adjacent formations, or of two consecutive stages of the same
succession. Mixed formations are really transitions in space or in time
between two distinct formations. Theoretically, they are to be referred
to one or the other, according to the preponderance of species.
Actually, however, they often persist in an intermediate condition for
many years, and it becomes necessary to devote considerable attention to
them. In some cases, there is good reason to think that the species of
two contiguous formations have become permanently associated, and thus
constitute a new formation. This is often apparently true in succession,
when the change from one stage to the next requires a long term of
years, but it is really true only of the very rare cases in which a
succession becomes stabilized in a transition stage. When the mixture is
due to development, the formations concerned are often quite dissimilar,
e. g., grassland and thicket, thicket and forest. If it is the result of
position, the formations are usually similar, i. e., both are grassland,
thicket, or forest, since the plants of the lower level are regularly
assimilated or destroyed, when invasion occurs at two levels. The term
_mictium_ (μικτόν, mixture) is here proposed for the designation of all
mixed formations, whether they arise from succession or from
juxtaposition. Thus, the _Mentzelia-Elymus-mictium_ is the transition
between the _Mentzelia-Pseudocymopterus-chalicium_ and the
_Elymus-Muhlenbergia-chalicium_. Similarly, the _Populus-Picea-mictium_
and the _Pinus-Pseudotsuga-mictium_ are transition stages in the
development of the _Picea-hylium_. On the other hand, the
_Andropogon-Bulbilis-mictium_ is a mixture produced by the mingling of
two contiguous prairie formations. In the future development of this
subject, it will probably become desirable to name mixed formations on
the basis of origin, but at present this is unnecessary. Both in
classification and in description they should be considered between the
formations which give rise to them, and this will at once indicate their
origin.
[Illustration: Fig. 82. A mixed formation of aspens and spruces
(_Populus-Picea-mictium_), preceding the final spruce forest of a burn
succession.]
Puzzling cases of mixture resulting from position occur toward the
limits of facies which occupy extensive areas. _Bouteloua oligostachya_,
and _Andropogon scoparius_ extend from the prairies through the
sand-hills and plains, and into the foot-hills of the Rocky mountains.
Their abundance at once raises a question as to the validity of the
prairie, sand-hill, plain, and foot-hill formations. If these two
grasses were controlling, and equally characteristic throughout, then
the entire stretch would have to be regarded as a single formation.
Since they are often absent, or mixed with other facies of greater
importance, they can not be considered the sole tests of the formation.
This view is reinforced by the fact that prairie, sand-hill, plains, and
foot-hill all have their characteristic principal and secondary species,
in addition to facies that are more or less typical. In certain
formations, doubtless, _Bouteloua_ and _Andropogon_ are relicts, in
others invaders, while in the formations actually constituted by them
they are dominant. The final solution of such problems is quite
impossible, however, until the comparative study of large areas can be
based upon the accurate detailed investigation of the component
formations.
EXPERIMENTAL VEGETATION
=356. Scope and methods.= The experimental study of the formation as a
complex organism rests upon methods essentially similar to those
discussed under experimental evolution. The scope of the two fields is
practically the same, moreover, in that both deal with the experimental
development of an organism and the structures that result. The actual
problems are naturally very different, since the formation is a complex
of individual plants, but the fundamental basis of habitat, function,
and structure is common to both. However, the functions now to be
considered are aggregation, invasion, competition, etc., and the
structures, zones, consocies, societies, communities, and families. The
latter may properly be regarded as adaptations called forth by the
adjustment, i. e., aggregation, migration, ecesis, etc., of the
formation to the physical factors of the habitat. As consequences of
measured factors, formational adjustment and adaptation must themselves
be carefully measured and recorded. For these purposes, the methods of
quadrat and transect, of chart, photograph, and formation herbarium are
used. Invaluable as they are for any scientific inquiry into vegetation,
such methods form the very foundation of experimental study in which
accuracy is the first desideratum.
It has already been shown that nature’s own experiments in the
production of new forms furnish the best material for experimental
evolution. This statement is equally true of experimental vegetation.
The formation of new habitats by weathering and transport, and the
denuding of old ones, yield experimental plots of the greatest value.
This is likewise the case in the great majority of formations, where
invasion or competition is active. These are the phenomena that must be
considered in any careful study of vegetation, but in taking them up
from the experimental standpoint, greater attention must be paid to
detail, and the changes must be followed closely for a longer time. The
method that makes use of existing changes in vegetation is designated
the _method of natural habitats_. In contrast with this is the _method
of artificial habitats_, in which the habitat itself is definitely
modified, or a group of species actually transferred to a different
habitat. Many problems of vegetation can be attacked with greater
success under control than in the field. This is particularly true of
competition, in which results can be obtained most readily by means of
the _method of control habitats_, as carried on in the plant house.
METHOD OF NATURAL HABITATS
=357. Natural experiments.= Every family as well as every community
constitutes an experiment in competition; the same statement necessarily
holds for the larger groups, society, consocies, and formation, which
are composed of families and communities. The last also make it possible
to study competition in two typical instances, viz., in the family,
where the individuals are of one kind, and in the community, where they
belong to two or more different species. The community, moreover, is a
product of invasion, and it furnishes material for the study of this
function, as well as for that of aggregation and competition.
Practically every formation shows some invasion, but as a rule stable
formations contain so few invaders that they are relatively unimportant
in this connection. Invasion is most active in transition areas and in
mixed formations, whether produced by juxtaposition or by succession,
and its study in these places yields by far the largest number of
valuable results.
As typical complete invasion, a succession is the best of all natural
experiments in aggregation, migration, ecesis, and competition. This is
especially true of the initial stages in which changes in the number and
position are most readily followed. The methods used in studying
successions have been given elsewhere. In addition, it should be pointed
out that one of the first tasks in taking up the ecological
investigation of a region is to make a careful search for all new and
denuded areas, as well as for those in which succession is taking place.
The phenomena in these areas can not be explained until the habitats and
formations have been worked over critically, but the facts must be
collected at the earliest possible moment, since the stages of the
succession are constantly changing, while the stable formations are not.
METHOD OF ARTIFICIAL HABITATS
=358. Modification of habitat.= As the final factors in ecesis and
competition, water, light, and temperature control the grouping of
plants into vegetation. An efficient change in one of these, or in all
of them, brings about a visible adjustment in the structure of the plant
group concerned. Modifications of water-content and light are readily
produced in the field by drainage, irrigation, shading, clearing, etc.
In fact, all the changes of habitat indicated under experimental
evolution serve equally well to initiate experiments in experimental
vegetation; indeed, the same experiment covers both fields. It is
impracticable, however, to modify the temperature of a habitat without
changing its water-content or light, and consequently the influence of
temperature can not be determined through experiment by modification.
The extent of the area modified should be as large as convenience will
permit, in order that the number of individuals may be large enough to
indicate clearly the resulting adjustment in position and arrangement.
The best results can be obtained where a small separate area of a
formation can be modified, e. g., where a small swamp can be drained, or
a depression flooded. In the case of light, however, it is usually
impossible to clear or to shade a large area, and the study must be
restricted to a relatively small group of plants. In regions where
lumbering is actively carried on, the consequent clearing initiates
invaluable experiments over large areas, and this is likewise true of
forest plantations. Modification of a large area has decided advantages
in bringing out the changes in the more prominent structural features,
but the causes and the details of the adjustment can be worked out much
more satisfactorily in a small area.
=359. Denuding.= The modification of the habitat by denuding is the sole
method of initiating succession by experiment. It is consequently of the
most fundamental importance in investigating aggregation, ecesis, and
competition, as well as the reactions exerted by the invaders of the
different stages. The possibilities of denuding an entire habitat or an
extensive area are not great, and the investigator must content himself
with denuded quadrats, transects, and migration circles, which are small
enough to permit a critical study of all the factors in succession. It
is of course unnecessary that the denuding be done by the ecologist
himself, provided he is able to follow the succession from the very
beginning. Accordingly, it becomes possible for him to make the very
best use of all those changes wrought by man in which the vegetation is
destroyed over considerable areas. These are essentially natural
experiments, and at this point the methods of natural and artificial
habitats merge.
The manner of denuding depends in a degree upon the nature of
vegetation, but, when time, convenience, and safety are all taken into
account, the actual removal of the vegetation as indicated under the
denuded quadrat is by far the most satisfactory. Under certain
conditions, flooding or burning can be used to advantage, but cases of
this kind are infrequent. The purpose of the experiment determines the
kind of area to be denuded. Quadrat, transact, and migration circle are
equally valuable for ecesis and competition. The quadrat is best adapted
to work in a homogeneous area, while the transect is suited to a
heterogeneous one characterized by zones, societies, or communities. It
is an advantage to replace the denuded transect by a series of denuded
quadrats, one for each zone or society, when the transect would be too
long for convenience. The denuded migration circle is invaluable for
aggregation and ecesis, since it makes possible the study of migration
as a distinct function. A series of denuded quadrats, consisting of one
or more in the different stages of a succession, furnishes important
evidence concerning the development of each stage. By far the best
method, however, for making a comparative study of the stages of a
succession is the quadrat sequence. A quadrat is denuded each year, thus
yielding a complete sequence of miniature stages through the whole
course of succession. This method is especially valuable when a
succession is represented by a single example, and there is no
opportunity of reconstructing it by the comparison of various stages. A
quadrat sequence is naturally of the greatest value if begun at the time
when the first invaders appear.
=360. Modification of the formation by transfer.= The study of partial
and intermittent invasion into an established vegetation is made through
the transfer of a species or group of species by means of seeding or
planting. The process differs in no way from that described for
experimental evolution, except in so far that an endeavor is made to
establish a family or a community, and not merely a few individuals.
Transfer makes possible the critical investigation of ecesis under
conditions of intense competition, as well as the study of aggregation
and the origin of plant groups under these conditions. Perhaps its
greatest value is in the experimental study of alternation and zonation,
especially the former. It is practically impossible to determine whether
alternation, especially when corresponsive, is due to physical or
historical causes, i. e., migration and competition, except by means of
the reciprocal transfer of the species concerned.
Field cultures for the careful study of ecesis and competition are made
by transferring seeds or plants to new or denuded soils. This is
practically a combination of the methods of modification and transfer.
It has a unique value in making it possible to initiate artificial
successions of almost any character that is desired, and to carry them
out with the reactions more or less under control. This opens up an
extremely important field of experimental inquiry, which promises to put
the study of succession upon a much more exact basis. Competition
cultures in the field are not essentially different from those under
control, and they will be considered under the next method.
METHOD OF CONTROL HABITATS
[Illustration: Fig. 83. Simple culture of floating ecads of _Ranunculus
sceleratus_.]
=361. Competition cultures.= Although it is quite possible to carry on
experiments in invasion and succession in the planthouse, the limited
space usually available makes this undesirable, except in a few problems
where control is necessary. Competition cultures, on the other hand,
yield better results in the planthouse than in the field, since the
physical factors and the appearance of unwelcome migrants are much more
easily controlled. The possibilities of the culture method in the study
of competition seem inexhaustible, and the author has found it necessary
to confine his own investigations to a few of the fundamental problems.
In this work, he has distinguished several kinds of cultures, based
chiefly upon the species concerned and the arrangement of the
individuals. _Simple_ cultures are those in which a single species is
used. The resulting group is a family, and the competition is between
like individuals. In such cultures, the problem of the factors in
competition is reduced to its simplest terms. _Mixed_ cultures are based
upon two or more species, and the problem is correspondingly
complicated. As a rule, all the seeds have been sown at the same time in
both simple and mixed cultures, but it has been found desirable to make
some _heterochronous_ cultures, in which seeds are also sown after the
plants have appeared. Mixed cultures are distinguished as _layered_
cultures, when the species are of very different height. Thus, rosettes
have been grown with stemmed plants, tall slender forms with low
branching ones, erect plants with twining and climbing plants, etc.
Further evidence as to the nature of competition has been sought by
means of _ecad_ cultures, and _factor_ cultures. In the former, plants
of different response to water and light are grown together under the
same conditions, in order to evaluate the part played by the nature of
the plant. In a factor culture, the area is divided into two or more
parts which are given different amounts of water or of light, in order
to determine the influence of slight variations upon the same
competitors. In somewhat similar fashion, an attempt has been made to
ascertain the bearing of biotic factors upon competition. Cultures are
easily made in which _Cuscuta_ or parasitic fungi are used to place
certain species at a disadvantage. _Permanent_ cultures are obtained by
allowing the plants to ripen and drop their seeds for several
generations, just as in nature. They are indispensable for determining
the final outcome of the competition between different species.
[Illustration: Fig. 84. Mixed culture of _Solidago rigida_ and _Onagra
biennis_.]
=362. Details of culture methods.= All competition cultures have been
made 1 meter square. In other words, they are quadrats, and they are
treated exactly as denuded quadrats in the field with respect to factor
readings, charts, and photographs. In the writer’s studies, germination
tests were made of a large number of species, and those selected which
showed a high per cent of germinability. Since this was the first
experimental study of competition, this test was deemed necessary, but
it is quite evident that no such selection is made in nature.
Consequently, when the seeds used are known to be fresh, a germination
test is usually superfluous. Considerable care was taken also to select
species known to be vigorous growers, with the result that practically
all the species used for experiment were ruderal or subruderal. The
species employed, and the kinds of cultures in which they were grouped
were as follows:
[Illustration: Fig. 85. Heterochronous culture of _Helianthus annuus_
and _Datura stramonium_. Family culture of _Datura_, _Verbascum_, etc.,
in the foreground.]
1. _Simple culture of Helianthus annuus._ The culture plot was divided
into four equal parts; 12 seeds were planted in one, 25 in another, 50
in the third, and 100 in the fourth.
2. _Mixed culture of Helianthus annuus, Panicum virgatum, and Elymus
canadensis._ Twenty-five seeds each of _Helianthus_ and _Panicum_ were
planted alternately at equal distances in one-half of the plot, while
the other half was planted similarly with _Helianthus_ and _Elymus_.
3. _Mixed culture of Solidago rigida and Onagra biennis._ Over
one-half of the plot were scattered 50 seeds of _Solidago_ and 100 of
_Onagra_; over the other, 100 and 200 seeds respectively.
4. _Layered culture of Laciniaria punctata, Bidens frondosa, Salvia
pitcheri, Cassia chamaecrista and Kuhnia glutinosa._ Fifty seeds of
each species were scattered more or less uniformly over the entire
plot.
5. _Layered culture of Silphium laciniatum, Datura stramonium and
Lactuca ludoviciana._ Fifty seeds of _Datura_ and _Lactuca_, and 25 of
_Silphium_ were sown uniformly in one-half of the plot. In the other
half, 25 holes were made at equal intervals, and one seed of each of
the three planted in each hole.
6. _Ecad culture of Oenothera rhombipetala (xerophytic), Verbascum
thapsus (mesophytic), and Penthorum sedoides (hydrophytic)._ One
hundred seeds of _Oenothera_ and 200 each of _Verbascum_ and
_Penthorum_ were scattered over the plot.
7. _Heterochronous culture of Helianthus annuus and Datura
stramonium._ One hundred seeds of Helianthus were scattered over one
half, and the same number of Datura seeds over the other half of the
plot. In both, also, 50 seeds were sown in one 4–inch circle, and 25
seeds in a second circle at some distance. A month later, 100 seeds of
_Helianthus_ were sown in the _Datura_ plot, and _vice versa_.
8. _Family culture of Helianthus, Kuhnia, Panicum, Bidens, Onagra,
Datura, Penthorum, Solidago and Verbascum._ The plot was divided into
9 squares and in each were sown 50 seeds of one of these plants.
9. _Community culture._ The sowing was made exactly as for the family
culture, except that 20 seeds of each plant were used. In the middle
of each square, 5 seeds of a different species were planted. For the
_Helianthus_, _Kuhnia_, and _Panicum_ groups, _Onagra_ was used; for
_Bidens_, _Onagra_, and _Datura_, _Helianthus_ was used, and for
_Penthorum_, _Solidago_, and _Verbascum_, _Panicum_.
At the time the cultures were started, check plants were sown in pots.
The most vigorous seedlings were transplanted singly to large pots, and
grown under conditions of water, light, and soil as similar as possible
to those of the competition plots. Photographs of check plants and plots
were made at the proper intervals, and the plots were charted in
quadrats to show the course of competition. The factors which control
competition were sought in a critical study of water-content and light
values, which is still in process. This work has gone far enough to
indicate the correctness of the view[45] that competition is purely
physical in character. It has, moreover, been demonstrated that “room”
in competition is merely a loose expression for the relation between the
number of individuals in a given space, and the amount of water, light,
and temperature available in the same space.
GLOSSARY
NOTE: Last terms frequent in compounds are found in their proper place
alphabetically. The accent is indicated only in those words accented on
the penult; all others are accented on the antepenult, or recessively.
=abundance=, the total number of individuals in an area.
=acospore= (ἀκή, point), a plant with awned disseminules.
=acrophyti´um= (ἄκρον, peak), an alpine plant formation.
=acti´um= (ἀκτή, rocky coast), a rocky seashore formation; =actad=,
plant of a rocky seashore.
=-ad= (-αδης, patronymic suffix), suffix for denoting an ecad.
=adaptable=, able to originate ecads; =adaptation=, the structural
response to stimuli.
=adjustment=, the functional response to stimuli.
=adventicious= (_adventicius_, foreign), invading from distant
formations.
=adventive= (_adventivus_, accidental), established temporarily.
=aggregation=, the coming together of plants into groups.
=agi´um= (ἀγή, beach), a beach formation; =agad=, a beach plant.
=agri´um= (ἀγρός, field), a culture formation; =agrad=, a cultivated
plant.
=aiphyti´um= (ἀεί, permanent), an ultimate formation.
=alsi´um= (ἄλσος, grove), a grove formation; =alsad=, a grove plant.
=alternation=, the heterogeneous arrangement of plant groups and
formations universally present in vegetation.
=amathi´um= (ἄμαθος, sand of the plain), a sand-hill or sandplain
formation; =amathad=, a sand-hill plant.
=ammochthi´um= (ἄμμος, sand, ὄχθη, bank), a sand bank formation;
=ammochthad=, a sand bank plant.
=anci´um= (ἄγκος, mountain glen), a canyon formation; =ancad=, a
canyon plant.
=anemi´um= (ἄνεμος, wind), a blowout formation; =anemad=, a blowout
plant; =anemochore=, a plant distributed by wind.
=-anum= (locative suffix), a suffix denoting a layer.
=apostrophe= (ἀπό, away from, στροφή, a turning), the arrangement of
the row of chloroplasts parallel to the rays of light.
=apparent noon=, the time when the sun crosses the meridian, i. e.,
sun noon as distinguished from noon, standard time.
=-ard= (ἄρδον, water of the land), combining term for water-content;
=ardium=, a succession due to irrigation.
=ardesiacus=, slate colored.
=-are= (locative suffix), suffix denoting a community.
=aspect= (_aspectus_, appearance), the seasonal impress of a
formation, e.g., the spring aspect.
=association=, the arrangement of individuals in vegetation.
=atmometer= (ἀτμός, vapor), an instrument for measuring evaporation.
=atropurpureus=, dark purple.
=atrovirens=, dark green.
=autochore= (αὐτός, self), motile plants, or those with motile spores;
=autochthonous= (χθών, ground), native.
=avellaneus=, drab.
=barrier=, a physical or biological obstacle to migration or ecesis.
=bathyphyti´um= (βαθύς, low), a lowland plant formation.
=blastochore= (βλάστη, growth), a plant distributed by offshoots.
=-bole= (βολή, a throw), combining term for propulsion; =bolochore=, a
plant distributed by propulsion.
=broti´um= (βροτός, mortal), a succession caused by man; =brotochore=,
a plant distributed by man.
=caeruleus=, pale blue.
=caesius=, eye-blue.
=camni´um= (κάμνω, cultivate), a succession due to cultivation.
=carphospore= (κάρφος, scale), a plant with disseminules possessing a
scaly or chaffy pappus.
=carpostrote= (καρπός, fruit), a plant migrating by means of fruits.
=centrospore= (κέντρον, spur), a plant with spiny disseminules.
=chalici´um= (χάλιξ, gravel), a gravel slide formation; =chalicad=, a
gravel slide plant.
=cheradi´um= (χέραδος, a sandbar), a wet sandbar formation;
=cheradad=, a wet sandbar plant.
=chersi´um= (χέρσος, dry barren waste), a dry waste formation;
=chersad=, plant of a dry waste.
=chioni´um= (χιών, όνος, snow), a snow formation; =chionad=, a snow
plant; =chionophyti´um=, a niveal plant formation.
=chledi´um= (χλῆδος, rubbish), a ruderal formation; =chledad=, a
ruderal plant.
=chlorenchym= (χλωρός, greenish yellow, ἐνχύμα, infusion), the
chlorophyll tissue of the leaf.
=-chore= (χωρέω, to spread abroad), combining term to denote agent of
migration.
=chresard= (χρῆσις, use), the available water of the soil, the
physiological water-content.
=clitochore= (κλίτος, slope), a plant distributed by gravity.
=clysi´um= (κλύσις, a flooding), a succession in a flooded soil.
=-colus= (κόλος, dwelling in), combining term for habitat forms.
=community=, a mixture of the individuals of two or more species, a
group of families.
=comospore= (κόμη, hair) a plant with hairy or silky disseminules.
=competition=, the relation between plants occupying the same area,
and dependent upon the same supply of physical factors.
=consocies=, that subdivision of a formation controlled by a facies.
=copious=, used of species in which the individuals are arranged
closely but uniformly.
=coryphi´um= (κορυφή, peak), an alpine meadow formation; =coryphad=,
an alpine meadow plant.
=creatospore= (κρέας, ατος, meat), a plant with nut fruits.
=cremni´um= (κρημνός, crag, cliff), a cliff formation; =cremnad=, a
cliff plant.
=creni´um= (κρήνη, spring), a spring formation; =crenad=, a spring
plant.
=crymi´um= (κρυμός, frost), a polar barren formation; =crymad=, a
polar plant; =crymophytic=, pertaining to polar plants.
=crystallochore= (κρύσταλλος, ice), a plant distributed by glaciers.
=cyaneus=, azure.
=cyriodoche= (κύριος, regular), a normal succession.
=dendri´um= (δένδρα, fruit trees), an orchard formation; =dendrad=, an
orchard plant.
=derived=, coming from other formations or regions, not native.
=diphotic= (δι-, two), the two surfaces unequally lighted;
=diphotophyll=, a leaf differentiated into palisade and sponge
tissues owing to unequal illumination.
=diplophyll= (διπλόος, twofold), an isophotic leaf with water-storage
cells in the middle.
=disseminule= (semen, seed), a seed fruit modified for migration.
=dissophyte= (δισσός, double), a plant with xerophytic leaves and
stems, and mesophytic roots.
=-doche= (δοχή, succession), succession.
=drimi´um= (δριμύς, biting, pungent), an alkaline habitat, and the
corresponding formation; =drimad=, a plant of such a formation.
=driodi´um= (δρίος, thicket), a dry thicket formation; =driodad=,
plant of a dry thicket.
=dysgeogenous= (δυς-, bad, γῆ, soil), weathering with difficulty to
form soil.
=ecad= (οἶκος, home), a habitat form due to origin by adaptation;
=ece´sis= (οἰκῆσις, act of coming to be at home), the germination
and establishment of invaders; =ecograph=, an instrument for
measuring a physical factor of a habitat; =ecotone= (τόνος,
tension), the tension line between two zones, formations, consocies,
etc.
=ecballi´um=, (ἐκβάλλω, cut down forests), a succession due to
lumbering.
=echard= (ἔχω, to withhold), the non-available water of the soil.
=edobole= (οἶδος, swelling), a plant whose seeds are scattered by
propulsion through turgescence.
=efficient difference=, the amount of a physical factor necessary to
produce a change in the response.
=enauli´um= (ἔναυλος, hollow channel), a sanddraw formation;
=enaulad=, a sanddraw plant.
=ende´mic= (ἐν, within, δῆμος, district), occurring in a single
formation, or natural region; =ende´mism=, the condition of growing
in but one natural area.
=epistrophe= (ἐπί, towards, στροφή, a turning), the arrangement of the
row of chloroplasts at right angles to the incident light.
=eremi´um= (ἔρημος, desert), a desert formation; =eremad=, a desert
plant.
=estival=, pertaining to summer.
=-etum= (locative suffix), suffix used to denote a consocies.
=eugeogenous= (εὖ-, well, γῆ, soil), weathering readily to form soil.
=facies=, a dominant species of a formation: a distinct area
controlled by it is a consocies.
=family=, a group of individuals belonging to one species.
=fixity=, the condition characterized by little or no response to
stimuli.
=flavovirens=, yellow green.
=forewold=, equivalent to the German “vorwald,” the thicket zone
bordering a forest.
=-genous= (γένω, to produce), producing.
=geotome= (γῆ, earth, τομή, edge), an instrument for obtaining soil
samples.
=gloeospore= (γλοιός, sticky stuff), a plant with viscid disseminules.
=-graph= (γραφή, a writing), combining term for a recording
instrument.
=gregarious= (gregarius, grouped in herds), used of species in which
the individuals occur in groups.
=habitat=, a definite physical area characterized by a formation;
=habitat form=, the impress given the plant by the habitat.
=harmosis= (ἅρμοσις, an adapting), response to stimuli, comprising
both adjustment and adaptation.
=hedi´um= (ἕδος, a sitting, base), a succession in a residuary soil.
=heliad= (ἥλιος, sun), a heliophyte; =heliophyll=, the leaf of a sun
plant; =heliophyte=, a sun plant; =heliophyti´um=, a sun plant
formation; =heliophilous=, sun-loving.
=heli´um= (ἕλος, marsh), a marsh formation; =helad=, a marsh plant;
=helodi´um= (ἑλώδης, marshy), a swampy open woodland formation;
=helodad=, a marsh plant; =helodrium= (δρίος, thicket), a thicket
formation; =helodrad=, a plant of a marshy thicket; =helohyli´um=
(ὕλη, forest) a marsh forest formation; =helohylad=, a marsh forest
plant.
=hepodoche= (ἕπω, follow), a secondary succession.
=hizometer= (ἵζω, to sink), an instrument for measuring gravitation
water.
=holard= (ὅλος, whole), the total water-content of the soil.
=hydrad= (ὑδρο-, water), a hydrophyte; =hydrochore=, a plant
distributed by water; =hydroharmose=, response to water stimuli;
=hydrophyll=, the leaf of a hydrophyte; =hydrophyte=, a water plant;
=hydrophyti´um=, a water plant formation; =hydrophilous=,
water-loving; =hydrosta´tic= (στατικός, standing), completing the
succession under hydrophytic conditions; =hydrotropic= (τροπικός,
turning), applied to successions which become mesophytic.
=hygrome´tric= (ὑγρίς, wet), measuring or absorbing water;
=hygroscopic= (σκοπέω, look), measurable only by a hygroscope; able
to absorb moisture.
=hyli´um= (ὕλη, forest), a forest formation; =hylad=, a forest plant;
=hylocolum=, dwelling in a forest; =hylodi´um= (ὑλώδης, wooded), a
dry open woodland formation; =hylodad=, a plant of this formation;
=hylophyte=, a forest plant.
=hypsi´um= (ὕψος, elevation), a succession caused by elevation.
=-ile= (locative affix), suffix denoting a society.
=immobile=, without effective devices for migration.
=indigenous= (_indigena_, sprung from the land), native.
=insolation=, exposure to intense heat and light.
=isabellinus=, leather-colored.
=isolation=, separation by barriers.
=isopho´tic= (ἴσος, equal), equally illuminated; =isophotophyll=, a
leaf in which both halves of the chlorenchym are alike, due to equal
illumination.
=-ium= (-εῖον, locative affix), suffix denoting a formation.
=labile=, plastic, easily modified.
=lauri´um= (λαύρα, drain), a drain formation; =laurad=, a drain plant.
=limni´um= (λίμνη, lake), a lake formation; =limnad=, a lake plant;
=limnodium= (λιμνῶδες, marshy ground), a salt marsh formation;
=limnodad=, a plant of a salt marsh.
=lochmi´um= (λόχμη, thicket), a thicket formation; =lochmad=, a
thicket plant.
=lophi´um= (λόφος, crest, hill), a hill formation; =lophad=, a hill
plant; =lophospore=, a plant with plumose disseminules.
=mastigospore= (μάστιξ, ιγος, lash), a plant with ciliate or
flagellate disseminules.
=melleus=, honey-colored.
=meridian=, used chiefly as a synonym for apparent noon; also an
imaginary line of longitude.
=mesad= (μέσος, middle), a mesophyte; =mesophilous=, growing in moist
soils; =mesophyll=, the leaf of a mesophyte; =mesophyte=, a plant of
moist soils; =mesophyti´um=, a mesophytic formation; =mesosta´tic=
(στατικίς, standing), completing the succession under mesophytic
conditions; =mesotro´pic= (τροπικός, turning), applied to
successions which become mesophytic.
=-meter= (μέτρον, measure), combining term for instrument.
=micti´um= (μικτόν, mixture), a mixed formation.
=migrant=, a plant that is migrating or invading.
=migration= (_migratio_, removal), the movement of plants into new
areas; =migration circle=, a circle employed to measure migration.
=mobile=, able to be moved, i. e., modified for migration.
=monochronic= (μόνος, single, χρόνος, time), arising but once;
=monogenesis= (γένεσις, origin), the origin of a new form at a
single place or time; =monophyle´sis= (φῦλον, race), origin from a
single ancestral type; =monoto´pic= (τόπος, place), arising at one
place only.
=motile=, able to move by growth, by means of cilia, etc.
=mutable=, able to produce mutants; =mutant=, a form arising by
mutation; =mutation=, the sudden appearance of new forms.
=namati´um= (νάμα, ατος, brook), a brook formation; =namatad=, a brook
plant.
=nomi´um= (νομός, pasture), a pasture formation; =nomad=, a pasture
plant.
=occupation=, possession of the ground by plants.
=oceani´um= (ὠκεανός, ocean), an ocean formation; =oceanad=, an ocean
plant; =oceanophyte=, an ocean plant; =oceanophilous=,
ocean-dwelling.
=ocheti´um= (ὀχετός, drain), a succession due to artificial drainage.
=ochroleucus=, yellowish white.
=ochthi´um= (ὄχθη, bank), a bank formation; =ochthad=, a bank plant.
=oligope´lic= (ὀλίγος, little, πηλός clay), containing little clay;
=oligopsam´mic= (ψάμμος, sand), containing little sand.
=olisthi´um= (ὄλισθος, slip), a succession in a landslip.
=ombrometer= (ὄμβρος, a rainstorm), a rain gauge.
=-on= (-ών, locative suffix), suffix used to denote a family.
=oncospore= (ὄγκος, hook), a plant with hooked disseminules.
=orgadi´um= (ὀργάς, άδος, meadowland partially wooded), an open
woodland formation; =orgadad=, an open woodland plant.
=orophyti´um= (ὄρος, mountain), a subalpine plant formation.
=oxodi´um= (ὀξώδης, sour), a humus marsh formation; =oxodad=, a plant
of a humus marsh.
=pagi´um= (πάγος, rocky hill, glacier), a succession in a glacial
soil; =pagophyti´um=, a foot-hill plant formation.
=pediophyti´um= (πεδίον, plain), an upland plant formation.
=pelagi´um= (πέλαγος, surface of the sea), a surface sea formation;
=pelagad=, a plant of the sea surface.
=pelochthi´um= (πηλός, mud, ὄχθη, bank), a mud bank form;
=pelogenous=, producing clay; =pelopsammic= (ψάμμος, sand), composed
of mixed clay and sand; =pelopsammogenous=, producing clay and sand.
=permobile=, extremely mobile.
=perquadrat=, a quadrat of 16 square meters or more.
=petasospore= (πέτασος, sunshade), a plant with parachute-like
disseminules.
=petri´um= (πέτρα, rock, stone), a rock formation; =petrad=, a rock
plant; =petrochthi´um= (ὄχθη, bank), a rock bank formation.
=petrodi´um= (πετρώδης, abounding in boulders), a boulder field
formation; =petrodad=, a plant of a boulder field.
=phelli´um= (φελλεύς, stony ground), a rock field formation;
=phellad=, a rock field plant.
=-philous= (φίλος), loving, dwelling in.
=-photic= (φῶς, φωτός, light), pertaining to light; =photoharmose=,
response to light stimuli; =photometer=, an instrument for measuring
light.
=phreti´um= (φρητός, tank), a tank formation; =phretad=, a tank plant.
=phyad= (φυή, form of growth), a vegetation form, e. g., tree, shrub,
etc.
=-phyll= (φύλλον, leaf), combining term for leaf.
=-phyte= (φυτόν, plant), combining term denoting plant; =phyteris=
(ἔρις, strife), plant competition; =-phyti´um= (φυτεῖον, place
covered with plants), combining term for formation; =phytostrote=, a
species migrating by means of the plant body.
=pladobole= (πλάδος, moisture), a plant whose seeds are scattered by
propulsion due to moisture.
=plasticity=, the condition characterized by ready response to
stimuli.
=pnoi´um= (πνοή, blast), a succession in an aeolian soil.
=poi´um= (πόα, meadow), meadow formation; =poad=, a meadow plant;
=poophyte=, a meadow plant.
=polyan´thous= (πολύς, many, ἄνθος, flower), producing many flowers;
=polychro´nic= (χρόνος, time), arising at two or more times;
=polyde´mic= (δῆμος, district), occurring in two or more formations
or natural regions; =polygenesis= (γένεσις, origin), the origin of a
new form at two or more places or times; =polyphyle´sis= (φῦλον,
race), the origin of a form, species, or genus from two or more
ancestral types; =polyspermatous= (σπέρμα, seed), producing many
seeds in each flower; =polyto´pic= (τόπος, place), arising at two or
more distinct places.
=ponti´um= (πόντος, deep sea), a deep sea formation.
=potami´um= (ποταμός, river), a river formation; =potamad=, a river
plant.
=potometer= (ποτόν, drink), an instrument for measuring absorption.
=prevernal=, pertaining to early spring.
=prior=, earlier, used of alpine aspects.
=prochosi´um= (πρόχωσις, a deposition of mud), a succession in an
alluvial soil.
=prodophyti´um= (πρόοδος, pioneer), an initial formation.
=protodoche= (πρῶτος, first), a primary succession.
=proximity= (proximitas, nearness), nearness to the area invaded.
=psamathi´um= (ψάμαθος, sand of the seashore), a strand formation;
=psamathad=, a strand plant; =psammogenous= (ψάμμος, sand),
producing a sandy soil.
=psili´um= (ψιλά, land without trees), a prairie formation; =psilad=,
a prairie plant.
=psychrometer= (ψυχρός, chill), an instrument that measures humidity
by means of a fall in temperature; =psychrograph=, a psychrometer
that records automatically.
=ptenophyti´um= (πτηνός, passing), an intermediate formation.
=pterospore= (πτερόν, wing), a plant with winged disseminules.
=purpureus=, purple.
=pycnophyti´um= (πυκνός, thick), a closed formation.
=pyri´um= (πῦρ, fire), a burn succession.
=quadrat= (_quadratum_, a square), a square meter of vegetation marked
off for counting, mapping, etc.; =major=, a quadrat of 2–14 square
meters.
=reaction=, the effect of the formation upon the habitat.
=relict= (_relictus_, left), a species belonging properly to an
earlier type of succession than the one in which it is found.
=repi´um= (ῥέπω, sink), a succession due to subsidence.
=rhoi´um= (ῥόος, stream), a creek formation; =rhoad=, a creek plant.
=rhoptometer= (ῥοπτόν, something absorbed), an instrument to measure
absorption of water by the soil.
=rhyaci´um= (ῥύαξ, ακος, mountain torrent), a torrent formation;
=rhyacad=, a torrent plant.
=rhysi´um= (ῥυσίς, a flowing of fire), a succession due to volcanic
action.
=ruber=, red.
=saccospore= (σάκκος, sack), a plant with sack-like disseminules.
=sarcospore= (σάρξ, σαρκός, flesh), a plant with fleshy disseminules.
=sciad= (σκιά, shade), a sciophyte; =sciophyll=, the leaf of a shade
plant; =sciophyte=, a shade plant; =sciophyti´um=, a shade plant
formation; =sciophilous=, shade-loving.
=selagraph= (σέλας, light), an instrument for recording light values
automatically.
=serotinal=, late, pertaining to autumn.
=social=, used of plants in which the individuals are compactly
grouped; =exclusive=, excluding individuals of other species;
=inclusive=, permitting the entrance of individuals of other
species.
=society=, a subdivision of the formation, characterized by a
principal species.
=sparse=, scattered singly.
=spermatostrote= (σπέρμα, ατος, seed), a plant migrating by means of
seeds.
=sphyri´um= (σφύρον, ankle, talus), a succession in a talus soil.
=spongophyll= (σπόγγος, a sponge), a leaf consisting of sponge tissue.
=sporadophyti´um= (σποράς, άδος, scattered), an open formation.
=-spore= (σπορά, seed, fruit), combining term for migration
contrivance; =sporostrote=, a plant migrating by means of spores.
=stability=, the condition in which the plant makes little or no
response.
=stabilization=, the tendency typical of succession, in which the
successive stages become more stable.
=stasi´um= (στάσις, a standing), a stagnant pool formation; =stasad=,
a plant of stagnant water.
=staurophyll= (σταυρός, a pale), a leaf consisting of palisade tissue.
=sterrhi´um= (στερρός, barren), a moor formation; =sterrhad=, a moor
plant.
=-strote= (στρώτος, strewn), combining term for means of migration.
=subcopious=, scattered somewhat loosely.
=subgregarious=, arranged in loose groups.
=subquadrat=, a quadrat of 1–8 decimeters.
=succession=, complete and continuous or repeated invasion, in
consequence of which formations succeed each other.
=symmetry=, used of topography when it shows uniform changes;
=radial=, a condition in which the different areas are concentric;
=bilateral=, where the areas occur in two similar rows.
=syrtidi´um= (σύρτις, ιδος, sandbar), a dry sandbar formation;
=syrtidad=, a plant of a dry sandbar.
=taphri´um= (τάφρος, ditch), a ditch formation; =taphrad=, a ditch
plant.
=telmati´um= (τέλμα, ατος, water meads), a wet meadow formation;
=telmatad=, a wet meadow plant.
=testaceus=, pale brick colored.
=thalassi´um= (θάλασσα, sea), a sea formation; =thalassad=, a sea
plant.
=thallostrote= (θαλλός, shoot), a species migrating by means of
offshoots.
=theri´um= (θήρ, wild animal), a succession due to animals.
=thermi´um= (θέρμη, hot spring), a hot spring formation; =thermad=, a
hot spring plant.
=thini´um= (θίς, θινός, a dune), a dune formation; =thinad=, a dune
plant.
=tiphi´um= (τῖφος, pool), a pool formation; =tiphad=, a pond plant.
=tiri´um= (τείρω, rub away), a bad land formation; =tirad=, a bad land
plant.
=tonobole= (τόνος, tension), a plant whose seeds are scattered by
projection from calyx or involucre.
=transect= (_transectus_, cut through), a cross section of vegetation.
=trechometer= (τρέχω, to run off), an instrument for measuring
run-off.
=tribi´um= (τρίβω, wear or rub away), a succession in an eroded soil.
=umbrinus=, umber.
=variable=, able to produce variants; =variant=, a form arising from
origin by variation; =variation=, the origin of new forms by the
action of selection upon minute differences.
=vegetation form=, a characteristic plant form, e. g., tree, rosette,
etc.
=vernal=, pertaining to spring.
=vicine= (_vicinus_, neighboring), invading from adjacent formations.
=viridis=, green.
=vixgregarious=, arranged in small or indistinct groups.
=water-content=, the water of the soil or habitat; =physiological=,
the available soil water; =physical=, the total amount of soil
water.
=xenodoche= (ξένος, strange), an anomalous succession.
=xerad= (ξηρός, dry), a xerophyte; =xerasi´um= (ξηρασία, drought), a
succession due to drainage or drought; =xeriobole= (ξηρία, dryness),
a plant whose seeds are scattered by dehiscence due to dryness;
=xerohyli´um= (ὕλη, forest), a dry forest formation; =xerohylad=, a
dry forest plant; =xerophyll=, the leaf of a xerophyte; =xerophyte=,
a dry soil plant; =xerophyti´um=, a xerophytic formation;
=xerophilous=, dwelling in a dry habitat; =xeropoi´um=, a heath
formation; =xeropoad=, a heath plant; =xerosta´tic= (στατικός,
standing), used of successions which are completed under xerophytic
conditions; =xerotro´pic= (τροπικός, turning), applied to
successions which become xerophytic.
=zonation=, that condition in which plant groups or formations appear
in belts or zones.
=zone=, a belt of more or less uniform vegetation.
=zoochore= (ζῶον, animal), a plant distributed by animals.
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-----
Footnote 1:
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Bot. 2:514. 1890.
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du milieu. Ann. Nat. Sci. 6:7:93. 1878.
Footnote 2:
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Soils. Bull. Div. Soils, 12:12. 1898.
Footnote 3:
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Certain Plants, Principally Mesophytes. Rep. Bot. Surv. Nebr., 6:48.
1902.
Footnote 4:
BUNSEN, R., AND ROSCOE, H. Photometrische Untersuchungen.
Poggendorff’s Annalen., 117:529. 1862.
WIESNER, J.
Photometrische Untersuchungen auf pflanzenphysiologischen Gebiete.
Sitzb. Akad. Wiss. Wien., I, 1893. II, 1895.
Untersuchungen über das photochemische Klima von Wien, Cairo, und
Buitenzorg (Java) Denksch. Kais. Akad. Wien., 64. 1896.
Untersuchungen über den Lichtgenuss der Pflanzen im arktischen
Gebiete. Sitzb. Kais. Akad. Wien., 109. 1900.
Footnote 5:
REINKE, J. Bot. Zeit., 41:713. 1883.
Footnote 6:
MEYEN, F. J. F. Grundriss der Pflanzengeographie, 12. 1836.
Footnote 7:
Instructions to Field Parties and Descriptions of Soil Types, 35.
1903.
Footnote 8:
PFEFFER-EWART. Physiology of Plants, 1:13. 1900.
Footnote 9:
The Relation of Leaf Structure to Physical Factors. 1905.
Footnote 10:
JEVONS, W. A. The Principles of Science, 2:137. 1874.
Footnote 11:
Die Transpiration der Pflanzen, 14. 1904.
Footnote 12:
Recherches sur le Rôle Physiologique de l’Eau dans la Végétation. Ann.
Nat. Sci., 7:20:65. 1895.
Footnote 13:
Observations on Stomata by a New Method. Proc. Camb. Phil. Soc.,
9:303. 1897.
Footnote 14:
Zur Kenntnis des Pflanzenlebens schwedischer Laubwiesen. Beih. Bot.
Cent., 18:311. 1904.
Footnote 15:
Lehrbuch der Oekologischen Pflanzengeographie. 2d ed., 196. 1902.
Footnote 16:
Physiologische Pflanzenanatomie. 3d ed., 537. 1904.
Footnote 17:
SACHS, J. Ein Beitrag zur Kenntniss der Ernährungsthätigkeit der
Blätter. Gesammelte Abhandlungen über Pflanzenphysiologie. 1:355.
1892.
Footnote 18:
CLEMENTS, E. S. The Relation of Leaf Structure to Physical Factors.
1905.
Footnote 19:
DE VRIES, H. Die Mutationstheorie, 1:6. 1901.
Footnote 20:
POUND AND CLEMENTS. A Method of Determining the Abundance of Secondary
Species. Minn. Bot. Studies, 2:19. 1898.
Footnote 21:
The Development and Structure of Vegetation, 84. 1904.
THORNBER, J. J. The Prairiegrass Formation in Region I. Rep. Bot.
Surv. Neb., 5:29. 1901.
Footnote 22:
Pound and Clements. The Vegetation Regions of the Prairie Province.
Bot. Gaz., 25:381. 1898.
Footnote 23:
Clements, F. E. and E. S. Herbaria Formationum Coloradensium. 1902.
Footnote 24:
Lehrbuch der Ökologischen Pflanzengeographie, 97. 1896.
Footnote 25:
Phytogeography of Nebraska, 1st ed., 101. 1898.
Footnote 26:
Pflanzengeographie auf physiologischer Grundlage, 208. 1898.
Footnote 27:
Die Vegetation der Erde. Engler Bot. Jahrb., 17:b55. 1893.
Footnote 28:
Thornber, J. J. The Prairiegrass Formation in Region I. Rep. Bot.
Surv. Nebr., 5:36, 46. 1901.
Footnote 29:
Grundzüge einer allgemeinen Pflanzengeographie, 157. 1823.
Footnote 30:
Essai de phytostatique, etc. 1849.
Footnote 31:
_l. c._, 116. 1896.
Footnote 32:
_l. c._, 3. 1898.
Footnote 33:
Uber mechanische Shutzmittel der Samen gegen schädliche Einflüsse von
aussen. Engler Bot. Jahrb., 5:56. 1883.
Footnote 34:
Essai Elémentaire de Géographie Botanique, 45. 1820.
Footnote 35:
Stationes Plantarum Amoen. Acad., 4:64. 1754.
Footnote 36:
Die Vegetation der Erde, 4. 1872.
Footnote 37:
When this word was first proposed, the author did not know that
Briquet had already applied the term _polytopism_ to this concept
(Ann. Conserv. Bot. Gen., 5:73. 1901). Since polygenesis expresses the
idea of origin, and applies to multiple origin in time as well as in
space, it is retained as the name of this concept. Polytopic and
monotopic are adopted for multiple and single origin in space
respectively, and _polychronic_ and _monochronic_ are proposed for
similar origin in time.
Footnote 38:
The Origin of Species, 186. 1859.
Footnote 39:
Rocks, Rock-weathering, and Soils, 300. 1897.
Footnote 40:
Relation d’un Voyage du Levant. 1717.
Footnote 41:
Clements, F. E. The Development and Structure of Vegetation. Rep. Bot.
Surv. Nebr., 7:163. 1904.
Footnote 42:
Clements, F. E. The Development and Structure of Vegetation, 24, 27.
1904.
Footnote 43:
Clements, F. E. A System of Nomenclature for Phytogeography. Engler
Jahrb., 31:b70:1. 1902.
Footnote 44:
The terms, _oceanad_, _hylad_, _poad_, _eremad_, etc., are proposed in
place of _oceanophyte_, _hylophyte_, etc. They are much shorter and
make consistent groups under the general term, _ecad._, i. e., habitat
form.
Footnote 45:
CLEMENTS, F. E. The Development and Structure of Vegetation, 166.
1904.
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TRANSCRIBER’S NOTES
Page Changed from Changed to
84 144 square inches of water, with 144 cubic inches of water, with
an opening ¼ × 12 inches at the an opening ¼ × 12 inches at the
base base
234 as many readily be seen from the as may readily be seen from the
fact that migration varies fact that migration varies
inversely as the inversely as the
319 ochthi´um (ὄχφη, bank), a bank ochthi´um (ὄχθη, bank), a bank
formation; ochthad, a bank plant formation; ochthad, a bank plant
1. Typos fixed; non-standard spelling and dialect retained.
2. Enclosed italics font in _underscores_.
3. Enclosed bold or blackletter font in =equals=.
4. The caret (^) serves as a superscript indicator, applicable to
individual characters (like 2^d) and even entire phrases (like
1^{st}).
5. Subscripts are shown using an underscore (_) with curly braces { },
as in H_{2}O.