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parative anatomy of thorns of Hawaiian Lobeliaceae. Amer. Jour. Bot. 49(4): 413-419. Illus.
1962.-Species of Rollandia and Cyanea (sections Uenuinae and Palmaeformes), endemic Hawaiian genera of Lobeliaceae, are unique in the familyin possessing thorns and thorn-likestructures on leaves, and in some cases, on stems and flowers.These thorns always originate in con-
junctionwith a unicellular,non-glandular
trichomewhichterminatesthe thorn.Ontogenetic
studies show that divisions leading to the formationof the thorn occur in the ground meristem
as soon as the trichomeis differentiated.Periclinal divisions predominate at first,but anticlinal
IN LOBELIACEAE,
thorns on leaves, stems or largethornand a simpleunicellulartrichomemay
flowersare presentin only2 genera,the Hawaiian be found.Delbrouck (1875) admits.alltheseinterendemicsCyanea and R?ollandia(Wimmer,1956). grades as "Stacheln." For purposesof simplicity,
The writerstudiedthornsof these species as one "thorn," as used in Lobeliaceae, refersto a triphase of an anatomicalinvestigationof Hawaiian chome raised upon a conical base formedfrom
Lobeliaceae, the anatomy of whichhas not been groundmeristem.
studied previously. In addition, a survey of
Thorns of Lobeliaceae are limited to some
literatureon spines and thorns(e.g., Delbrouck, species of Rollandia and species of 2 sectionsof
1875; Schnee, 1939) reveals a remarkablelack of the genus Cyanea. Several species of Rollandia
anatomical studieson thornsof various kinds. A bave no thorns,althoughsome (R. calycina)have
numberofpapersdeal withthornsor spineswhich small ones. Thorns are especially prominentin
are modifiedleaves, leaf-portions,
or branch-tips. Rollandia St. Johnii,but they are comparablein
The thorns described in the present paper are size onlyto the smallerones in the thornyspecies
distinctive in being, basically, unicellular tri- of Cyanea. Species of Clyaneawhich bear thorns
chomes raised on a conical mound of tissue de- belong to the sectionsPalmaeformesand Genurived fromthe groundmeristemof stems,leaves, inae, as recognizedby Hillebrand (1888), Rock
or flowers.Delbrouck (1875) described thorns (1919) and Wimmer(1956). The most prominent
whichresemblethose of Lobeliaceae in a number thorns occur in species of Genuinae such as
of Boraginaceae,such as Echium vulgare.He also C. ferox,C. Marksii, and C. nolimetangere,
alshows that leaf teeth of Pandanus gramineus, though in some of the Palmaeformes,such as
Agave americana, and Lomatophyllummacrum C. aculeatiflora
and C. hamatiflora,
theyare almost
possess a terminaltrichomeearly in their on- as conspicuous. In C. Marksii and C. nolitogeny.The term"thorn" does not have a precise metangere
thornsmay be a centimeteror morein
botanical usage. The term "spine" seems to be length.For a detailedsurveyofthorndistribution
applied mostly to portionsof leaves or to spe- in Cyanea,see Rock (1919) or Wimmer(1956).
cialized leaves. "Thorn" as used in this paper has
MATERIALS
AND METHODS-Materials of Loa topographical,not a morphologicalsignificance. beliaceae were collectedin the fieldand preserved
The problem of definingthorns and thorn-like in formalin-acetic-alcohol.Voucher specimens
structuresin Lobeliaceae is complicatedby the were also prepared. Specimensdocumentingthe
fact that every intergradebetween a relatively presentstudyare as follows(herbariumabbreviations accordingto Lanjouw and Stafleu, 1959):
Cyanea aculeatiflora
Rock, Carlquist 571 (RSA);
1 ReceivedforpublicationOctober14, 1961.
This investigation
was aidedby a grantfromClaremont C. atra Hillebr.,Carlquist569 (RSA); C. Marksii
Rock, Carlquist 593 (RSA); C. tritomantha
A.
University
College.
2
Withouttheassistanceof a numberofindividuals,
the Gray, Carlquist 601 (RSA); Rollandia calycina
materialsfor this study could not have been collected. G. Don, Carlquist 575 (RSA); R. St. Johnii
Sincereappreciationis expressedto Mr. Edwin Bonsey, Hosaka, Carlquist H2 (UC). Additional species
of Makawao, Maui; Mr. Norman Carlson, of Kona, could have been studied,but sectionsof thornsin
Hawaii; Dr. Harold St. John,of Honolulu,Oahu; and
Mrs.Ella StephensofVolcano,Hawaii. The writerwishes the above reveal the same basic structureand
observationson thornsin otherspecies
also to thankDr. JosephF. Rock,authority
on Hawaiian superficial
Lobeliaceae,forhis interestin thisstudy.
suggestedno differences
fromthis pattern.
413
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CARLQUIST-THORNS
fIllJU?L%L~~~~~~12
IN
LOBELIACEAE
415
1314
Material was sectionedby means of the usual suitable stages forillustration.Cyanea Marksii is
paraffintechniquesand stained with a safranin- more densely covered with spines than other
fastgreencombinationcorresponding
to Northen's species; Fig. 8 shows the appearance of foliar
modification
ofFoster'stannicacid-ferricchloride thornsin this species.
technique(Johansen,1940).
As seen in Fig. 1, 9, formationof a foliarthorn
ANATOMICAL
DESCRIPTIONS-Ontogeny-Cyaneabegins with differentiation
of a protodermalcell
Marksii (Fig. 1-8; 9-14) was selected for illus- as a unicellulartrichome.In species with large
trating ontogenetic studies because: (1) the thorns,such as C. Marksii,divisionsinitiatingthe
numerousthorns,on both surfacesof leaves in spinebase occursimultaneously
withthe originof
this species,are moreeasily studiedfromsections the trichome.Althoughanticlinal divisionsmay
of vegetativeshoot apices than thornson stems be seen at any stage, periclinalones predominate
could be; and (2) this species has many stems, duringearlystages. Numerouspericlinaldivisions
providingmore shoot apices than the palmiform may be seen in Fig. 1-6, as interpretedin Fig.
species. Sectionsof shootapices of the palmiform 9-14. As the mound of tissue underlyingthe trispecies, C. aculeatiflora and C. tritomantha,chome expands, cell divisions occur in various
showed the same features but provided fewer planes, increasingthe volume of the thornbase.
Fig. 1-8. Thorndevelopment
in CyaneaMarksii.-Fig. 1-6. Stagesin expansionoftrichomeand underlying
tissues.
Fig. 1 showsearlydevelopmentof trichome.Shape of trichomes
in Fig. 2-6 is probablyalteredby fixation,
because
maturestages (Fig. 7) are invariablyconical,and someconicalyoungtrichomes
may be found.Activecell divisionin
celllayersis occurring
subepidermal
in thesectionsshownin Fig. 1-6 (see interpretive
drawingsand legend,Fig. 9-14).
-Fig. 7. Trichome-tipped
thornon abaxial surfaceofyoungleaf,suchas thatshownin Fig. 8. Laminabelow.-Fig. 8.
Youngleaf,seenfrombelow.Note largerthornson petiole;thornsare mostlyrestricted
to petioleand zonesalongveins.
Fig. 1-6, X600. Fig. 7, X66. Fig. 8, X4.
416
AMERICAN
JOURNAL
OF BOTANY
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April,19621
CARLQUIST-THORNS
IN LOBELIACEAE
417
injury by decay-causingmicroorganismsor by
epiphytes.With damage of the epidermis,formation of a peridermoccurs (Fig. 17). This periderm
developmentis only moderatelyactive, and although several layers of phellem may be developed, peridermaction never producesnotable
corkyexerescences.In very old thorns(Fig. 16),
the peridermis partlylost,presumablyby decay.
At no time do thornsdevelop internalsclerenchyma. Firmness of the old thorns can be attributedonly to succulenceof the groundtissue
parenchyma.In thisrespect,thornsofLobeliaceae
are quite unlike the thornsand spines of thornscrub and desertplants. Such thornsand spines
are composed mostly of sclerenchyma(Schnee,
1939).
Comparativedata-Studies were undertakento
determinewhat, if any, is the range in thorn
anatomyamongHawaiian Lobeliaceae. The most
obvious variation is one of size, for thornsmay
grade from relatively large structuressuch as
those describedabove to trichomesonly slightly
raisedupon an eminence.In all species examined,
unicellular trichomesnot accompanied by any
evidence of thorn formationare present (viz,
Fig. 21).
Unicellular,non-glandulartrichomeshave been
reportedin a number of Lobeliaceae (Metcalfe
and Chalk, 1950; Wimmer,1956). Such trichomes
were the only sort observed in Hawaiian Lobeliaceae of the presentstudy.Close groupingsof
unicellular trichomes,which might be termed
tufted hairs, were occasionally observed in
Rollandia calycina.
Thorn-likestructuresmay be presenton the
exteriorsurfaceof calyxand corolla,as in Cyanea
aculeatiflora(Fig. 18, 19). These "thorns" are
oftenvery softand delicate, and are characteristically long and narrow. Moreover, the unicellulartrichomewhichterminatessuch thornsis
thin-walled,as are the epidermal cells which
surroundit.
Structureswhich are not thorn-like,but are
mound-likeor knobby in shape occur on leaves
and stemsof Cyanea atra. Sectionsrevealed that
these structures(Fig. 20) appear to result from
relatively late periderm action beneath unicellular trichomes.Whether this peridermis a
normal phenomenonor resultsfrominjury (e.g.,
fractureof the trichome) could not be determined.However,emergencesof the natureof the
one figuredin Fig. 20 mightbe termedcorkwarts
rather than thorns. True cork warts (Fig. 21)
were observed in Hawaiian Lobeliaceae on the
midribof a leaf of Rollandia calycina.Periderm
418
AMERICAN
JUORNAL
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Fig. 20-23.-Fig. 20. Cyanea atra. Transection of leaf, showing cork wart formationat base of trichome remnant
of lea.f miArib. Uniserinat trichomes
Fi.
-1 Conrkwart. frnmtfransaection
(below). Fig.- 91-99. Rdlln."i. caycna
April,1962]
CARLQUIST-THORNS
419
IN LOBELIACEAE
CITED