacional
de Colombia for administrative support.
Thanks to D. Verdcourt (K) for sending me
a copy of his article and commentaries on
the nomenclature changes of the African
species of Morella. I am also indebted to
A. Macdonald, Lakehead University, Ca-
nada; A'. Bornstein (SEMO); and an anon-
ymous reviewer for their valuable com-
ments on the manuscript.
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