Reinstatement of Bonellia (Theophrastaceae)

Author(s): Bertil Ståhl and Mari Källersjö

Source: Novon, Vol. 14, No. 1 (Mar., 2004), pp. 115-118
Published by: Missouri Botanical Garden Press
Stable URL: http://www.jstor.org/stable/3393447 .
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 Reinstatementof Bonellia (Theophrastaceae)
BertilSt&hl
Departmentof Science and Technology,GotlandUniversity,SE 62167 Visby,Sweden.
bertil.stahl@hgo.se

MariKallersjo
MolecularSystematicsLaboratory,
SwedishMuseumof NaturalHistory,P.0. Box 50007,
SE 10405 Stockholm,Sweden.mari.kallersjo@nrm.se

ABSTRACT. Followingthe resultsof a recentphy- tablishedto accommodatea species previouslyin-

logeneticstudyshowingthatthe genusJacquiniais cluded in Jacquinia.
paraphyletic, the followingnewcombinations in the Followinga newanalysisbasedon a combination
genusBonelliaare made(all by Stahl& Kallersjo): of morphologicaland cpDNA data (Kallersjo&
B. albiJYora (Lundell),B. brevifolia(Urban),B. Stahl, 2003), it has become clear that the genus
ffammea(Millspaugh ex Mez),B.frutescens(Miller), Jacquiniais not monophyleticand has to be divid-
B. Iippoldii(Lepper),B. Iongifolia(Standley),B. ed intotwogenera.Accordingto thatanalysis,Jac-
Ioeffingii(Carrasquel), B. macrocarpa (Cavanilles), quinias. str.shouldincludespecies withpseudov-
B. macrocarpa(Cavanilles) Stahl & Kallersjo erticillateleaveslackingorwitha poorlydeveloped
subsp.pungens(A. Gray),B. macrocarpa(Cavan- mucro,lepidoteyoungshoots,whitishflowers,glo-
illes) Stahl & Kallersjosubsp.panamensis(Lun- bose seeds completelyimmersedin placentaltis-
dell), B. montana(StahI),B. mucronata(Roemer& sue, and sunkenabaxialfoliarsclerenchyma.This
Schultes),B. nervosa(Presl),B. nitida (Stahl),B. circumscription makes Jacquiniaa chiefly Carib-
paludicula(Standley),B. pringlei(Bartlett),B. pau- bean genus of 13 species. Only one of these spe-
cifora (Stahl& F. Axelrod),B. shaferi(Urban),B. cies, J. armillarisJacquin,occurs outside of the
seleriana(Urban& Loesnerex Mez), B. sprucei Caribbeanislands,in northerncoastalSouthAmer-
(Mez),B. stenophylla(Urban),andB. umbellata(A. ica and easterncoastalBrazil(Stahl,1992, 1995).
DC.).The genusBonelliais distinguishedfromJac- JacquiniaarboreaVahl,witha wide Caribbeandis-
quinia s. str. by its mostlyorangeflowersand al- tribution,has also been reporteda few times from
ternateleaves,flattenedseeds thatareincompletely Mesoamerica,but then only from islands off the
coveredby placentaltissue, and two-layeredtesta. easterncoast of Mexicoand Honduras.
A key to the generaof the Theophrastaceae is pre- The reinstatedgenus Bonelliais definedby al-
sented. ternateleaves with a well-developedmucro,gla-
Keywords: Bonellia,Theophrastaceae. brous or puberulousyoung shoots, mostlyorange
flowers,flattenedseeds only partlyimmersedinto
The Theophrastaceae form a small Neotropical placentaltissue, and abaxialfoliar sclerenchyma
family of about 100 species of shrubsand small locatedadjacentto the lowerepidermis.Twospe-
trees. Recent phylogeneticresearch(Kallersjoet cies, B. Iongifoliaand B. paludicola,deviate by
al., 2()00)has shownthatSamolusL., a genustra- havingwhitishflowers,and it is thereforeinterest-
ditionallyincluded in the Primulaceae,is sister ing to note that bothappearat the base of the Bo-
groupto the Theophrastaceae and thereforeshould nellia clade in the phylogenyproposedby Kallersjo
be includedwithinthe latterfamilyor recognized and Stahl (2003). Of these, B. paludicolais the
as a separatefamily. mostproblematical species since it, like mostmem-
The Theophrastaceae s. str.(excludingSamolus) bers of Jacquinias. str. (exceptJ. keyensisMez),
weremonographed by Mez(1903),whorecognized has pseudoverticillate leaves.It shouldalso be not-
fourgenerabasedmainlyon floralmorphology, viz. ed that one additionalspecies, B. albiflorayhas
Clavija,Jacquinia,Deherainia,and Theophrasta.white flowers,but that species is morphologically
Votsch(1904), in a leaf anatomicalwork, trans- very similarto B. macrocarpa,indicatinga close
ferredone of the species of Deherainiato a new relationshipbetweenthese two taxa.
genus,Neomezian and duringmorerecenttaxonom- The genus Bonellia consists of 22 species dis-
ic work(Stahl,1993), the genus Votschiawas es- tributed in Mesoamerica,northernand western
NOVON
14: 115-118. 2004.

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116
South America,and the GreaterAntilles. Within
the genus two moFphologically and geographically
distinctgroupscan be discerned.Oneof thesecom- lb. Branched
prises the South Americanspecies B. mucronata
(northernPeru),B. sprucei(Ecuador),B. frutescens
(northern Colombia-Venezuela), B. Ioefingii (north-
ern Venezuela),and the Mesoamerican B. nervosa.
All species in this grouphave ratherlarge,thick-
walled and opaque fruits, comparativelyflexible
leaves (deciduousin B. nervosa),and whitened
youngshoots.The secondgroupconsistsof species
fromCuba,Hispaniola,and PuertoRico (B. brevi-
folia, B. Iippoldii,B. paucifora,B. stenophylla,B.
umbellata)with small or very small spine-tipped
leaves, small, thin-walledfruits,and tiny flowers.
Theremaining,Mesoamerican speciesaremoredif-
ficult to define as a naturalgroup,althoughsome
species appearto be closely related(Stahl,1989;
Kallersjo& Stahl,2003).
The name Bonellia was published by Colla
(1824) in a catalogof plantsfromhis privategar-
den. The namewas evidentlyproposedby C. G. L.
Bertero,whosematerialof a plantfromSantaMar-
tha in northernColombiawas distributedunderthe
name of Bonellia cavanillesiito herbariaand bo-
tanical gardensin Europe.However,since Colla
(1824) cited JacquiniamacrocarpaCavanilles,a
differentMesoamerican species, as a synonym,the
nameB. cavanillesiihas to be treatedas an obligate
synonymof the latterspecies.
The name Bonellia honorsthe Italianzoologist
FrancoAndreaBonelli (1784-1830), evidentlya
close friendof Berteroand Colla, but whose con-
tributionto botanyis obscure.
KEYTOTHEGENERAOFTHEOPHRASTACEAE
S. STR.
la. Unbranchedor sparselybranchedshrubsand
trees; fully developedleaves 20-120 cm long,
marginsoftenserrateor serrulate.
2a. Corollarotate,tube smoothon inside;stam-
inodes ovoid-gibbous;flowersoftenunisex-
uaLmale or functionallymale flowerswith
stamen filamentsfused into a permanent
tube; anther not or inconspicuouslypro-
ducedat apex;leaf marginsentire,serrulate,
or serrate ........ ClavijaRuiz& Pav6n Bonellia brevifolia (Urban)Stahl & Kallersjo,
2b. Corollacampanulateor bowl-shaped,irreg-
ularly puckeredon inside of tube; stami-
nodesnot obovoid;flowersbisexual,stamen
filamentsfusedat base only;antherconnec-
tive distinctlyprolongedat apex;leaf mar-
gins usuallyspinose-dentate.
3a. Plants erect, the stem spiny; corolla
deeply campanulate,pale buff turning
black upon drying;staminodestrans-
verselyoblong,insertedwithinthe co-
rollatube .... Theophrasta L.
3b. Plants semi-decumbent,the stem not
spiny;corollabowl-shaped,orange,not
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.................
....................
.................
................
...................
................
Deherainia
Votschia
Bonellia
Decaisne
Stahl
Colla
Novon
blackeningupondrying;staminodestri-
angular,minute,insertedat the mouth
of the corollatube .............NeomeziaVotsch
shrubsor small trees;leaves usually
less than20 cm longemarginsentire.
4a. Corollagreen, > 2 cm diam. at anthesis;
staminodesobtrullateto linear;fruitsovoid
4b. Corollaorange,whitish,or pale yellow,< 2
cm diam.at anthesis;staminodesflattened,
petaloid.
5a. Corollabowl-shaped,pale yellow;style
shorterthan the ovary;foliar scleren-
chymaof the lowerside arrangedin a
continuousor subcontinuous layer . . .
5b. Corollacampanulateor urceolate,or-
angeorwhitish;styleof the samelength
as the ovary;foliar sclerenchymaar-
rangedin distinctbundles.
6a. Youngshoots + lepidotefromir-
regularlybranched,thick-walled
hairs;leavesmostlypseudoverticil-
late (alternatein J. ke^ensis),with-
out or witha poorlydevelopedapi-
cal mucro;corollawhiteor whitish;
seeds + globose,coveredby pla-
centaltissue;abaxialfoliarscleren-
chyma often somewhatimmersed
intothe mesenchyma. . . JacquiniaL.
6b. Youngshootspuberulous fromshort
uniseriatehairsor glabrous;leaves
alternate(pseudoverticillate in B.
paludicola),usuallydistinctlymu-
cronate;corollaorange,sometimes
white or whitish;seeds flattened,
incompletelycoveredby placental
tissue;abaxialfoliarsclerenchyma
adjacentto lowerepidermis . .
TAXONOMY ANDNEWCOMBINATIONS
BoneHia Colla, HortusRipul. 21. 1824. TYPE:
BonelliacavanillesiiBerteroex Colla(-Jac-
quinia macrocarpa Cavanilles).
Bonellia albiflora (Lundell)Stahl & Kallersjo,
comb.nov.Basionym:JacquiniaalbifloraLun-
dell, Wrightia2: 60. 1960. TYPE:Guatemala.
Peten:Tikal,29 July 1959, Contreras 45 (ho-
lotype, LL; isotypes,S, TEX, US).
comb.nov.Basionym:JacquiniabrevifoliaUr-
ban, Symb.Antill. 1: 379. 1899. TYPE:Cuba.
Sine loco, Wright2913 (lectotype,designated
in Stahl(1995), GH;isotypes,G, GOET,M).
Bonellia flammea (Millspaughex Mez) Stahl &
Kallersjo, comb. nov. Basionym:Jacquinia
J/ammea Millspaughex Mez, Pflanzenr.IV.
236a:40. 1903.TYPE:Mexico.Yucatan: Silam,
Gaumer531 (lectotype,designatedin Stahl
(1989), G; isotypes,A, BM, BR, C, CAS,DS,
E, F, FHO,GH,K, LE,MO,NY,UPS,US, W).
Volume 14, Number 1
2004
Bonellia frutescens (Miller)Stahl & Kallersjo,
comb.nov.Basionym:Ruscusfrutescens Miller,
Gard.Dict. ed. 8: unpaginated(species no. 8
underRuscus). 1768. TYPE:Colombia.Bolf-
var:vic. of Cartage, no date, Heriberto279
(neotype,designatedin Stahl(1995), US).
Bonellia lippoldii (Lepper)Stahl & Kallersjo,
comb.nov.Basionym:Jacquinia lippoldii Lep-
per, Wiss. Z. Friedrich-Schiller-Univ. Jena,
Math.-Naturwiss. Reihe32: 876. 1983. TYPE:
Cuba.Guantanamo: SanAntoniodel Sur,Abra
de Mariana,E of Abra,100400 m alt., 9 Feb.
1979, Bisse et al. s.n. (holo-andisotype,HAJB
no. 39068).
Bonellia loeflingii (Carrasquel) Stahl& Kallersjo,
comb.nov.Basionym:Jacquinia loeJ8ingiiCar-
rasquel,ActaBot.Venez.4: 348. 1970. TYPE:
Venezuela.Anzoategui:about10 kmfromPir-
itu on roadto Clarines,Jan. 1968,Aristeguieta
et al. 6524 (holotype,VEN).
Bonellia longifolia (Standley)Stahl & Kallersjo,
comb. nov. Basionym:Jacquinia longifolia
Standley,J. Wash.Acad. Sci. 14: 241. 1924.
TYPE:E1 Salvador.San Vicente:vic. of San
Vicente, alt. 350-500 m, 2-11 Mar. 1922,
Standley 21159 (holotype,US; isotypes,F, G,
GH, K, NY).
Bonellia macrocarpa (Cavanilles)Stahl & Kal-
lersjo,comb.nov.Basionym:Jacquinia macro-
carpa Cavanilles,Icon. 5: 55, tab. 483. 1799.
TYPE:Mexico.Acapulco,Feb. 1791, lVee's.n.
(lectotype,designatedin D'Arcy (1980), B-
WILLD;isotypes,HBG,MA).
Bonellia macrocarpa (Cavanilles)Stahl & Kal-
lersjosubsp.pungens (A. Gray)StahldzKal-
lersjo, comb. nov. Basionym:Jacquinia pun-
gens A. Gray,Proc.Amer.Acad. Arts5: 325.
1854. TYPE:Mexico. Sonora:hills between
Rayonand Ures, 10 Oct. 1851, Thmrber903
(lectotype,designatedin Stahl (1989), GH;
isotypes,GH, NY).
Bonellia macrocarpa (Cavanilles)Stahl & Kal-
lersjo subsp. panamensis (Lundell)Stahl &
Kallersjo, comb. nov. Basionym:Jacquinia
panamensis Lundell, in Woodson& Schery,
Ann. Missouri Bot. Gard. 27: 329. 1940.
TYPE:Panama.Panama:vic. of Bejuco, 18
Oct. 1938, Allen 985 (holotype,MICH;iso-
types, GH, MO,US).
Bonellia montana (Stahl) Stahl & Kallersjo,
comb. nov. Basionym:Jacquinia montana
Stahl,NordicJ. Bot. 9: 27. 1989. TYPE:Nic-
aragua.Jinotega:Cerrode la Cruzand vic., W
of Jinotega,alt. 1200-1400 m, 7 July 1947,
Standley 10968 (holotype,F).
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Stahl & Kallersjo
Reinstatement of Bonellia 1 17
Bonellia mucronata (Roemer& Schultes)Stahl
& Kailersjo,comb. nov. Basionym:Jacquinia
mucronataRoemer& Schultes,Syst. Veg. 4:
802. 1819. TYPE:Peru. Colasay,Humboldt
3581 (holotype,B-WILLD,seen on microfiche;
isotypes,HAL,P).
Bonellia nervosa (Presl)Stahl& Kallersjo,comb.
nov.Basionym:JacquinianervosaPresl,Reliq.
Haenk.2: 67. 1835. TYPE:Mexico.Sine loco,
Haenkes.n. (holotype,PR).
Bonellia nitida (Stahl)Stahl & Kallersjo,comb.
nov.Basionym:Jacquinianitida Stahl,Nordic
J. Bot.9: 24. 1989. TYPE:Honduras.Comay-
agua:vic. of Comayagua,alt. 600 m, 12-13
Mar. 1947, Standley& Chacon6025 (holo-
type, F).
Bonellia paludicula (Standley)Stahl& Kallersjo,
comb. nov. Basionym:Jacquiniapaludicola
Standley,Field Mus. Nat. Hist., Bot. Ser. 11:
138. 1932. TYPE:Belize. PuntaGorda,"For-
est Home,"alt. 60 m, 11 Sep. 1932, Schipp
1028 (holotype,F; isotypes,A, BM,G, GH,K,
MICH,MO,NY, S).
Bonellia pauciflora (Stahl& F. Axelrod)Stahl&
Kallersjo, comb. nov. Basionym:Jacquinia
pauciJYoraStahl& F. Axelrod,Brittonia50: 1.
1998. TYPE:PuertoRico. MunicipioQuebra-
dillas, BarrioCharcas,0.7 kmSSEof Charcas,
200-350 m alt., 23 June 1994, Proctor et al.
49406 (holotype,SJ;isotypes,GB, NY).
Bonellia pringlei (Bartlett)Stahl & Kallersjo,
comb.nov.Basionym:Jacquinia pringlei Bart-
lett, Proc. Amer.Acad. Arts 44: 630. 1909.
TYPE: Mexico. Guerrero:Iguala Canyon,8
Oct. 1908, Pringle 10337 (holotype,GH;iso-
types, BH, BM,C, CM,E, F, G, GOET,HBG,
K, L, LE, M, MICH,MO, MSC,NY, P, PH,
PR, US, VT,W).
Bonellia seleriana (Urban& Loesnerex Mez)
Stahl& Kallersjo,comb.nov. Basionym:Jac-
quinia seleriana Urban & Loesnerex Mez,
Pflanzenr.IV.236a: 43. 1903. TYPE:Mexico.
Oaxaca:above San CarlosYauhtepec,5 Jan.
1896, Seler& Seler1759 (lectotype,designat-
ed in Stahl(1989), GH;isotypes,A, NY).
Bonellia shaferi (Urban)Stahl& Kallersjo,comb.
nov. Basionym: Jacquinia shaferi Urban,
Symb.Antill.7: 320. 1912. TYPE:Cuba.Hol-
guin: vic. of Holguin, 7 Apr. 1909, Shafer
1231 (lectotype,designatedin Stahl (1995),
NY;isotypes,A, F, GH, HAC,K, MO,US).
Bonellia sprucei (Mez)Stahl & Kallersjo,comb.
nov. Basionym:JacquiniaspruceiMez, Pflan-
zenr. IV. 236a: 43. 1903. TYPE: Ecuador.
Guayas:Chanduy,Spruce6478 (lectotype,des-
ignatedin Stahl (1990), K; isotypes,BM, BP,
CGE,G, LE, OXF,P, W).
 Novon
1 18

Bonellia stenophylla (Urban)Stahl & Kallersjo, , G. Bergqvist& A. A. Anderberg. 2000. Generic

comb. nov. Basionym:Jacquiniastenophylla realignmentsin primuloidfamiliesof the Ericaless.l.:
A phylogeneticanalysisbasedon DNAsequencefrom
Urban, Symb. Antill. 1: 378. 1899. TYPE: threechloroplastgenes and morphology. Amer.J. Bot.
Cuba. "CubaOcc.," 1865, Wright2912 p.p. 87: 1325-1341.
(lectotype,designatedin Stahl(1995), G; iso- Mez, C. 1903. Theophrastaceae. Pp. 1-21 in A. Engler
types, BM, GH, GOET,HAC, LE, M, NY, S, (editor),
Das Pflanzenreich 15 (IV,236a).
Stahl, B. 1989. A synopsisof CentralAmericanTheo-
US, W). [See Stahl(1995) for comments.] phrastaceae.NordicJ. Bot.9: 15-30.
Bonellia umbellata (A. DC.) Stahl & Kallersjo, . 1992. On the identityof Jacquiniaarmillaris
comb.nov.Basionym:JacquiniaumbellataA. (Theophrastaceae) and related species. Brittonia44:
DC.,Prodr.8: 150. 1844. TYPE:PuertoRico. 5g60.
. 1993. Votschia,a new genus of the Theophras-
Sine loco, Berteros.n. (holotype,G-DC,seen taceae fromsoutheasternPanama.Brittonia45: 204-
on microSche;isotypes,H, MEL,MO,P). 207.
. 1995. A synopsisof Jacquinia(Theophrastaceae)
Literature
Cited in the Antillesand SouthAmerica.NordicJ. Bot. 15:
493-511.
Colla,L. A. 1824. Hortusripulensis.Torino. Votsch,W.1904. Neue systematische-anatomischeUnter-
Kallersjo,M. & B. Stahl.2003. Phylogenyof the Theo- suchungenvon Blattund Achse der Theophrastaceen.
phrastaceae (Ericaless.l.). Int.J. P1.Sci. 164:579-591. Bot.Jahrb.Syst.33: 502-546.

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