Horotely, Bradytely, and Tachytely
by George Gaylord Simpson
t is abundantly evident that rates of evolution vary. They vary greatly from
group to group, and even among closely related lineages there may be strikingly
different rates. Differences in rates of evolution, and not only divergent
evolution at comparable rates, are among the reasons for the great diversity
of organisms on the earth. Among the living primates there are, for instance,
some rather unspecialized or primitive prosimians (i.e., little changed from
Eocene progenitors), a larger number of divergently specialized prosimians,
many monkeys of different degrees of progression and divergence, a few apes,
and the unique species of man. Important as is the purely divergent
evolution, it is also clear that differential rates are involved. At the
extremes, the lineages of the more primitive living prosimians have evolved
less rapidly as regards the whole of their structure and adaptive position
than has the lineage of man.
Anyone group, large or small, seems to have a fairly
characteristic mean or modal rate of evolution and a certain range of less
common rates on each side of this. The rates have a distribution pattern
which is open to investigation and which is an important element in
evolution. It has long been noticed, further, that there are a few lines in
most large groups that seem to evolve at altogether exceptional rates. Some
hardly change at all over long periods of time. Others change, over short
periods of time, at rates so rapid as to be almost beyond comparison with
those usual among their relatives. It must be considered whether these
extraordinary rates are merely the extremes of normal variation in rates or
whether they reflect special circumstances different from those in the
distributions of usual rates. In either case they must involve different
intensities, combinations, or both as regards evolutionary determinants,
and it would be of extreme interest, perhaps even of considerable
practical value, to identify these.
Distributions of Rates of Evolution
In Chapters I and II the examples of how rates may be
estimated and represented showed that: (a) related lines of descent
commonly differ in evolutionary rates; (b) within larger taxonomic groups
such as orders or classes there is generally an average or modal rate
typical for the group; and (c) the average rates may differ greatly from
one group to another. Since rates do vary broadly around a group mode,
rate distributions of different groups tend to overlap widely even though
the modes are quite distinct. Mammals seem surely to have a much higher
model rate of evolution than molluscs, but some mammals have evolved more
slowly than some molluscs. It even seems clear that some mammals have
evolved at rates below the mode for molluscs and some molluscs at rates
above the mode for mammals.
Such comparisons are hazardous because there is no
absolute way to measure equivalent amounts of evolution in two groups
anatomically so different and changing in such different ways. The two
cannot even both be studied authoritatively by one specialist. This sort
of objection is often made, and of course it carries a great deal of
force. It is nevertheless not as strong or as fatal to comparisons as
sometimes claimed. For instance, it is often said (e.g., again by
Hutchinson, 1945, in criticism of the predecessor of this book) that
molluscs may have evolved very rapidly in the soft parts without having
this reflected in the shell. But in molluscs the whole skeleton is usually
available and studied in fossils (which is very rarely true of, say,
fossil mammals) and their skeleton reveals a great deal about the soft
parts, directly (muscle scars, etc.) or indirectly (growth rates,
thickness and chemical nature of shell, twisting of body, etc.).
Percentagewise a mollusc shell probably tells more about the soft parts
than do the usually available parts of a vertebrate. It is extremely
unlikely that any marked progressive change in soft parts could occur
without obvious changes in the shell.
We need not, then, bend over backward in denying
validity to comparisons of rates between groups, even such very
disparate groups as molluscs and mammals, and of course this applies all
the more strongly to comparisons between groups that are similar, such as
different sorts of molluscs or of mammals. There is, too, another sort of
comparison that is really very little affected by this particular
difficulty. The rate distributions can be compared in any two groups
without regard for the absolute values of the rates. It was seen in
Chapter II that survivorship curves are quite comparable even when the
absolute rates involved are altogether different. There is little reason
to doubt that both the resemblances and differences in shapes of such
curves can be significant facts that are independent of our ability (or
inability) to equate individual rates in the groups compared.
Survivorship tends to be negatively correlated with
rate of evolution. By and large, the longer a genus, say, endures the
more slowly its included populations are evolving. The correlation cannot
be exactly -1.00 and it cannot be well measured, but it is a reasonably
safe approximation when we can average out fairly large bodies of data.
This provides a very convenient way to approach the problems of the
distributions of evolutionary rates. By assuming that evolutionary
rates are the reciprocals of survivorship, it is possible to calculate
from any survivorship curve (or tabular data) what percentage of the
population has evolved at any particular rate. For purposes of
comparing different distributions, it is further possible and desirable
to divide the scale of rates not in absolute but in relative terms,
e.g., in decile classes of range.
Such rate distributions for pelecypod and mammalian
land carnivore genera have been calculated from the survivorship data of
Figure 5, Chapter II, and are graphically presented in Figure 39. For
reasons that will become clear, these rate distributions are based on
extinct genera, only. The two curves have a characteristic difference:
the carnivore mode is more strongly peaked and classes just below the
mode are correspondingly lower. Nevertheless the two are strikingly
similar in these respects: (1) both have a single mode from which
frequencies drop in both directions; (2) both are more peaked
(leptokurtic) than a normal curve; (3) both are strongly asymmetrical
with the mode much nearer the upper (faster rate) end of the
distribution (negatively skewed). It happens also that in both the
mode is in the ninth decile class, and that the amount of skew is
comparable in the two.
FIGURE 39.
FREQUENCY DISTRUBUTIONS OF
RATES OF EVOLUTION IN
GENERA OF PELECYPODS AND LAND
CARNIVORES. Histograms based on survivorship
of extinct genera (see Figs. 5-6) and on the postulate of—1.00
correlation of survivorship and rates of evolution. Ranges (abscissal
scale) divided into deciles to produce histograms strictly comparable
in form although absolute rates are very different. Normal curves equal
in area to the histograms drawn for comparison.
Rate distributions of this sort have so far been
made for only a few groups, although the number has increased since
1944 when these were first published. Those that have been studied all
agree in the general characteristics (1)-(3), above. It is probable
that these are generalizations true of most distributions of
evolutionary rates. Distributions may however, differ markedly in
degree of leptokurtosis (as do the pelecypod and carnivore generic
distributions to some extent) and of negative skew (as these do not
to any marked degree). Figure 40 shows a distribution that is near
the extreme in both respects, rates of evolution in the 1,374 extinct
species of centric diatoms, calculated from raw data published by Small
(1946). Kurtosis is very great, with no less than 73 percent of the
total frequency in a single decile class as against about 26 percent in
each of the two middle decile classes of the normal curve. Skewness is
also extreme, with the mode in the tenth decile class (it is, of course,
at the fifth decile, between fifth and sixth decile classes, in the normal
curve). If decile classes, only, are shown, such a curve seems to be
J-shaped and an exception to (1), above, but the frequency does fall off
to the right of the mode, as suggested schematically at the top of the
figure. Finer subdivision would show that the mode is near the ninth
decile and that frequency does fall rapidly from it to zero at the tenth
decile. There is also some slight irregularity in the dropping off of
frequencies to the left (in the fourth and sixth decile classes), but
this is evidently mere sampling fluctuation here where frequencies are
very low. The distribution is of the same type as those of Figure 39,
but more extreme.
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FIGURE 40.
EVOLUTIONARY RATES IN
EXTINT SPECIES OF
CENTRIC DIATOMS. (Calculated from
tabulation of raw survival data compiled by Small, 1946.)
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The peakedness of these distributions shows that among
related lineages, most of them tend to evolve at about the same rate. In
extinct pelecypod genera about a third, in extinct carnivore genera about
half, and in extinct centric diatom species about three-fourths evolved
at rates included in only one-tenth of the whole range of variation in
rates. These concrete data confirm the vague and subjective estimation of
many paleontologists that each group of organisms does have a
characteristic rate of evolution approximated by most lineages in the
group.
The asymmetry of the distributions shows that the
characteristic (modal) rate is decidedly nearer the maximum for the group
than the minimum. Rates above the modal class do not extend far beyond it
(are not much faster) and are, moreover, exhibited by relatively few
lineages. In pelecypods and carnivores only 9 to 10 percent of the genera
had rates faster than the modal decile class; none of the diatom species
have rates above the modal decile class. Rates below the mode are more
numerous than those above and also extend farther (are much slower relative
to the mode than those above the mode are faster). In pelecypods more than
half the rates are below the modal decile class, in carnivores more than a
third are, and in centric diatoms more than a quarter. Although it was less
clearly noticed before rate distributions were objectively calculated, this
tendency, too, agrees with opinions occasionally expressed on the basis of
subjective impression by earlier paleontologists (e.g., Matthew, 1914).
These distributions, based on taxonomic units that had
run their full span and had become extinct, show that there is a definite
range and pattern of such rates. The pattern is similar in different groups
(so far as yet demonstrated), but it also has characteristic peculiarities
in each. It gives a picture of a sort of evolutionary metabolism, one might
say, for each group as a whole. (Of course I do not mean that it is an
internal thing, like individual metabolism; it reflects the evolution of
adaptation and the turnover of taxonomic units in the course of this.) It
is this distribution that I have called "horotelic" (Simpson, 1844a).
Organisms involved in it have horotelic rates, and evolution at rates so
distributed is horotely.[1]
Further study of a number of groups, including the
pelecypods and diatoms, shows that some of their lineages have evolved at
rates much slower than any in the horotelic distribution as calculated
from fossil records of extinct taxonomic units, only. Further studies show
that these extremely low rates are part of a statistical excess of low rates
in general in comparison with the horotelic distribution. This low-rate,
nonnhorotelic excess is bradytely and the rates involved in it are bradytelic.
Other considerations strongly suggest that some phases of evolution have also
involved rates higher than any in distributions obtained in this way. That
exceptionally fast evolution is tachytely and it moves at tachytelic rates.
Bradytely and tachytely are to be separately discussed in
the remainder of this chapter. Before their detailed consideration, however,
it may be pointed out that, although these concepts are derived mainly from
analysis of the fossil record, it is sometimes possible to apply them with
reasonable probability to groups inadequately or not known as fossils. Thus
Ross (1951) has made a careful analysis of a tribe of caddis flies
(Hydrobiosini) and its probable zoogeographic history. He adduces cogent
evidence that generic splitting of the group began in the late Cretaceous
and that of its 13 genera: (a) one has hardly changed at all since that time;
(b) eleven have changed conspicuously and to varying degrees without becoming
extremely different, and (c) one has undergone extreme change and
reorientation of characters. His reasonable conclusion is that (a) is
bradytelic, (b) represent together the horotelic distribution, and (c) is
tachytelic (or it might be better to say, its lineage has been tachytelic at
some time since the late Cretaceous-persistence of tachytely for so long a
period is unlikely).
The Phenomenon of Bradytely
Evolution occurs at a great variety of rates and the lower
limit of rate is obviously zero. In view of the constant flux of the
environment and constant mutation and genetic recombination in organisms, a
sustained rate of precisely zero would hardly be expected. It would, however,
be expected that rate distributions like those of Figures 39 and 40 would
tail off to the left to a point not far from zero and would include a few,
rare lineages that had not changed appreciably for long periods of time or
had fluctuated without appreciable additive effects. Such lineages do occur,
as has long been recognized. They are of great interest and the extreme
slowness of their evolution calls for explanation.
Further analysis of rate distributions has, however,
revealed that there is something more to explain than merely the slow rate of
an occasional lineage in the tail of a falling distribution curve. In certain
groups, although not in all, it turns out that rates near zero are actually
more frequent than are slow rates in horotelic distributions calculated as
above. It also appears that there may be a discontinuity in survivorship. In
some groups, if the history of lineages arising at a given time is followed,
their extinction rises to an early peak and then tapers off gradually, as
expected; but there finally comes a time when no more extinctions occur over
geologically long periods of time. This point should be reached when the
number of existing lineages drops to zero. But in some cases lineages do
continue to survive; their number has not reached zero. Survivorship then has
not dropped gradually to zero but reached a definite point in the history of
each group where a discontinuity in survivorship occurred.
FIGURE 41.
FREQUENCY DISTRUBUTIONS OF
RATES OF EVOLUTION IN
GENERA OF PELECYPODS AND LAND
CARNIVORES. Histograms based on survivorship
of extinct genera (see Figs. 5-6) and on the postulate of—1.00
correlation of survivorship and rates of evolution. Ranges (abscissal
scale) divided into deciles to produce histograms strictly comparable
in form although absolute rates are very different. Normal curves equal
in area to the histograms drawn for comparison.
This odd phenomenon appears and can be analyzed in various
ways. For instance, of the 54 broad genera of pelecypods first appearing in the
Devonian, 30 became extinct in that period, 10 in the next, and so on down to
1 in the Jurassic. None became extinct thereafter, but 4 survive today, and
will for some unmeasurable time longer (see Table 10 in Chapter II). Even when
such a gap does not appear in our often rather crude data, an effect of what is
surely the same phenomenon is seen in discrepancies between rate distributions
calculated in different ways.
If the rates of evolution of living genera are distributed
in the same way as rates of extinct genera, then a cumulative curve of ages of
recent genera will be exactly the same as a survivorship curve for extinct
genera, aside from fluctuations of sampling. The examples of such curves given
in Chapter II showed that this definitely is not true for curves based on
pelecypod genera. Schindewolf (195Gb), although critical of those pelecypod
data (and with some reason), obtained the same sort of discrepancy with better
data on prosobranch gastropods. Lest it be thought that this result depends on
use of genera or is limited to molluscs, it is noted also that calculation of
the same curves for Small's species of centric diatoms produces the same sort of
discrepancy in still greater degree (Fig. 41).
The discrepancy arises from the fact that more genera or
species have survived from remote times in the past than would be expected
from the horotelic rate distribution for extinct genera. In other words, there
have been more lineages with rates near zero than that sort of distribution
indicates. The same discrepancy is clearly shown by comparison of actual age
composition of the recent fauna with expectation on the basis of survivorship
among extinct forms (Table 24, here, also Table 12 in Chapter II).
| Time of Appearance (in millions of years previous to present) | |
A. Percentage of Expected Survival from Survivorship in Extinct Genera | |
B. Percentage of Realized Survival | |
Difference B—A |
| 25 | |
92 | |
78 | |
–14 |
| 50 | |
63 | |
62 | |
–1 |
| 100 | |
24 | |
39 | |
15 |
| 150 | |
8 | |
30 | |
22 |
| 200 | |
2 | |
24 | |
22 |
| 250 | |
1 | |
19 | |
18 |
| 300 | |
0 | |
13 | |
13 |
| 350 | |
0 | |
8 | |
8 |
| 400 | |
0 | |
4 | |
4 |
| 450 | |
0 | |
0 | |
0 |
In carnivores actual survival is somewhat less than
expectation throughout (Table 12), but this has affected genera of all
ages in about the same proportion. As a result, the recent carnivore
fauna is less varied than would have been expected from the Tertiary
record, but its age composition, as proportions of genera surviving from
various times in the past, is almost exactly as expected (Fig. 42). Rate
of extinction was higher than expectation from late Pliocene to Recent,
but it did not tend to affect later genera more than earlier, or earlier
more than later. Moreover no living genus is older than the maximum span for
extinct genera. In this group the discrepancy under consideration docs not
occur, and the inference is that there are no bradytelic carnivores.
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FIGURE 42.
REALIZED AND EXPECTED
AGE COMPOSITIONS OF
RECENT PELECYPOD AND
LAND CARNIVORE FAUNAS.
Percentage of living genera arising (first appearing in fossil
record) in stated epochs and periods as observed in the recent fauna
(counting only genera that are known as fossils) and as would have
been predicted on the basis of survivorship in extinct genera.
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In pelecypods and gastropods, on the other hand, actual
survival is greater than expected survival (Table 12, Chapter II) and
these groups are today much richer, more varied, than expected on the
basis of their horotelic distribution. Survival of late genera, those
arising in the Tertiary, is near expectation: a little low for pelecypods
and high for gastropods but in both cases as near as would be probable
with sampling effects. Survival of pre-Tertiary genera, from the
Ordovician for the very broad pelecypod genera and from the Triassic for
the narrower gastropod genera, is, on the other hand, much greater than
horotelic expectation. The result in terms of percentage composition of the
recent fauna is that there is a much smaller proportion (not absolute
number) of young genera and a much higher proportion of older genera than
the expectation, as is well shown in Figure 42 for pelecypods.
Moreover, even though their spans have not yet ended,
some living genera have already had much longer spans than any extinct
genera of the same group. Some broadly defined living genera of pelecypods
have been in existence for upward of 400 million years, but the
longest-lived of extinct genera, defined equally broadly, endured no more
than 275 million years. In the centric diatoms, there are living species on
the order of 75 million years of age but no extinct species more than about
50 million years old. It is such facts that indicate a discontinuity in
survivorship. On the other hand, in the prosobranch gastropods
(Schinndewolf's data as previously noted) the oldest living genera are
about 200 million years old and there is at least one extinct genus that
endured for 250 million years or somewhat more. This shows that there are
complicating factors,[2] but does not alter the fact
that this group, too, shows clear-cut excess of survivorship of old genera
in the recent fauna.
The phenomenon here noted has of course long been
discussed in a more general way or in other terms and techniques. T. H.
Huxley (e.g., 1870) pointed out as early as 1862 that certain types of life
are unusually "persistent." The phenomenon of "arrested evolution"
(Ruedemann, 1918) has long engaged the attention of paleontologists. Their
earlier analysis was, however, even more unsatisfactory than at present
because the usual criterion of "persistence" was quite arbitrary and it
was not shown whether the rates of evolution involved were really slow for
the particular group in which they appeared. Thus Ruedemann considered as
"persistent" and cases of "arrested evolution" any genera, from protistan
to reptilian (there are no avian or mammalian examples) that appeared in
more than two geological periods, in spite of the fact that this might mean
survival for anything from 30 million years upward and that in some groups
survival of broadly defined genera for such spans is the rule rather than
the exception.
Rensch (1947) has demonstrated that the mean age of
living higher categories (families to orders) is generally greater than the
mean span of extinct units of the same categorical rank and in the same
group, often two or three times as great. Since the long persistence of a
family or an order does not preclude a fairly high rate of evolution within
the group, these examples do not necessarily represent essential arrest of
evolution, but the latter phenomenon is probably involved in such averages.
It is quite certainly involved in the elaborate series of studies by Small
(e.g., 1946), begun in the 1920's and still continuing, on "short" and
"long" species, mostly in diatoms. Diatoms do show an unusually high degree
of bradytely, as demonstrated above by a different analysis of some of Small's
data, and Small seems to be quite right in concluding that two sorts of
species, different in some quantitative way as regards rate of evolution,
can be distinguished among these organisms.[3]
Thus there is nothing really new in the observation of
what I have called "bradytely" (in print since 1944), but it does seem that
rate distribution and survivorship methods put its study on a more nearly
objective basis and permit clearer analysis of it.
FIGURE 43.
BRADYTELY IN PELECYPODS. Survivorship in
extinct genera, continuous, unshaded curve, and analogous curve based on
recent genera, broken curve, see Figure 5. The shaded area represents the
excess of realized over expected survivorship caused by the nonextinction
of bradytelic phyla and so is approximately proportional to the rise of
bradytelic lines at the stated times in the past. (The apparent absence
of bradytelic lines arising in the last 50,000,000 years may be real, but
is probably an artifact caused by differential extinction of horotelic
genera in this period, the extent of which cannot be estimated from the
available data.)
A next step is suggested in Figure 43. In this particular
group, the pelecypods, it appears that any living genus more than about 250
million years old, i.e., dating from about the Mississippian or earlier, is
almost surely bradytelic. Such genera include Nucula, Nuculana
(or Leda), Volsella (or Modiolus), and Pteria (or
Avicula).[4] This is sufficiently clear and
these individual genera can be labeled "bradytelic" on a reasonably objective
basis. But the situation in, say, the Jurassic, around 150 million years ago,
is different. The curves and tabular survivorship data give good reason to
believe that perhaps three-fourths of the genera that survive from Jurassic
to Recent and perhaps about one-fourth of the genera that arose in the
Jurassic are bradytelic. But this is a statistical result, and like many such
results it cannot be directly applied to the individual. There is no possible
way from these data to tell which of the genera first appearing in the Jurassic
are bradytelic and which are horotelic in the terms of this analysis. Such
designation could only be made if some separate and individual criterion of
bradytely could be found. If some Jurassic-Recent genera were found not to
have changed at all and others merely to have changed very slowly, the former
would include the bradytelic genera, but such a difference cannot be very
clear-cut among lineages all of which have changed so little as not to warrant
calling the change generic in broad, conservative classification. Presumably
some other features might particularize bradytelic genera but, as will appear
in the next section, such criteria really diagnostic as to individual genera
are not available. It is, indeed, likely that the statistical result does not
depend on any inherent differences uniform in each individual line.
It thus appears that some of the oldest genera or species
in some groups and after analysis of their rate distributions can be
identified as bradytelic. Among younger forms, however, the analysis may
clearly show that bradytelic lines were still arising and yet give no way of
designating them individually. In other groups, as among prosobranch
gastropods, the analysis indicates that bradytelic lines are present, but
none can be individually designated, even among the oldest, because the same
analysis shows that none is older than some horotelic lines. We can only go
so far as to say that lines that have shown no appreciable change while most
allied lines did show change must include the braclytelic lines, perhaps
along with some that are slow horotelic, and may be more likely bradytelic
than not.
Since this concept of bradytely was first published, many
of those who have discussed it either have assumed that no distinction was
intended between bradytely and very slow horotely or have denied that such a
distinction exists. Most of these students have overlooked, or at least have
made no reference to, the fact that bradytely is defined by a difference
between rate distributions, a difference that does objectively exist. (There
are, of course, noteworthy exceptions, e.g., Schindewolf, Carter.) It seems
pointless to discuss whether a difference exists, unless it can be shown to
be an artifact of the method, which has not been demonstrated or even
suggested.[5] It is also somewhat beside the point to
designate some particular slowly evolving line as bradytelic, to show that
its evolution had slowed down from a more usual, presumably horotelic line,
and to deny any distinction between the two on that basis. The impossibility
under some circumstances of designating particular lines as bradytelic has
been pointed out above. It is also quite clear that all bradytelic lines are
derived ultimately from horotelic or often from tachytelic ancestors. It is
not to be assumed that they became abruptly bradytelic, although that may
sometimes occur. Discontinuity in that sense, in application to change of
rate within one line, is not involved in the evidence for or concept of
bradytely.
It is very much to the point, however, to inquire what the
difference between bradytely and horotely is, whether, for instance, it is
quantitative and statistical, with intergradation and yet with mean
differences and possible discontinuity of average outcome, or whether it is
qualitative, with inherent discontinuity of sort and of underlying factors.
Of these two alternatives, the evidence strongly suggests that the former is
correct. In this sense, I agree with, for instance, Westoll (1949)
that bradytely and very slow horotely are indistinguishable as seen in
individual instances, but it remains true that in certain groups the
prevalence of such exceedingly slow rates has produced a statistical
discontinuity in rate distributions which is a collective phenomenon
distinguishable from collective horotely. An opposing view is taken by Small
(1946 and numerous other papers in the same series), who thinks his "shorts"
and "longs" are inherently different as applied to individual cases, the short
or long duration being "very largely determined by what happened to one
nucleus" when the species arose by saltation. Extensive and conclusive
evidence against such a view is given throughout this book, and the
phenomenon of bradytely neither demands nor warrants the postulate of
predetermined life span or any other factors than those normally involved in
the complex evolution of adaptation.
Factors of Slow or Arrested Evolution
Bradytely is essentially a statistical effect produced by
the prevalence in some groups of organisms of lines with extremely low rates
of evolution, or with changes fluctuating on a small scale and not
appreciably cumulative. The first point, then, is to consider the
circumstances of very slow or arrested evolution in general.
Certain genetic and other factors in organisms set limits
to rates of evolution. It is therefore natural to suppose that slow evolution
is primarily conditioned by these factors, and this has often been suggested.
Nevertheless there is much evidence: (a) that the limits set by these factors
in real populations are generally permissive of average or even high rates,
and (b) that very slowly evolving lines are not particularly likely to have
these factors in unusual degree.
Low mutation rate is one such factor often suggested (e.g.,
Dobzhansky, 1941, but
the suggestion was practically withdrawn in 1951). But what appear to be low
rates are sufficient to account for sustained and even rapid evolution (see
Chapter IV). Mutation rates have not been directly determined in many slowly
evolving lines, but the evidence tends to oppose the postulate that they have
low rates. Drosophila, itself, may now be a very slowly evolving group,
but seems to have rates of mutation comparable with those of any other group
studied in this respect. The facts that very slowly evolving groups often
arise by rapid evolution, that they also often split to give rise to rapidly
changing branches, and that they do not have particularly low variation
(below) all suggest indirectly that their mutation rates are more or less
normal. Stebbins
(1949, 1950) points out that slowly evolving plants may be rich in genetic
variability, speciating luxuriantly and (e.g., Sequoiadendron)
readily yielding horticultural varieties.
Factors that reduce genetic variability, notably apomixis,
self-fertilization, and reduction of crossing over, certainly limit the
possibilities for change and therefore may characterize groups that change but
little ("blind alley" groups, Chapter IX). In the longer run, however, such
groups generally do finally change or become unusually liable to extinction.
Limitation of variation may, indeed, be a factor in bradytely of such groups
as diatoms, but even in such cases it is all insufficient explanation and it
does not apply at all to the majority of slowly evolving animals and plants.
Most of these, even in lines surely bradytelic and with essentially no
progressive evolution over very long spans, are sexually reproducing and show
usual or even high degrees of variability. Opossums, which have evolved more
slowly since the Cretaceous than any other living mammals, have quite as
variable populations as the average of rapidly evolving mammals. Within the
established adaptive type they also have undergone repeated, wide speciation,
which implies a considerable store of variability. Stebbins (1949) also
notes that slowly evolving plants may speciate freely. Selaginella,
one of the most slowly evolving of all plants (little changed from
Selaginellites in the early Mississippian) has over two hundred
living species (Manton, 1950).
Long life spans or long generations, also frequently
suggested as correlated with slow evolution, may be ruled out even more
conclusively. As noted in Chapter V, there is surprisingly little evident
correlation between lengths of generations and rate of evolution. Here it
may be added that most of the outstanding examples of slow or arrested
evolution occur among organisms with relatively short generations. Bradytelic
lines do not seem to have generations consistently different in length from
their horotelic and tachytelic allies, and again it is pertinent that
bradytelic lines may arise from and give rise to tachytelic lines. Surely
change of length of generations is not a significant factor.
Another common approach, exemplified by Ruedemann
(1918, 1922a, 19220), who devoted much attention to this subject a
generation ago, is to seek particular morphological and physiological
characteristics associated with low-rate lines. Small size, nocturnal
habits, and carrion-feeding are among the many suggestions of this sort
that have been made. Some such characters may have a bearing (although it
is not likely that those just mentioned have much if any), but obviously
they are not really explanatory. Among related lines fully comparable in
such respects, some may evolve very slowly and others rapidly. Groups
almost wholly different in characters of this sort, such as some plants
and some animals, may have very similar slow—or fast—rates of
evolution. It is perhaps true that sessile animals have evolved more
slowly on an average than mobile animals, but this is a slender and
probably misleading clue in view of the facts that similarly sessile
animals have evolved at very different rates and that most plants are
sessile but that they have run as broad a gamut of rates as have
animals.
In such comparisons as that of sessile and mobile
animals a certain confusion may arise. It is one problem that whole groups
(say, corals) may have an average rate that is lower than that of another
group (say, fishes) and quite a different problem that one line (among
corals or among fishes) may have a very much lower rate than another
allied and similar line. Both problems call for study, and they are not
wholly unrelated, but it is stultifying to confuse the two. The frequently
repeated generalization that primitive organisms have evolved more slowly
than advanced ones lends itself to that confusion and also involves a
howlingly circular argument as applied to groups now living: we call some
of them "more primitive" precisely because they have evolved more slowly.
If the statement is applied more correctly to older, hence literally more
primitive, and younger members of the same group, it simply is not true.
There is no definite tendency for average rates within given groups to
accelerate. Changes in such averages are highly variable and if there is
an over-all balance it may be on the side of deceleration of rates. For
instance, the later lungfishes evolved much more slowly than the primitive
members of the group (Chapter II, and Westoll, 1949), and this seems to be
a rather common pattern in long-lived groups.
As applied to individual lines, the idea that more
primitive forms evolve more slowly is also a misconception. It is an
interesting fact that bradytelic lines, as far as identifiable as such, or
any extremely slowly evolving lines are usually seen to have been
progressive, relatively advanced types up to the time when their evolution
was arrested. It is their long persistence without much change after their
allies have gone on to explore other avenues of modification that gives them
an archaic aspect or makes them primitive in comparison with contemporaries
of later phases of their history. Lingulids had a fairly high type of
invertebrate structure for the Ordovician and limulids were advanced for the
Triassic. Sphenodonts were about as advanced as any reptiles in the Triassic
and so were crocodiles in the Cretaceous. Lingula, Limulus,
Sphenodon, and Crocodylus are primitive as of now just because
they are (with little question in these cases) bradytelic and have been for
long periods. But such groups have been passed in the race; they did not have
an initial handicap. Stebbins (1949) has pointed out that the same
generalization is true of plants: the lines with arrested evolution were
specialized and advanced when their evolution was arrested.
Another approach has been to designate certain places or
broad environmental situations as conducive to slow evolution: oceans,
islands, tropical forests, and so on. It is probably correct that sustained,
slow, or arrested evolution is more likely to be found in some such
environments than others. (Islands happen to be one of the places where it
is less likely, contrary to some statements, see Chapter IX.) But the
approach to the problem is still obviously inadequate, for very rapid
evolution may occur in exactly the same situations (many marine groups,
epiphytes in tropical forests, etc.).
It is, in fact, clear once more that the key to this
evolutionary problem is not in the organisms or in their environments but
in the relationship between the two. When lines persist with arrested
evolution, they have maintained adaptation without appreciable change. For
this to occur, there must be a balance between breadth of adaptive zone and
its variation or deviation in time. If organisms are very broadly adapted,
if they can, for instance, live on a variety of foods or grow on a variety
of soils and in a variety of climates, environmental factors may fluctuate
greatly without requiring any adaptive change for survival, as long as some
part of the broad zone persists. If adaptation is narrow, the zone must be
a very uniform and stable one in space and time for evolution in it to be
arrested and its occupying populations to persist.
Arrested evolution may thus occur in a great variety of
environments and organisms and with quite different sorts of adaptation and
specialization. It is, however, more likely to occur with broad adaptation
than with narrow, because adaptation in a broad zone is more likely to persist
than in a narrow one. Both broad and narrow may persist, however. The lungfish
zone would seem to be quite narrow, and yet it has persisted without marked
change since the Devonian and all lungfishes have apparently been bradytelic
in it for perhaps 200 million years. It may well be that if a major zone
("major" in being quite distinctive from all others) is rather narrow, only
bradytelic groups will eventually be left in it. On the other hand, the zone
of the opossums is quite wide, and the opossums have survived repeated and
radical environmental change without themselves changing much.
Evolutionary change is so nearly the universal rule that a
state of motion is, figuratively, normal in evolving populations. The state
of rest, as in bradytely, is the exception and it seems that some restraint
or force must be required to maintain it. This force undoubtedly exists and
is the same as the force that usually orients evolutionary change: selection.
Selection on groups with arrested evolution must be centripetal and must
be more effective than mutation, cyclic or fluctuating environmental change,
or any other factor making for change. Whether adaptation is broad, as it
usually is, or narrow, such a group must be perfectly adapted to its zone
and almost any change must be quickly detrimental and effectively checked by
selection.
The most slowly evolving groups do seem all to be very
highly and specifically adapted to a particular zone. Their typical history
is one of rather rapid shift into a new, stable and persistent zone, still
rapid adjustment (postadaptation) in the zone, with weeding out of less
adaptive characters and lineages, and then a long, eventless course of
relatively unchanging continuation. That they are actually held by force, so
to speak, is confirmed by the fact that many of them (not all) have given off
branches that rapidly evolved into different zones when the opportunity, or
perhaps it is better to say the occasion, arose. Thus bradytelic Ostrea
(sensu lato) repeatedly gave rise to relatively very rapidly evolving
coiled (Gryphaea) and spiral (Exogyra) lineages, among others. Opossums gave
rise rapidly and early to a great variety of divergent groups in South
America: Caroloameghiniidae, Borhyaenidae, Polydolopidae, Caenolestidae,
each family except the first (as far as known) highly varied. Typically these
rapid side-branches, or most of them, become extinct while the parent group
continues unchanged.
With this background, let us turn back briefly to the
characteristics of the populations and environments involved. As was found
true of extinction, the balance maintained here, lost in extinction, is
compounded of such extremely numerous, varied, and complex factors That it
may he quite impossible to designate them explicitly in a specific instance.
For the populations, the prime essentials are that these be extremely well
integrated genetically and that any deviation be subject to effective
counter-selection. It might be expected that such groups would have
especially elaborate systems of modifiers or polygenes, but as far as I
know no concrete comparisons pertinent to this relationship have been made
or are yet possible.
If selection, and apparently quite feeble selection in
some cases, is to be effective, gene drift must be excluded. This requires
population above a minimal size, but that size need not be very large in
all cases. Westoll (1949) has given some reason to believe that early
lungfishes, evolving rapidly, had larger populations than the bradytelic
living forms, but the population size in the latter is apparently quite
large enough to enable selection to hold them in a narrow zone—the
fact that the zone is narrow probably means that centripetal selection is
relatively strong and so is still effective in relatively small populations.
Most groups with arrested evolution have large, even extremely large,
breeding populations. Outstanding exceptions seem to be Sphenodon and
Latimeria, but Sphenodon apparently had large populations
until human disturbance banished it from the mainland of New Zealand and no
one has any idea how large a breeding population Latimeria may
have.[6]
As Stebbins (1949, 1950) has emphasized for plants,
there is no reason to think that large population size in itself promotes
slow evolution. It makes for more effective selection, and also within
limits for longer duration, and hence is propitious for arrested evolution
when selection is centripetal. That selection may tend not toward change
but toward stabilization is a thesis developed with skill and at great
length by Schmalhausen,[7] whose work cannot be
summarized here (but see also brief comment in Chapter VI). Others, e.g.,
Heuts (1949), have also given specific examples of mechanisms developed by
selection that inhibit rather than promote change. One additional point
that might be mentioned is that intraspecific selection must also be
centripetal if evolution is to be arrested while a lineage continues.
The group must be so organized as to minimize genetically effective
competition within it, or always to favor the modal type, or both. Strong
dimorphism in secondary sexual characters would not be expected in such a
group among animals and in fact does not occur in any example known to
me.
As to the environment, the principal requirement is
that it persist and do so with less fluctuation than the maximum that can
be met by existing and expressed, not potential, variation in the
populations conncerned. How much fluctuation may be so met depends of
course on the amount and sort of expressed variation and the details of
the adapptation involved but, in general and in terms of the environment
itself, the more stringent it is in requirements for life, the less
variation can occur without requiring evolutionary change or causing
extinction. Arrested evolution is therefore less likely to occur in very
difficult environments, such as deserts, impermanent environments, such
as salt lakes, or highly variable environments, such as the alpine zone.
It is more likely to occur (although of course rapid evolution may also
occur) in the easier, more permanent, and less variable environments such
as the ocean, its strand, the shifting but long enduring major lowland
rivers, the more slowly shifting and also long enduring great forest
belts, and particularly in the most nearly permanent of climatic zones,
subtropical to tropical.
It now seems to me possible to explain the phenomenon
of bradytely in general terms and in the light of these factors of slow
or arrested evolution. Any series of related lineages or even one lineage
in the course of its history is subjected to a great many environmental
changes and to many changes in its own populations. Some of these changes
are concomitant with or in a sense produce progressive adaptive changes
in the populations. Others shunt development off into another directions,
or move it into a different adaptive zone. Still others lead to loss of
adaptation, hence extinction. In the case of single lineages, in most
instances one of these three results will ensue. Sometimes, however,
early change may, in a manner of speaking, have oriented and aimed the
group down a line where adaptation is retained indefinitely without
further change.
In terms of multiple lineages the process might be
visualized as movement of the various sorts of organisms down a series
of corridors with many baffles. Most corridors turn, forcing the
organisms to turn, too (evolve new adaptation). In a straight corridor
the baffles (changes in environment and population) deflect some into
side corridors (new adaptive zones) and stop some literally dead
(extinct). After a certain time (a long time) all those seriously
liable to deflection or stopping have been weeded out. There remain
only those sorts of organisms that have run through the whole repertory
of baffles and are no longer subject to deflection or stopping.
Suddenly at the last baffle that does stop or deflect, there is a
discontinuity of outcome. The remaining sorts of organisms now
persist indefinitely.
The statistical result actually observed would arise
from such a situation. It is seen, too, that one of the survivors, the
bradytelic lines, need not be evolving more slowly than some of the
others, the horotelic lines, and might have no characters that would
reveal its bradytelic nature, before the final event, the disappearance
of the last horotelic line of the same age as the bradytelic line. The
distinction of the bradytelic line is an adaptive relationship that
makes it miss the baffles, that permits indefinite survival without
change in the face of all vicissitudes that do eventuate in its
adaptive zone. Bradytelic lines are merely the residuum of a process
that regularly reduces the percentage of unchanged groups but that stops
short of reduction to zero, if bradytely does occur.
The baffles are so numerous in nature that one would
expect bradytely to be in general less common than horotely and to be
absent altogether in some groups. That expectation is confirmed by
observations, such as the examples discussed earlier in this
chapter.
Tachytely
It is my opinion that tachytely is a usual element in
the origin of higher categories and that it helps to explain systematic
deficiencies of the paleontological record. It will therefore be
discussed in some detail in the following chapter, which is devoted to
those topics. At this point, the need is only to try to clarify the
relationships of tachytely to horotely and bradytely (a subject that I
personally find both abstruse and difficult but highly significant and
enlightening as regards major evolutionary processes).
It is in the nature of the method that the study of
survivorship cannot reveal whether there is also a nonstandard
distribution of rates generally faster than horotelic, analogous to the
nonstandard distribution of low rates that it does reveal. Evolution at
exceptionally high rates cannot long endure. A tachytelic line must soon
become horotelic, bradytelic, or extinct. For this and other reasons,
which will later be suggested, tachytelic lines, or tachytelic phases in
the evolution of lines horotelic or bradytelic at other times, are poorly
recorded by fossils and often not recorded at all. There are occasional
glimpses in the form of fossils that happen to be found at some point in
the tachytelic phase (e.g., the Devonian amphibians from Greenland) and
some complete sequences of a rather marginal sort, but for the most part
a given tachytelic phase is an inference from ancestral and descendant
forms in a record with important gaps. Often the inference is perfectly
clear and its correctness is confirmed by the scattered records that do
show it directly, but the incidence of tachytely cannot be counted or
calculated, even on a sampling basis, as can that of bradytely.
What distinguishes tachytely from horotely is not
only that it occurs at exceptionally fast rates but also that it occurs
while populations are shifting from one major adaptive zone to another,
and especially when a threshold is crossed (see Chapter VI). Such a
phenomenon differs from horotely only quantitatively and not sharply.
It grades into horotely both in the sense that a tachytelic line
gradually becomes horotelic and in the sense that the differences in
degree of adaptive change involved, in rate of evolution reached, and
in clarity of threshold are all on continuous scales. There is no sharp
point where one can say, "Below this is horotely and above it is
tachytely." Nevertheless there is a difference that may be very
appreciable and significant in less marginal cases.
Well-recorded but marginal instances occur in such
sequences as that of the Equidae (Chapter VIII). The changeover from
browsing to grazing has a threshold effect and involves a definite,
temporary increase in evolutionary rate. This verges, at least, on
tachytely. The even more rapid changes from one stable type of foot
mechanism to another are still more clear-cut examples of this sort of
phenomenon and might well be designated as rather small-scale tachytely.
Another well-recorded example of small-scale tachytely, leading to
extinction in this case as tachytely often does, is that of
Gryphaea (Chapter IX). One of the most remarkable known examples
is that of the snail Valenciennesia, formerly classified in
another suborder than the notably different Limnaea, but shown
by Gorjanovic-Kramberger (190l, 1923) to have evolved from the latter
so rapidly that the whole process occurred while a horse, Hipparion
gracile, on adjacent lands showed no appreciable change. Here,
too, change to a distinctly different adaptive zone is involved: from
clear and fresh to muddy and brackish water.[8]
As Stebbins (H)50) has suggested, tachytelic lines
are likely to be, although they are not necessarily, narrowly adapted.
They include the forms that are forced into rapid change by changes in
their adaptive zones. Their adaptive evolution tends to be the opposite
of that seen in bradytely, and the factors involved tend to be inverse.
In the figure of speech utilized above, tachytelic lines hit the baffles
and are deflected into sharply divergent corridors.
Tachytely is defined as a phylogenetic phenomenon
and is to be distinguished from rapid speciation or splitting. Rapid
evolution in the sense of high origination rates, such as occur on
invasion of new and open habitats (e.g., in the Galápagos or Hawaiian
Islands, as discussed before, or rapid development of endemics in
African lakes, see Worthington, 1937, 1940) is a phenomenon quite
distinct from tachytely. Nevertheless, when adaptive radiation occurs,
and especially if this is of a basic or major sort, the shift of each
divergent line into the zone it comes to occupy is commonly
tachytelic.
FIGURE 44.
DIAGRAMMATIC REPRESENTATION OF
CASES OF HOROTELY, BRADYTELY,
TACHYTELY. This particular sequence, one of many that
occur, shows a bradytelic group from which is split off a tachytelic
branch which in turn gives rise to a horotelic group.
Horotely represents a sort of normal or average
turnover or metabolism in the evolution of a group of organisms. To
the extent that evolution is defined as change, bradytely is a
cessation of evolution without extinction. Tachytely is an episodic
acceleration, figuratively a brief fever imposed on the normal
metabolism. Tachytelic lines arise from horotelic or bradytelic lines,
in the latter case usually (perhaps always) by splitting. A tachytelic
line must soon become horotelic, bradytelic, or extinct. Horotelic
lines usually arise from other horotelic lines, occasionally from
tachytelic lines (but these occasional events are unusually important)
or from bradytelic lines, in the latter case by splitting and usually
through a tachytelic phase. Horotelic lines may remain horotelic, or
become bradytelic or tachytelic. Bradytelic lines develop from
horotelic or often rather directly from tachytelic lines. The
bradytelic lines themselves remain bradytelic and rarely if ever
become horotelic or tachytelic again, but it is rather common for
them to give rise to tachytelic branches, which in turn become
horotelic or extinct. A simplified pattern of one of several possible
relationships between horotely, bradytely, and tachytely is shown
in Figure 44.
Notes
When I proposed the terms horotely, bradytely, and
tachytely, I apologized for adding to the technical terminology of evolution
and explained that I had been quite unable to discuss rates clearly without
this invention. As expected, at least one critic did object to the use of new
terms, but apparently most others agreed that they make for clarity as they
have come into rather general use.
A really detailed analysis of this group cannot
be attempted here, and should be done as a separate study by a specialist in
fossil gastropods, but the suggestion may be made that high Triassic
mortality (related to the Permian-Triassic crisis for marine vertebrates)
may in this case have caused the unusual extinction of bradytelic lines. The
five genera surviving from Ordovician to Triassic were probably bradytelic
and the three then surviving from the Silurian were possibly so. Four of the
former and two of the latter became extinct in the Triassic, and the last
Ordovician and Silurian genera disappeared in the Jurassic. Except for these
few genera, the maximum span for extinct genera is under 200 million years,
hence somewhat, at least, less than for the oldest living genera.
It is necessary to add that Small's criterion for
"shorts" and "longs" is arbitrary and does not really separate the two rate
distributions. His conclusions as to the bearing of these data on
evolutionary theory further seem to me beside the point and not really
supported by the data. These personal opinions should not be allowed to
obscure the real value of Small's laborious compilations.
Shimer and Shrock (1944), although reflecting
fairly modern nomenclature and constriction of genera, still record Nucula
and Nuclllana from the Silurian, Volsella and Pteria
from the Devonian. Of course there is some difference between the earliest
species and any living species in each of these genera, and sooner or later
someone is sure to call that difference "generic." This will obscure but
not alter the fact that the genus, as here recognized, has not shown
appreciable progressive change since the Silurian or Devonian, although it
has continued to undergo continual, noncumulaative speciation, as do all
long-continued or widespread groups.
Hutchinson (1945) did suggest an artifact of
materials rather than of method: that the bradytelic genera are merely those
in which evolution at normal rates affected the soft parts only, and so did
not appear in fossils. This seems quite impossible, for reasons already
mentioned in part. It may be added that this explanation cannot possibly
apply to the diatoms, which include much bradytely in spite of the fact that
living and extinct species are commonly based on identical criteria.
It has also been remarked (e.g., Carter, 1951)
that the monotremes and Peripatus have small populations and so do
not agree with requirements for bradytely. It is now clear that a large
population is not a requirement but only a probability. What some students
would consider a small population in subjective and relative terms may yet
be such as to make centripetal selection effective, which is the essential
point. As regards the monotremes, they are extremely specialized in many
respects. It is improbable that they are bradytelic and their evolution may
not even have been unusually slow—they merely happen to have some
reptilian characters lost in true mammals. Their breeding populations were
also moderate if not large before they felt the effects of man and his
works. Peripatus is quite likely to be bradytelic, but I know of no
estimates as to its breeding populations; forms so obscure in habitat are
likely to appear rare when not so in fact.
This is the same man occasionally cited as
"Shmalgausen," which is an English transliteration of the Russian
transliteration of his originally German name.
Basse (1938) has cast doubt on
Gorjanovic-Kramberger's interpretation, but Basse's opposing evidence is
weak.
