A Glossary of Plant Hair Terminology

Author(s): Willard W. Payne

Source: Brittonia , Apr. - Jun., 1978, Vol. 30, No. 2 (Apr. - Jun., 1978), pp. 239-255
Published by: Springer on behalf of the New York Botanical Garden Press

Stable URL: https://www.jstor.org/stable/2806659

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Brittonia, 30(2), 1978, pp. 239-255.
( 1978, by the New York Botanical Garden, Bronx, NY 10458

A GLOSSARY OF PLANT HAIR TERMINOLOGY

WILLARD W. PAYNE

Payne, Willard W. (The Cary Arboretum of the New York Botanical Garden,
Millbrook, NY 12545 and Fairchild Tropical Garden, Miami, FL 33156). A
glossary of plant hair terminology. Brittonia 30: 239-255. 1978.-This is a
compilation of English terms having to do with hairs and hairy surfaces of
plants. The glossary is divided into two sections. Part I includes terms for types
and attributes of individual hair kinds and hair clusters. Part II deals with kinds
and characteristics of induments.

Plant hairs, or trichomes, are of great interest to descriptive and experimental

botanists, and data about them and about induments are routinely included in studies
of many kinds. As simple tools of morphology, trichomes are useful because of the
ease with which they may be examined, and because of their almost universal oc-
currence, particularly in the ferns and flowering plants. Beyond purely descriptive
use, comparative data may be important for studies of evolution and relationships,
and for the roles of hairs in various aspects of physiological and ecological adaptation.
Until recently, our interest in plant hairs has been quite superficial, but scientists
increasingly realize that a great deal is still to be learned about the importance of
individual trichomes and of induments in the life cycle, about their influence on
development of organs upon which they mature (often at very early stages), and
about their roles as waste repositories and as primary recipients of environmental
stimuli and forces. These considerations, plus the historical use of hair data, make
the terminology of hairs and induments very important. Until the present work,
however, there has been no readily available dictionary of terms in the English
language. It is to this need that the glossary presented here is addressed.
The terms provided are divided among two groups: Part I of the glossary deals
with those descriptors that only or usually differentiate particular kinds or attributes of
individual trichomes. Part II is devoted to terms having to do with the indument
as a whole, or to masses of hairs with special appearance or function. Some special
terms related to particular families or genera have been omitted intentionally from
the glossary, especially when they are derived from the names of taxa (e.g., "arundo-
typve"). Such are especially abundant in the Poaceae and Bromeliaceae. Further, no
attempt has been made to search out or include terms from the voluminous literatures
of languages other than English. The list of references from which terms have been
garnered is presented at the end of the glossary.

ACKNOWLEDGMENTS

I am grateful to several students who contributed data for this compilation, in-
cluding Joseph D. Galsley, Marvin L. Bowles, Jere N. Brunken, Liwayway M. Engle,
E. Guhardja, Boonkun Kurmarohita, Alan D. Parker, Kathleen M. Peterson, Robert
R. Robbins, Kathleen A. Schemske, Anastasia L. Stemler, and Thomas A. Striker.
Dr. Terry W. Lucansky, University of Florida, Gainesville, read the manuscript and
made many helpful suggestions. The work was facilitated in part by National Science
Foundation grant GB-30477.

BRITTONIA 30: 239-255. April-June, 1978.

239

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 240 BRITTONIA [VOL. 30

PART I. TERMS FOR TRICHOMES

A bapical. Situated at the lower pole or related to it.

Abietiform hair. A kind of branched, monopodial hair resembling a fir tree in overall
configuration; candelabra hair (Fig. 1).
Abjoint. To delimit by a septum or joint; to separate at a joint.
Acantha. A spiny process; a spine, prickle, or thorn.
Acarodomatia hairs. Hairs found in cavities; hairs found on leaves in the axils of
the veins of the first and sometimes second or higher orders of ramification.
Accessory cell. A subsidiary cell; a cell with special relationship to the trichome, but
not part of it.
Acephalous. Without a head.
Acerate. Having the shape of a needle; acicular (Fig. 2).
Acicula. The bristle-like continuation of the rachilla of certain grasses; needle-
shaped spines or prickles.
Acicular. Needle-shaped; acerate (Fig. 2).
Acinaciform (acinasiform). Scimitar-shaped (Fig. 3).
Acroscopic. Oriented toward the sky or shoot tip (contrast basiscopic).
Aculeate. Prickly; thorny; with rudimentary, strongly silicified, tufted hairs or
papillae.
Aculeus. A sharp epidermal emergence, such as a prickle; a rigid, hair-like projection
growing from the epidermis.
Acuminate. Having a long, slender, sharp point; tapering gradually toward the apex;
drawn out to a long point; tapering at the end (Fig. 4).
Adpressed. Appressed.
Adspersed. Scattered or widely distributed.
Aduncate. Crooked, twisted, bent, hooked (Fig. 7).
Aigret. A feathery crown or tuft of hairs attached to the diseminule (as the pappus of
a thistle fruit, or coma of a milkweed seed).
Anchor hairs. Serving to assist in support of climbing or reclining shoots (e.g., in
Humulus), often with short, terminal barbs (e.g., in Avicennia) (Figs. 5, 6).
Ancistrous. Barbed (Figs. 5, 6).
Anfractuose. Wavy, sinuous, twisted; full of turnings and windings; serpentine
(Fig. 7).
Angler hair. Fish-hook-shaped (Fig. 8).
Anticlinal. With walls placed at right angles to the terminal surface; often used to
. describe the segmentation of heads of glandular trichomes (contrast periclinal).
Antler hairs. Antler-like; antler-shaped; pedate (Fig. 9).
Antrorse. Directed forward or bent toward the tip (contrast retrorse).
Anvil hairs. Shaped as an anvil, short-stalked with short, opposite, terminal, lateral
projections; a form of malpighiaceous hair; dolabriform (Fig. 10).
Apicicircinatus. Terminating circinately; coiled at the apex (Fig. 11).
Apiculate. With a short, sharp, flexible point.
Apiculus. Tip.
Apodial. Without a foot-stalk or podium; sessile (Figs. 9, 23, 34, 37).
Appendage. An attached, secondary part.
Appressed. Lying close against the surface; adpressed.
Arrect. Stiffly erect, directed upward from an inclined base (Fig. 12).
Arthrodactylous. Joined or jointed as a finger (Fig. 13).
Articulate. Jointed, as a multicellular, uniseriate trichome.
Articulation. A joint; a node; a septum; a segment which is marked or separated
by a joint (Fig. 3 8).

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1978] PAYNE: PLANT HAIR TERMINOLOGY 241

1 ~~~~~~~~3
9

6 ~~~~~~~~~~~~~15

10 121

FIGs. 1-15. Trichome types. 1. Abietiform; candelabra; orthocladous. 2. Acerate; acicular.

3. Acinaciform. 4. Acuminate terminal cell; belemonoid. 5. Anchor hairs, uncinate and with
terminal fluke cells; barbed. 6. Ancistrous; anchor; barbed; glochids. 7. Aduncate; anfractuose;
curly; ribbon-hair; serpentine. 8. Angler; hamate; hooked; uncinate. 9. Antler; pedate (some-
times improperly termed stellate). 10. Anvil; malpighiaceous. 11. Apicircinatus; circinate. 12.
Arrect. 13. Arthrodactylous. 14. Asciiform; hatchet-shaped. 15. Attenuate.

Asciiform. Hatchet-shaped (Fig. 14).

Ascus-shaped. Pointed at the end, attaining its greatest breadth above the middle and
from there gradually narrowed toward the base; club-shaped.
Aseptate. Having no cross walls, partitions, or septa; unicellular; eseptate.
Asperous. Scabrous, asperate, rough, harsh.
Aspersed. Scattered; sprinkled over; adspersed.
Attenuate. With a long, gradual taper (Fig. 15).
Axial. Pertaining to the axis.
Axis. The central part of the hair to which branches are attached.

Barb. A sharp, reflexed point on a hair, awn, or other process.

Barbed. With short, rigid, reflexed processes (Figs. 5, 6).
Barbellate. With small, laterally placed as well as terminal barbs (Fig. 6).
Basal cell. Lowermost cell of a trichome.
Basilatus. Arising from a broad base.

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 242 BRITTONIA [VOL. 30

Basipetal hairs. With a tardily maturing basal portion; while the lower cells are
still growing, the terminal one has reached its final stage of development (as the
pappus hairs of certain Compositae).
Basiscopic. Oriented toward the ground or shoot base (contrast acroscopic).
Belemnoid. Dart-shaped (Fig. 4).
Bicellular. Of two cells.
Bifid. Two-parted; cleft; forked; divided into two lobes; split into two segments
(Figs. 16, 18, 21).
Bifurcate. Divided or forked into two branches; having two prongs; Y-shaped (Figs.
16, 21).
Biseriate hairs. Hairs which consist of two rows of cells (Figs. 6, 17, 18, 41).
Bladder hairs. Swollen, bladder-like cells of the epidermis.
Bootjack hairs. Forked, with one arm apically forked again (Fig. 18).
Boraginaceous hairs. Unicellular to multicellular, conical, calcified and/or silicified
bristles.
B3osselated. Covered with knobs; beset or covered with small protuberances or bosses
(Fig. 19).
Brachiate. With spreading or widely diverging branches; with pairs of opposite
branches spreading at right angles.
Bracket hairs. Hairs which are hooked or curved or bent at the apex.
Brevicollate. Short-necked (Fig. 20).
Brevifurcate. Repeatedly forked with a short stalk segment between bifurcations
(Fig. 21).
Breviramose. Short-branched.
Bristle. A stiff or rigid, hair-like process.
Brush hairs. Branched with ray cells radiating in all directions; the hairs in Com-
positae that are found on the upper part of the style.
Bud-shaped hair. A form of peltate hair with all the cells of the shield elongated in
the direction of the stalk so that the marginal cells do not appear as ray cells in
the surface view.
Bulbous hairs. With a rounded or bulb-like basal portion (as the stinging hairs of
Urtica) or cushion.
Bullate. With a blistered or puckered surface; bubble-like; having one or more
hemispherical outgrowths (Fig. 19).
Bulliform cells. Cells in leaves of grasses that have a swollen, bladder-like appearance.
Bullose. Swollen; blistered.
Bursiculate (bursiform). Shaped like a bag or purse; pouch-like.

Calcified. Incrusted with carbonate of lime.

Callus. A hard prominence or protuberance; the plug of substance that may fill the
tip of a broken trichome.
Callus hairs. The hairs at the base of the floret of some grass genera (e.g., Cala-
magrostis).
Calvitium. An area lacking hairs; a bald spot.
Calyptriform hairs. Glandular hairs in which the young terminal cell splits into
quadrants, then each of these cells divides by means of a periclinal wall to form
two tiers of cells. The four cells of the upper tier are called the lip, those of the
lower tier the center.
Candelabra hairs. Monopodially branched with the branches whorled; abietiform
(Fig. 1 ).
Capitate. With an enlarged head (Figs. 14, 17, 20, 28).
Chaeta. A spine, bristle, or seta.

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 1978] PAYNE: PLANT HAIR TERMINOLOGY 243

Chaetotaxy. The pattern of bristle arrangement.

Chalk glands. Multicellular glands which deposit calcareous matter, the secretion
escaping through a special channel, the water pore, as found in some Saxifragaceae.
Chartaceous. Having a stiff, papery texture; parchment-like.
Cilia. Conspicuous hairs inserted along the margins of leaves, bracts, petioles, or
other organs.
Ciliola. Minute or secondary cilia.
Circinate (circinal). Coiled as a watch spring (Fig. 11).
Cladose. Branched (Figs. 1, 9, 16, 21, 40, 41, 46).
Clavate. Club-shaped; elongate, largest near the tip and rounded (Fig. 22).
Clavellate. In the form of a little club.
Climbing hairs. Hairs with tubercles or barbs that assist in support of climbing shoots
(Figs. 5, 6).
Clothing hairs. The hairs giving the indument its characteristic appearance and
texture; usually contrasted with the glandular hairs of an indument.
Collecting hairs. Of stigmatic surfaces, papillose or multicellular trichomes which
collect pollen. Collecting hairs have various morphologies and are often secretory.
The collecting hairs of Campanulaceae may be contractile, drawing adherent pollen
grains into cavities formed by their invagination.
Colleters. Glandular hairs having a multicellular head on a multicellular stalk (Figs.
17, 28).
Columella. A cell at the apex of the stalk of a glandular hair that extends upward
into the body of the gland.
Coma. A tuft of fine, spreading hairs, as with the seeds of Asclepias.
Compound. Of several parts; resembling a compact tuft of individual hairs.
Compressed. Flattened laterally.
Contorted. Twisted; irregularly sinuate; spiral (Figs. 7, 47).
Corolline hairs. Hairs found on various parts of flowers.
Crenate. Of peltate hairs, with scalloped margins due to projection of the ray cells.
Cruciate (cruciform). Having the form of a cross with equal lateral arms (Fig. 23).
Cup-shaped hair. A form of peltate hair in which cells of the central area of the shield
form the bottom and those of the margin the walls of a cup (Fig. 24).
Curly. With several bends or curvatures; tortuous; sinuous (Figs. 7, 47).
Cushion hairs. Geniculate hairs with cushion-like cell masses at the convex side. In
Clematis vitalba, such hairs function as tactile hairs (Fig. 25).
Cystolith hairs. With concretions occurring usually in the basal cells; in hairs with
no distinct stalk, cystoliths may be situated on a lateral wall of the usually bulbous
base, or may be attached to a terminal portion (Fig. 26).

Declinate. Oriented downward; directed downward from the base.

Decurved. Bent down; deflexed; curved downward.
Deflexed. Decurved; bent sharply downward or outward.
Dendroid (dendriform; dendritic) hairs. Branched to resemble a tree in form; having
a stemlike part arising immediately from the broadened base and dividing to
produce diverging branches which may be repeatedly forked in their turn.
Depilation. The natural loss of the hairy covering by plants as they mature.
Depressed. Sunk in the surface.
Depressed glands. Glands which are flattened vertically.
Dichotomous. Forked as a "Y" (Figs. 16, 18, 21).
Dickotomously ramified. Branching by means of development of similar projections
from the apex of the stalk (contrast monopodially ramified) (Fig. 21).
Dilepidous. Consisting of two scales.

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 244 BRITTONIA [VOL. 3 0

16

0
1 19 21

23
jX x ;2~~~~8 o3 225 02
22 24

21
26
29

31

28 30 32

FIGS. 16-32. Trichome types (continued). 16. Bifid; dichotomous; forked; furcate. 17. Biseriate;
colleter; gland. 18. Bootjack. 19. Bosselated; nodose; pustulate. 20. Brevicollate. 21. Brevi-
fiircate. 22. Clavate. 23. Cruciate; stellate. 24. Cup-shaped. 25. Cushion hair. 26. Cystolith hair
(with cystolith in basal cell). 27. Dolabrate. 28. Doliform head of colleter. 29. Falcate. 30.
Flagelliform; whip-like. 31. Fusiform. 32. Geniculate.

Divaricate. Widely spreading or diverging.

Divergent. Spreading broadly, but less so than divaricate.
Dolabrate (dolabriform). Shaped like the head of a pick, with two divaricate or
opposed, terminal branches; axe-shaped; malpighiaceous (Fig. 27).
Doliform. Barrel-shaped (Fig. 28).
Dove-tailing papillae. Papilloid projections from the surfaces of appressed organs,
serving to hold them together, as for some bud scales, cone scales, petals, and
connate anthers; dove-tailing papillae may be forked (e.g., as for the cone scales
of Picea excelsa).

Elaphinous. Pronged like a deer's antlers; fawn-colored.

Emarginate. Having notched margins or tips.

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 1978] PAYNE: PLANT HAIR TERMINOLOGY 245

Ensiform (ensate). Shaped like a sword with a straight blade.

Equisetoid hairs. Hairs having the appearance of miniature Equisetum plants, found
in several fern genera; the appearance is due to regular, deep corrugations in the
transverse septa of stiff, multicellular hairs.
Erectopatent. Intermediate between spreading and erect.
Eseptate. Without septa or cross walls; aseptate.
Excoemum. A fringe or tuft of hair at the base of the glumes in some grasses.
Explosive hairs. For many Cucurbitaceae, hairs on the floral organs that contract
forcefully when the easily detached tips are touched, expelling the contents on the
touching object. In some cases (e.g., Cyclanthera pedata) the protoplasmic sub-
stance quickly hardens, and might immobilize small insects; in other cases the
trichome contents may moisten the legs and bodies of visiting insects and aid
pollen adhesion.
Extrafloral nectiary. A nectary, often of secretory hairs, appearing on or in some
part of a plant other than the flower.
Extranuptial. Said of nectaries or honey glands which are not part of the floral organ

Falcate (falciform). In the shape of a falconi's talon; curved to one side as a sickle;
sickle-shaped (Fig. 29).
Fasciculate. Occurring in bunches or clusters; of compound hairs with contiguous
basal or stalk cells.
Faucial. Situated in the throat or mouth of the tube of the perianth.
Felt kairs. Dense trichomes produced early in the development of a leaf, floral part,
etc., as a protective covering.
File hair. A trichome covered with small knobs formed by the cuticle or by small
crystals.
Filiform. Having the shape of a thread or filament; thread-like.
Fimbrilla. A single trichome from a fringe of hairs.
Flagelliform hairs. Typically multicellular, uniseriate trichomes with the distal end
of the terminal cells delicate and much elongated; whip-like (Fig. 30).
Flocci. Small tufts resembling cotton; any tufts of woolly hairs.
Fluke cell. Any of the lateral, projecting cells of an anchor hair (Fig. 5).
Foveolate. Pitted.
Fugaceous. Falling away early, as the indumentum of some tree leaves (e.g., Aesculus,
Quercus).
Furcate. Forked (bifurcate, trifurcate, etc.).
Furcellate. Minutely or slightly furcate or forked.
Furfur. Bran or scurfy particles.
Fusiform. Spindle-shaped; broadest at the middle and tapering toward both ends
(Fig. 31).
Geniculate. Bent abruptly, as at the knee (Fig. 32).
Gland. A secreting cell or group of cells, sessile or stalked; most glandular trichomes
have only the apical cell or apical cluster of cells (head) secretory (Figs. 14, 17,
20, 21, 34, 45).
Gland, chalk. A gland which exudes salt solutions that give whitish deposits on
drying.
Gland, internal. One or more secreting cells inside the plant, such as those containing
essential oils which form the translucent dots in leaves of the orange.
Gland, salt. A gland which excretes solutions of hygroscopic salts which are dry in
daytime and deliquesce at night.

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 246 BRITTONIA [VOL. 30

Glandular. With a secretory or excretory function; with certain cells or regions

obviously modified for secretion.
Glochid. A barbed hair, especially when barbed at the tip; for cacti, serving a
primarily protective function; for fruits and seeds serving functions of dispersal
and anchorage in the soil (Fig. 6).
Glochidium. A hair-like appendage with a hooked tip formed on the spore masses of
the water fern, Azolla.

Hair. A typically elongate, slender outgrowth from the epidermis; any hair-like
structure; a trichome.
Hair cystoliths. Structures resembling cystoliths which occur in trichomes (cystolith
hairs), often in the basal cell or cells (Fig. 26).
Hamate (hamose). Hooked or curved at the end; sickle-shaped (Figs. 5, 8).
Hamulus. A small hook or hooked process.
Head. An enlarged, terminal portion; the glandular portion of a glandular trichome.
Heliciform. Coiled like a snail shell (Fig. 33).
Helminthoid. Worm-shaped, vermiform.
Hippocrepiform. Horseshoe-shaped.
Hooked. With an incurved apex (Figs. 5, 6).
Hydathode. A pore which excretes liquid water; water gland; water-excreting hairs
may serve as hydathodes.

Inflated. Swollen and bladder-like.

Integumental glands. Peltate glands that have an abundant secretion which causes
the integument to be raised like a bladder.
Intercellular hairs. Hairs which occur in the intercellular spaces of the rhizome, the
base of the petiole, and the leaf parenchyma of some ferns (e.g., Aspidium spp.).
Intramural glands. Having secretion deposited not only between the cuticle and the
underlying walls of the cells of the head, but also between the longitudinal walls
of these cells.

Jointed. With real or apparent articulation; nodose (Figs. 13, 35, 38).

Labriform. With a terminal projection to one side; asciiform.

Laccate. Appearing as if varnished.
Lacinif orm. Fringe-like.
Lactiferous. Producing or containing latex; laticiferous.
Lageniform. Gourd-shaped; flask-shaped (Fig. 34).
Latex hairs. The hairs continuous with latex tubes; these often break easily and
liberate drops of latex.
Laticiferous. Lactiferous.
Lepides. Scales.
Lepra. A white mealy matter extruded from the surface of some plants.
Lever hairs. Pappus bristles of Aster and certain other Compositae that become
reflexed at maturity and press against the receptacle and other surrounding objects
to tear the attached fruits loose.
Ligulate. Strap-shaped.
Limaciform. Shaped as a garden slug (Fig. 36).
Lingulate (linguiform). Tongue-shaped; shorter, broader and thicker than ligulate.
Lumbrical (lumbricous). Having the shape of an earthworm.
Lunate. Crescent-shaped, as the new moon; selenoid (Fig. 35).
Lymphatic hairs. Non-glandular hairs with living contents, with the exception of
those that are inserted on the various parts of flowers or on their margins.

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 1978] PAYNE: PLANT HAIR TERMINOLOGY 247

Lyrate hairs. Lyre-shaped; bifurcate with the branches curved upward and toward
one another.

Malpighiaceous hairs (malpighian hairs). Unicellular, two-armed, dolabriform tri-

chomes, the arms opposite one another and oriented parallel to the epidermis; anvil
hairs (Figs. 10, 27).
Mammila. A nipple; a nipple-like projection.
Mammilliform. Applied to those papillate protuberances on a petal that give it a
velvety appearance; nipple-shaped.
Mammose. Breast-shaped; mastoid.
Mammulose. Minutely mammose.
Many-fluked. With many lateral barbs, as some types of anchor hairs (Fig. 5).
Marcescent. Withering but persistent.
Mastoid. Resembling a nipple or breast; mammose.
Meniscoid (meniscate). Bent into a half circle; shaped as a watch crystal; crescent-
shaped.
Microlepis. With small scales.
Middle cell. A cell between the distal and basal cells.
Miliaris (miliarius). A minute, glandular spot on the epidermis.
Miliary glands. Stomata.
Moniliform (moniliferous). With regular constrictions, thus resembling a string of
beads or necklace.
Monopodially ramified. Branched with the branches attached to a central stalk
(contrast dichotomously ramified) (Fig. 1).
Mucilage hairs (mucilaginous hairs). Trichomes which secrete slimy or mucilaginous
substances, often serving to glue together the structures upon which they are borne;
mucilage glands.
Multifurcate. With many forks or divisions (Fig. 21).
Multigyrate. With many turns or gyres.
Multiseriate. With several rows or columns of cells (biseriate, triseriate, tetraseriate,
etc.); pluriseriate.
Muricate. Said of a surface roughened with short, stiff points.
Muriculate. Finely muricate.
Muscariform. Having long hairs toward the tip, like the ancient fly whisk (e.g., the
styles of some Compositae).
Muticous. Blunt; lacking a pointed tip.

Neck cell. The intermediate cell of the stalk of a uniseriate, glandular hair; a middle
cell.
Nitid (nitidous). Glossy; lustrous; smooth and clear.
Nodose. Knobby; with irregular, knot-like swellings (Figs. 19, 47).
Nodulose. Minutely nodose.
Non-septate. Without cross partitions or walls.

Obliterated. Erased; deciduous, leaving no trace.

Ornithorhynchous. Shaped as a bird's bill (Fig. 37).
Orthocladous. With straight branches (Figs. 1, 16, 40).
Osteolate. Shaped as a thigh bone, as for many cylindrical, subterminal cells with
somewhat swollen ends (Fig. 38).
Ovoid. Shaped as an egg in form and attached at the large end.

Pachycladous. With thick branches.

Palea. In ferns, a scale which bears numerous glandular hairs.

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 248 BRITTONIA [VOL. 3 0

Papilla. An ordinarily blunt projection from the surface of a single epidermal cell
and usually only a portion of that cell.
Pappo. The down of thistles.
Parachute hairs. Trichomes attached to seeds or fruits to form comose or umbrella-
like aggregations to facilitate wind dispersal; coma hairs.
Pearl glands. Usually spherical, short-stalked structures, the cells of which are rich
in glistening contents which are highly nutritious; pearl glands serve as food
sources for ants.
Pedate. Branched, with successive branches lateral and arising near the base of
earlier branches to give a fan-like arrangement; branched to resemble the antlers
of a deer, as in many Cruciferae; antler hairs (Fig. 9).
Pedestal. The unicellular or multicellular raised base to which hairs or prickles may
be attached.
Peltate hairs. Scales or lepides; sessile or stalked trichomes with flattened heads
which consist either of a single cell developed as a flat structure, or of a varying
number of cells arranged in one or more layers (Fig. 39).
Penicillate. With a terminal cluster of close-set branches; resembling conidial branches
of Penicillium in form; brush-like tuft (Fig. 40).
Periclinal. Of walls placed parallel to the terminal surface; often used to describe
the segmentation of heads of glandular trichomes (contrast anticlinal).
Pit. A small surface depression or opening.
Pitted. Marked with pits.
Plumose. Featherlike, as the pappus bristles of many Compositae (Fig. 41).
Pluricellular. Multicellular.
Pluriseriate. Multiseriate.
Prickle. A sharp, rigid, epidermal outgrowth, often relatively massive, as for Rosa;
frequently and erroneously termed spine.
Pulvinate. With an abruptly swollen basal portion (Fig. 34).
Punctation. A minute spot or depression; an interior gland or external gland head
seen with the unaided eye.
Pungent. Tipped with a sharp, rigid point.
Pustulate. With a surface covered with short, blunt projections; minutely blistered
(Fig. 19).
Pyrenodeous. Wart-like.

Quadricellular. Of four cells.

Radiate. Spreading from a common center, as the arms of many stellate hairs, or
the shield cells of scales.
Recurved. Curved toward the base or backwards.
Resin warts. Glands which secrete resin.
Reticulate. With a net-like arrangement of ridges or depressions.
Retrorse. Directed or bent toward the base (contrast antrorse).
Ribbon hairs. Shaped as a flowing ribbon (Fig. 7).
Root hairs. Hairs of the root epidermis that absorb water and nutrients.
Rosette gland. A circular cluster of glands.
Rugose. With a creased or wrinkled surface.

Saccate. Bag-shaped; bag-like.

Scale. Any flattened trichome, but especially those that are relatively large.
Scarious. Dry and papery, as the scales and hairs on the petioles of many ferns.
Sciuroid. Curved and bushy as a squirrel's tail.

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1978] PAYNE: PLANT HAIR TERMINOLOGY 249

FIGS. 33-48. Trichmetyes(cnt

31

39

Q~~~~~~~~~) ~~~~43
40 K

45

44 6
48 4647
FIGS. 33-48. Trichome types (continued). 33. Heliciform; snail-shaped. 34. Lageniform;
pulvinate. 35. Lunate; selenoid. 36. Limaciform. 37. Ornithorhynchous. 38. Osteolate cells of
uniseriate hair. 39. Peltate. 40. Penicillate. 41. Plumose. 42. Spiral. 43. Stellate. 44. Subulate.
45. Surculate. 46. Sympodial. 47. Torulose. 48. Trochlear; trochleariform.

Sclerenchymatous trichome. Any epidermal hair of uniformly thick-walled, dead

cells; a supportive or protective hair, composed of thickened cells from which
the protoplasm has been removed.
Scutate. Shield-shaped; having horny plates.
Selenoid. Crescent-shaped; lunate (Fig. 35).
Semi-malpighian hairs. One-armed hairs; half-malpigheaceous hairs; asciiform hairs;
labriform.
Sensitive hairs. Trigger hairs; stimulators.
Septate. Divided by cross partitions. True septa of hairs are usually taken to be cell
walls; false septa are wall outgrowths which close off portions of a hair.
Serpentine. Curving and re-curving in form; sinuate; anfractuose (Fig. 7, 47).
Sessile. Not stalked, as for glands or scales (Figs. 10, 12, 34, 37).
Seta. A stiff, coarse bristle.
Setiform. Shaped as a bristle.

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 250 BRITTONIA [VOL. 3 0

Shag hairs (shaggy hairs). Multiseriate, coarse trichomes as the inflorescence hairs
of Portulaca, or the petiolar hairs of many Begonia species.
Sheatlh. A cluster of subsidiary cells that covers and supports the hair stalk or its
basal portion.
Shield. The expanded, flattened portion of a peltate hair.
Sigmoid. "S"-shaped.
Siliceous (silicified). Impregnated with or containing particles of silica; encrusted
with silica.
Simple. Unbranched; uniseriate, or unicellular.
Sinuate. See anfractuose and serpentine.
Smooth. Without surface irregularities.
Snail-shaped. Curved or coiled as a snail shell; heliciform (Fig. 33).
Spindle hairs. Hairs which resemble malpigheaceous hairs, attached centrally and
with the two arms hooked.
Spinous. Having a spine-like appearance; stiffened and sharply pointed.
Spiral. Shaped as a corkscrew; contorted (Fig. 42).
Squamiform. Flattened or scale-like and of irregular outline.
Stalk. The supporting part of a hair or gland.
Stellate. Branched as a star; star-shaped (Figs. 23, 43).
Stellular. Compound or fasciculate, with the branches spreading as rays.
Stimulators. Tactile hairs or bristles which transmit stimuli to the sensitive motor
tissue; trigger hairs.
Stinging hairs. The urticating or irritant hairs of nettles and other stinging plants;
stinging bristles.
Strict. Straight and upright; unbranched or little-branched.
Subcespitase. Inclined to grow in bunches; somewhat tufted.
Subramose. With a tendency to branch, but with few or small branches.
Subsidiary cells. Neighboring cells; differentiated cells that surround the base of
a hair.
Subulate. Awl-shaped, tapering from a broad or thick base to a sharp point (Fig. 44).
Sunken. Situated below the level of the epidermal cells.
Superficial cells. Cells of the surface.
Surculate. Having the form of a sucker (Fig. 45).
Swollen. Enlarged; with enlarged areas. Swellings may be apical, basal, or inter-
calary, and there may be several per trichome.
Sympodial. Branched hairs in which the basal portion of each branch participates in
formation of the stalk (Fig. 46).
Syncladous. Growing in tufts from the same point.

Tactile hairs. Stimulators; trigger hairs.

Tanniferous (tanniniferous). Containing tannin or tannin-like substances.
Tentacle. One of the sensitive, thigmotropic, glandular hairs of the leaf of Drosera,
the sundew.
Tentaculiform. Tentacle-shaped.
Tortuous. With several bends or curvatures; sinuous; curly (Figs. 7, 47).
Torulose. Twisted or knobby; irregularly swollen at close intervals.
Trap hairs. Trichomes that fill the constrictions of tubular perianths (e.g., Aristo-
lochia) or inflorescences (e.g., Ficus) to trap pollinators within specially developed
chambers; in certain insectivorous plants (e.g., Utricularia, Dionaea, Aldrovanda)
the marginal trichomes or setae that reinforce the trapping function.
Trichiform. Bristle-shaped.
Trichoblast. Any cell of the protodermis from which a trichome will arise.

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1978] PAYNE: PLANT HAIR TERMINOLOGY 251

Trichocysts. The cells in seed coats of Lythraceae that project hair-like appendages
when wetted.
Trichome. Any epidermal outgrowth which prevents classification of a plant surface
as glabrous; hair.
Trichome hydathode. A hair-like appendage, especially in young leaves, which releases
and absorbs water.
Trichosanthous. With hairy flowers.
Trichospermous. With hairy seeds.
Trigger hairs. The sensitive hairs of the Venus fly trap, Dionaea muscipula, that
stimulate closure of the trap; stimulators.
Trochal. Wheel-shaped.
Trochlear. Shaped as a pulley; also trochleariform (Fig. 48).
Tubercle. A wart-like or knob-like excrescence; a nodule.
Tufted. Closely clustered; stellate but branched from the base; cespitose.
Turbinal. Spirally coiled or rolled.
Turbinate. Conical.
Twinned hair. A form of reduced stellate hair with only two, single-celled rays
oriented opposite one another and separated by an anticlinal wall.

Unciform. Shaped like a barb or hook.

Uncinate (uncate). Hooked at the tip (Fig. 8).
Uniarticulate. With one joint.
Uniseriate. With a single column or row of cells.
Urticating hairs. The stinging hairs of nettles and other plants.

Ventricose. Inflated; swelling on one side or unequally.

Vermiform. Worm-shaped; helminthoid.
Verrucate (verrucose). Warty.
Vesiculate. Bladder-like.
Vibrissa. A sensitive hair, as in Dionaea.
Villus. A fine, straight hair.

Wart. A massive, blunt outgrowth; a plant protuberance from the surface of a

cell wall.
Water bladders. Hairs acting as water reservoirs.
Wicker hairs. Trap hairs; elements of a weel.
Whip-like hair. A very long, slender hair which has an enlarged base (a kind of
"whip handle"), and a long, lash-like terminal portion; a flagelliform hair (Fig. 30).

Y-shaped. With two arms diverging at an angle of less than ninety degrees; furcate;
bifurcate (Fig. 16).

PART II. TERMS FOR INDUMENTS

Acanaceous. Having thorns or prickles, said of such prickly plants as thistles; aculeate.
Acanthocomous. Spiny-haired or crowned.
Acervate. Occurring in tufts, heaps, clusters, or cushions.
Aculeate. Prickly; thorny; armed with spines or aculei; acanaceous.
Acuminiferous. With pointed tubercles.
Albopilose. White-shaggy.
Alepidate. Having no scales or scurf; esquamate.
Arachnoid. Covered with long and loosely entangled hairs, longer and fewer than in
tomentose; cobwebby.
Argyrocomous. Silver-haired.

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 252 BRITTONIA [VOL. 3 0

Asperous (asperate). Scabrous; rough; harsh to the touch.

Atrichous. Lacking trichomes, flagella, or cilia.
Auricomous. Golden-haired.

Barbate. Bearded; with long, weak hairs.

Bearded. Provided with long trichomes; with a tuft, line, or zone of hairs, as some
corollas; barbate.
Brackytrichous (brachycomous). Short-haired.
Brevisetous. Short-bristled.
Bristly. Bearing stiff, strong hairs or bristles.

Canescent. With fine, short, whitish or hoary pubescence; having a hoary appearance
due to a covering of short, inconspicuous hairs.
Canous. Whitened with pubescence; greyish-white.
Chaetomallous. Having flowing hair; thick-maned.
Ciliate. Fringed with conspicuous hairs on the margin.
Ciliate-dentate. Having teeth fringed with hair.
Citiatifolious. Having ciliate leaves.
Ciliolate. Inconspicuously ciliate; with small cilia.
Comose (comate). Having a coma or tuft of hairs, especially on the fruits or seeds;
tufted.
Contematose. With a covering which is described as being between bristly and
aculeate.
Crinate. With long, weak hairs; crested; bearded.

Downy. Covered with short, weak, soft hairs.

Echinate. Covered with stout prickles.

Eciliate. Lacking cilia.
Epilose. Destitute of hairs.
Eriantherous. Having woolly anthers.
Erianthous. Having woolly flowers.
Ericanthous. Having woolly spines.
Esquamate. Without scales; alepidote.

Farinaceous. Mealy; furfurate.

Felted. Matted with intertwined hairs.
Fimbriate (fimbricate). With a fringe of long, coarse trichomes along the margin, the
hairs more closely set than for ciliate.
Flavicomous. Yellow-haired.
Floccose. With scattered patches of hairs that usually rub off easily.
Furfurate. Covered with bran or scurfy patches; mealy; farinaceous.

Glabrate. Nearly glabrous.

Glabrous. Lacking trichomes; not hairy.
Glandular. With an indument comprised of or including glandular trichomes.
Glandular-hispid. Rough-hairy, the hairs tipped with glands.
Glandular-pubescent. Covered with fine, gland-bearing hairs, or with glandular and
non-glandular trichomes intermixed.
Glaucescent. Somewhat glaucous.
Glaucous. Covered with a waxy bloom that rubs off; pruinose.
Glochidiate. With glochids (barbed hairs).
Glutinous. Covered with a sticky or gummy exudate.
Gossypine. Cottony; Gossypium-like.

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1978] PAYNE: PLANT HAIR TERMINOLOGY 253

Granular (granulate, granulose). Covered with small granules; finely mealy or

farinaceous.
Granuliferous. Minutely granular; sparsely granular.

Hamulate (hamulous). Beset with small, hooked processes.

Ilebecarpous. Having fruiting structures covered with a downy pubescence.
Hebephyllous. With downy leaves.
Heteroleptic. Variable-scaled.
Hirsute. With long, rather stiff trichomes.
Hirsutellous. Minutely hirsute; hirtellous.
Hirsutulous (hirsutulose). Approaching hirsute.
Hirtellous. Softly or minutely hirsute; hirsutellous.
Hirtiflorous. With hairy flowers.
Hirtipes. Hairy-stalked; hairy-stemmed.
Hispid. With long, very stiff bristles or trichomes.
Hispidulous. Approaching hispid.
Histoid (histioid). Arachnoid; cobwebby.
Hoary. Covered with short, fine, white or whitish hairs; canous; incanous; canescent.
Hypoglaucous. Glaucous beneath.
Hypotrichous. Having pubescence mainly restricted to the undersurface.

Imberbate (imberbis). Without a beard or other hairs.

Imbricate. Flat and lying over one another as the shingles or tiles of a roof.
Incanescent. Becoming grey-pubescent.
Incanous. Hoary with white pubescence.
Indument (indumentum). The hairy covering of a plant surface.
Indumentum, bist;'ate. A two-layered, hairy covering, the outer layer becoming ca-
ducous.
Indumentum, unistrate. A one-layered, hairy covering.
Inuncate (inuncant). Having the surface covered with glochidia or hooked hairs.

Jaculiferous. With dart-like spines.

Lanate. Covered with fine, long hairs; having a woolly surface; lanose.
Lanipes. Woolly-footed; woolly stalked.
Lanose. Woolly; lanate.
Lanuginose. Woolly or cottony, the hairs shorter than lanate.
Lanulose. Very short-woolly.
Lasiacanthous. Pubescent-spined.
Lasiandrous. With pubescent stamens.
Lasianthous. Woolly-flowered.
Lasiocarpous. With pubescent fruit.
Lasiodontous. Woolly-toothed.
Lasioglossous. With a rough, hairy tongue.
Lasiolepous. With woolly scales.
Lasiopetalous. With rough, hairy petals.
Lepidote (lepidioid). Scaly.
Leprate. Covered with white, mealy particles.
Longicomous. With long hair.

Mallococcous. With downy fruit.

Malpighiaceous. With an indumentum of dolabriform trichomes, as many species of
Malpighiaceae or Cornus.

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 254 BRITTONIA [VOL. 3 0

Mealy. Covered with minute particles that tend to come free when touched, as the
leaves of Chenopodium; farinaceous.
Microlepidote. With small scales.
Microtrichous. Having a pubescence so minute that a microscope must be used to
see it; microtrichal.
Mucedinous. White and cottony; appearing moldy.
Mucic. Gummy.

Nitidous. Having a smooth or glossy surface.

Pannose. With the texture of felt; ragged; tattered.

Papillose (papillate, papillar, papilloid). With papillae.
Papillose-pilose. With stiff hairs arising from papillae.
Pappose. Downy or covered with feathery hairs; having a pappus.
Pellucid-dotted. Applied to leaves which contain internal oil glands which are trans-
lucent; punctate or glandular-dotted.
Phoenicolasious. With purple hair.
Pilose. With long, soft trichomes.
Pilosism. Abnormal hairiness in plants.
Pilosism, deforming. Excessive hairiness which completely disfigures the plant.
Pilosism, pkysiological. Hairiness occasioned by circumstances, as growth in dry soil.
Plumulaceous (plumulate). Downy; having a downy covering.
Pollinosous. As though dusted with pollen.
Polyadenous. Having many glands.
Pruinose. Conspicuously glaucous.
Puberulent. With very short hairs; minutely pubescent.
Pubescent. With trichomes; hairy; with soft hairs.
Pulverulent. Covered as if with powder.
Punctate. Dotted as if by punctures; with translucent or pigmented dots or glands.
Puncticulate. Minutely punctate.

Resinous. Coated with a sticky, resin-like exudate, as the bud scales of Aesculus
hippocastanum or Populus deltoides.
Rufous. Reddish; dull red; rusty.

Scaberulent (scaberulous). Slightly scabrous.

Scabrous. Rough and harsh to the touch, as with coarse papillae or trichome bases.
Scurfy. Rough with scales, exfoliations, small shreds, or scaly incrustations.
Sericate. Covered with fine, closely-appressed, silky hairs.
Sericeous. With long and silky hairs.
Setaceous. Bearing setae; with coarse bristles.
Shagreen. Roughly pebbled or minutely cobble-stoned in appearance; papillate.
Smootk. Not scabrous.
Sparse. Thinly scattered or distributed.
Squamose. Coarsely covered with small scales.
Stellipilous (stellate-pilose). Covered with soft, stellate hairs.
Stimulose. Covered with stinging hairs.
Strigose. With short and stiff, appressed hairs, often with swollen bases.
Subcanous. Somewhat hoary.
Subhirtellous. Somewhat hairy.
Supracanous. Grey-pubescent above.

Thelephorous. Covered with nipple-like prominences.

Titanous. Clipped or sheared.

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 1978] PAYNE: PLANT HAIR TERMINOLOGY 255

Tomentose. Densely woolly or pubescent with soft, matted, wool-like trichomes.

Tomentulose. Approaching tomentose.
Trichocarpous. With hairy fruit or ovary.
Tuberculate. With small, rounded protrusions from the surface; warty.

Ulotrichous. Having woolly or curly hair.

Unciferous. Bearing hooks or hook-like processes.
Unisetose. With one bristle.

Velutinous. Velvety with a covering of short, silky hairs.

Verrucose (verrucate). Warty.
Vesicular. Composed of or covered with vesicles; inflated and bladder-like.
Vestiture. Any hairy covering causing a surface to be other than glabrous; indu-
mentum.
Villous (villose). With long, soft, curly but not matted hairs.
Viscid. Clammy; moistly sticky.

Weel. An arrangement of hairs in a flower that keeps out undesirable insects.

Woolly. With long, soft, matted hairs; lanate.

LITERATURE CONSULTED

Arber, A. 1941. On the morphology of the pitcher leaves in Heliamphora, Sarracenia, Cephalotus,
and Nepenthes. Ann. Bot. (London) II 5: 563-576.
Belling, J. 1914. Inheritance in plant hairs. J. Heredity 5: 348-360.
Cannon, W. A. 1870. Studies in heredity as illustrated in the trichomes of species and hybrids
of Juglans, Oenothera, Papaver, and Solanum. Publ. Carnegie Inst. Wash. 117: 1-67.
Carlquist, S. 1961. Comparative plant anatomy. Holt, Rinehart and Winston, Inc., New York.
Esau, K. 1953. Plant anatomy. John Wiley and Sons, Inc., New York.
Gardner, W., & T. Ito. 1887. On the structure of the mucilage secreting cells of Blechnum and
Osmunda. Ann. Bot. (London) 1: 27.
Hemenway, A. F., & M. J. Allen. 1936. A study of the pubescence of cacti. Amer. J. Bot. 23:
139-144.
Kaussman, B. 1954. Die Trichomhydathoden von Mulhenbeckia platyclados. Wiss. Z. Univ.
Rostock, Reihe Math. 3: 231-236.
Linsbauer, K. 1930. "Die Epidermis", Vol. 4, in K. Linsbauer, ed., Handbuch der Pflanzen-
anatomie.
Metcalf, C. R. 1960. Anatomy of the monocotyledons. I. Gramineae. Clarendon Press, Oxford.
1971. Anatomy of the monocotyledons. V. Cyperaceae. Clarendon Press, Oxford.
& L. Chalk. 1950. Anatomy of the dicotyledons. 2 vol. Clarendon Press, Oxford.
Roe, K. E. 1971. Terminology of hairs in the genus Solanum. Taxon 20: 501-508.
Sayre, J. D. 1920. The relations of hairy leaf coverings to the resistance of leaves to transpiration.
Ohio J. Sci. 20: 55-86.
Schwartz, D. 1971. Collegiate dictionary of botany. Ronald Press Co., New York.
Solereder, H. 1908. Systematic anatomy of the dicotyledons. (English ed., translated by L. A.
Boodle and F. E. Fritsch.) 2 vol. Clarendon Press, Oxford.
Tomlinson, P. B. 1961. Anatomy of the monocotyledons. II. Palmae. Clarendon Press, Oxford.
. 1969. Anatomy of the monocotyledons. III. Commelinales-Zingiberales. Clarendon
Press, Oxford.
Uphof, J. C. Th. 1962. Plant hairs. Vol. 4, sect. 5, in K. Linsbauer. Encyclopedia of plant
anatomy. Histology. Gebr. Bortraeger, Berlin.
Williams, S. 1925. Equisetoid hairs. Ann. Bot. (London) 39: 655-656.

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