Epidermal characteristics of toxic plants for cattle

41 Pages·2006·2.63 MB·English

ISSN 0373-580 X
Bol. Soc. Argent. Bot. 40 (3-4): 241 - 281. 2005
Epidermal characteristics of toxic plants for cattle from
the Salado river basin (Buenos Aires, Argentina)
SUSANA E. FREIRE1,2, ANA M. ARAMBARRI1, NÉSTOR D. BAYÓN1, GISELA SANCHO1,2,
ESTRELLA URTUBEY2, CLAUDIA MONTI1, MARÍA C. NOVOA1, and MARTA N. COLARES1
Summary: One hundred and eighty species belonging to 41 families inhabiting the Salado River
Basin of the province of Buenos Aires (Argentina) were previously reported to be toxic for cattle.
The purpose of this study was to provide a tool to distinguish the taxa when the plant material is
desintegrated. In this way, an approach to the identification of these taxa through leaf epidermal
features (anticlinal epidermal cell wall patterns, cuticular ornamentation, stomata, and hair types) is
performed. A key to the 180 species as well as illustrations of diagnostic characters are given.
Key words: Buenos Aires, Salado River Basin, toxic plants, anatomy, epidermal characters,
stomata, hairs, Dicotyledons, Monocotyledons.
Resumen: Caracteres epidérmicos de las plantas tóxicas para el ganado de la Depresión del
Salado (Buenos Aires, Argentina). Las plantas tóxicas para el ganado están representadas en
la Depresión del Salado (provincia de Buenos Aires, Argentina) por 180 especies pertenecien-
tes a 41 familias. El objetivo del presente trabajo es determinar estos taxa a partir de material
desintegrado, utilizando caracteres epidérmicos foliares (paredes anticlinales de las células
epidérmicas, ornamentación de la cutícula, tipos de estomas y pelos). Se brinda una clave para
la determinación de las especies e ilustraciones de los caracteres diagnósticos.
Palabras clave: Buenos Aires, Depresión del Salado, anatomía, caracteres epidérmicos,
estomas, pelos, Dicotiledóneas, Monocotiledóneas, plantas tóxicas.
Introduction
ThepoliticalprovinceofBuenosAiresissituated
incentraleasternArgentina.Itiscoveredinmostof
its surface by a herbaceous grassy steppe, called
(cid:147)pampas.(cid:148)Thespeciesstudiedarefromaplainwitha
poorlydevelopeddrainagesystem.Thisareaisknown
asSaladoRiverBasinwhichrepresentsapproximately
80,000km2(Fig.1).Theprincipaleconomicactivityin
this area is the cattle breeding, based on natural
pastures. The knowledge of the vegetation of this
areaisrelevantforhumandevelopment.
Numerousfloristicstudieshavebeencarriedout
inthisarea(Cabrera,1963-1967;Vervoorst,1967;
Cabrera & Zardini, 1978; Cabrera et al., 2000).
WithinthediverseplantfamiliesinhabitingtheSa-
lado River Basin, 180 species belonging to 41
familiesshowtoxicityforcattle(Cas(cid:243)s,1935;Rate-
ra,1945;Tokarnia&Dobereiner,1982;Ragonese&
Milano,1984;Gallo,1979;Pertusi,1987).Epidermal
traits, i.e. epicuticular wax deposition, cuticular
1 Área de Botánica, Departamento de Ciencias Biológi-
cas, Facultad de Ciencias Agrarias y Forestales, UNLP,
calle 60 y 119, C.C. 31, 1900 La Plata, Argentina).
Fig. 1. Map of Buenos Aires province showing the Salado
2 División Plantas Vasculares, Museo de La Plata, Paseo
River Basin (area in gray).
del Bosque s/n, 1900 La Plata, Argentina.
241
Bol. Soc. Argent. Bot. 40 (3-4) 2005
ornamentation,epidermalcells,stomata,andhairs,have patterns used in (Table 1) was adapted from Stace
provedtobeanimportanttoolintaxadelimitationin (1965):Stace(cid:146)stypes1and2correspondtotype1
many plant families (Metcalfe & Chalk, 1950-1979; here; Stace(cid:146)s types 3 and 4 correspond to type 2
Uphofetal.,1962;Sinclair&Sharma,1971;Lackey, here; Stace(cid:146)s types 5 and 6 correspond to type 3
1978;Arambarri&Colares,1993;Ditschetal.,1995;
here; Stace(cid:146)s types 7 and 8 correspond to type 4
Barthlottetal.,1998;Stengleinetal.,2003)andalso
here.Theanticlinalepidermalcellwallpatternsare
in distinguishing fragmented vegetables from feces
indicated in Table 1 as: adaxial surface / abaxial
andstomachcontentsbeingresistanttothedigestive
surface.
process(Yagueddœ&Cid,1992;Pellizaetal.,1997;
Stomata types were classified according
Cid&Sierra,2004).Therefore,itwouldbeinterestingto
Metcalfe & Chalk (1950, 1979) and Van Cotthem
seekmorphologicaltraitsthatledtodiagnoseoncauses
(1970), however, to establish the monocotyledons
ofanimalmortalitythroughfecesandstomachcontents.
types, we followed Fryns-Claessens & Van
In order to achieve this goal, we apply herein histo-
morphologicalcharacters,suchasleafepidermalfeatures. Cotthem(cid:146)s(1973)classification.
Leaf margin (visible in transparent leaves) was
Material and Methods onlyusedtodistinguishtwogroupsofspecieswith
glabrous leaves.
Plant material studied ThenomenclaturefollowsZuloagaetal.(1994)
The study was performed using fresh leaves andZuloaga&Morrone(1996,1999).
(collected in Buenos Aires province), and dried EpidermalcharactersinPoaceaeweredescribed
leaves taken from herbarium specimens belonging accordingtotheterminologyofMetcalfe(1960)and
to LP, LPAG, LPS, and SI (acronyms according Ellis(1979),andhairterminologyfollowsMetcalfe
Holmgren et al., 1990). The 180 taxa investigated &Chalk(1950,1979)andUphof(1962).Forglandu-
andvouchersaredetailedinAppendix1. larhairs(Table1)thenumberinbracketsindicates
Methods the number of head cells, and for non-glandular
Fully expanded leaves were selected for the hairsthenumberofhaircellsabovetheepidermis.
study. Data were obtained from the central area of Theobservedhairsandstomatatypesaredescribed
the midlamina on both surfaces. For reconstitution according to Metcalfe & Chalk (1979), Ramayya
of dried leaves we followed D(cid:146)Ambrogio de (1962), andUphof(1962).
Arg(cid:252)eso (1986). Then, the material was fixed in
Results and Discussion
formalin, glacial acetic acid, and 50% ethanol at a
5:5:90 ratio (F.A.A.). Most of the epidermal
Fromthe41studiedfamiliesinhabitingtheSala-
microcharacters were studied by peeling and/or in
do River Basin, the most representated were:
samples cleared using the technique of Dizeo de
Asteraceae(41spp.),Poaceae(16spp.),Solanaceae
Strittmatter (1973). However, the replica method
(14spp.),Fabaceae(13spp.),andBrassicaceae(10
(accordingtoFreeman,1984)wasusedinsometaxa
spp.). TheresultsarepresentedinTable1.
where it was not possible to get epidermis by Even if stomata and indumentum types of the
peeling or chloral hydrate clearing. To study the speciesstudiedarebasicallyinagreementwiththose
epidermal characters of the species belonging to describedbyMetcalfe(1960),Metcalfe&Chalk(1979,
PoaceaewefollowedthetechniqueofMetcalfe(1960). 1989)andUphof(1962)forallfamiliesstudied,amore
Thesemipermanentslideswerestainedusingsaffranin detailed discussion of some families is included,
in80%ethanolandmountedingelatine-glycerine. because they have some microcharacters that would
Observations, and original drawings were made be cited for the first time and/or they present some
withalightmicroscope,LeitzSMluxwithcameraluci- specialtraits.Theyareasfollows:
da.Measurementsofstomata(lengthandwidth)and
hairs were taken using a Nikon light microscope Apiaceae (6 spp. surveyed): We found
equippedwithanocularmicrometer.Theaveragesize anomocytic, diacytic, and paracytic stomata types;
of hairs and stomata were determined based on all species showed glabrous epidermal surfaces,
measurementsperformedon15-20replicatespersample. although papillae were found at midvein level in
Cuticularornamentationwascitedonlywhenit two species of Ammi. It is the first time in which
wasconspicuous(Table1) diacytic stomata type has been cited for Apiaceae,
whereas anomocytic and paracytic have been
Theclassificationofanticlinalepidermalcellwall
242
S. E. Freire et al., Epidermal characteristics of toxic plants for cattle
Table 1. Epidermal characters of the toxic plants of Salado River Basin. Numbers in brackets by the species refer
to bibliography cited below the table.
MONOCOTYLEDONS
Taxa Stomatype Stomata Stomata Anticlinalcell Trichometype Shortcells Silicabody H P Pa
subsidiarycells lengthxwidth wallpatterns
(shape) ((cid:181)m) (adax/abax)
Amaryllidaceae
Habranthustubispatus Anomocytic 30-40x20-25 1/1
Rodophialabifida Anomocytic 30-35x12-20 1/1
Cyperaceae
Cyperusrotundus Paracytic Dome 34-44x29-33 4/4
Juncaginaceae
Triglochinpalustris Tetracytic 25-35x15-20 1/1
Poaceae
Brizaminor Paracytic Parallel-sided 46-50x20-26 1/1 1-3 Quadrangular +
Cortaderiaselloana(1) Paracytic Dome 31-33x22-28 1/1 Macrohairs 1-3 Rectangular +
Triangularto Macrohairs;microhairs
Cynodondactylon(1-4) Paracytic 18-21x13-18 2/2 1-many Saddle +
dome (hemispherical)
Triangularto Macrohairs;microhairs
Cynodonhirsutus(4) Paracytic 20-25x14-21 2/2 1-many Saddle +
dome (hemispherical)
Macrohairs;microhairs
Digitariasanguinalis Paracytic Dome 34-41x22-30 2/2 (3)5-many Dumb-bell + +
(rod-like)
Triangularto Macrohairs;microhairs
Echinochloacrusgalli(1,5)Paracytic 27-37x18-25 1/1 1-many Dumb-bell + + +
dome (rod-like)
Saddle,
Eleusineindica(1,2,6) Paracytic Triangular 31-42x20-30 2/2 Microhairs(pear-like) 1
rectangular
Elymusbreviaristatus Paracytic Parallel-sided 41-62x19-27 1/2 2 Quadrangular + +
Triangularto Macrohairs;microhairs
Eragrostiscilianensis Paracytic 23-27x16-26 2/2 2-many Saddle +
dome (hemispherical)
Quadrangular
Holcuslanatus(1) Paracytic Parallel-sided 22-40x18-26 1/1 Macrohairs 1-3 +
torectangular
Hordeummurinumssp. Quadrangular
Paracytic Parallel-sided 28-45x12-17 1/1 Macrohairs 1 + +
murinum(1) torectangular
Triangularto Microhairs
Leptochloachloridiformis Paracytic 18-22x15-18 2/2 1-many Saddle + +
dome (hemispherical)
Parallel-sidedto
Loliummultiflorum(1) Paracytic 49-56x19-26 1/1 1-4 Rectangular +
dome
Parallel-sidedto
Loliumperenne Paracytic 49-56x20-22 1/1 1-2 Rectangular
dome
Parallel-sidedto Quadrangular
Loliumtemulentum Paracytic 40-49x22-25 1/1 2-3 + +
dome torectangular
Dumb-bellto
Sorghumhalepense(1) Paracytic Triangular 30-32x18-23 2/2 Microhairs(rod-like) 1-many +
cross
DICOTYLEDONS
Taxa Cuticular Stomatatype Stomata Anticlinalcell Trichomestype Trichomes Special
ornament lengthxwidth wallpatterns ornament characters
ation ((cid:181)m) (adax/abax) ation
Amaranthacecae
Amaranthushybridus Anomocytic 12-18x22-24 1/3 Conical(many)
Conical(many);papillae
A.viridis Anomocytic 20-25x10-15 3/3
onmidvein
Apiaceae
Anomocytic;
Ammimajus + ca.25x15-20 1-2/1-2 Papillaeonveins
diacytic
A.visnaga + Anomocytic 25-30x20 1-2/1-2 Papillaeonveins
Coniummaculatum + Anomocytic 25-30x20 3/3
Cyclospermum
+ Diacytic ca.20x15 2-3/2-3
leptophyllum
Stomata
Eryngiumpaniculatum + Paracytic 15-20x10-15 1/1 parallelto
veins
Anomocytic;
Foeniculumvulgare + ca.25x20 1-2/1-2
diacytic
Apocinaceae
Conical(1-3,rarely
Vincamajor(7) Paracytic ca.35x25 2/3
many)
243
Bol. Soc. Argent. Bot. 40 (3-4) 2005
Table 1. Continuation
Asclepiadaceae
Asclepiascurassavica(8) + Paracytic ca.25x15-20 1/2 Conical(2-many) +
Paracytic;
A.mellodora(8) + 25-30x20-25 1/1 Conical(2-many) +
paralellocytic
Morreniabrachystephana
+ Paracytic 20-25x15-20 1/1 Conical(2-many) +
(8)
M.odorata(8) + Paracytic 20-25x15-20 1/1 Conical(2-many) +
+(apical
Oxypetalumsolanoides(8) + Paracytic 25-30x15-20 1/2 Conical(2-many)
cell)
Asteraceae
Achyroclinesatureioides Whip(3-4);glandular(2-
Anomocytic 20-22x18-20 1/2
(11,12) seriate)
Conical(3-many);
Ambrosiatenuifolia Anomocytic 24-30x20-30 1-2/1-2 +
glandular(2-seriate)
SymmetricalT-shaped;
Anthemiscotula Anomocytic 21-30x21-24 1/2
glandular(2-seriate)
Whip(3-8);glandular(2-
Arctiumminus Anomocytic 18-21x15-15.5 1/2
seriate)
Astersquamatusvar.
Anomocytic 20-30x20-24 1/1-2 Whip(8-9);conical(2-5)
squamatus
Baccharisartemisioides
Anomocytic 24-26x24-26 1/1 Whip(3)
(13,14)
Pilosenest:conical(4-
B.articulata(13,14,15) Cyclocytic 40-46x36-40 1/1 5)andglandular(2-
seriate)
B.coridifolia(13,14) Anomocytic 33-36x24-27 1/2 Bulbiferous(4-5)
Pilosenest:non
B.notosergila Cyclocytic 54-66x39-48 1/1 glandularandglandular
(2-seriate)
Pilosenest:non
B.rufescens(13,14) Cyclocytic 45-66x39-57 1/1 glandularandglandular
(2-seriate)
Pilosenest:non
B.trimera(13,14) Anomocytic 24-42x24-33 1/1 glandularandglandular
(2-seriate)
Conical(2-8);glandular
Bidenspilosavar.pilosa Anomocytic 30-38x20-26 3/3
(2-seriate)
Whip(ca.11);shaggy
Carduusacanthoides Anomocytic 21-30x18-21 1/2
(onmargins)
Whip(10-11);glandular
Centaureacalcitrapa Anomocytic 21-24x18-21 3/3
(1-seriate)
Whip(8-10);glandular
C.melitensis Anomocytic 24-30X15-21 1/1-2
(1-seriate)
Whip(2-3);glandular(1-
C.solstitialis Anomocytic 24-30x18-24 1/2
seriate)
Cichoriumintybus Anomocytic 30-36x18-24 3/3
Whip(3-5);shaggy;
Cirsiumvulgare Anomocytic 21-30x18-21 1/2-3
glandular(2-seriate)
Conyzabonariensisvar.
Anomocytic 24-30x15-21 2/2 Conical(3-6)
bonariensis
Whip(2);glandualr(2-
Cynaracardunculus Anomocytic 24-27x19-21 1/1
seriate)
Gaillardiamegapotamica
+ Anomocytic 24-48x21-30 1/1 Conical(2-6)
var.megapotamica
Glandular(2-seriate)
Hymenoxysanthemoides Anomocytic 20-38x16-30 2-3/2-3
sunken
H.cabrerae Anomocytic 27-30x21-27 2/2 Glandular(2-seriate)
Lactucaserriola Anomocytic 20-24x16-20 2/2 Shaggy
Whip(2-4);glandular(2-
Onopordonacanthium Anomocytic 15-18x12-15 1/2
seriate)
Seneciobonariensis Anomocytic 36-45x30-36 1/1
Seneciobrasiliensisvar.
Anomocytic 30-39x21-30 1/2 Whip(3-4)
tripartitus
S.grisebachiivar.
Anomocytic 36-45x27-30 1/3 Whip(3-5)
grisebachii
S.madagascariensis Anomocytic 30-33x24-30 1/3
S.tweediei Anomocytic 45-54x27-42 1/2 Whip(ca.9)
244
S. E. Freire et al., Epidermal characteristics of toxic plants for cattle
Table 1. Continuation
S.vulgaris Anomocytic 30-36x24-30 1/3 Whip(ca.9)
Silybummarianum Anomocytic 27-33x24-30 1/1 Shaggy(onmargins)
Solidagochilensisvar.
Anomocytic 20-28x18-22 1/1 Conical(3-many)
chilensis
Sonchusoleraceus Anomocytic 24-28x20-24 1/1
Conical(many),very
Tagetesminuta(16) Anomocytic 18-28x14-20 3/3 rarely;glandular
glandular(2-seriate)
Taraxacumofficinale Anomocytic 20-30x18-22 2/2
Verbesinaencelioides Anomocytic 22-30x18-24 2/3 Conical(3-5) +(apical
cells)
AsymmetricalT-
Vernoniarubricaulisvar.
rubricaulis(17) +slightly Anomocytic 27-33x21-24 1/3 shaped;glandular(2- +
seriate)
Conical(2-4);glandular
Wedeliaglauca Anomocytic 28-34x20-22 1-2/1-2 +
(2-seriate)
Conical(3-5);glandular
Xanthiumcavanillesii Anomocytic 22-30x14-24 2/2
(2-seriate)
Xanthiumspinosumvar. Conical(2-5);glandular
Anomocytic 24-38x20-26 3/3
spinosum(15) (2-seriate)
Boraginaceae
Falcate(1);glandular Pedestal
Anchusaofficinalis(9) Anomocytic 50-55x30-35 3/3
capitate(1),rarely trichomes
Pedestal
Echiumplantagineum Anomocytic 20-25x15-20 1-2/2 Conical(1)
trichomes
Conical(1);glandular
Heliotropiumamplexicaule Anomocytic 30-35x15-20 3/3 capitate(1)withlong
stalk
Brassicaceae
Brassicanigra Anisocytic 15-20x15 2/2-3 Conical(1)
Anisocytic;
B.rapa 25-30x20-15 2/2 Conical(1)
anomocytic
Anisocytic; Stellatesessile3-
Capsellabursa-pastoris 30-25x25-20 1-2/3
anomocytic armed;conical(1)
Cardariadraba Anisocytic 25x25-20 1/1-2 Falcate(1);conical(1)
Anisocytic; Conical(1);barrel-
Raphanusraphanistrum ca.25x20 1-2/2-3
anomocytic shaped
Conical(1);barrel-
R.sativus Anisocytic 30-20x15 2/3
shaped
Rapistrumrugosum Anisocytic 20-30x15 1/2-3 Falcate(1)
Roripanasturtium-
Anomocytic 25-30x15-20 1-2/2-3 Conical(1)onmargins
aquaticum
Sisymbriumaltissimum Anomocytic; Longtoverylong
20-25x15-20 2-3/3
(18) anomocytic conical(1)
Anisocytic;
S.irio 20-25x15-20 1-2/3 Longconical(1)
anomocytic
Cactaceae
Opuntiaarechavaletae Hexacytic 30-45x25-45 1/1
Parodiaottonis(10) Parallelocytic 30-45x25-45 2/2 Moniliform
Wigginsiatephracantha Thickanticlinal
Parallelocytic 35-50x25-40 1/1 Moniliform
(10) walls
Caprifoliaceae
Sambucusaustralis + Anomocytic 45-50x30-45 1/3 Shortbristles(1),rarely
Caryophyllaceae
Agrostemmagithago Diacytic 42-60x34-46 1/1 Bristles(1)
Mesophyllwith
Polycarpontetraphyllum Anomocytic 24-42x14-24 3/3
drusas
Saponariaofficinalis Anomocytic 30-45x24-30 3/3 Moniliform(5-7)
Anomocytic; Shortconical(1);
Spergulaarvensis 27-30x15-18 1/1
diacytic glandularcapitate(1)
Chenopodiaceae
Bassiascoparia Anomocytic 20-25x20 2/2 Conical(3)
245
Bol. Soc. Argent. Bot. 40 (3-4) 2005
Table 1. Continuation
Chenopodiumalbum Anomocytic 20-25x15-20 1/2 Vesicular
C.hircinumvar.hircinum Anomocytic 25-35x15-25 2/2 Vesicular
C.murale Anomocytic 20-30x20 2/2 Vesicular;whip(3-5)
Mesophyllwith
Holmbergiatweedii Anomocytic 25-35x20-25 1-2/1-2
drusas
Salsolakalivar.kali Paracytic 20-25x20 1-2/1-2 Conical(1)
Convolvulaceae
Conical(1);glandular
Convolvulusarvensis + Paracytic 30x20-25 1-2/1-2
capitate(many)
Euphorbiaceae
Euphorbialathyrus Anomocytic 22-30x14-24 1/1-2
E.peplus Anomocytic 20-24x14-24 3/3
Mesophyllwith
Manihotgrahamii Paracytic 24-30x12-16 2/3
drusas
Ricinuscommunis + Paracytic 16-22x12-20 1-2/1-2 Shortconical(1)
Fabaceae
Adesmiabicolor(19) Anisocytic 25-30x20-30 2/2 Conical(2) +
Anisocytic;
Galegaofficinalis 17-25x15-20 1-2/3 Bristles(2)
anomocytic
Anisocytic;
Lotuscorniculatus(20) 20-30x15-25 1/2 Bristles(2)
anomocytic
Anisocytic;
L.glaber(20) 15-20x10-15 1/2 Bristles(2)
anomocytic
Anisocytic;
Lupinusgibertianus 25-40x20-30 2/2 Conical(2)
anomocytic
Anisocytic; Bristles(2),glandular
Melilotusalbus 20-25x15-20 2/3 +
anomocytic capitate(4)
Anisocytic; Bristles(2);glandular
M.indicus 15-20x12-17 2/1 +
anomocytic capitate(1)
Anisocytic;
M.officinalis 20-25x15-20 2/3 Conical(2) +
anomocytic
Parkinsoniaaculeata Anomocytic 20-30x15-25 1-2/1-2 Bristles(1) +
Sennacorymbosa Paracytic 20-30x20-25 2/2 Bristles(1)
Anisocytic;
Trifoliumrepensf.repens 20-30x20-25 1/2 Glandular(1-seriate)
anomocytic
Viciagramineavar. Bristles(2);glandular
Anomocytic 35-40x25-30 2/3
graminea capitate(1)
Anisocytic; Bristles(2);glandular
V.sativassp.nigra 25-30x20-25 3/3
anomocytic capitate(1)
Fumariaceae
Crystals
Fumariacapreolataf.
Anomocytic 25-30x20-25 1-2/3 aroundthe
capreolata
veins
F.officinalis Anomocytic 25-30x20-25 1/1-2
F.parviflora Anomocytic 30-35x25-30 1-2/1-2
Geraniaceae
Bristles(1);glandular Mesophyllwith
Erodiumcicutarium Anomocytic 25-30x20-25 2/3 capitate(1) +(rarely) drusas
Bristles(1);bottle-
E.malacoides Anomocytic 25-30:20-25 2/3 shaped;glandular +(rarely)
capitate(1)
Conical(2);glandular
Geraniummolle Anomocytic 20-30x15-25 1/3
capitate(1)
Lamiaceae
Conical(1-2);stellate
Lamiumamplexicaule Diacytic 25-35x20 3/3 sessile;glandular +
capitate(4)
Porrectstellate;
Marrubiumvulgare(15) Diacytic 25-30x15-20 1/1-2
glandularcapitate(4)
Malvaceae
Anisocytic; Conical(1);glandular
Malvasylvestris 25-35:15-20 3/3
anomocytic capitate(2-4)
246
S. E. Freire et al., Epidermal characteristics of toxic plants for cattle
Table 1. Continuation
Stellatemultiangulate;
Anisocytic;
Sidarhombifolia ca.20x10-15 2/2 glandularcapitate(2-
anomocytic
many)
Martyniaceae
Glandularcapitate
Ibicellalutea Anomocytic 15-20x15-20 2/2-3 (manyandradially
arranged)
Menyanthaceae
Nymphoidesindica(21) Anomocytic 20-25x15-20 1/1 Hydropoten
Nyctaginaceae
Boerhaviadiffusavar. Anisocytic;
ca.30x25-30 1-2/2 Glandularcapitate(1)
leiocarpa anomocytic
Anisocytic;
Mirabilisjalapa 35-40x25-30 1-2/2-3 Glandularcapitate(2)
anomocytic
Oxalidaceae
Oxaliscorniculatavar. Bristles(1);glandular
Anomocytic 23-28x13-18 2/2 +
corniculata capitate(1)
Passifloraceae
Passifloracaerulea(15) Anomocytic 20-30x20-25 1/2
Phytolaccaceae
Shortconical(1-2),
Phytolaccaamericana(22) Anomocytic 30-40x20-25 1/1
rarely
P.dioica(22) Anomocytic 30-40x20-25 1/1
P.tetramera(22) Anomocytic 35-45x25-30 1/1
Plantaginaceae
Plantagoaustralisssp. Polocytic;
20-30x15-25 1/1 Glandularcapitate(2)
australis anomocytic
Plumbaginaceae
Limoniumbrasiliensevar.
+ Anomocytic 26-32x22-30 1/1 Licopoliglands
brasiliense(23)
Polygonaceae
Polygonumaviculare(24) Anisocytic 25-30x15-20 1/1 Waxes
P.convolvulus(24) + Anomocytic ca.25x15-20 1-2/1-2 Shortconical + Vesicularcells
Shaggy;conical- Vesicularcells
Anisocytic;
P.lapathifolium(24) 20-25x15-20 1/1 flagellate(1);glandular (nearthe
paracytic
capitate(many) margins)
Vesicular
Shaggy;glandular
P.persicaria(24) Anomocytic 25-30x15-20 1/1-2 cells;
capitate(many)
hydropoten
Anisocytic; Mesophyllwith
Rumexcrispus 45-50x25-30 1/1-2
anomocytic drusas
Anisocytic;
R.obtusifolius 35-40x20-30 2/2 Papillae
anomocytic
Anisocytic;
R.pulcher 25-30x15-20 1/1 Papillaeonveins
anomocytic
Portulacaceae
Portulacaoleracea Parallelocytic ca.25x15 1/1 Papillae
Primulaceae
Anagallisarvensis Anomocytic 34-50x24-30 3/3 Glandularcapitate(1-2)
Ranunculaceae
Anemonedecapetalavar. Bristles(1);glandular
Anomocytic 34-45x28-36 2/3
decapetala clavate(1)
Bristles(1);glandular
Clematisbonariensis Anomocytic 33-45x30-33 2-3/2
clavate(1)
Bristles(1);glandular
C.montevidensis Anomocytic 30-48x27-33 1/1
clavate(1)
Ranunculusapiifolius Anomocytic 30-45x24-33 1-2/3
Mesophyllwith
R.cymbalaria + Anomocytic 27-36x27-30 1-2/2
crystalsand
R.muricatus Anomocytic 54-66x36-51 1-2/3 Bristles(1)
R.repensvar.repens Anomocytic 42-51x30-39 3/3 Bristles(1)
247
Bol. Soc. Argent. Bot. 40 (3-4) 2005
Table 1. Continuation
Rhamnaceae
Mesophyllwith
Discariaamericana(25) Anomocytic 40-45x20-30 1/1 Bristles(1)
drusas
Rubiaceae
Borreriaverticillata Paracytic 30-32x22-25 2/3 Barrel-shaped
Galiumrichardianumssp. Mesophyllwith
Paracytic 35-45x20-30 3/3 Bristles
richardianum raphides
Rutaceae
Fagarahyemalis Anomocytic 20-24x10-12 2/2 Stellate,rarely
Sapindaceae
Bristles(1);peltate
Dodoneaviscosa Anomocytic 30-40x20-25 1/1-2
scales
Scrophulariaceae
Stellatestalked;
Verbascumthapsus(15,
Anomocytic 25-28x20-28 1-2/2 glandularcapitate(2-
26)
many)
Solanaceae
Conical(2-many);
Cestrumparqui(27) Anomocytic 35-45x30-35 1-2/1-2 glandularcapitate(2-
many)
Conical(2-many);
Anisocytic;
Daturaferox 20-30x15-20 1/2 glandularcapitate(2- +
anomocytic
many)
Anisocytic; Mesophyllwith
Nicotianaglauca(27) ca.25x15-20 1-2/2-3
anomocytic crystalsand
Mesophyllwith
N.longiflora(26) Anisocytic 30-35x15-20 1-2/2-3 Conical(3)
crystalsand
Dendritic;Y-shaped;
Physalisviscosa(27,28) Anisocytic 20-30x15-20 2/2 glandularcapitate(2- +/+
many)
Conical(2-many);Y-
Anisocytic; shaped(rarely);
Salpichroaoriganifolia(27) 30-35x20-25 2/2-3 +
anomocytic glandularcapitate(1-
many)
Anisocytic; Stellate;glandular
Solanumbonariense ca.25x15-20 1/2
anomocytic capitate(1-many)
S.chacoensessp. Anisocytic; Conical(1-many);
ca.25-30x20 2-3/3 +
chacoense anomocytic glandularcapitate(1-2)
Conical(1-many);
S.commersoniissp. Anisocytic;
ca.30x20-30 2/2-3 glandularcapitate(1- +
commersonii(29) anomocytic
many)
Y-shaped,rarely;
Anisocytic;
S.diflorum(24) 15-25;15-25 1-2/2-3 dendritic;conical(2), +/+(slightly)
anomocytic
rarely /+
Anisocytic; Stellatestalked;
S.elaeagnifolium(29,30) 20-25x15-20 1/2
anomocytic glandularcapitate(1)
Anisocytic; Mesophyllwith
S.glaucophyllum(29) 30-35:20-25 1/1
anomocytic crystalsand
S.pygmaeumvar. Anisocytic; Conical(2-many);
25-30x20-25 1-2/2 +
pygmaeum anomocytic glandularcapitate(1)
Conical(2-many);Y-
Anisocytic; shaped(onebranch
S.sublobatum(29) 20-30x15-20 1-2/3 +/+
anomocytic shorter);glandular
capitate(3)
Turneraceae
Turnerasidoidesssp. Anisocytic; Bristles(1);glandular
30-35x25 1/1
pinnatifida anomocytic shaggy
Urticaceae
Parietariaofficinalis Anomocytic 20-25x10-20 2/3 Conical(1) +
Urticaurens(15) Anomocytic ca.25x20 2/3 Bristles(1);stinging Cystoliths
1) Metcalfe (1960); (2) Prat & Vignal (1968); (3) Caro & SÆnchez (1969); (4) SÆnchez (1971); (5) Gould & Shaw
(1992); (6) SÆnchez (1974); (7) Westerkamp & Demmelmeyer (1997); (8) Bay(cid:243)n & Arambarri (1999); (9) Selvi
& Bigazzi (2001); (10) Di Fulvio (1976); (11) Amat (1988); (12) Gattuso & Gatusso (1998); (13) Ariza Espinar
(1973); (14) Pertusi (1987); (15) Barboza et al. (2001); (16) Sim(cid:243)n et al. (2002); (17) Cabrera (1944); (18)
Arroyo (1986); (19) Yagueddœ & Cid (1992); (20) Arambarri & Colares (1993); (21) Gatusso & Gatusso (1989);
(22) Gatusso (1996); (23) Ragonese & Covas (1947); (24) Gatusso (2000); (25) MedÆn (1986); (26) Metcalfe &
Chalk (1950); (27) Colares et al. (1999); (28) Cabrera (1979); (29) Stenglein (2001); (30) Bruno et al. (1999). H
= hooks; P = prickles; Pa = papillae
248
S. E. Freire et al., Epidermal characteristics of toxic plants for cattle
Fig. 2. Epidermal characters of Poaceae. a, stoma with parallel-sided subsidiary cells: Briza minor; b, stoma with
triangular subsidiary cells: Cynodon dactylon; c, stoma with dome subsidiary cells: Leptochloa chloridiformis; d,
micro-hair with almost hemispherical distal cell: Eragrostis cilianensis; e, dumb-bell silica-body: Echinochloa
crusgalli; f, saddle-shaped silica-body: Eragrostis cilianensis; g, rectangular silica-body: Lolium multiflorum; h,
cross-like silica-body: Sorghum halepense; i, micro-hairs with pear-like distal cell: Eleusine indica; j, micro-hair
with rod-like distal cell: Digitaria sanguinalis. Scale bar = 50 m m.
previouslymentionedbyMetcalfe&Chalk(1979). nalandbasalcells(similartobristlehair,according
Types of stomata varied notably in such a small totheillustration);whiphairs(i.e.aseptateflagellate
group of genera and species. This variability was according to Ramayya, 1962); two-armed (i.e. T-
previouslynoticedbyForcone&Ayestaran(1996). shaped); stellate; candelabra hairs; shaggy hairs,
Eryngium epidermal cells showed parallel and peltate scales. Uniseriate like bristles, peltate
distributionandalsoparacytic stomata, which was scales, and candelabra hairs, were mentioned by
notedbeforebyYagueddœ&Cid(1992). theseauthorsforgeneranotincludedinthepresent
study, whereas stellate or branched hairs cited by
Asclepiadaceae(5spp.analyzed):Resultsagree them for the genus Baccharis were found in other
withMetcalfe&Chalk(1950,1979)andalsowith species notincludedinthispaper(Freire,inprep.).
Bay(cid:243)n & Arambarri (1999) who mentioned that All other hair types, i.e. whip hairs, two-armed or
paracytic was the most frequent type of stomata. T-shaped, and shaggy hairs, agree with those
Bay(cid:243)n & Arambarri (l.c.), also illustrated the described in this study. In addition, we found
epidermalandtrichomeornamentation,speciallyfor conical hairs in nine species, in two of which
Oxypetalum solanoides which present hairs, (Ambrosia tenuifolia, Aster squamatus), they were
exclusivelywaxesornamentatedontheapicalcell. previously reported by Yagueddœ & Cid (1992),
whereas Barboza et al. (2001) illustrated conical
Asteraceae(41spp.analyzed):Metcalfe&Chalk hairsinonespeciesofXanthium(X.spinosum).In
(1950,1979)characterizedAsteraceaebyhavingthe agreementwithMetcalfe&Chalk(1950,1979),glan-
following seven hairs: uniseriate consisting of dularhairsarewidelydistributed(onlyfewspecies
uniformcellsapartfrommodificationofthetermi- of the total species analyzed have not glandular
249
Bol. Soc. Argent. Bot. 40 (3-4) 2005
Fig. 3. Hairs of Asclepiadaceae, Boraginaceae, Brassicaceae, Cactaceae, Caprifoliaceae, Caryophyllaceae, Chenopodiaceae,
Convolvulaceae, Geraniaceae, and Lamiaceae. a, conical: Morrenia odorata; b, conical: Oxypetalum solanoides; c, falcate, c,
1
glandular capitate: Anchusa officinalis; d, conical: Echium plantagineum; e, conical, e, glandular capitate: Heliotropium
1
amplexicaule; f, moniliform: Parodia ottonis; g, short bristle: Sambucus australis; h, bristle: Agrostemma githago; i,
moniliform: Saponaria officinalis; j, vesicular: Chenopodium album; k, whip: Ch. murale; l, conical: Bassia scoparia; m,
conical: Salsola kali; n, glandular capitate, n, conical: Convolvulus arvensis; o, conical: Brassica nigra; p, 3-armed: Capsella
1
bursa-pastoris; q, falcate: Cardaria draba; r, barrel-shaped: Raphanus sativus; s, falcate: Rapistrum rugosum; t, long-conical:
Sisymbrium altissimum; u, glandular capitate bottle-shaped: u, bristle: Erodium malacoides; x, conical: Geranium molle; y,
1
conical, y, glandular capitate: Lamium amplexicaule; z, porrect stellate, z, glandular capitate: Marrubium vulgare. Scale
1 1
bars:a,b,c,g,j-o,q,z 50(cid:181)m;c,d=300(cid:181)m;e,e h,i,p,r,s,x,y=100(cid:181)m;f,t=200(cid:181)m;y,z =25(cid:181)m.
1 = 1, 1 1
250