Biological Journal ofthe Linnean SociCp (1982), 18: 145196. With 6 figures
Bryophyte evolution and geography
H. A. MILLER
Department of Biological Sciences, Uniuersip of Central Florida, Orlando, Florida 32816,
U.S.A.
Accepted for publication November 1981
The three extant Divisions comprising the bryophytes extend, as fossils, well back into Palaeozoic time.
Bryophyte origin is part of the rise of terrestrial, vascularized, plants with sporopollenin-walled spores
in the Silurian. Before the end of Carboniferous time, bryophyte lines were widely present. Separation
of Gondwana and Laurasia by the Permian Tethys Sea and subsequent widespread desert episodes
fragmented an already diversified bryoflora subjecting it to intense selective pressure. The cool, mesic
climate ofsouthern Gondwana provided a refugium for austral bryophytes. Warmer and drier climates
of the Permo-Triassic Laurentian-Laurasia favoured drought-adapted or niche-specific groups
creating marked systematic discontinuities. The Angaran wet, probably cool, temperate region
provided refuge for basic stock for much of today’s rich holarctic and wet ‘tropical’ bryofloras. Climatic
changes, correlated with tectonic events and the rise of angiosperms, opened habitats favourable for a
diversity explosion. Despite demonstrated potential for long-distance dispersal, modem distributions
are mostly linked with total floras or establishment on islands prior to niche saturation. Remnants of
Gondwanan bryoflora persist in high southern latitudes as disjunctions with the possibility that the
folded ranges of the African Cape have been an insular fragment at higher latitudes becoming attached
shortly after angiosperm diversification.Floras ofsouthern India and east Africa have common features
but the Himalayan flora shows evidence that the Gondwanan flora of the Indian plate was lost during
the movement through desert and tropical latitudes; neotropical and palaeotropical floras are
distinctive. Much of the northern Australian bryoflora is recently Malesian-derived while the southeast shows strong austral influence and commonality with New Zealand. Tropical Pacific island floras
are mostly Malesian-derived but with both holarctic and austral elements present as in Hawaii and the
Society Islands. Holarctic bryoflora is circum-polar with temperate areas of Euro-American and far
eastern elements floristically bound by disjunct and vicariad species. Kroeber Coefficients of
Correlation difFer as Pacific island floras are compared and Guttman-Lingoes Smallest Space
Coordinates indicates floristic subgroups within Polynesia. Although these and other mathematical
treatments yield potentially promising results, the methods are yet unrefined and there is some
uncertainty whether characteristics of numbers or of organisms are implicit in the summations.
KEY WORDS:-Bryophyta
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evolution - fossils - geobotany - historical biogeography.
CONTENTS
Introduction . . . . . . . . . . . .
( I ) The bryophytic habit . . . . . . . . .
(a) Hornworts
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(b) L’iverworts . . . . . . . . . .
(c) Mosses . . . . . . . . . . .
(2) Bryophyte dispersal . . . . . . . . .
(3) The fossil record . . . . . . . . . .
(a) Pre-Permian bryophytes and not-quite bryophytes .
(b) Permian and Triassic fossils . . . . . .
(c) Jurassic and Cretaceous fossils . . . . . .
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H . A. MILLER
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(4) Bryophyte evolution and systematics
(a) Origin and macroevolution . . .
(d) Tertiary fossils
(e) Quaternary fossils .
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(b) Taxonomy. systematics and microevolution
Major features of bryophyte distribution. . . . .
(1) Antarctic Kingdom . . . . . . .
(2) Australian Kingdom . . . . . . .
(3) South African Kingdom . . . . . .
(4) Neotropical Kingdom . . . . . . .
(5) Paleotropical Kingdom . . . . . .
(6) Boreal Kingdom . . . . . . . .
Some problems and approaches for bryophyte geobotany.
Summary . . . . . . . . . . . .
Acknowledgements. . . . . . . . . .
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INTRODUCTION
The last serious attempt to embrace all the bryophytes within a single worldwide geographical treatment was Theodor Herzog’s ( 1926) now classic
‘Geographie der Moose’ completed in 1925. He used 452 pages and eight plates to
do it but not once provided a map showing the limits of the areas characterized . I
have created a map From his descriptions (Fig. 1) and maps of representative taxa
in order to get a quick overview of how the bryophytic world looked to him 55
years ago. It was a good time to undertake such a project-Max Fleischer
(1904-1923) had just updated his revisions of the systematics of mosses in a manner
that took into account the great tropical diversity, Viktor Brotherus (1924-1925)
I
...
Figure 1. World map of floristic kingdoms as extrapolated from Henog’s (1926) descriptions and
based upon bryophyte distributions.
BRYOPHYTE EVOLUTION AND GEOGRAPHY
147
had summarized with brief description at the familial and generic level (with brief
distributional notes) about all there was to know about moss taxonomy, and Franz
Stephani’s (1900-1924) monumental coverage of the liverworts of the world
together provided a stupendous data base. This is not to say that Herzog created
his book from three sources, his fine bibliography and detailed commentary
indicate the contrary, but having the Brotherus and Stephani touchstones was
helpful for consistent nomenclature and in other ways.
It is not my purpose to update Herzog’s lists of species area by area, albeit that
some of that is appropriate later on, but to place the present day diversity of
bryophytes relative to their evolutionary history in context of modem and ancient
distributions to the extent possible. In a few words, then, this is a review of the
geobotany of bryophytes as we understand it today.
Bryophytes have usually been recognized as comprising a single Division
Bryophyta containing the Classes Musci and Hepaticae, with the Anthocerotae
sometimes as third class. However, so much new information has been developed
from many sources that this simplistic, monophyletic view is untenable. Most
bryologists agree that at least three Divisions are properly recognized amongst the
bryophytes and, although not frequently so noted, that the bryophytic concept
represents a level of evolution paralleling roughly the concept as understood for
‘algae’ or ‘fungi’. Levels ofclassification above the generic are in such a state offlux
that the most useful indication of the taxonomic extent of the bryophytes can be
gained from approximate numbers of genera and species : mosses, 795 genera and
15800 species; liverworts, 340 genera and 6890 species; and hornworts, five
genera and 270 species. In recent years, the number of genera has tended to
increase while the number of species has remained more or less constant, about the
same number being described as that placed in synonymy. Many of the old *
‘geographical species’ are being placed under earlier names and new species are
emerging from remote areas of the tropics and southern hemisphere now being
bryologically explored for the first time.
Numerous previously unknown genera, and families, will probably yet be found
in South America with fewer instances from Australia through the Malesian area
and still fewer from Africa and the holarctic regions. Only western Europe, Japan
and extra-tropical North America are well enough known that most (not all) alpha
taxonomy is complete. Thus, world-wide distributional data are imperfect, both
because the naming may be faulty and because vast areas remain bryologically
unknown. Enough is known, however, for bryofloristic regions to be sketched with
considerable confidence that future data will serve to sharpen the resolution of the
regions and their floristic affinities
( 1 ) THE BRYOPHYTIC HABIT
Bryophytes are diminutive plants from one to a few millimeters (mm) up to a
decimeter (dm) or more, with a few larger taxa. Most have slender stems and
unistratose leaves which lack a waxy cuticle. They dry out rather quickly when
subject to conditions favouring evaporation. Even though the richest diversity of
mosses and hepatics occurs in tropical latitudes, it must be stressed that they are
temperate plants best suited for cool, wet, high mountain forests or other sites
where evaporative stress is low during the growing phase. Comparatively few
bryophyte families comprise the floras of lowland tropical and warm, dry climates.
148
H.A. MILLER
Some species are strictly ephemeral, others annual, still others perennial. Many
bryophytes occupy specialized niches to which they seem to be specially adapted,
such as the renowned ‘goblin gold’, Schistostega, in caves, or Fontinalis in running
water, Sphagnum in acid bogs and Andreaea on granite rocks at high
latitudeslaltitudes. Some are restricted to disturbed soils-Anthoceros, Blasia and
Physcomitrium. Mnium and Lophocolea are usually associated with humus, and a great
diversity of forms occur at the bases and on the branches of angiospermous trees.
Some leafy liverworts complete their life history as epiphylls. Correlation between
specific substrate and the presence of particular bryophytes is very close in some
cases, but generalized in others. The most significant underlying factor is the
necessity for moisture during growth and sexual reproductive stages. The
biflagellate sperm must swim through a film of water to reach the egg. The
resulting sporophyte generation remains attached and derives significant nutrition
from gametophyte throughout its temporary existence (Pate & Gunning, 1972).
( a ) Hornworts
The Anthocerotophyta are characterized by a thallose, parenchymatous,
chlorophyllose gametophyte which may have a unistratose wing (Dendroceros). The
sporophyte, unlike all other bryophytes, is indeterminate with a basal intercalary
meristem which receives mechanical support from the surrounding conicocylindric, gametophytic involucre. The basal meristem assures a continual
production of spores during active growing periods (Thomas et al., 1978). Three
genera, Anthoceros, Phaeoceros and Notothylas have rather thick-walled spores, yellow
to blackish, with considerable drought resistance. Spores of these genera have been
successfully germinated more than 3 years after collection and dry storage under
ordinary room conditions, and Proskauer (1957) reported 13 year viability. Some
species in these genera also form drought-resistant tubers in soil, which persist after
the surface portion of the thallus has succumbed. The remaining genera,
Dendroceros and Megaceros, are essentially tropical forest plants. They have thinwalled, transparent, hence green-appearing, spores. Both are normally found on
humus or as epiphytes in contrast to the other genera which usually occur on
disturbed mineral soils. The pioneer species are often quite seasonal.
The Hepatophyta exhibit a wide range of habits and adaptive strategies: ( 1 )
lea@liverworts dependent upon high humidity levels through most of the growing
period, with few ephemeral; (2) thallose liverworts of the metzgeriopsid type, with
an unspecialized thallus and lacking mechanical or physiological drought
resistance; and (3) thallus liverworts of the marchantiopsid type, with internally
differentiated thalli and often mechanical or physiological adaptations to drought.
Leafy liverworts are thought to be derived from an ancestral type with 3-ranked,
unistratose leaves. Morphological studies of contemporary species and the meagre
fossil record suggest that the ancestral plants had a prostrate stem, perhaps
thallose, with numerous ascending branches with terminal archegonia and distal
axillary antheridia. The archegonia and developing sporophyte were protected by
a whorl of specialized leaves forming the perianths as in Herbertus. For many types,
the ascending habit is considered to have given way to a prostrate life-form with
BRYOPHYTE EVOLUTION AND GEOGRAPHY
149
the lower leaves somewhat reduced, as in Lepidozia or Lophocolea, or even to loss of
lower leaves entirely, as in Plectocolea. The greatest number of extant species are
epiphytic in large genera such as Plagiochila, Radula, Frullania or the Lejeuneaceae.
This is mainly associated with moist forests and great diversity of liverworts occurs
on angiospermous trees. In cloud forests or in otherwise very humid, cool sites in
the tropics, epiphylls may also occur. Those thallose forms which are
anacrogynous, such as Pellia or Pallavicinia, with essentially parenchymatous thalli
require high moisture levels during the growing season. Species of Riccardia grow
over shaded humus in forest and MetLgeria species tend to be epiphytic. In
mediterranean climates genera such as Fossombronia appear over soil in the seasons
of highest humidity and rainfall and then survive unfavourable seasons as droughtresistant spores. A similar strategy exists for Sphaerocarpos and the semi-desert aquatic
Riella. The distribution of the ephemeral anacrogynous hepatics is poorly known.
The familiar liverwort of folklore and basic biology books is Marchantia or its
allies. Marchantiopsida are mostly soil-inhabiting, prostrate forms with numerous
rhizoids and internally complex thalli bounded by a well-defined epidermis with a
cuticle and (often) pores for crude control of transpiration. Many can persist over
long periods of desiccation and resume growth upon wetting. Long-dried thalli of
Asterella palmeri collected in 1955 from Guadelupe Island, Mexico, grew vigorously
in culture dishes where placed in 1958 to soak for sectioning and study. Perhaps the
tuberous Ricciae are equally desiccation resistant. Almost glass-brittle colonies of
Turgionia hypophylla *ill resume growth upon wetting. By contrast Dumortiera and
Monoclea require moist sites.
( c ) Mosses
Mosses also must be hydrated to survive and numerous strategies have evolved
where conditions are but occasionally favourable for growth. Flowers ( 1973)
discussed these strategies in relation to the mosses of Utah, where the climate is
generally dry and temperate continental. Great diversity has evolved in areas
where evaporative stress is minimal. Thus, the montane rainforests of the tropics
and the summer cool, winter wet, high latitude regions have an abundant
bryovegetation.
Peatmosses (Sphagnum spp.) are well-known to gardeners and potted plant
enthusiasts because of the tremendous water-holding capacity of the porose
hyalocysts and the presence of phenolic compounds which inhibit growth of
bacteria and fungi (Dickinson & Maggs, 1974). Peatmoss is often thought of as
strictly an acidic bog plant, where it is constantly soaked in water; several species
in Florida, however, occur in normally dry ditches or small depressions in the pine
flatwoods. Even severely dried and brittle patches of Sphagnum recover very quickly
when rain comes. The water-holding capacity serves to extend the growing period.
Few species are tolerant of shade, so for most, potential evaporative stress remains
high. The tiny trilete spores are exploded a metre or more into the air, enhancing
dissemination even if wind is still.
Soil mosses may be ephemerals, such as Aphanorrhegma, Micromitrium, Eccremidium
or a host of other so-called ‘pygmy mosses’. Pioneer turf-forming genera include
Polytrichum, Dicranum and Bryum, among others, in northern latitudes. Pottiaceous
mosses form close carpets over steep soil banks and other exposed sites. Whereas the
soil-dwelling mosses are mostly erect, acrocarpous types, Eurhynchium, Zsopterygium
8’
150
H.A. MILLER
and Ectropothecium are representative of pleurocarpous genera with terricolous
species. Under extreme conditions of evaporative stress no mosses may exist, or the
flora may be limited to ephemeral types which survive by virtue of long-viable,
desiccation-resistant spores. Soil mosses are effective in preventing erosion,
concentrating minerals and holding water, making them an ideal medium for seed
germination and subsequent overgrowth by other vegetation.
Rock mosses as considered here include all normally growing on rock, not just
the genus Andreaea. Some taxa occur most frequently on acidic rocks. New lava
flows in Hawaii are first populated by Grimmia, Racomitrium or Campylopus,
depending upon altitude and moisture levels ;these and others also occur elsewhere
on acidic rocks. Basic rocks, usually all or part limestone, carry distinctive floras.
Tufa forms around Didpodon tophaceus in temperate regions, and raw coral
limestone in Guam and Palau supports closely adnate mats of Racopilum or
Macromitrium. Much has been written about rock substrate specificity (e.g.
Nagano, 1972; Brown & Buck, 1978) even including special ‘copper mosses’
(Gams, 1974) and concentrators of heavy metals (Crundwell, 1976; Miller, H.,
1971).
Extra moisture-holding capacity and mineral content of decaying logs and
humus makes them sites amenable for development of mesophilic mosses, such as
Mniwn, Leucophanes and Hypnum, which form close mats or tufts over the organic
matter. The Splachnaceae have gone so far that most occur only on dung, bones or
other animal remains-they attract insects by odour and colour of the capsule and
these insects spread the spores (Koponen & Koponen, 1978) to a fresh animalderived substrate. It is possible that some mosses have a mycorrhizal association
similar to that found in the liverwort Jenseniu which grows on the sides of open peat
cracks. The organophilic bryophyte colonies eventually disappear with total
substrate decay or as a result of being out-competed by larger plants. Even so, they
are an integral and important part of the delicate balance of nature.
Buttresses, boles and branches of trees support an incredible variety of both
mosses and liverworts. Few seem to be ‘host’ specific. Instead, bark texture, acidity,
moisture capacity and competition seem to influence establishment. Vertical
zonation of epiphyte communities has been studied from both physiological and
community composition standpoints. Tobiessen, Mott & Slack ( 1978) summarized
recent work in north temperate forests, which shows that species higher on the tree
reach optimum photosynthesis at lower humidity and higher light intensities than
species lower on the trunk. In tropical cloud forests where epiphytes form thick
cushions on branches or hang as festoons several decimetres long (Russell & Miller,
H., 1977), the green colours characteristic of most forest floor or tree base species
give way to golden brown or blackish (sometimes reddish) colours which
predominate among such moss genera as Macromitrium, Acroporium, Phyllogonium,
Trachypodopsis, Aerobvopsis, Papillaria and Meteoriwn, as well as hepatics like
Herbertus, Pleurocia, Matigophora and Baccania. Although gymnospermous forests of
the Pacific north-western U.S.A. support tremendous quantities of bryophytes, the
diversity of epiphytes is comparatively low. Angiospermous trees on the other
hand, support a great melange of taxa. In Florida, Taxodium, Acer, Liquidambar,
Sabal and Quercus grow admixed on low flood plains. A few hepatics grow on the
cypress but few mosses are found there, in contrast to the abundance (e.g
Forsstroaia, Cryphaea, Thelia, Octoblepharum, Neckeropsis, Syrrhopodon, Schlotheimia and
Leucodon) on angiospermous trees. Tree fern trunks support a limited, often rather
BRYOPHYTE EVOLUTION AND GEOGRAPHY
1.5 I
specific, bryoflora. In east Africa, Rhizofabronia sphaerocarpa is a tree fern epiphyte
as is Hymenodontopsis in New Zealand, Hymenodon, Lopidium and Eriopus in tropical
America (Gams, 1932) and Calomnion in the tropical Pacific Islands.
Epiphyllous mosses are few with the curious, essentially leafless, Ephaeropsis
from the Malesian tropics being perhaps best known. However, pendant mosses
(Barbella) or small quickly maturing mosses (Leskeodon and Daltonia) are
occasionally epiphyllous as well as being pioneers on young twigs. Propagation of
Barbella is mostly by fragmentation whereas Daltonia produces many spores. Both
types of diaspores are probably trapped in leaves of the adnate hepatics which,
with crustose lichens, seem always to be the first invaders of the leaf surface.
Aquatic mosses, including Fontinalis, Scouleria and Wardia, may form large
masses attached to stones, logs or roots in running water. Other genera may have
some aquatic species, such as Fissidens manateensis, F. debilis (Hiltunen, 1966),
Sciaromium tricostatum and Leptodictyon rbparium. ‘Moss balls’, small spherical colonies
of mosses much modified in gross appearance and other bryophytes are
occasionally recovered from considerable depths in clear freshwater lakes (Baba &
Iwatsuki, 1973; Light & Smith, 1976; Luther, 1979). Marine bryophytes are
unknown and only a few species like Grimmia maritima, Phaeolejeunea spp. and some
Fuegian hepatics have developed tolerance to coastal sea spray (Engel & Schuster,
1973).
(2) BRYOPHYTE DISPERSAL
The small spores of bryophytes seem well-suited for long distance dispersal by
wind. Indeed, establishment of bryophytes on remote islands and the
disharmonious nature of their bryofloras is good evidence for the phenomenon.
Transport by sea must be ruled out because both the mature plants and diaspores
are intolerant of sea water. It might be expected, then, that bryophytes would be
broadly and rather uniformly distributed wherever a suitable niche might exist.
However, as Crum (1972) carefully documented, mosses seem to have migrated as
members of a flora “not as individuals and not aimlessly, but along natural
migratory routes”. The hypothesis of whole scale movement of a flora with the
complex interaction of its numerous components and the resulting diversity of
niches, each with its particularly well-adapted bryophytic occupant, is both
reasonable and probable for continuous continental floras. Slowly migrating floras
cannot, however, account for many distributional disjunctions nor for the presence
and composition of island floras.
Invasion of newly exposed land, such as the Pleistocene-glaciated Great Lakes
region, surely followed retreat of the ice. Crum (1972) addressed details of
bryophytic revegetation of northern Michigan in particular and eastern North
America in general. Beach pools of Michigan lakes have associated with them such
northern disjunctions as Cinclidium stygium, Catoscopium nigritum, Meesia spp.,
Amblyodon dealbatus and Scorpidium scorpioides. Some of the mosses present in
northern Michigan occur also in the Rocky Mountains. Both sites are calcareous
and the presence of such species as DitrichumJlexicaule, Tortellafragilis, T. tortuosa, T.
inclinata, Thuidium abietinum, P1agiop.r oederiana, Encahpta spp. and Saelania
glaucescens may represent elements which migrated eastward along the glacial
front. Acid peatlands are outliers of the boreal taiga, with numerous Sphagnum
species, Splachnum ampullaceum and Calliergon stramineum among others, which seem
1.52
H.A. MILLER
to have persisted since the Pleistocene. On a broader scale, the moss flora of eastern
North America has few endemic genera and species and most are widely
distributed in the eastern temperate region. Even an area so comparatively isolated
as the ancient Ozark Mountains (U.S.A.), has a flora of about 90% eastern
temperate species and some 70% of these alsb occur in the old world. Disjuncts in
northern Michigan mostly occupy habitats similar to those prevalent at the glacial
margin or on shores of post-glacial lakes and so provide evidence for the history of
the flora.
Although Crum discussed various bryophytic diaspores, he considered that most
lacked obvious adaptation for long distance carriage, pointing out the lack of
wings, sparse food reserves, limited or unknown viability, and questionable
resistance to extreme temperatures or ultra-violet radiation. Subsequently, van
Zanten (1976, 1978) undertook a series of experiments on 139 species of New
Zealand mosses. The strategy behind the experiments was laboratory replication of
the environmental stresses to which diaspores would be exposed in the course of
long distance (over2000 km)transport. Air-dried spores were shipped to Holland
from New Zealand by surface mail and stored at ambient indoor temperatures.
Spores had been dry for 12-14 months when exposed to simulated high altitude
atmospheres as follows: (1) spores wet frozen at - 30 "Cand 0.25 atm for 24 h; (2)
as (1) for 4 days; (3) spores dry frozen at - 30°C and 0.25 atm for 4 days; and (4)
no treatment, equivalent to low altitude dry winds. After exposure to experimental
conditions, the spores were sown on agar plates where germination and nongermination was scored after about 4 weeks. Photoperiod, light intensity and
temperature of the culture plates was unreported and exposure to UV was not
incorporated into these initial experiments. After being dried for a year, 97 (69%)
of 139 species germinated; about 58% of species survived as wet frozen spores but
only about 35% ofthe species could withstand the dry freezing treatment. The best
survivors were mostly species of wide geographical distribution. It is apparent from
van Zanten's experiments that at least some moss spores can be expected to survive
the long journey, but mature plants must develop in the new territory and be able
to reproduce year after year if establishment is to be accomplished.
Some mosses produce incredible numbers of spores of the order of 10-25 pm
diameter and these can become airborne in a light wind. That some can be
effectively transported at least 1000km is attested by the discovery of Tortulu,
Funariu and Murchuntiu in a newly formed crater on Deception Island, Antarctica,
9 months after the eruption (Young & Klay, 1971). Ultimate establishment of the
newly arrived colonists is unconfirmed.
Spore production in hepatic sporangia varies from a few hundred (Ricciu and
Sphuerocur/~os)to a few thousand (thallose marchantiopsid species and Pelliu) to tens
of thousands (some foliose genera) (Watson, 1971). Overall, numbers of spores
produced are consistent between hepatics and mosses but the condition of the
spores upon release is not always comparable. The Radulaceae and Porellaceae
and to a lesser degree Lejeuneaceae (which comprise about 25% of all liverwort
species) release spores which are multicellular by virtue of precocious germination
prior to the usually explosive discharge into the air. It is thought that some spores
rupture the spore wall soon after release, thus becoming vulnerable to desiccation.
The very thin-walled spores of epiphyllous Lejeuneaceae may live less than an
hour in dry air once released from the capsule (Fulford, 1951). We know less about
spore longevity among the leafy liverworts than among the mosses.
BRYOPHYI’E EVOLUTION AND GEOGRAPHY
153
Much has been made of the high incidence of dioecism in the bryophytes (e.g.
Schuster, 1979a)which necessitates the deposition oftransported spores so that male
and female plants are within range to accomplish fertilization. Sperm are shortlived once released from the antheridium so the distances cannot be more than a
few centimetres. Some (e.g. Atrichum or Mnium,) are dispersed in rain-splash or in
run-off. In the liverwort Sphaerocarpos, for example, spores often remain in tetrads
so that upon germination two male and two female plants are in a single rosette,
but this is an exceptional case. If spore formation were an absolute requisite to
establishment of a dioecious bryophyte, then few would be expected on remote
islands of the Pacific or to be limited to high peaks often separated by great
distances from similar habitats. This is not the case. In Hawaii, species which are
both dioecious and broadly distributed in the Indo-Pacific region include Leucoloma
molle, Hyophila involuta, Breutelia arundinifolia, Trachypus bicolor, Aerobryopsis longissima
and Haplohymenium triste. Each may have become established from the chance
arrival of a single spore to a fortuitous spot but the expansion of the range in
Hawaii was probably dependent mainly upon vegetative propagation.
Non-meiosporic, i.e. vegetative, diaspores are frequent and varied among the
mosses and liverworts. Some of these specialized asexually produced propagules
are illustrated in Fig. 2. The absence of gemmiparous structures on dioecious
mosses such as Barbella and Aerobryopsis or the hepatics Herbertus or Barzania does
not mean that they cannot be dispersed over considerable distances. The fragile
nature of plants during normal dry phases of growth results in fragmentation at
exactly those times when maximum opportunities for transport exist. A
temporarily dried clump on rock or soil may be crushed by a passing animal or
have small pieces blasted off by wind-borne sand particles. Epiphytic species,
especially pendant forms, are constantly abraded by wind action, as evidenced by
great numbers of moss and liverwort leaf fragments trapped in the cloud-bathed
elfin forest of Puerto Rico (Howard, 1968 and unpubl. data). Fragments, no
matter how produced, are totipotent so long as survival tolerance limits are not
exceeded (e.g. temperature limits are much broader in a less than fully hydrated
condition). In other words, any living bryophyte cell has the potential to establish
a new plant and normally will do so in an amenable niche. In his review of asexual
reproduction, Watson ( 1971) observed “that the exceptional regenerative powers
of bryophytes cannot fail to be of use in propagation” but minimized the
significance of vegetative diaspores for any but very local dispersal. The atoll floras
of Micronesia (Miller H., Whittier & Bonner, 1963) seem to be partly established
from vegetative diaspores. For example, Calymperes tenerum, the most common atoll
moss, is apparently dioecious but produces great numbers of propagula. Some of
these were laboratory germinated many months after being collected, pressed and
dried along with vascular plant specimens. Calymperes hyophilaceum propagula
collected at the same time also survived. The abundance of sterile Calymperaceae
and Leucophanes, which produce nearly identical propagula, on the atolls strongly
suggests effective vegetative dispersal. Other species such as Pelekium velatum,
Thuidium plumulosum, Taxithelium lindbergii, Trichosteleum hamatum and Ectropothecium
spp. and the hepatics, probably were transported as spores since they fruit freely in
contrast to Calymperaceae.
Diaspores of the bryophytes are quite varied. True spores are mostly small with
irregular surfaces which enhance wind transport; others are large (200 pm) and
were confused in at least one case with seeds borne in a pyxis of an angiosperm by
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H. A. MILLER
Figure 2. Asexual diaspores ofbryophytes. Mosses: A, Tetraphic with terminal gemma cup, A’, gemma;
B, Pohlia ‘tuber’; C, Aulacmnium with gemma-stalk, C’, gemma; D, Tmkcla leaf with gemmae, D’,
gemma; E, Plalvgniun with axillary gemmalings; F, Calympcrcs leaf with distal propagula, F‘,
propagulum; G, Calyptothccium axillary gemma. Hepatics: H, Blaria with gemma ‘bottle’, H’, gemma;
I, Marchutilia gemma cup, with gemmae, on upper surface of the thallus, 1’, gemma; J, Mcrrgnin
thallus with surface gemmae, J’, gemma; K, Radula leaf with marginal discoid gemmae, K’,young
gemma; L, Odontoschisma rect gemmiparous branch, L’, unicellular gemmae; M, Plagiochila with
gemmalingson leaf surface, M’, gemmaling; N, Rcctolejeunca showing remnants of caducous leaves. The
scale bar with each gemma is approximately 50 pm.
Ferdinand von Muller, who described Trianthema humillima (Aizoaceae) for fertile
Gigaskermum repens, an Australian moss (Black, 1943-1957). Spores may be
waterborne (as in Fontinalis, Scouleria, Riccia and Sfihaerocarfios) or carried in mud on
the feet of waterfowl (as postulated for Riella; Persson & Imam, 1960). Gemmae,
propagula, tubers, bulbils, deciduous branchlets and variously derived fragments
are clearly of high importance for localized dissemination, with medium to long
distance dispersal being accomplished in at least a few cases.
(3) T H E FOSSIL RECORD
Fossils of bryophytes so far recognized and reported are few when compared to
vascular plants but they have been adequate to indicate an early existence of
BRYOPHYTE EVOLUTION AND GEOGRAPHY
155
mosses and liverworts. Because of their rather delicate structure and the necessity
for examination of cellular detail for determination, some fossils thought to be
bryophytes cannot be confirmed. Application of maceration techniques for
isolation of small fossils has recently detected pieces of bryophytes sufficient for
good approximation of their systematic positions. Jovet-Ast ( 1967) provided a
comprehensive update of fossil Bryophyta as reported to the end of 1964. Schuster
(1966) reviewed reported hepatic fossils up to the same period with his
interpretation of the significance of each. Subsequent publications have reported
fossil taxa, with preservation methods, from diverse strata and localities.
Compression-preserved cellular detail was found for Diettertia montanensis (Brown &
Robison, 1974) making possible estimates of systematic affinities. Spores in fossil
deposits (e.g. Grubb’s (1978) report of Anthocerotalean spores in Australian
Quaternary; the discovery of Notothylacites and Oxymitra-like spores in Upper
Cretaceous; NEmejc & Pacltovi, 1972) serve to illustrate continuing refinement of
palynological evidence for bryophytes: The appearance of An Atlas of Recent
European Moss Spores (Boros & Jirai-Komlbdi, 1975) has clarified bryophyte spore
variation with either light or scanning electron microscope photographs of over
200 species and by comparative studies with extant spores. Growing availability of
the scanning electron microscope to palynologists and bryologists portends rapid
expansion and refinement of information derived from fossil spores which prove to
be bryophytic.
( a ) Pre-Permian bryophytes and not-quite bryophytes
Bryophytes represent a level of evolution correlated with transmigration to
terrestrial environments. Critical were the origin of sterile-jacketed gametangia,
retention of an embryo within the archegonium, mineral absorbing and anchoring
structures, adaptation to evaporative stress and, perhaps most significantly, the
formation of trilete, sporopollenin-walled, protected spores (Gray & Boucot,
1977, 1979).Trilete spores and cuticle fragments are found only in the shoreline deposits of Silurian age laid down some 30 million years before Cooksonia, the oldest
vascular plant presently known. It is not unlikely that some plants which today would
be classed as bryophytes created those tantalizing traces embedded in Silurian fine
mud. No one has yet claimed to have found unambiguous mosses or liverworts
among Silurian fossils, but it seems reasonable to expect that bryophytes had
emerged along with other vascularized plants by that time (Miller, H., 1979) and
that presently questionable remains are indeed bryophytic. For now, however, the
oldest known liverwort is Pallaviciniites devonicur ( =Hepaticites devonicus Hueber,
1961) from the Lower Upper Devonian of New York. The earliest probable fossil
moss recognized is Muscites plumatus from the Lower Carboniferous of
Gloucestershire (Thomas, 1972). The oldest hornwort in the fossil record is
uncertain. Jovet-Ast ( 1967) accepted some Oligocene spores as being the oldest ;
Jarzen (1979) reported spores of Phaeoceros (an extant genus) from the Cretaceous;
the late Johannes Proskauer held the private opinion that some of the Devonian
and pre-Devonian spores were of Anthocerotophyta rather than from Horneophyton,
which does have similar spores. Certainly, the organization of the hornworts
suggests a plant group of great antiquity.
Schuster (1966) referred to “two Metzgerialean fossils from the Lower
Carboniferous (Mississippian) of Scotland” but I have been unable to trace his
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H.A. MILLER
precise reference unless it was to Walton’s ( 1949)Hepaticites from Clackmannanshire.
The middle coal measures (Upper Carboniferous) of Shropshire and
Nottinghamshire yielded Hepaticites kidrtoni, H . lobatus, H. langii and H . metzgerioides
(Walton, 1925, 1928).Schuster created the genus Treubiites for H . kidstonii, Blasiites
for H. lobatus and redefined Hepaticites typified by H . langii. Other Upper
Carboniferous possible hepatics are Thallites willsii and T. lichenoides. According to
Lacey (1969), Oschurkova has found Hepaticites metzgerioides in the Upper
Carboniferous of the Karanganda Basin, U.S.S.R., indicating a broad
distribution. DiMichelle & Phillips ( 1976) described Thallites dichopleurus from the
Middle Pennsylvanian (Upper Carboniferous) of Illinois but the affinities are
uncertain. From the Lower Devonian in Poland, Zdebska (1979) described a
Spongiophyton for a liverwort-like fossil thallus. Earlier Chaloner, Mensah & Crane
(1974) reported two species of Spongiophyton as possibly hepatic from the Middle
Devonian of Ghana. Only the cuticle remained of the organism but that was up to
80 ym thick! Remy & Remy (1980a, b) found Lynophyton rhyniensis in the Lower
Devonian Rhynie cherts as structures which appear to be weakly vascularized,
stomatate gametangiophores bearing archegonia and antheridia. Another Lower
Devonian (Siegenian) gametophyte, Sciadophyton sp. has been reported in detail by
Remy et al. (1980) from the Westphalia region of West Germany. Both of these
gametangium-bearing organisms have simple vascular systems. A S’haerocarpos- or
Riccia-like spore tetrad, Tetrapterides, is known from among fragmentary remains of
a thalloid plant from Lower Carboniferous of Wales and Gloucestershire (Sullivan
& Hibbert, 1964; Hibbert, 1967). But it is the discovery of Torticaulis transwalliensis
from the late Silurian Old Red Sandstone that particularly piques the imagination
(Edwards, 1979). This still problematical fossil resembles an hepatic sporophyte
which, if confirmed by subsequent studies and discoveries, will further strengthen
the case for independent origins of mosses, liverworts and hornworts. Trilete
spores, e.g. Ambitisporites, are known from early Silurian deposits in Virginia (Pratt,
Phillips & Dennison, 1978).
As noted, the oldest probable moss is Muscites plumatus from Lower
Carboniferous shales of the Forest of Dean, Gloucestershire, with some
compression tissue present having sufficient cellular detail to show it comparable to
true mosses (Thomas, 1972). Muscites polytrichaceus and M . bertrandii are prePermian mosses being from the StCphanien strata (Upper Carboniferous) of
France. Plumstead (1966) reported a probable moss with preserved cellular detail
from the “Upper Carboniferous Stage, Protoglossopteris Zone” of the Transvaal but
the age seems uncertain (Lacey, 1969). A questionable Lower Devonian specimen
from the Rhynie cherts was identified by Lemoigne (1966) as a moss sporangium,
but M e s h considered the specimen to be Sporogonites and called it S. lemoignei
(1966) in his review of the original report. It may be Horneophyton, but even if it is
Sporogonites, it is not a true moss.
Fossils attributed to bryophytes but based upon other organisms were not fully
accounted for in Jovet-Ads compendium. Eohepatica dyfriensis ( = Thallomia
llandyfriensis) from the very early Devonian is now known to be a crustacean
(Miller, H., 1973). Kozlowski & Greguss (1959) described Hepaticaephyton and
Musciphyton presumed to be from the Ordovician. Lacy (1969) indicated that most
paleobotanists considered the materials in question to be no more than
contaminating fragments of recent plants-perhaps roots of Carex, a sedge. Still
another problematical plant once considered bryophytic is Protosalvinia, but Niklas
BRYOPHYTE EVOLUTION AND GEOGRAPHY
157
& Phillips (1973) favoured a place for it among the brown algae. Unnamed
fragments of cuticle were isolated by maceration of Autunien (Lower Permian)
rocks from the Saar and figured by the late Rolf Busche ( 1968). The remains were
compared to several mosses but the large size of the cells and wall configurations
are much closer to those of fern stipe scales than to mosses. Unless future research
develops additional evidence, these cuticle fragments should not be considered as
bryophytes. A report by Fleischer ( 1919) of a Devonian fossil of Andreaea was based
upon Sporogonites according to Dixon ( 1927), the specimen coming from
Sporogonites-bearing strata.
We can expect that the next 20 years will bring to light numerous Silurian and
Devonian plants of bryophytic habit and Carboniferous discoveries will confirm
the integrity of major extant groups. Never before have so many fine tools and
techniques been available, and surely never before have so many been involved in
prising information from the fossil record. From the bryologist’s viewpoint,
whereby the bryophytic habit is defined by the presence of a partially dependent
sporophyte borne on an independent gametophyte, at least two potential problems
remain: (1) Rhynia has been assumed to be a vascularized diploid plant-i.e. a
‘normal’ land plant; Merker ( 1958), and subsequently Lemoigne ( 1970),
presented evidence that the horizontal axis is a gametophyte and the erect axis an
haustorial sporophyte. If this is so, although many palaeobotanists are sceptical,
then could Rhynia be a bryophyte? I think not, simply because such an
arrangement is not alien to extant plants such as Actinostachys which all agree is a
perfectly good fern (Bold, Alexopoulos and Delevoryas, 1980). (2) Sporogonites has
been variously placed with psilophytes (now rhyniophytes) somewhere near
Horneophyton, in the mosses as an ancient Andreaea, or most recently as a bryophyte
something like a thallose liverwort. One species, S.exuberans (Lower Devonian of
Belgium and Norway) and a second, S. chupmani (Lower Devonian of Australia), is
tentatively placed in the genus. Andrews ( 1960) thoroughly studied specimens
from the Emsian (Upper Lower Devonian) beds of Belgium and found a dark
thallus-like layer at the base of each erect sporangiophore bearing a single erect
sporangium. The plan and scale of Sporogonites is consistent with the bryophytesi.e. the ‘seta’ is about 50mm long with the 6-9mm long capsule having a central
columella covered by a domed archesporium which produced tetrahedral to
globose spores 20-25 pm diameter; the ‘thallus’ appeared to be non-vascular and
at least 150 x 50mm. IfAndrews’ (1961) interpretation of the plant as bryophytic
is correct then Sporogonites represents a long extinct experiment about the overall
size of modern Monoclea.
( b ) Permian and Triassicfossils
Except for the land covered by Carboniferous glaciers, many suitable cool, moist
sites were apparently available for bryophytes and they were widely distributed
prior to the Permian. Some were predominantly Laurasian and others, including a
majority of the groups today considered evolutionarily primitive were
Gondwanan. These floras fixed the background upon which the modern
biogeographic mosaic rests even though the fossil record to date is meagre.
Hepatics have not been strongly represented in Permian deposits. Zalessky
(1937) described Marchantites lorea from Bardien strata in the Urals, but Lacey
(1969) did not consider the assignment unambiguous. I have seen a specimen
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H.A. MILLER
superficially like Hepaticites wonnacotti from Permo-Triassic deposits in Arizona but
it has not been formally reported. Townrow’s ( 1959) Hepaticites cyathodoides was
described from Middle Triassic shales of Natal, South Africa and Australia
(Plumstead, 1966). Anderson (1976) recognized two species, Marchantites cyathodoides (Townrow) Anderson and M. tennantii, as well as a single unnamed Thallites
from the Molten0 Formation which had yielded Townrow’s specimens. Takahashi
( 1960)reported an undescribed “Hepaticites oishii” from Triassic of western Japan
but I have been unable to locate a confirming description.Jain & Delevoryas (1967)
reported Thallites sp. from the Triassic of Argentina. A comparative abundance of
hepatics is known from Rhaetic-Liassic (Triassic-Jurassic) deposits. The most famous
of these is Naiadita lanceolata from the English Rhaetic. Harris (1937, 1939) found
leaves, stems, rhizoids, gemma cups, gemmae, archegonia, sporophytes and spores,
making Naiadita the best known of fossil bryophytes. It was apparently an aquatic
plant allied to modern Riellaceae which are mostly plants of ephemeral desert
ponds. Thallites rostajnskii is from the Lower Liassic of Poland and T. uralensis from
the Rhaetic-Liassic of the Urals. Strata of the same age in Iran have yielded
Ricciopsis iranica described by Fakhr (1977) as allied to R. scanica. An unnamed
species of Hepaticites has been reported from the Liassic of Germany (Krausel,
1958). Marchantiolites porosus, Ricciopsis Jlorinii and R. scanka were found in the
Liassic of Scania, Sweden. Ricciisporites tuberculatus from the same strata is
considered of uncertain affinities (Lundblad, 1959). Hepaticites solenotus is based
upon a thallus from Lower Liassic rocks near Bristol, England. Additional thalloid
forms described from eastern Greenland include Metqeriites glebosus, Hepaticites
laevis, H . rosenkrantzi and H. amauros. Save the special case of Naiadita, all preJurassic hepatics are thus of thalloid morphology.
Permian mosses were unknown, and pre-Permian records questionable because of
the modes of preservation, until Marion Neuberg of the Institute of Geology of the
Academy of Sciences of the U.S.S.R., reported the discovery of true mosses in
1956. This report was followed in 1958 by an abbreviated presentation in English
announcing new genera and species, to be described in her forthcoming
monograph, along with four plates of photographs representative of the incredibly
well-preserved material in hand. The monograph, complete with 52 drawings and
hundreds of photographs arranged in 78 plates, appeared in 1960 entirely in
Russian and summarized details from some 212 specimens obtained from drill
cores, mines and outcrops (but the monograph did not cite the 1958 English
language synopsis which was overlooked by Jovet-Ast and Lacey) . Both bryopsid
and sphagnopsid mosses were preserved in sediments of the Kuznetzk, Tungus and
Pechora basins of Angara. Two Lower Permian taxa, Junjagia glottoplylla and
Vorcutannularia plicata, plus the Upper Permian Protosphagnum recurvatum comprise
the order Protosphagnales and form a graded morphological series approaching
modern Sphagnum. Bryopsids from the Lower Permian are Intia vermicularis, I.
variabilis, I.falcifonnis, I. angustifolia and Salairia longifolia. Upper Permian taxa are
Uskatia conferta, Polyssaieuia spinulifolia, P . deJexa, Bajdaievia linearis, Bachtia ovata and
Muscites unifonne which is known only from a fragment of a leaf blade. A second,
possibly distinct, Protosphagnum with a strongly perfoliated leaf has been found in
the Upper Permian of the Priuralia region of the southern Urals (Meyen, 1966).
True mosses have been reported from northern European Permian strata of the
U.S.S.R. by Fefilova (1978). An extremely well-preserved Permian moss from the
BRYOPHYTE EVOLUTION AND GEOGRAPHY
159
Antarctic, not yet formally published but previously noted (Miller, H., 1979),
confirms the presence of mosses in the Glossopteris flora.
Reports of Triassic mosses are few, indeed, but Anderson’s (1976) discovery of
numerous specimens from several sites of Muscites guescelinii including “eight moss
cushions (clusters) which were perfectly preserved in situ” gives some cause for
optimism for future discoveries. Townrow (1 959) developed sufficient cellular
detail to be able to compare M. guescelinii with Leucodontaceae on many points
and Anderson found probable antheridia and archegoniophores. A moss with
Sphagnum-like areolation and lacking a costa was reported from Nidpur, India, by
Pant & Basu (1978) as Sphagnophyllites triassicus. Spores presumed to be
sphagnopsid were described from the Rhaetic strata near Thuring, Germany, as
Sphagnumsporites apolaris. True Sphagnum leaves have been reported from the Liassic
near Nuremberg (Lacey, 1969).
( c ) Jurassic and Cretaceousfossils
The position of Rhaeto-Liassic deposits in the boundary zone between Triassic
and Jurassic strata is recognized, but for uniformity Liassic taxa have been treated
above as if they were Triassic. Thallose hepatics continue to dominate the fossil
record so far known in the middle and upper Mesozoic but mosses have been
reported with increasing frequency since broader application has been made of
maceration techniques on embedded small plant remains.
Hepatics recognized by Jovet-Ast ( 1967) from the Jurassic include Hepaticites
wonnacotti, H . haiburnensis, H. hymenoptera and H . arcuatus all from the Middle
Jurassic in Yorkshire (Harris, 1961), the last being the same as Marchantites erectus
from Victoria, Australia, and similar to a Liassic specimen from northern Iran.
Hepaticites plicatus was found in Lower Jurassic strata near Donetzk in the Ukraine.
Several species of Thallites are distributed in the northern hemisphere-T. erectus
from Yorkshire, T. zeilleri from England’s Upper Jurassic, T. marchantiaefrmis from
Portugal, T.polydichotornus from the Emba region of the U.S.S.R. and T..yabei from
both the Jurassic and Cretaceous of Angara, Korea and Japan. Marchantites
oolithicus was described from Nancy, France, a great distance from Marchantites
barwoni from Victoria, Australia. Krassilov (1973) found a rich and diversified
bryophyte flora in the Upper Jurrassic and Lower Cretaceous beds along the
Bureja River, a tributary of the Amur. Bulk macerations revealed structural details
of hepatics with stems and leaves as well as of thalloid forms and mosses. Cheirorhiza
brittae has Lejeuneaceous complicated bilobed leaves and underleaves similar to
lobules (much as for Porella) but rhizoids are scattered on the stem. The description
of Cheirorhiza almost suggests that several taxa may be mixed in the matrix or that a
generalized pre-Porellalean plant is at hand with some metzgeriopsid features still
retained in the late Jurassic. The same mix offeatures is found in Laticaulinapapillosa
which has, however, massive, sometimes bifid, underleaves and reduced leaf
lobules. Also found in the Late Jurassic beds was Aporothallus la@zhenskajae, a
Cyathodium-like thallus with possible sex organs. Gondwanan Jurassic deposits on
the west side of the Palmer Peninsula of Antarctica yielded Schizolepidella gracilis, a
fragment with transversely inserted, scale-like, shallowly bifid, obovate, sometimes
subopposite, leaves about 2.0 x 1.5 mm. Preservation seems to have been such that
further structural details cannot be revealed and so exact systematic placement is
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H. A. MILLER
impossible, Nonetheless, its is possible that it is an ascending, bifid-leafed, hepatic
similar to numerous primitive taxa which characterize the liverwort flora today
known from high southern latitudes.
Hepatics show greater diversity in the Cretaceous than in any previous period.
Several probable leafy liverworts occur as well as thalloid forms. Thallites sewardii
was found in the Lower Cretaceous of Maryland and T. jimboi in Sakhaline;
Marchantites hallei came from the Lower Cretaceous of Patagonia, M . yukonensis
from Yukon Upper Cretaceous and M . baicalensis of apparently inexact Mesozoic
age is Angaran. Thallites blainnorensis of Jovet-Ast’s (1967) summation has been
transferred to Marchantiolites by Brown & Robison (1976) on the basis ofadditional
material from the Kootenai Formation of Montana which yielded rhizoids and
cellular details of air pores. Jovet-Ast noted two species of Jungermannites (foliose
hepatics) 3. vetustior from Portugal and 3. cretuceus from Alabama. Each species is
imperfectly known but hepatic affiliation seems most appropriate. Krassilov has
found well-preserved marchantiopsid thalli of Striatothallus adnicanicus with cellular
detail in the early Cretaceous as well as Riccardia- and Riccia-like thalli which have
not yet yielded morphological details upon maceration. Lower Cretaceous beds of
Victoria, Australia, contained eight hepatics including an unnamed Fossombronialike Thallites as well as two probable foliose hepatics, Thallites spp. b and c
(Douglas, 1973). Two seeming anthocerotophytes allied to Notothylas, Notothylucites
filiformis (Ntmejc & Pacltovi, 1972) and Shuklanites deccanii (Singhai, 1973), have
been reported from the Upper Cretaceous. Spores resembling Oxynitra were found
with the Bohemian Notothylucites. Preservation was so good with the Deccan
material that elater morphology could be studied.
Traces of Sphagnales were found in Queensland Jurassic deposits with
Sphagnumsporites adnatus, S. tenuis and S. clavus, the latter also reported from the
Lower Cretaceous of the Perth Basin. In the Upper Jurassic of British Columbia,
Sphagnum punctaesporites is based upon spores only. Krassilov ( 1973) described
Tricostium pupillosum from late Jurassic beds of the Bureja Basin as being
superficially like Coscinodon but with three longitudinally ribbed costae. Of equal
age is Muscites fontinaliodes which is ecostate with carinate, apparently 3-rankedY
leaves and an erect capsule (details unknown) immersed in a perichaetium. Some
small leaves in the bulk macerations with Tricostium may be perichaetial leaves of
that genus or perhaps of still another taxon. Filatoff (1975) reported several
bryophytic genera of “Sphagnaceae-type spores” in Jurassic well cores taken from
the Perth Basin of Australia. In addition to Stereisporites (synonyms are Sphagnites
and Sphagnumsporites) psilatus and S. antiquarporites, Rogalskaisporites cicatricosus and
R . canaliculus are described and figured, including SEM photographs. Other spore
taxa from the Perth Basin are Po&cingulatisporites crenulatus, P . striatus, Antulsporites
varigranulatus, A . clavus, A . saeuus and Foveosporites moretonensis. It is my impression
that the diversity represented by figured spores exceeds that expected for extant
Sphagnaceae and that eventually both true mosses and hepatics may be sorted out
of these form taxa.
The first moss plant remains reported for the Cretaceous were of Muscites
lesquereuxii from Tennessee. Brown & Robison (1974) found an exceptionally wellpreserved moss, Diettertia montanensis, in the Lower Cretaceous of Montana which,
in my judgment, most resembles some Ptychomitriaceae such as Dichelodontium now
known from New Zealand-a pattern consistent with the distribution of Gunnera
pollen Uarzen, 1980). Krassilov (1973) found Yorekiellapusilla in Aptian deposits in
BRYOPHYTE EVOLUTION AND GEOGRAPHY
161
the Bureja Basin as a series of stem fragments bearing clasping 2- and (sic!) 3ranked leaves with eroded margins. Sphagnumsporites (now Stereisporites)
antiguasporites was found near Vancouver and S. psilatus came from western
Canada. Three species of Sphagnum based on fossil spores are attributed to the
north-eastern U.S.S.R., S. subflavum, S. pedatiformis and S . europaeum also known
from the Crimea. Other Sphagnum-like spores have been reported, Phillips & Felix
(1971) listed four species from the Lower and Middle Cretaceous of the southeastern United States, Romans (1975) found three types in Upper Cretaceous
Black Mesa Coals of Arizona and Hopkins & Sweet (1976) identified Stereisporites
antiquaesporites and Cingutriletes clauus from the Lower Cretaceous Mattagami
Formation near James Bay, Ontario. Only Marsypiletes cretacea seems to be different
in that the spore is monolete and resembles that of Plagiopus oederi Uarzen, 1976).
(d) Tertiary fossils
Angiosperms dominated the vegetation by the beginning of the Tertiary. Some
groups of bryophytes which had diversified greatly in the Cretaceous adapted to
the abundance of mesic to humid niches associated especially with the hardwood
forests. Partyka (1976) listed 75 species offossil bryophytes from the Tertiary ofjust
the U.S.S.R. including seven hepatics, five sphagna and 63 mosses with most being
referred to extant species or at least to extant genera. Wu & Feng (1978) described
Neckera shanwanica from the Cenozoic of China but I have not seen the report to
determine the age or type of preservation. N. Miller’s (1980a) list of North
American fossil mosses notes the small number of discoveries in North America to
date as compared, for example, to the record known for the U.S.S.R.
The Palaeocene-Eocene epoch moss and liverworts fossils so far known confirm
the presence of several species from most major groups. Marchantiopsid hepatics
have been reported from France, Texas, Arkansas and Montana and the
Notothylas-like Shuklanites from India is known from late Cretaceous- early Tertiary
deposits Uovet-Ast, 1967). Both Jungennannites eophilus from Colorado and 3 .
bryopteroides from Texas are foliose with a possible resemblance to Porellales.
Sphagnales so far recorded include Stereisporites (as Sphagnumsporites) stereoides and S.
megasteroides from Germany Uovet-Ast, 1967) and S. concepcionensis from central
Chile (Takahashi, 1977). Partyka (1976) lists six taxa assigned to Sphagnum from
Russian Palaeocene-Eocene strata. Acrocarpous genera of mosses include Dicranites
australis from Kerguelen, Mnium montanense from Montana Wovet-Ast, 1967),
Ditrichitesfylesii (Kuc, 1974d) and Aulacomnium heterostichoides Uanssens, Horton &
Basinger, 1979) both from British Columbia. Drepanocladus aff. sendtneri as well as
Calliergon trifarium and C. stramineum are listed by Partyka from the Russian Eocene.
Among other established pleurocarpous mosses represented, the much reduced
(primarily protonematal) Ephemeropsis has been found in Germany, Hypnites
haeringianus from Tyrol Uovet-Ast, 1967), H. arkansana from the Arkansas Wilcox
formation (Wittlake, 1968), with Palaeohypnum jovet-astii and P . steerei from the
Allenby Formation of British Columbia (Kuc, 1974d). Fossil mosses of unknown
habit are placed in Muscites as M . thuidioides from Kerguelen Uovet-Ast, 1967), M .
wilcoxensis in which the habit “suggests a Fontinalis-type of moss” (Wittlake, 1968)
and M . maycokii and M. ritchiei from the Allenby Formation which has also
produced some unnamed fragments of moss gametophyte (Basinger & Rothwell,
1977).
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H.A. MILLER
Most Oligocene bryophytes are reported from the famous Baltic Ambers
(Magdefrau, 1957; Grolle, 1980) in which the plants appear to be almost perfectly
preserved in a transparent matrix. Jovet-Ast (1967) listed many genera and species
attributed to the amber flora. Hepatics were assigned to extent genera such as
Frullania, Phragmicoma (now distributed among several genera of LejeuneaceaeHolostipae), Lejeunea, Madotheca ( =Porella), Radula, Lophocolea and Jungermannia
but three species of uncertain position were retained in Jungermannites. Grolle
(1980) has added Cephaloziella and Bazzania to the list of genera. Mosses
represented are mostly from acrocarpous genera4.e. Dicranum, Dicranites,
Dichodontium, Trichostomum, Phascum, Grimmia and two uncertain Muscites. A capsule
from brown coal at Yallourn, Victoria, Australia, has been called Muscites
yallournensis by Clifford & Cookson (1953), but its bryophytic origin is ambiguous.
An impression of a pleurocarpous moss has been described from Montana by
Steere (1972) as Palaeohypnum beckeri, the lack of anatomical detail making more
precise placement impossible. Russian Oligocene plants @de Partyka, 1976)
include reports of Riccia magna, R . ricciaellifonnis, R . tenera, Ricciites leiodorsales, R .
euricciafonnis and Marchantites globosus among hepatics and no other taxa. The
predominance of aridity-adapted Ricciae suggests a mediterranean climate at the
sites studied.
Miocene bryophytes reported by Jovet-Ast ( 1967) were from Poland, France,
western U.S.A. and one from Burma amber as listed:
Hepaticae
Marchantites sinuatus, France
Marchantia coloradoemis, Colorado
Plagiochila saportana, France
Jungmannites cockerellii, Colorado
Musci
Polytrichites spokanensis, Washington
Trachycystis Jagellaris, Poland
T. szaferi, Poland
Hypnodendron sp., Burma
Plagiopodopsis scudderi, Colorado
P . cockaelliae, Colorado
Fontinalis sismondana, Piemont
? Papillaria sp., Poland
Thamnium alopecurum, Poland
T. sp., Poland
Cluopodium sp., Poland
Heterocladium squarrosulum, Poland
Amblystegium schrotzburgme, Poland
? Muscites joursacensis, France
M. Jorissanti, Colorado
Palaeohypnum arnoldianum, Oregon
P. brittoniae, Washington
P . brownii, Colorado
P . patens, Washington
P . knowltoni, Washington
Since then, Selkirk ( 1974) has found in Australia the characteristic epiphyllous
haptera of Ephemeropsis known previously from Eocene brown coal in Germany
and today known only from the Malesian tropics. A capsule resembling that of
Desmatodon heimii and accompanying Pottiaceous spores found in brown coal in the
Lvov region of the Ukraine (Shchekina, 1959) was overlooked in Partyka's list.
Discovery of hepatic spores in Neogene sedimentary rocks in Tunisia (van Campo,
1978) extended that bryoflora from previously known localities in central Europe
(Jovet-Ast & Huard, 1966; Nagy, 1968). Jaehnichen (1974) found additional
material of Thamnites marginatus in Lower Lusatia, G.D.R. A moss from MiocenePliocene lignite found in Derbyshire possesses a combination of characters
BRYOPHYTE EVOLUTION AND GEOGRAPHY
163
unknown today ; Boulter ( 1974) has suggested that Muscites lanceolutu, originally
reported as Hypnodendron sp. (Boulter, 1971), represents the remains of an extinct
family of European mosses. Such a fossil contrasts to mosses of similar age reported
from the Canadian Beaufort Formation as found on Banks Island at 74”N with 13
species in eight genera (Kuc & Hills, 1971) and Meighen Island at 80”N with
Porellu sp. and 42 moss species. All fossils are of extant species of the boreal forest
except Culliergon aftonianum known only as a fossil (Kuc, 1973, 1974b). Sphagnumtype spores have been reported from the Mackenzie Delta region of Canada
( S t a p h , 1976).
( e ) Quaternary fossils
Post-Pliocene palaeobryology in the U .K. in relation to Europe, especially, was
admirably summarized by Dickson ( 1973) wo provided an extensive bibliography
of Pleistocene bryology. The most recent summary of Quaternary fossil bryophytes
in North America is that of Miller, N. (1976) who has worked extensively on
glacial and inter-glacial deposits. A full catalogue and an annotated bibliography
followed (Miller, N., 1980) in which minor corrections were made and additional
data recorded. Of particular note are comments about ccsupposedextinct moss
species” the validity of which is considered doubtful. Evidence is substantial that
the questionable taxa fall within the range of phenotypic variability of extant
species. The small number of hepatic fossils, only five species recognized in four
genera and two genera additionally represented without assignment of specific
epithet, is equivalent to only about 1o/o of today’s flora. Mosses are represented by
172 species (about 15%) in 82 genera (about 3104) of North American mosses.
The Quaternary bryophytes of northern Asia, U.S.S.R. were listed in considerable
detail by Partyka (1976). Nine hepatics, 26 Sphugnu and about 130 mosses reported
are comparable in number to those found in North America and, generally, many
of the same taxa were represented. Sharma (1978) reported some Sphagnum taken
from sediments C14 dated at c. 3500 years BP. Janssens (1977) studied subfossil
material from 10 sites in Belgium and one in France. All material could be
identified to modern species of holarctic and arctic zones although some taxa are
no longer extant in the vicinity of their discovery. No attempt is made here to
update Pleistocene and Holocene palaeobryological literature for it is so widely
dispersed (e.g. Goswami, 1957; Forsyth, 1961; Kirk & Godwin, 1963; Dickson,
1963, 1964, 1967; Birks, 1965; de Vries & Bird, 1965; Terasmae, Webber &
Andrews, 1966; Pilous, 1968; Conolly & Dickson, 1969; Miller, N. G. &
Benninghoff, 1969; Barry & Synnott, 1970; Berdowski & Wilczynska, 1973;
Burrows, 1974; Fredskild, Jacobsen & Raen, 1974; West et al., 1974; Zenkovitch et
al., 1975;Dickson,Jardine & Price, 1976; Seward & Williams, 1976; Brassard &
Blake, 1978; Hulme, 1979 and, as Miller, N. (1980a) noted, mention of bryophytes
is often so incidental that they can be easily overlooked in literature far out of the
bryologist’s usual range of search.
(4) BRYOPHYTE EVOLUTION AND SYSTEMATICS
Even though considerable agreement exists on which taxa are ‘closer’ to which,
the schematic concepts of overall relationships can, and do, vary widely from one
investigator to the next. However, as evidence accumulates, possibilities are
eliminated and concepts become more homogeneous. Applicable evidence may
take many forms ranging from fossil discoveries to biochemical linkages or the
164
H.A. MILLER
ultrastructure of spermatozoids. The array of techniques now being applied to
bryophyte study and the ever increasing number of investigators attuned to highly
specialized investigations based on sophisticated technology has brought a
quantum leap in understanding of the total biology of the bryophytes. This is true
for every field of life sciences, but the impact is especially obvious in bryology
which historically was the province mainly of amateurs holding an essentially
philatelic philosophy on exchange of specimens. Few of the most mentioned
bryologists prior to 1950 were mainly employed for bryology. In the U.K., for
instance, H. N. Dixon was headmaster of a school for the deaf, S. M. Macvicar was
a physician, William Mitten an apothecary and so on. In North America, W. S.
Sullivant was an Ohio banker and land developer, A. W. Evans trained as a
physician but subsequently trained in botany becoming professor at Yale, A. J.
Grout taught at a high school in Brooklyn and E. B. Bartram was an early retired
Pennsylvania businessman. It seems generally agreed that A. J. Sharp of the
University of Tennessee was the first American hired actually to be a bryologist
along with other academic duties-and ‘Uncle Jack’, as he has come to be
respectfully known to hundreds of botany graduate students, is still actively
preparing a moss flora for Mexico. O n the continent, Max Fleischer, who devised
the basic system applied to the mosses was a famous Dutch painter; V. F.
Brotherus was lecturer in natural history and mathematics in a girls’ school in
Helsinki for 40 years; Franz Stephani retired early from the publishing business,
and other famous names followed careers in farming, the military, civil
engineering, wine-making, pharmacy, geology, medicine and a variety of other
professions. It was, therefore, largely a band of dedicated hobbyists that set
bryology in motion and laid the taxonomic-systematicfoundation. Bryology is thus
only now emerging as a multi-faceted biological discipline and serious gaps remain
in our understanding of the organisms.
( a ) Origin and macroevolution
Bryophytes are polyphyletic; they represent a level of evolution and their origins
seem to be clearly linked to both transmigration to the land and the beginning of
vascular plants. Evidence mounts that evolutionary experiments with
vascularization were well along in Silurian time, with development of a central
strand of specialized elongate water-conducting cells surrounded by a ring of foodconducting cells rich in plasmodesmata. Lowry, Lee & Hibant (1980) have
suggested that ultra-violet radiation limited terrestrial life until adequate oxygen
had formed in the atmosphere, and that an ozone shield developed such that
expanded foliar organs were evolved by mid-Devonion plants in several divisions.
Compounds synthesized by the land plants, but not true algae, include the
phenolics which absorb UV radiation. Palaeophytochemical analysis of
Eohostimella fossils has shown the presence of phenylpropanoids chemically allied to
lignin which is comprised of polymerized phenolics formed in an oxygenated
atmosphere. Many possible combinations exist; exact make-up and chemical
configuration of lignin remain unknown despite the fact that it is the primary
stiffening agent in most plant cell walls. Polyphenols are found in bryophytes as are
many flavonoid compounds, but reports of true lignin in the large mosses, e.g.
Dawsonia (Siegel, 1969) have been demonstrated to be spurious (Erikson &
Miksche, 1974; Hibant, 1977; Miksche & Yasuda, 1978). Thus, even though
BRYOPHYTE EVOLUTION AND GEOGRAPHY
165
lacking lignin, bryophytes do approximate a near-lignin as well as having a diverse
flavonoid chemistry (Markham, Porter & Miller, 1976, Markham & Porter, 1978,
1979). Whether the bryophytes have ‘true tracheids’ or just elongate, stiffened,
pitted cells-the ‘hydroids’-the fact is that in extant forms both tracheids and
hydroids have similar structure and function in the plant (Scheirer, 1980). The
correspondence is greater for mosses than for liverworts (Crandall-Stotler, 1980),
however, and augers strongly for a separate and perhaps delayed origin for mosses.
The hornworts are of quite another ilk but Proskauer (1960) detailed structure of
the Dendroceros columella and the incomplete cellular thickenings suggestive of
vascularization. Renzaglia ( 1978) compared the Anthocerotophyta with
pteridophytes and found that gametophytic organization was very similar but that
the sporophytes were markedly different. Other differences cited previously were
also noted.
This matter of vascularization and implied evolutionary connections to other
land plants has loomed large in phylogenetic thinking because of the philosophical
fixation on bryophytes either as separately derived from green algae, and thus alien
to vascular plants, or as degenerate rhyniophytes. Evidence, of sorts, exists for both
points of view and it is unnecessary to review the classical morphological
observations (e.g. Meeuse, 1967; Mehra, 1967, 1969) mustered for each case. Ifwe
focus attention onto similarities of bryophytes to other land plants, great
commonality is found. Photosynthetic pigments are the same, the gametangia have
a common plan, the same biosynthetic pathways seem to exist in their phenolic
chemistry and flavonoid chemistry so far known for mosses shows “levels of
biosynthetic evolution which far surpass the range of morphological
diversity”(Suire & Asakawa, 1979),stilbenoids occur in hepatics and angiosperms,
sesquiterpenes of hepatics are like those of higher plants and triterpenes and sterols
of bryophytes seem to be identical to those offerns and higher plants (Suire, 1975).
Thus, as Suire and Asakawa put it, “possibility of a direct line from algae to
bryophytes is not supported by any of the chemical data. On the contrary,
bryophytes seem more closely related to higher plants than to algae”. The view
that bryophytes generally are far removed from the algae was also put forth by
Smith (1978b) who viewed their origin as a result of evolutionary morphological
reduction. Steere ( 1969) considered bryophytes a ‘dead-end’ group derived from
the archegoniates; others, (e.g. Zerov, 1966) hold to direct algal origin. New
information derived from previously unexplored sources has permitted new
insights on the possible origin of bryophytes and conventional proposals, as above,
cannot hold. For now, the bryophytes seem best considered as non-generate
vascular plants (Miller, H., 1977) of diverse origin. Hepatics were probably
derived from the same early rhyniophytic stock that led to Cooksonia, with the
hornworts quite removed toward Horneophyton but also from rhyniophytic stock.
Mosses seem to have originated later from early ancestors of zosterophyllophytesperhaps from pre-microphyllophyte stock among the zosterophyllophytic complex.
Bold et al. (1980) have recognized the Divisions Hepatophyta,
Anthocerotophyta and Bryophyta in concert with Crandall-Stotler ( 1980), who
called special attention to the basic differences in apical organization between
mosses and hepatics and the sharp contrast in gametangial development between
hepatics and hornworts. Smith (197813) recognized only a single Division with five
equal classes placing Sphagnopsida and Andreaeopsida as taxa equivalent to
mosses, liverworts and hornworts. Although peat mosses and rock mosses are
166
H.A. MILLER
morphologically distinct, particularly the sporophyte, recognition as being coequal
with hepatics seems beyond evidence now in hand. The higher categories of
bryophytes can be summarized as follows:
Division Anthocerotophyta (1 Order)
Division Hepatophyta :
Class Metzgeriopsida (4 Orders)
Class Marchantiopsida (3 Orders)
Division Bryophyta :
Class Sphagnopsida (1 Order extant; 1 Order fossil)
Class Andreaeopsida (1 Order)
Class Bryopsida (12 Orders)
( b ) TaxonomJ, systematics and microevolution
Cytogenetics and diverse approches lumped under the umbrella of
biosystematics have been applied with varying intensity to bryophytes. Smith
(1978b) provided a very thorough review of the status of such studies and then
followed up (Smith, 1979) with an outline of the need for experimental work in
bryophyte taxonomy. Specialised reviews with evolutionary implications will be
found in Clarke & Duckett (1979). Taylor (1980) provided an intelligent overview
of evolutionary implications as well as a synopsis of a series of papers on the biology
of bryophytes. The significance of blepharoplast ultrastructure in bryophyte
spermatozoids (Carothers & Duckett, 1980) provided strong evidence for the
separation of hornworts from other bryophytes and Taylor viewed this work as a
more broadly applicable line of inquiry. She stressed, too, the need for better
understanding of bryophyte niches and the manner in which closely related species
occur within physically similar communities. It may be that subtle influences
confer a slight advantage for minor, but fixed, morpho-physiological adaptations.
Approximately 10% of bryophyte species have reported chromosome counts
(Fritsche, 1972); the numbers are generally low. The Anthocerotophyta have
reported base numbers o f N = 5 or 6 ; most counts in Marchantiopsida are N=9, or
a multiple, except in Ricciu, where N=8, 16 or 24 in all but a few cases. The
Metzgeriopsida have a basic number of N = 9 with few polyploids although the
enigmatic Tukakiu has N = 4 and 5. Sphagnopsida show N=21 (19+2m), whilst
Andreaeopsida are poorly known cytologically, the three species counted yielding
N = 10 and 11 ; Bryopsida show several cytological lines-i.e. the Polytrichales
with N = 7, 14 or 21, the Aplolepideae with N = 12, 13 and 14 (a probable basic
number of x = 7) ;acrocarpous Diplolepideae with N = 6 and 10 (the basic number
x =6), possibly excepting the Mniaceae; the pleurocarpous Diplolepideae with
N = 10 and 11 with a possibility that few are haploid if the the basic number is
x=6. Smith (1978b) has summarized cytological relationships ofBryopsida (Fig. 3).
Cytology, then does provide a basis for some broad systematic lines but for others
the case is not clear.
Being small plants with subtle phenotypic variation, and the persistent leafy
generation being gametophytic (haploid), bryophytes have not received great
attention for genetic studies. Neither of the two most recent major symposia (Suire,
1978; Clarke & Duckett, 1979) have dealt with genetics as such. Wettstein (1932)
covered principally the results of experiments in hybridization of mosses from the
Funariaceae in which he was able to recognize six characters which assorted in
BRYOPHYTE EVOLUTION AND GEOGRAPHY
Hopolepideae
Nematodontae
I
I
167
Arthrodontaao
Diplolepideae
I
Acrocorpae
I
I
Bryales
I
Pleurocarpoe
--
Figure 3. A phyletic scheme for the Bryopsida based on chromosome numbers. Heavy lines indicate
strong trends and light lines possible trends. Adapted from Smith, 1978.
spore-produced plants. Also found was evidence for maternal cytoplasmic
influence on the spores resulting from hybrid crosses of haploid gametophytes. The
few viable spores produced gametophytes with greater similarity to the maternal
parent.
Lewis (1961) provided a “geneticist’s view of bryophytes” in which he reviewed
sex determinative mechanisms, mostly not an XY chromosome system even though
that system was first discovered in plants in the hepatic Sphaerocarpos. The tendency
of bryophytes with one gametophytic chromosome complement (x) to be dioecious
and those with a gametophytic number of 2x to be monoecious was shown. With
loss of a chromosome to 2x - 1, however, dioecism is frequently established with a
resulting ratio, for an unknown reason, ofmore female than male plants. While it is
not difficult to see evolutionary implications from such phenomena, nothing was
reported of phenotypic variation of ordinary morphological characters.
The tendency towards very low spore viability from interspecific crosses has
been demonstrated by Anderson & Lemmon (1972, 1974), who reported that no
spores germinated from reciprocal crosses of Weissia (Astomum) ludoviciana x W .
controversa; earlier Reese & Lemmon (1965) had obtained a few viable spores for
such an apparent cross in nature, but whether the protonemata formed actually
matured to sexually functioning gametophytes was apparently unreported. It
seems clear that such interspecific hybrids are rare and that they form when gene
flow distances are very short. For hybrids of populations of Weissia controversa,
about 40mm was maximum with most lOmm or less, and dependent upon
contiguous colonies. Both Weissia and Climuium colonies produced no sporophytes
if bare earth existed between the plants. The Weissia hybrids were recognized from
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H. A. MILLER
the varying number of m-chromatids at meiosis. Phenotypic characters of the type
normally employed for studies of mendelian inheritance were not used because
“environmental influences on phenotypic expressions in mosses is not understood”
(Anderson & Lemmon, 1974). Many of the properties of higher plants used to
assay their genetic heritages cannot be (or have not been) applied to bryophytes
with success. The cytological approach to the genetics of bryophytes has so far
yielded most information.
Most textbooks indicate that bryophytes are ancient plants, as indeed they are,
and that they are so evolutionarily conservative as to be coming to the brink of
extinction. Some bryophytes, to be sure, have very restricted niches and are very
isolated phylogenetically from other taxa. Among mosses such genera as
Schistostega, Mittenia and Bryoxiphium seem vulnerable to extinction because of their
highly disjunct ranges and narrowly limited habitat. Parallels can be found among
the hepatics such as Haplomitrium, Takakia, Phyllothallia, Ascidiota and Riella.
Overall, however, mosses and liverworts have demonstrated a remarkable
adaptability to environmental extremes which exceeds that of the angiosperms.
Great phenotypic diversity is found among the Fissidentaceae, Dicranaceae,
Pottiaceae, Bryaceae, Hookeriaceae, Sematophyllaceae and Hypnaceae among
the mosses, and hepatics of the Lepidoziaceae, Plagiochilaceae, Radulaceae,
Porellaceae, Jubulaceae, Lejeuneaceae and to a somewhat lesser extent
Marchantiaceae, Ricciaceae and the hornworts, Anthocerotaceae. Because the
largest bryophyte families are mostly associated with, and adapted to, the many
niches that are peculiarly angiosperm linked-e.g. shade of broad-leafed, rapidly
transpiring trees; circum-neutral bark; rapidly decaying leaf litter ; rich
invertebrate fauna creating enhanced edaphic conditions-it is likely that
bryophytes are today at the greatest generic and specific diversity. Some few
isolated evolutionary lines seem doomed unless new adaptive strategies evolve, as
seems to have been the case with Sphagnum, but taken as a whole, the bryophytes
have proven highly adaptable over time and eminently successful components of
the earth’s vegetation.
MAJOR FEATURES OF BRYOPHYTE DISTRIBUTION
Limited floras and distribution of component species have been studied by
several contemporary bryogeographers. These studies have been reviewed or
summarized in the proceedings of the recent international bryological symposia in
Boulder (Anderson, 1973), Lille (Bonnot, 1974), Leningrad (Gadstein, 1976),
Bordeaux (Suire, 1978) and Bangor (Clarke & Duckett, 1979) in which most
attention is directed towards details of holarctic distributions and disjunctions.
Floras of New Zealand, Tasmania and circum-Antarctic islands, especially, have
been considered both from the standpoint of generic and specific disjunctions and
the presence of numerous ‘primitive’ hepatics (e.g. Grolle, 1969; Schuster, 1979a).
Among the tropical south Pacific islands, only Tahiti’s moss flora has been
subjected to analysis of floristic elements (Whittier, 1974, 1976). P6cs (1976a)
compared the east African bryoflora with that of Asia (1976b). Subsequently, he
(1978) analysed epiphyllous communities in east Africa and compared them with
epiphyllous floras of other continents. Recent floristic analyses of broad scope are
limited or wanting for Latin America, tropical Africa and Asia, Malesia, Australia,
New Zealand and most tropical Pacific islands.
BRYOPHYTE EVOLUTION AND GEOGRAPHY
169
From almost the first attempts at recognizing more or less floristically
homogenous geographical areas, the term applied to the most inclusive territory
was Kingdom. Lesser areas within a Kingdom have been designated as Regions,
portions of Regions usually being districts, although sometimes they are not
specifically designated, e.g. Good (1974). Herzog’s ( 1926) major floristic areas are
summarized as presented with inconsistencies of terminology preserved but with
Florenreich numbers corrected :
Die Florenreiche (after Hercog, 1926‘)
1. Holarktisches Florenreich
(a) Arktis
(b) Eurasisch-silvestres Vegetationsreich
(c) Ostasiatisches Vegetationsreich
(d) Zentralasiatisch-Pontisches Vegetationsreich
(e) Sindisch-Nordafrikanisches Vegetationsreich
( f ) Mediterrangebiet
(g) Makaronesien
(h) Nordamerikanisches Vegetationsreich
2. Neotropisches Florenreich
(a) Mittelamerika
(b) Die Hylaea
(c) Das Bergland von Guyana
(d) Das sudbrasilische Bergland
(f) Die sudlichen und westlichen Randgebiete
(g) Das chilenische ubergangsgebiete
3. Palaotropisches Florenreich
(a) Das tropische Afrika
(b) Die Indomalaya
(c) Ozeanien
4. Australisches Florenreich
5. Austral-antarktisches Florenreich
(a) Ostaustralien-Tasmanien-Neuseeland
(b) Die Notohyle Westpatagoniens
(c) Die antarktischen Inselgruppen und der antarktische Festlandsrand
6. Sudafrikanisches Florenreich
Rembering that Herzog’s system was based mainly on bryophyte distributions as
known over 50 years ago, the similarity of natural floristic areas he recognized
compared to those of, e.g. Good (1974), or Takhtajan (1969) established to
accommodate flowering plants is striking. Good’s system is summarized here for
comparison and general reference in discussions to follow.
Classijcation of the World in Floristic Units (after Good, 1974)
Boreal Kingdom
1. Arctic and Subarctic Region
2. Euro-Siberian Region
3. Sino-Japanese Region
4. Western and Central Asiatic Region
9
5.
6.
7.
8.
Mediterranean Region
Macaronesian Region
Atlantic North American Region
Pacific North American Region
I70
H.A. MILLER
Palaeotropical Kingdom
A . African Subkingdom
9. North African-Indian Desert Region13.
14.
10. Sudanese Park Steppe Region
1 1. North-east African Highland and 15.
16.
Steppe Region
12. West African Rain Forest Region
B. Indo-Malaysian Subkingdom
19.
17. Indian Region
18. Continental South-east Asiatic
Region
C. Polynesian Subkingdom
22.
20. Hawaiian Region
21. Region of New Caledonia (with
Lord Howe and Norfolk Islands) 23.
Neotropical Kingdom
28.
24. Caribbean Region
25. Region of Venezuela and Guiana 29.
30.
26. Amazon Region
27. South Brazilian Region
South African Kingdom
31. Cape Region
Australian Kingdom
32. North and east Australian Region 34.
33. South-west Australian Region
Antarctic Kingdom
37.
35. New Zealand Region
36. Patagonian Region
East African Steppe Region
South African Region
Madagascar Region
Region of Ascension and St
Helena
Malaysian Region
Region of Melanesia and
Micronesia
Region of Polynesia
Andean Region
Pampas Region
Region of Juan Fernandez
Central Australian Region
Region of the South Temperate
Oceanic Islands
The six kingdoms are derived from floras as they existed following the origin of
angiosperms and subsequently isolated during late Cretaceous and Tertiary time.
Individual regions correlate mainly with climatic regions, terrestrial physiognomy
and, to varying degrees, tectonic history. In a sense, then, today’s regions are
defined by an indigenous flora which is sufficiently diverse in its elements to create
a vegetational mosaic capable of saturating available niches within a contiguous,
or once contiguous, geobiotic zone. Thus, a floristic region may exist today on a
different terrestrial base than it once occupied or it may exist on the same
terrestrial base at a somewhat different latitude and relative position from other
floristic regions or it may exist on separated fragments of a once single terrestrial
base. As a flora migrates, some very niche specific, strongly adapted species may
not be eliminated by changing conditions and become relicts within another flora.
These relicts may be useful for tracing floristic history. Alternatively, as a flora
migrates some taxa may be lost and new taxa evolved having a superior adaptation
to changed conditions. If new land is created which is not contiguous to a flora, i.e.
within normal diaspore range, pioneers may be introduced by chance from several
sources so that a distinctive but disharmonious flora may develop as in the case of
Hawaii. For the most part, before pioneers become successful colonists they must
find niches within their biotic limitations. Genera are often only within a
latitudinal/altitudinal zone as well as a range of moisture and edaphic conditions
BRYOPHYTE EVOLUTION AND GEOGRAPHY
171
which broadly define geobiotic parameters within a floristic region. Lowland areas
of Hawaii then, for example, are populated by old world tropical bryophytes and
very high altitudes by temperate to subarctic taxa. Other examples abound.
The customary phytogeographical scheme in which holarctic floras are first
treated, as in the summaries above, is set aside here in order to focus upon special
attributes of the austral bryofloras which seem to derive in some significant
measure from the cool temperate lake regions of Permo-Triassic Gondwana.
The broken distributions of austral hepatic genera were brought to general
attention with the appearance of Fulford's (1951) landmark paper on South
American taxa of leafy liverworts, as then understood, in relation to world
distribution. It is recognized today that some generic and familial concepts which
were current in 1950 have been markedly revised since, but Fulford's fundamental
bryogeographic insights were on target. Her own taxonomic works on South
America, those of Hodgson on New Zealand, Herzog on Latin America and New
Caledonia, Grolle and Schuster on austral floras and, more recently, Engel on the
circum-Antarctic hepatics have done much to resolve apparent inconsistencies and
seemingly incongruous genera and families. The c. 180 genera of leafy hepatics
recognized in 1950 grew to c. 265 by 1968 (Schuster, 1969) and to c. 310 by 1980
(Schuster, 1979b; Schuster & Engel, 1981; Engel & Schuster, 1981). Fulford's
paper, first given at the Seventh International Botanical Congress (1950), stood
alone until articles by Schuster (1969) and Grolle (1969) appeared nearly two
decades later. In that span, the idea of moving continents became accepted in
contrast to the concept of fixed land masses which held sway into the early 1960s.
Distributional discontinuities did not change but their explanation became
decidedly easier with a basic understanding of plate tectonics (Gray & Boucot,
1979). Schuster has updated his initial paper several times (1972, 1976, 1979a,
1980) adding data from other groups or organisms and incorporating his on-going
systematic revisions and new genera of austral hepatics.
Except for limited floristic studies or occasional monographs, distributional
phenomena among austral mosses have not been broadly summarized since
Herzog's book appeared. Thus, distributional data presented here to sketch a
crude outline of the situation have been selected from taxonomic publications in a
very limited way. Total floristic analyses will be of value for interpreting floristic
dynamics of temperate and high altitude austral floras as the remote islands
become better known.
( I ) Antarctic Kingdom
The Antarctic botanical zone, comprising all land south of 60"s latitude plus the
South Sandwich Islands and Bouvet (Greene, 1964), is unique in that much of the
land is ice covered and the vegetation almost entirely cryptogamic. Antarctic
mosses include both strictly Antarctic species and some known from holarctic
regions. As a practical matter, the Antarctic floristic kingdom is much more
broadly defined by plant geographers to include New Zealand, the Patagonian
Region and the south temperate oceanic islands including Norfolk and Lord Howe
(Takhtajan, 1969; Good, 1974). Herzog (1926) included south-eastern Australia
and Tasmania in the Antarctic Kingdom on the basis of bryophytic distributions.
The limited land mass south of 45"s and free of permanent ice means that most
climates are oceanic, cool to cold temperate and exceptionally favourable for
development of substantial bryophytic vegetation.
I72
H.A. MILLER
Liverworts have not been found, to the best of my knowledge, on the Antarctic
mass itself but Marchantia p o h o r p h a was found in the 1969 crater on Deception
Island (South Shetlands) near the Antarctic Circle. Mosses have been collected at
numerous sites around the perimenter of Antarctica proper (Greene et al., 1970).
The hardiest known moss, the endemic Sarconeurum glaciale, occurs well above 70”s
and one station has been found in the Transantarctic mountains at nearly 85”S!
The best known of the moss floras of the Antarctic south of 50”sappears to be that
of South Georgia (Greene, 1974) with 51 species recognized. Among the South
Georgian mosses, Polytrichum alpestre, P . alpinum, P . juniperinum and P . piliferum are
all widely distributed in the holarctic. Among the other 23 moss genera reported
for the island, some have ‘bipolar’ species, endemics or circum-austral types first
described from New Zealand, Fuegia or other circum-Antarctic sites. The moss
flora of the Antarctic peninsula includes the same species of Polytrichum as known
from South Georgia. Three Andreaea species are known: A . defiessinervis is endemic
to the Antarctic up to the South Orkneys (Schultze-Motel, 1970); A . regularis is
similarly localized with one station in the South Sandwich Islands; and A . gainii
extends up to the South Sandwich Islands with a report from Kerguelen (as A .
parallela) by Roth (1911). Both Pohlia cruda and P. nutans are holarctic as is B y u m
argenteum which I have seen from McMurdo.
High latitude hepatics from the southernmost extension of the South American
continent, the Brunswick Peninsula, have been very thoroughly reviewed by Engel
( 1978) who recognized several phytogeographical categories. No species were
reported from the Antarctic (sensu stricto) and only four species from the subantarctic zone--Cephalozia badia, Herzogobryum erosum, Leptoscyphus abditus and
Riccardia georgiensis. The distinction between the subantarctic zone which may be
found at high elevation at lower latitudes and the cool south temperate region is
sharply drawn so that the latter includes the majority of species considered as
“antarctic” (Fulford, 1951), “antipodal” (Schuster, 1963) and “subantarctic”
(Grolle, 1969), by other authors. Only 24 species of the 193 recognized for the
Brunswick Peninsula were considered to be extra-American temperate forms and
only five as being pan-temperate : Acrobolbus ochrophyllus, Bazzania nitida, Crytochila
grandgora, Jamesoniella colorata (bipolar) and Marchantia berteroana. Hassel de
Menendez (1977) brought the known hepatic flora of South Georgia to 59 species
including seven additional species common with Kerguelen, three common with
New Zealand, several pan-austral species and one, Schistochila alata, extending to
Table Mountain, South Africa, representative of the disjunctive nature of the cool
south temperate hepatic flora. Examples of disjunct pan-austral taxa are
Blepharidophyllum densifolium, Cryptochila acinacifolia, Lepidoria laevifolia, Leptoscyphus
abditus, Noteroclada conzuens and Pachyglossa j s s a . On a broader scale, Gackstroemia
has five species in southern South America, one in New Zealand and Tasmania
and one restricted to Campbell Island ; Phyllothallia has two closely related species,
one in New Zealand and one in Fuegia; Clasmatocolea is comprised of numerous
austral species (Engel, 1980); and Acrolophozia is comprised of a disjunct species
pair. Further examples are numerous with many being discussed by Schuster
( 1979a).
Mosses illustrate similar phenomena to those noted for hepatics in the far
southern hemisphere with Acrophyllum (Pterygophyllum) (several species) and
Goniobryum subbasilare in each of south-eastern Australia, New Zealand and the
Chilean rainforest. A similar pattern, somewhat expanded, can be seen for
BRYOPHYTE EVOLUTION AND GEOGRAPHY
I73
Rhacocarpus (22 species) which additionally occurs in Madagascar, east African
mountains and in montane regions of Latin America. The monotypic
Dendroligotrichum dendroides occurs in the wet forests of Chile, Juan Fernandez and
New Zealand. Two of three species of Weymouthia occur in Chile, New Zealand,
Tasmania and Australia with the third being limited to South America.
Endemism is not infrequent on the major land masses of the cool south
temperate region at both the generic and specific level. For both the mosses and
hepatics, many of these area-limited forms manifest morphological characters
generally considered to be ‘primitive’. Thus, many of the leafy liverworts have
underleaves nearly as large as the lateral leaves as well as being overall somewhat
more robust than holarctic Jungermanniideae. The largest mosses known,
Dawsonia spp., are found most abundantly and with maximum diversity in
south-eastern Australia and New Zealand, with other taxa well represented in cool,
oceanic temperate climates extending northward into high elevations of New
Guinea with few species extending further, Examples of large mosses include
Dicnemon, Mesotus, Eucamptodon, Psilopilum, Dendroligotrichum, Polytrichadelphus,
Porothamnium and Breutelia, not all of which are restricted to the circum-Antarctic
temperate regions.
At least for the bryophytes, the limits of the Antarctic Kingdom cannot be
sharply defined on the basis of total floras. Every bryoflora within the geoclimatic
limits of the oceanic, cool, south temperate to Antarctic land masses has bipolar,
warm temperate, subtropical and tropical (high elevation) elements present
further complicated by the east-west elements among South America, New
Zealand, Tasmania, Australia, Kerguelen, the South African Cape and the
southern islands. In practice, the diverse floristic elements can be recognized and
set apart in a geograpically centred floristic analysis. There is however, a
substantial, perhaps majority, residue of geographically limited and often circumAntarctic, genera and species (even a few families) which is unmistakably whole
and distinctive in its character.
(2) Australian Kingdom
Within the geographical limits of the Australian Kingdom, i.e. forested eastern
and northern Australia, Tasmania, south-west Australia and central Australia,
some elements of the Antarctic bryoflora are clearly well represented. The
distinctiveness of the angiospermous flora and the marsupial fauna of Australia set
the continent well apart from the rest of the world. Limits of an Australian
bryoflora are much less easily stated. I n Tasmania and the south-east, many
temperate antarctic taxa are abundant. The drier areas of Victoria, South
Australia and Western Australia have a limited bryoflora which seems to have
decided floristic affinities with South Africa (Scott & Stone, 1976). Pleurocarpous
moss taxa are few in these drier areas and ephemeral acrocarpous forms, several
seemingly endemic, are well represented. Only about 140 mosses are listed in Scott
and Stone for either South Australia or all ofWestern Australia where tropical forms
might be expected in the northern part. Species common with South Africa are
found in the genera Fissidens, Eccremidium, Pleuridium, Bruchia, Aloina, Bryobartramia
and Gigaspemurn. A new treatment of the mosses of South Australia (Catcheside,
1980) has been announced but was not available as this was written. The hepatic
flora of the southern and western regions is quite limited. Many Riccia species as
174
H.A. MILLER
well as several Asterellas and Targionia hypophylla comprise much of the flora
although the unusual ephemeral endemics Petalophyllum preissii and Fossombronia
intestinalis are known from the Swan River area. The low places where water
collects in temporary ponds, which may leave a salt pan upon evaporation, support
populations of the desert aquatic liverwort Riella and the monotypic Carrpos
sphaerocarpos-both
Riella and Carrpos occur in similar sites in South Africa
(Schelpe, 1969).
Palaeotropical floristic elements are strongly represented in northern Australia,
especially Queensland. Complete bryofloristic compilations are not available for
Queensland or any of the Malesian and Pacific islands. Hundreds of papers have
appeared over the past 30 years which make casual mention of one or a few species
making the assembly of even a preliminary flora a considerable task. The literature
on hepatics is particularly dispersed but some summaries are available for mosses.
Our experience with the tropical Pacific insular floras has been that distributional
phenomena are approximately equivalent for mosses and liverworts so the majority
analysis here, although moss based, is thought to be representative. Scott & Stone
(1976) stressed the flora of temperate southern Australia but did list other species
reported from the country without evaluation. A rough count of the Queensland
citations gives 130 genera and just over 300 species--surely well under the actual
moss flora of tropical Australia. Schultze-Motel (1963) compiled a list of New
Guinea mosses with brief notes on extra-New Guinean distribution. Of 832 taxa
listed, approximately 65 are attributed to Australia (some others are indicated only
as cosmopolitan or pantropical). The 20% of the tropical Australian flora shared
with New Guinea is clearly somewhat understated as the number of common
genera represents a much greater share of the flora. A crude tabulation of genera
noted by Scott and Stone for all Australia compared to the Schultze-Motel list for
New Guinea shows that 50-60°/,of the Australian genera occur in New Guinea.
This is consistent with the observations of Scott and Stone that the affinities of
Queensland are with the Malesian flora to the north.
Herzog proposed that Lord Howe Island and New Caledonia be included in the
Australian Kingdom. New Caledonia has about 550 specific taxa of mosses of
which some 60% are regarded as endemic (Miller, Whittier & Whittier, 1978).
About 20°4 of the Queensland flora also occurs on New Caledonia or, put the
other way, a little less than lo0{, of the New Caledonian flora is congruent with
Queensland. It is about equally matched to the Fiji flora. The 80 species on Lord
Howe are about 25% endemic with strong representation of south-eastern
Australian taxa and nearly 4OCY/,of the species are common with New Caledonia.
At the generic level, some 7O0In of the Queensland genera occur in New Caledonia
being 50°/, of the generic flora of that island. About 80°/, of the Lord Howe genera
are on New Caledonia or, alternatively 2514 of New Caledonian genera occur on
Lord Howe but that is not strikingly different from several other surrounding
areas. In all, Queensland and Lord Howe have some floristic relationships with
New Caledonia but, by numbers alone, the case for new Caledonia being
Australian is not especially strong.
(3) South African Kingdom
The geographical limits of the South African flora have been variously drawn.
Herzog had a very wide concept extending as far north as Rhodesia (Zimbabwe)
BRYOPHYTE EVOLUTION AND GEOGRAPHY
I75
and South West Africa. His limits seem to have been determined more by the
mediterranean climate of extra-tropical areas rather than by strictly floristic
considerations. However, even reasonably reliable data were not available on
specific localities at the time he was working-Sim’s (1926) “The Bryophyta of
South Africa” had not yet appeared, old lists were very unreliable and the
Brotherus treatment was very inexact about distributions. Since then, Arne11
(1963) has provided a new treatment of the hepatics which added new discoveries
as well as clarifying the status of a great many species. He treated 298 species with
the notation that if South Africa, South West Africa and Rhodesia were considered
together, then 153 species could be considered endemic.
A tabulation of the hepatics found on Table Mountain and in the east-west fold
ranges south of a line from Port Elizabeth to Clanwilliam, as proposed by
Takhtajan (1969), yields a picture of the austral nature of the Cape flora. More
than 30 of about 130 species reported from the region are very local endemics.
Several species are disjunct from temperate South America such as Qmphyogyna
podophylla, Clmatocolea vermicularis, Adelanthus sphalerus, Jamesoniella colorata, 3.
grandijora, 3. paludosa, J . oenops, Acrobolbus excisus, Lepicolea ochroleuca, Hyalolepidoria
bicuspidata and Sphaerocarpos stipitatus. Cape species from the temperate circumAntarctic islands, but not South America, include Lethocolea congesta, Adelanthus
uncgormis, and Marsupidium brevifolium. Grolle (1971) noted that four of 36 species of
hepatics on Marion and Prince Edward Islands occurred at the Cape. The unusual
Carrpos described from the salt pans of Australia has been discovered recently in
South Africa (Schelpe, 1969).
The catalogue of bryophytes of southern Africa by Magill & Schelpe (1979)
provides a checklist of proper names for South African bryophytes along with
corrections of erroneous reports and taxonomic synonyms. Genera of mosses
endemic to the southern African area include Nanobryum and Wardia, each the sole
genus in a family, Hypodontium of the Calymperaceae, as well as Hypnofabronia,
Zschyrodon and Leptoischyrodon all of the Fabroniaceae. In addition to the pygmy
mosses shared between South Africa and Australia, some species of Lethocolea,
Hedwigidium and Rhacocarpus are also shared. The relationship of the moss flora to
South America is shown by the presence of Eustichia, Rigodium and Dimerodontium.
Sibling species are described for the Sierra de la Ventana and Sierra del Tendil
mountains south of Buenos Aires and the fold mountains of the Cape. About a
dozen mosses of the 80 known from Marion and Prince Edward Islands also occur
in the Cape Region (van Zanten, 1971) indicative of the circum-Antarctic
elements in the Cape moss flora.
The peculiarities of the phanerogamic flora of the African Cape have been
interpreted repeatedly as the result of a long period of northward migration of the
African continent with a retreat of the Gondwanan-originated flora to the tip of
the continent. Somehow this doesn’t add up for bryophytes. If the flora was in
retreat, why weren’t more Gondwanan bryophyte relicts left behind? Why are the
east African mountains floristically more like India than the Cape fold mountains?
Why are the Sierra de la Ventana (South American) disjuncts mostly limited to
the Cape region? Why, even using Herzog’s and Arnell’s or Magill and Schelpe’s
“southern Africa’’ limits, is there such a high endemism rate? It cannot be entirely
an artifact of inadequate taxonomy. All things considered from the standpoint of
the organisms involved, the evidence can be interpreted to mean that the Cape
fold ranges south of a line from roughly Port Elizabeth to Clanwilliam represents
176
H.A. MILLER
an island which was left behingl and later caught up with the primary continental
block after the continental ecosystems were essentially closed. Thus out-migration
from the Cape Island would be severely limited resulting in the continuing insular
aspect of the flora and in-migration would be limited from the north because of
parallel niche saturation. Recent discoveries of island archipelagoes (now landlocked) in Idaho confirm the tectonic possibility for a long separated Cape Island.
On such a basis, the flora suggests an estimated arrival time in late Cretaceous or
early Tertiary.
( 4 ) Neotropical Kingdom
The Neotropical Kingdom includes the West Indies and all but the southern
extremities of the South American continent. On the continent itself, the three
major mountain complexes have somewhat distinctive floristic elements present
which essentially overlay the pan-tropical and generally neo-tropical generic flora
which is not especially useful as we presently understand it for reconstruction of the
history of a particular floristic area. The Latin American region is very large,
geographically complex, climatically diverse, and the closest extant nearly
continuous land mass to the ancient Gondwanan refuge. It was the last route
available for antarctic floras to escape to the warmth of the north. The result is that
the cool, mesic mountain forests of the Latin American tropics harbour a most
diversified flora resplendent with many relicts from pre-Cretaceous times. Their
distributions are not uniform and seem to reflect the variation in available
migration tracks and the origins of the mountain regions.
The flora of the West Indies and Central America have much in common in that
the tropical species represented are those from both the north and the south which
seem to have efficient diaspores. High volcanic peaks in Central America and the
highest elevations on Hispaniola have essentially holarctic bryofloras while the
composition of floras of the middle to lower altitudes, below about 1800m, is
decidedly neotropical. Mosses of Central America have been collected by many
North American botanists working in the area with the result that every country
except Mexico has had a checklist formally published within the past few years
(Crum, 1952; Winkler, 1965; Crum, 1967; Breen & Reese, 1971 ; Bowers, 1974;
Delgadillo, 1979; Steere, 1979a). Hepatics may have been equally collected but
their study has not been on a country by country basis and no one has specialized
in Central American liverwort floras. The Guatemalan moss flora is best known
with over 500 species and an equivalent number of hepatics would be expected.
Bartram (1949) noted that the lowland flora up to about 1500m was
representative of Caribbean regions. A zone from 1500 to 3500m has “surprising
vagrants from far distant northern and southern latitudes.” Mountain summits
3600m and higher are truly alpine. The role of the Cordilleran track and the
origin of its flora is suggested by Bartram’s observation that “strange mixtures
present a puzzling problem in phytogeography, especially when two species of the
same genus such as Ditrichum giganteum of northern United States and Yukon and
Ditrichum gracile of the South American Andes are found growing in close
proximity”. All indications are that the isthmus of Central America constitutes a
migration route for both North American and Andean taxa but that it has not
always been present.
BRYOPHYTE EVOLUTION AND GEOGRAPHY
177
With the exception of some high ranges on Hispaniola, the West Indies have no
mountains exceeding 1500m. The floras are accordingly typical lowland
neotropical or Caribbean with the best development of bryophytes on those islands
high enough to have a cap of cloud forest usually manifest as elfin forest (Howard,
1968). The floras of the West Indian islands are unevenly known. The Jamaican
moss flora of 328 species has been elucidated and keys provided by Crum &
Bartram ( 1958) and Crum ( 1965). A moss manual for Puerto Rico by Crum and
Steere (1957) provides diagnoses of 268 species and varieties and de Foucault’s
( 1977) Guadeloupe flora lists 221 mosses and 273 hepatics. The moss floras of these
islands are not only of about the same extent but are composed of a majority of the
same taxa, most of which also occur in Central America and in lower elevations of
northern South America.
The Guiana highlands of Venezuela, northern Brazil and the Guianas, the ‘lost
world’ country, remains imperfectly explored but has been opened recently by
mining interests. The table mountain ‘tepuis’ of interior Venezuela have yielded
distinctive primitive hepatics including the Cladomastigaceae and
Trabacellulaceae known only from that area (Fulford, 1967). Mount Roraima has
a distinctive high altitude bryovegetation including such species as Breutelia
scoparia, Rhizogonium lindigii and Eopleuroeia paradoxa. As for Central America, the
low altitude bryoflora is essentially Caribbean (Florschutz, 1964).
The Brazilian highlands south of the Amazon are drier than the tropical forest
regions of the great basin but isolated peaks rising above the plains frequently
support a bryophyte-rich forest cap (R. Harley, unpublished) of unknown floristic
make-up. Herzog considered in detail only the mountains of the wet southern
Brazilian provinces from the 20th to 30th parallels recognizing two floristic
elements-( 1) a wooded mouhtain forest element and (2) a subantarctic alpine
element at high elevations. Genera limited to, or characteristic for, the area
include Moseniella, Spiridentopsis, Philophyllum, Puiggarella, Meiotheciopsis and
Cladastomum. The Andean tropical rainforests from Venezuela ( Pursell, 1973),
Colombia (Florschutz-de Waard & Florschutz, 1979; Gradstein & Hekking,
1979), Ecuador (Steere, 1948; Robinson, Holm-Nieslon & Jeppesen, 1971) ,
Galapagos (Weber, 1966), Peru (Hegewald & Hegewald, 1975) and Bolivia
(Hermann, 1976) southward into northern Argentina (Kuhnemann, 1938)
contain a bryoflora of enormous diversity. Many genera are endemic to the region
including Schliephackea, Stenodictyon, Amblytropis, Stenodesmus, Uleobryum, Schroeterella,
Flabellidium and Herzogiella. Endemic hepatic genera include Myriocolea, Mytilopsis,
Chaetocolea and Stephaniella. The eastern slope of the Andes and the Amazonian
rainforest can be characterized by such systematically isolated and monotypic
mosses as Phyllodrepanium, Hydropogon and Hydropogonella and similarly distinctive
hepatics such as Anomoclada, Protocephaloeia, Pteriopsiella and Micropterygium. The
high mountains are somewhat drier with numerous xeromorphic forms including
Pseudocrossidium, Gertrudiella, Rhexophyllum, all Pottiaceae, as well as Polymerdon,
Aligrimmia and Mandoniella. Hepatics are fewer and tend to be drought-tolerant
marchantiopsids such as Riccia, Clevea, Sauteria, Asterella and Targionia.
Peculiarities of climate and geography of southern South America bring the
neotropical flora into proximity with Antarctic elements. The southern Chilean
moss flora is a combination of widely distributed families and distinctly austral
types (Mahu, 1979). Rainfall drops sharply south of Buenos Aires to the east of the
Andean ridge to create grasslands and steppe from about 35”S, roughly the
178
H.A. MILLER
latitude of San Francisco and Gibraltar, southward to the Straits of Magellan. The
only bryophytes present in this vast region of southern Argentina are those adapted
to near-desert conditions such as Bruchia, Archidium and several Pottiaceae. Some
high altitude tropical mosses extend southward along the Andes to the northern
reaches of the southern Chilean rainforest with its strongly Antarctic bryoflora.
Few temperate habitats are available which parallel those of western Europe or the
eastern U.S.A. The Sierra de la Ventana, north of Bahia Blanca, receives more
rainfall than the surrounding lowlands and supports a limited Antarctic bryoflora
with some noteworthy disjunctions from the African Cape region such- as
Dimerodontium.
(5) Paleotropical Kingdom
The geographical extent of the Palaeotropical Kingdom embracing Africa,
India, Malesia and the tropical Pacific islands is so great that three Subkingdoms
are properly recognized-African, Malesian and Pacific Island or Polynesian.
The great deserts of north Africa across the Arabian peninsula into central Asia
now present a formidable barrier to bryophytic migration and serve to isolate the
present palaeotropical and holarctic floras except along the Pacific coast of Asia.
Even so, the broad spectrum of common genera of both mosses and hepatics
reflects either a time of active floristic exchange or origin from a single floristic
area. Genera of mosses and liverworts in tropical Africa are mostly pan-tropical,
holarctic or palaeotropical. Diversity is significantly less in tropical Africa than in
either Latin America or Malesia despite extensive rainforests and numerous
mountain ranges. Jones (1980 et piior) has found many tropical African hepatics to
be conspecific with those of the Caribbean area. Pbcs’ (1976, 1978) large
collections from Tanzania and immediately adjacent areas of Kenya and Zambia
have shown substantial numbers of taxa in common with India. De Sloover’s
hepatic collections from Zaire, Rwanda, Burundi and RCunion (VBiia, Pbcs &
Sloover, 1979) included 18% species also in Latin America and 20% species also in
Malasia of which about 10% of the total species reported are best considered pantropical. The only strictly African hepatic genus reported was Sprucella. Mosses of
west Africa were catalogued by Schultze-Motel (1975) who listed 964 species for
the area south of the Sahara, east to Chad and south to the Congo basin. Of the
genera listed, only Nanobryum, Bryotestua, Tisserantiella, Jonesia, Rhachitheciopsis,
Rhizofabronia and Pylasiobryum are strictly African, although many species of the
predominantly holarctic and pan-tropical genera are limited to Africa.
The tropical forests of India, the Indochina peninsula and the Indonesian
islands from the Andamans to New Guinea and northward to Taiwan, collectively
comprising Malesia, are of the greatest extent in distance of any of the world’s
three great tropical angiospermous forest regions. The discontinuities owing to the
fragmentation into large islands, island groups and the widely separated
occurrence of very high mountain peaks have created conditions favouring
development of great systematic diversity, The richness of the moss flora can be
seen in Fleischer’s (1904-1923) massive Musci der Flora von Buitenzorg which,
in fact, covers more than just the mosses of a part of Java. It remains the single
most important reference for the Indonesian islands. A moss flora of eastern India
is being compiled by Gangulee (1969-1978 et seq.) incorporating an illustration and
distributional map for each species using the “Index Muscorum” (Wijk, Margadant
& Florschutz, 195S1969) base map for widely distributed taxa. Touw (1978) listed
BRYOPHYTEEVOLUTIONANDGEOGRAPHY
179
607 species of mosses for Borneo. Philippine mosses were treated by Bartram ( 1939)
with recent additions to the flora by Iwatsuki & Sharp (1968) and Iwatsuki &
Noguchi (1978). Wang’s (1970) study of Taiwan mosses lists 61 1 species, surely an
understatement when the flora becomes better known. Much attention has been
drawn to New Guinea as a result ofimproved access to its remote interior but so far
only Schultze-Motel’s ( 1963) checklist of 832 moss species, which antedates recent
expeditions into the area, is available. Thece floristic compilations suggest a
broadly distributed and diversified generic flora with a combination of generally
dispersed species at low altitudes, perhaps to as much as 2000m, and disjunctive
taxa or locally endemic species more numerous at higher elevations where
geographical isolation is greater and the area of suitable habitat is much restricted.
On the highest peaks and ridges above c . 3000m, north temperate to subarctic
species are well represented with typically antarctic species, or close allies, also
present as demonstrated by this list of examples from New Guinea with reported
altitudes noted:
Holarctic Species
Sphagnum subsecundum, 2400 m
Andreaea rupestris, 4250 m
Encalypta vulgaris, 4 150 m
Tayloria mnioides, 3000 m
. Meesia triquetra, 3225m
Marsupella revoluta, 4000 m
Acrobolbus ciliatus, 4100 m
Blepharostoma trichophyllum, 2500 m
Reboulia hemisphaerica, 3000 m
Austral Species
Sphagnum antarcticum, 3400 m
Tcyloria octoblepharum, 2750 m
Zygodon hookeri, 3400 m
Rhacocarpus humboldtii, 4250 m
Dawsonia superba, 2800 m
Gymnomitrion incompletum, 4000 m
Pseudocephalozia leptodicpon
(aJJ lepidzioides), 3800 m
East of New Guinea and the Philippines, the tropical Pacific islands of
Melanesia, Micronesia and Polynesia collectively have a flora mostly derivative
from Malesia but, as recognized for flowering plants, the various archipelagoes are
not homogeneous. The coastal plains of the high islands and the atolls have floras
comprised mainly of epiphytic Calymperaceae (Calymperes, Syrrhopodon,
Mitthyridium), Leucophanes, Taxithelium or other Sematophyllaceous genera and
Lejeuneaceae (mostly Cheilolejeunea, Lejeunea, Microlejeunea and Cololejeunea) . Coral
rocks support Pelekium and Thuidium with Ectropothecium being found mostly on
sand in lightly shaded mesic areas. All produce great numbers of diaspores.
Comparatively modest elevations of about 500 m are sufficient to generate
orographic rainfall adequate to support a mesic forest. Only slightly higher
elevations are required for development of an elfin cloud forest on peaks and
ridges. It is from the cloud forests that both the greatest diversity and bryomass are
obtained. It is here, too, in the cool, wet, mountain woods that the anomalous taxa
occur. Malesian types comprising the majority of taxa collectively show evidence
of chance dispersal, in the reduced numbers of genera and the presence of different
genera on isolated islands. In addition, austral elements occur which suggest a flora
which perhaps has existed on one island and then another for a very long time.
Skottsberg recognized this “Paleantarctic” floristic element years ago ( 1936)
among the angiosperms. Schuster ( 1979a) has discussed distribution of selected
antipodal hepatics as indicators of the unity of Gondwanaland, but did not
specifically address Pacific island floras, save a few species that reach New
I80
H.A. MILLER
Caledonia. The New Caledonian bryoflora is not quite as unusual as is the
angiosperm flora, but numerous genera present are otherwise only austral rather
than Polynesian-Malesian. Hawaii is the most isolated, topographically diverse
land mass in Polynesia. With few exceptions, the flora up to about 1500m is
comprised of Malesian genera and Polynesian or endemic species. Above that,
Holarctic species begin to appear and at the highest elevations on Haleakala,
Mauna Loa and Mauna Kea, the very limited bryoflora is almost wholly of taxa
also found in North America.
(s) Boreal Kingdom
The holarctic lands of Europe, North America and extra-tropical Asia comprise
today a nearly contiguous land mass long isolated from southern continents by a
combination of geographic and climatic barriers. Although these land masses have
been fragmented variously since Palaeozoic time and some floristic evidence for
these schisms remains, Quaternary glaciation has had a profound effect upon
modern distributions. Steere (1978, 1979b) estimated that 7580% of the arctic
bryophyte flora of North America is comprised of the circumboreal element and
some 15% is of arctic-alpine relicts in areas not glaciated. A few otherwise
temperate species and some temperate desert taxa occur disjunctly in small nonglaciated refugia. North American mosses have been catalogued by Crum, Steere
& Anderson (1973), those in Canada by Ireland et al. (1980); the hepatics have
been listed by Stotler & Crandall-Stotler (1977). Schofield (1980) analysed the
geography ofNorth American mosses as listed by Crum et al. (1973) recognizing 20
well-segregated geographical patterns based upon distributional data for about
1200 species. Ten of the patterns are strictly North American and 10 are of
disjuncts between North America and other regions.
Relationships between the temperate floras of North America and eastern Asia
have been reviewed by Iwatsuki (1958) for Japan and the southern Appalachians
and for North America by Iwatsuki & Sharp (1967) and Sharp (1972).Numerous
disjunct and vicariad mosses are known. The essentially holarctic nature of the
Himalayan bryoflora is considered by Sharp (1974) to be the result of the loss of
the ancient flora of India during its northward movement through desert and
tropical latitudes following separation from Gondwana. Although the bryoflora of
Japan is well known and Korea has been much studied as well, central Siberia and
nearby northern Mongolia eastward to Khabarovsk Territory and the Sikhote
Alin mountains are still quite imperfectly known bryologically. Some indication of
the present moss flora of the Lake Baikal and upper Amur basin was given by
Bardunov (1969) who reported only five genera not listed by Crum et al. (1973) for
North America-i.e. Cephalocladium, Erythrodontium, Indusiella, J a f f l i o b y m and
Thamnium. I have, in the vicinity of Khabarovsk, seen several genera which were
not listed in Bardunov’s flora of a region hundreds of kilometres to the west. The
forest region from Mount Belukha and the Altai Mountains north of the Gobi
desert, south from the 60th parallel, and eastward to the sea, is surely the least
known of the north temperate bryofloras and deserving of attention.
European and North American bryofloras are closely allied to the extent that
the new “Moss Flora of Britain and Ireland” (Smith, 1978a) can be used with
considerablesuccess to identify mosses of the north-eastern United States. The same
can be said for Miiller’s (1954-1958) treatment of European hepatics. This
BRYOPHYTE EVOLUTION AND GEOGRAPHY
18 I
considerable coincidence of taxa between the north Atlantic land masses surely
demonstrates their once greater proximity and common flora. All in all, the boreal
bryoflora is remarkably homogeneous above 60”N. In temperate latitudes the
flora of eastern Asia differs significantly from Euro-American floras, but the high
generic and specific commonality among them profoundly documents their single
heritage.
SOME PROBLEMS AND APPROACHES FOR BRYOPHYTE GEOBOTANY
Limitations of the fossil record will always be a problem but the almost
incredible progress in palaeobotany of bryophytes over the past 15 years can be but
a harbinger of what is to come. New methods have enhanced the possibility for
detecting bryophyte remains, although aroused interest and sensitivity of
palaeobotanists is probably even more significant. Expansion of information about
Permian (e.g. Fefilova, 1978) and Mesozoic bryophytes (e.g. Douglas, 1973;
Anderson, 1976) can only result in a better understanding of the evolution and
distribution of bryophytes than at present from known Cenozoic floras.
Most biogeography today is undertaken by taxonomists with interest limited to
either mosses or liverworts which comprise a delineated flora which, in turn, may
be compared to another on a species by species basis. This approach leads to lists of
shared species, of species expected but not found, or of species pairs followed by an
interpretation or reason for the observed phenomena. The explanation may be
climatic, tectonic, edaphic, ecological, historical or a mixture thereof. Limitations
of this approach become evident as both the geographical area and the correlated
flora is expanded. At some point, the usual procedure is to reduce the taxonomic
1971) or to select
coverage to a family or genus as monographers do (e.g. TOUW,
examples of one or more distributional patterns as indicative of special floristic
situations (e.g. Schofield & Crum, 1972; Steere 1979; Schofield, 1980). Clearly,
even an analysis based mostly upon the literature cannot be accomplished by these
conventional means on a world-wide basis with numerous geobotanic subunits,
over 1 100 genera and about 25 000 species of even the bryophytes.
Some statistical approaches have been made to floristic analysis, usually based
upon numbers or percentages of species or genera common between geographical
regions. In 1969, van Balgooy reported an analysis of Pacific island floras utilizing
Kroeber’s Coefficients of Correlation which provide a numerical value
representing a degree of similarity between two islands even if the floras are quite
unequal. In order to develop a Kroeber Coefficent it is necessary to count the total
number of equivalent taxa in each of the areas to be compared and the number of
shared species in each area. For small numbers of comparisons, hand sorting and
counting is possible as Whittier (1974) did for French Polynesia although he
calculated only percentage shared and not similarity coefficients. For large
numbers of areas and taxa, computer data manipulation becomes the method of
choice and allows several kinds of analyses from a single data bank in a minimum
of time.
Accordingly, a small group at the University of Central Florida (Whittier et al.,
1978) set up a program to convert taxonomic and geographical information from
the files for the “Prodromus Florae Muscorum Polynesiae” (Miller, H., Whittier &
Whittier, 1978) to IBM cards for analysis and rapid information retrieval. The
program allowed sorts at the familial, generic or specific level as well as
182
H.A. MILLER
geographical sorts for 105 insular areas and 58 extra-Pacific regions incorporating
the rest of the world. These 163 regions are also grouped in 40 coarser areas which
are more easily assimilated during the cross-checking and ‘debugging’ phase of the
project. A fringe benefit of this approach has been the ability to obtain a printout,
either in systematic or alphabetic order, of any or all of 105 floristic lists complete
to date and 58 lists of species known from the Pacific as they occur in diverse extraPolynesian (s. 1.) regions. The trial run of complete lists for each of the 163 areas
took the computer 600 seconds. However, over a year had gone into key-punching,
programming, checking and correcting both the data base and the program before
the 62 families, 254 genera and 1427 species could be sorted into each of the floras
where they belonged. Since then, the data bank has been extended geographically
to include more western Pacific mosses and some 53 families, 155 genera and 1516
species of Polynesian hepatics (Miller, H., Whittier & Whittier, 1982).
Before undertaking programming for data manipulation (save for “how many,
and which, species found on A occur also on B?”) a mathematician was brought
into the team to evaluate methods for development of correlation coefficients
including Jaccard’s, Soerensen’s, Exell’s, Van Steenis’ ‘Demarcation Knots’ and
Kroeber’s. For our purposes, Kroeber’s Coefficient of Correlation :
50C (A+B) ,
AB
where A=number of species at A; B=number of species at B; C=number of
species common to A and B; was recommended as the method of choice. The
computer was programmed to print ‘mileage chart’ matrices which show the
number of species for each area on the upper left to lower right diagonal with the
number of shared species above and Kroeber Coefficients below. The similarity
coefficient matrix for 40 x 40 geographical groups based upon 1427 Pacific island
moss species is reproduced in Fig.4. A coefficient of 100 indicates complete
similarity and zero indicates absolute dissimilarity. In the expanded matrix,
comparisons at the generic level have been illuminating for large floras as have
species comparisons with, e.g. endemics eliminated.
Guttman-Lingoe’s Smallest Space Coordinates show separation or clustering of
floras by presumed similarities by placement of the geographic code designation in
one of four quadrants of the xy-plane. Data on numbers of species in common from
the 40 x 40 matrix were thus computer plotted but the results made little senseprobably because of the uneven size of the floras and the presence of partial floras.
Raw species numbers common between pairs of all 2 1 consolidated islands groups
for which full known floras were in the data bank yielded more promising results.
However, the most encouraging Guttman-Lingoe’s plot to date is derived from the
21 x 21 (105 island areas grouped into 21) matrix based on Kroeber Coefficients of
Similarity as shown in Fig. 5. The significance of these two-dimensional clusterings
may be limited because the treatment and results are based upon numerical data
which may or may not reflect a biological reality. Introduction of a third
dimension incorporating taxonomic units in some manner may clarify the
situation.
Possible floristic relationships have been converted to dendrographic
representations by utilizing taxonomic units to characterize the islands to be
compared. In this system, the individual taxonomic units comprising the floristic
1
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@ 9
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35 8 6
Figure 4. Computer-generated 'mileage chart' showing shared species of mosses fmm 40 geographical areas relative to tropical Pacific island floras. Numbers 1-21
are island groups and 22-40 are extra-Pacific regions which have species also known for the Pacific. The number of shared species is in the upper right triangle
and Kroeber Coefficients are in the lower left triangle. Total known florasfor 1-21 only are indicated at the same number intercept. Courtesy of H. 0.Whittier.
H. A. MILLER
I84
0 Hawaii
.Easter, Pitcairn
Juan Fernandez
Tuomotuso
0 Gambier
OAustrals
*Society
Phoenix
Bismorcks~
I
*Marquesas
OFiii
‘New Hebrides
Solzmoni
Lord Howe.
Norfolk
Roturn’
OTanga
C
,,oaks, Niue
pool to be analysed are individually compared with equal units in all the pools
with which it is compared. The original flora is then divided into subunits of equal
amalgamated distance and these are further divided until all taxa are accounted
for. Generally, the longer the ultimate pair of branches on the ‘tree’, the closer the
affinities of those regions to the original flora. This can be seen in a dendrograph of
Society Islands mosses where Tahiti, Moorea and Raiatea are the major islands
from which the total flora is derived (Fig. 6). It is not surprising, either, that the
West Indies and Central America show little commonality with the Society
Islands.
Each of these programs for analysis is still comparatively crude, but the team of
bryologists, mathematicians and computer specialists is refining the techniques. All
being committed elsewhere to full time duties, progress is slower than it might be,
results now emerging are very promising. Details of data banking and
programming techniques used by the team will be indicated with publication of
refined results from the combined moss and liverwort floras. The next project is to
model migration tracks and filter bridges for known floras incorporating factors for
isolation, area, age and altitude of islands and island groups. In such a
biogeographical model, the biology of the organisms and their natural
relationships must be taken into account and somehow be factored into
quantitative data. The problem appears to have been identified but the solution
remains elusive.
Broad spectrum, correlative, statistically and biologically defensible approaches
to biogeographical problems are now possible with electronic assistance. However,
much remains to be learned about what to ask of the organisms so that meaningful
information comprises the data base. Because of comparatively small numbers of
58 I
I
,
I-
I
A H d V X 3 0 3 3 (INV NOILflTOAB 3.LAHdOAXff
186
H. A. MILLER
taxa and quantifiable isolation, islands rightfully continue to be the laboratories of
choice for development of biogeographical concepts. The ecological sensitivity of
bryophytes, along with their small size, niche specificity, general intolerance to sea
water, impalatability to animals and limited cultural significance (suggested by the
generalized Polynesian name of ‘mountain seaweed’ = limu mauku-Hawaiian) ,
combined, cause mosses and liverworts to be particularly good indicator
organisms. Perhaps their study will provide us with the means to achieve a modern
biogeographical paradigm.
SUMMARY
Bryophytes are the oldest extant terrestrial plants and represent the level of
evolution associated with transmigration to the land. They have been widely
distributed in the past but the fossil remains were mostly overlooked until
maceration and peel techniques became regularly employed in palaeobotany.
Thus, little pre-Tertiary fossil evidence has been available until now for
consideration of the historical geography of the bryophytes.
Except for the land covered by the Carboniferous glaciers, many suitable cool,
moist sites were apparently available for the bryophytes and they were widely
dispersed prior to the Permian. Some were predominantly Laurasian and others,
including the majority of those leafy hepatic groups today considered
evolutionarily primitive, were Gondwanan. These floras fixed the background
upon which the modern bryogeographical mosaic rests even though the rapidly
improving fossil record is limited. The warm Tethys Sea, as indicated by the
foraminifera1 fauna of Lower and Middle Permian (Gobbett, 1967) was a
formidable barrier between north and south both because it was marine and
because it was warm with bordering lands having a warm and somewhat arid
climate (Schwarzbach, 1961) across western Asia, Europe and North America. On
the other hand, north-eastern Asia (Angara) had a temperate to cold temperate
Permian flora which was separated from Europe by the Ural Sea (DuToit, 1937)
and thus must have served as a refugium for many Laurasian bryophytic lines. The
significance of old Angara has not been apparent to bryogeographers who seem to
have been preoccupied with Tertiary and Quaternary floras and the coincidence
borne of cohabitation of extant bryophyte and angiosperm floras. Persistence of the
separate Angaran continental mass until near the end of the Triassic, when it
became again joined to the Euro-American mass, is of consequence. The PermoTriassic desert episodes must have resulted in massive extinctions among the
bryophytes (Miller, H., 1974). Thus most, but not all, major systematic groups
extant today are derived from the Gondwanan and Angaran refugia or from desert
adapted lines. The fragmentation of Gondwana and the reduction of the moist
refugial area by drift of the pieces to drier and/or frigid climates has resulted in
remarkable disjunctions and retention of numerous relict types in the cool to cold,
moist, oceanic circum-Antarctic land masses. The Angaran bryoflora was not
subject to on-going constriction of range. Instead, new territories were opened in
various configurations at different times resulting in creation of the holarctic
bryoflora.
Bearing in mind, as one must, that these significant events in the history of the
bryophytes occurred before the rapid rise of the angiosperms, the existence of
distinctly Gondwanan and northern taxa becomes more understandable. Further,
BRYOPHYTE EVOLUTION AND GEOGRAPHY
187
the great systematic discontinuities which exist among bryophytes can be
appreciated in terms of extinctions of nearly entire floras as a result of climatic
change-mostly desertification or formation of dry-adapted coniferous woodlands
with their biostatic, tannin-laced bark and litter.
Origin of tropical rainforest bryofloras deserves notice because it has been
considered to be somewhat obscure from the seeming mix of origins. Most of the
palaeotropical bryoflora derives from Angara and has undergone a diversity
explosion into the new niches provided by the angiosperms. Essentially pantropical elements have the same history, in the main with their many diaspores
reaching the neotropics by several routes. Unique neotropical elements derive from
Gondwanan stock or from long-standing pre-Cretaceous groups of wide dispersal
as suggested by the fossil record. Gondwanan elements of the present
palaeotropical bryoflora are mostly limited to higher altitudes in cloud forests or
above in New Caledonia and New Guinea with very few taxa, indeed, penetrating
as far as Borneo and the Philippines on one hand and to the Pacific islands on the
other. Few Gondwanan elements survived the ‘rafting’ of India across desert and
tropical latitudes as witnessed by the Himalayan flora (Sharp, 1974).
The greatest single influence leading to present diversity of bryophytes and their
distribution is the origin and spread of angiospermous forests. Not only did the
climate become wetter over large areas at that time but the broadleaved trees
provided shade and elevated humidity. Further, their bark and litter are both
circum-neutral and retain moisture-an ideal combination for bryophytes (Miller,
H., 1974). Wherever the mesophytic angiosperms migrated, the bryoflora also
migrated and genetically plastic groups which today comprise the largest families
and orders diversified into the newly available habitats. Most bryophytes remained
sensitive to high evaporative stress, however, so the maximum diversity and
abundance was achieved in high tropical cloud forests and other continuously cool
and damp sites.
Present day distributions, with still only weak, but improving, supporting
evidence from Mesozoic fossils, suggest that some major groups such as the
Dicranaceae and Bartramiaceae occurred in both Permo-Triassic refugial areas. In
Gondwana, the Dawsoniales, Rhizogoniineae and Hypopterygiineae, especially,
diversified although today’s largest moss families were also present. The
Gondwanan Metzgeriopsida appear to have evolved mostly erect anisophyllous
forms in the suborders Lepicoleineae, Perssoniellineae, Balantiopsidineae,
Lepidolaenineae and Phyllothalliineae with strong representations of
Lepidoziineae,Jungermanniineae, Geocalycineae, Metzgeriales and Monocleales.
Angaran mosses include Bryoxiphiales, Schistostegales, Tetraphidales,
Fontinalales and Timmiineae with representation of all major families. Hepatics in
the Porellineae and Marchantiales were mostly northern with many
Jungermanniineae and Geocalycineae also present. Some bryophytic groups surely
survived in areas with a climate similar to that associated with steppes and
savannahs. In such areas today we find Riccia, Fossombroniaceae, Sphaerocarpos,
Fissidens, Pottiales and Bryales near seep zones or along steep banks of
watercourses-some as perennials and some as ephemerals. The terrestrial,
variously drought-tolerant, families now occur almost worldwide.
Modern distributions of terrestrial biota as indicated by the bryophytes derive
from a complex history of evolutionary vagaries, climatic vacillations and tectonic
phenomena. Ancient groups such as bryophytes and insects with emerging fossil
188
H . A. MILLER
records are especially valuable potential indicators of the nature of the biosphere
and the influences upon it. Application of modern computer technology to analysis
of heretofore impossibly large masses of raw data, by methods such as these now
being refined for Polynesian mosses and liverworts, promises to provide the basis
for insights into biogeographical cause and effect relationships now almost beyond
imagination.
ACKNOWLEDGEMENTS
The British Museum (Natural History) has provided herbarium and library
facilities by the generous cooperation of J. F. M. Cannon, A. Eddy and A. J.
Harrington, who also gave many constructive suggestions. J. H. Price and R. W.
Sims provided diverse assistance. H. 0. Whittier has generously allowed the use of
previously unpublished computer printouts and reviewed the manuscript. Special
thanks are given to many dozens of thoughtful colleagues worldwide who have so
generously assisted with their reprints without whose aid such a review would have
been much more difficult. The Systematics Association and the University of
Central Florida Foundation provided assistance for presentation of the paper.
REFERENCES
ANDERSON, H. M., 1976. A review of the Bryophyta from the Upper Triassic Molten0 Formation, Karoo
Basin, South Africa. Palaeontologia afiicana, 19: 21-30, 5 pl.
ANDERSON, L. E., (Ed.), 1974. Proceeding of Symposium on Taxonomy and Evolution of Bryophytes of the
First International Congress of Systematic and Evolutionary Biology, 1973. Journal of the Hattori Botanical
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