Bryology biogeography

Herzog 1929 - Geographie der Moose

Notices parallels in the distribution of bryophytes and vascular plants

Range of speciation in bryophytes Vs vascular plants

Similar

Is allopatric or sympatric speciation more frequent in bryophytes

Allopatric

Is anagenesis more common than cladogenesis

Anagenesis due to plasticity

Allopatric speciation

biological populations of the same species become geographically isolated from each other

Anagenesis

evolutionary process where a single species gradually accumulates genetic and physical changes over time, transforming directly into a new species without branching or splitting

ancestral form eventually disappears and is replaced by evolved form

cladogenesis

lineage splits into multiple, coexisting branches to drive biodiversity

what might prevent speciation in bryophytes

lineage splits into multiple, coexisting branches to drive biodiversity

Bryophyte ranges

generally much larger distribution ranges than vascular

Limiting factor of bryophyte range

fertilisation

fertilisation is dependent on

water and proximity of both sexes

fertilisation is more difficult for

dioecious species

dioecious vs monoecious ranges

dioecious should be smaller

lineage splits into multiple, coexisting branches to drive biodiversity

vast majority of spores fall within

a few tens of metres of the capsule

if bryophytes are not limited by migration barriers, distribution should

be driven only by environmental conditions/random

but this is not true

study of ice age return showed its influenced by barriers

and diversity depends on direction of wind/water

percent N American species not found in Europe

45%

percent European species not found in N America

30%

Why does N American have higher percent of species not in Europe

less extinctions in Pleistocene due to better refugia

European endemics are

neoendemics originating after separation from N America

Short Distance Dispersal occurs

especially in forests and heterogenous terrain,

Short Distance Dispersal is predominantly

clonal

types of short distance dispersal

zoochory

anemochory

hydrochory

anemochory

dispersal of seeds, spores, or fruits by the wind

Example of clonal propagules long distance dispersal

Sphagnum subnitens in North America using water birds – thousands of kilometers in decades.

Enormous ranges and short distance dispersal

Takes long time

Limitations of LDD (2)

low resistance of most spores to drying out and UV

therefore short germination capacity

Australia vascular vs liverwort endemics

26% vs 1%

Canary islands vascular vs bryophyte endemics

40% vs 2%

South america liverwort endemics

highest proportion of endemic liverworts (24% of species)

but have twice as much vascular endemics

centre of endemism unique to bryophytes

Pacific coast of N America

Why fewer bryophyte endemics (4)

bryophyte endemism may not be related to distribution limitations but rather the uniqueness of site in the world, very small niches

very effective LDD?

evolutionarily old, having more time to colonise available niches

troubles with identification and cryptic species

Troubles with identification / cryptic species

(there are probably up to 29% more species in Australia than previously

reported) [Renner et al. 2017]

vascular plants vs bryophytes overlap of centres of diversity and endemism

no overlap in bryophytes

highest number of endemic bryophytes is in

Madagascar

Endemic mosses and liverworts to Europe

150 mosses

60 liverworts

Species area relationship bryophytes, ferns and liverworts

flatter than vascular plants

basic principles of island biogeography

diversity traditionally considered to be governed by a balance between immigration and extinction

bryophytes principles of island biogeography (2)

do not fully apply

greater role played by island area, coastline complexity, elevation and environmental heterogeneity

Latitudinal diversity gradient

Similar to vascular plants, tropical areas are probably the center of diversity for liverworts and hornworts, but this is not so certain for mosses.

Latitudinal diversity hotspots moss (2)

Mosses also have diversity hotspots outside tropics.

For example, in Europe or South America, diversity increases with increasing latitude

Cosmopolitan genera (4)

Anthoceros, Chiloscyphus, Jungermannia, Orthotrichum

Cosmopolitan species (4)

not as many as expected

Marchantia polymorpha, Bryum argenteum, Ceratodon purpureus

Cosmopolitan families (7)

up to 60% of families

Buxbaumiaceae,

Fissidentaceae,

Grimmiaceae,

Orthotrichaceae,

Pottiaceae,

Plagiotheciaceae

Pangaean distribution (3)

in very old taxa

most genera and species too young

Pangea existed around 335-200 million years ago

Pangaean families (7)

Jungermanniaceae,

Marchantiaceae,

Bryaceae ,

Dicranaceae,

Hypnaceae,

Polytrichaceae

Sphagnaceae

Pangaean genera (5)

Phaeoceros

Frullania

Riccia

Andreaea

Racomitrium

Pangaean species (2)

Targionia hypophylla

Tortula atrovirens

Laurasian (Holoarctic) distribution (2)

occurrence outside the tropics

in europe, considerable extinction due to glaciation (fewer refugia)

Laurasian families (3)

Conocephalaceae,

Ephemeraceae,

Climaciaceae

Laurasian genera (3)

Dicranodontium,

Pleurozium

Ptilium

Laurasian species (2)

Pleurozium schreberi

Dicranodontium denudatum

Gondwanan distribution

Main range: Australia, South America, Antarctica

Gondwanan families (3)

Dicnemonaceae,

Hypnodendraceae,

Hymenophytaceae

Gondwanan genera (2)

Balantiopsis,

Pachyglossa

Gondwanan species (2)

Pachyglossa austrigena

Balantiopsis diplophylla

Circum-Tethyan distribution (2)

trace the borders of the ancient Tethys Ocean

that separated Laurasia and Gondwana

Circum-Tethyan genera (4)

Plagiochasma, Exormotheca, Crossidium, Timmiella

Circum-Tethyan species

Crossidium aberrans

Pantropic distribution (2)

Distributed across the tropical regions of Africa, America, and Asia;

not many species or genera

Pantropic genera (2)

Calymperes, Leucophanes

Pantropic species (3)

Octoblepharum albidum,

Frullania squarrosa,

Fissidens asplenioides

Disjunct distribution (3)

Separation of continents, which is not followed by speciation, in the case of bryophytes

it is not clear whether it is a remnant of historical ranges, introduction or LDD.

Environmental similarity of the areas is probably more important than the limits of spread.

Bipolar distribution species (3)

Ptilidium ciliare ,

Buxbaumia aphylla,

Conostomum tetragonum)

Laurasian disjunction:

eg. W North Amerika and W Europe

Laurasian disjunction species (3)

Antitrichia curtipendula, Dicranum tauricum, Plagiothecium undulatum

neophytes

introduced by humans

neophyte species (3)

Scleropodium purum

Orthodontium lineare

Campylopus introflexus

Scleropodium purum introduction

packaging material from EU To BC, Newfoundland, Japan, New Zealand

Orthodontium lineare introduction (5)

introduced to EU from AU

first recorded in 1911 Liverpool

1939 Germany

1964 CZ

1976 Sweden

Campylopus introflexus introduction (3)

introduced EU from South Africa/AU

first record 1941 England then mass spread to West

CZ first record 1988

Anthropogenic ranged genera (2)

Riccia

Anthoceros

Anthropogenic ranged species (2)

Bryum argenteum

Lunularia cruciata

Schofield (1992) Bryophyte distribution patterns (basic areas) (6)

Arctic

Paleotropic

Neotropical

Australian

South African

Antarctic

Index Muscorum

Wijk, R., Margadant, W.D. & Florschiitz, P.A. 1959-1969.

20 bryofloristic areas

characterised on typical representative families and endemic taxa typical of areas

Index Muscorum example areas (3)

Central Africa and Saint Helena Islands

Africa, Kergulen Islands

North America, Greenland, Allutian Islands, Bermudez

European species (holoarctic) area divisions

latitude (arctic, boreal, temperate, mediterranean)

sea proximity (atlantic, continental)

altitude (planar, colline, montane, subalpine, alpine, nival)

colline

transitional, low-elevation hilly zone just above the flat lowlands and below the mountain zones

Mateo et a. (2013).

Modeling species distributions from heterogeneous data for the biogeographic regionalization of the European bryophyte flora

Mateo et a. (2013). zones (8)

Alpine

Atlantic

Boreal

Macaronesian

Mediterranean

Pannonian

Black Sea

Continental

First list of threatened bryophytes

1969 Belgium

- lnternational Association of Bryologists:

created ECCB- European Committee for Conservation of Bryophytes in 1990

IUCN

established within the framework of the Species Survival Commission Bryophyte Specialist Group in 1991

The first Czech list of endangered bryophytes

Váňa J. (1993)

Czech legislative protection (3)

Act No. 114/1992 Coll. on Nature Conservation and Landscape Protection

no bryophyte species covered

proposed update 67 spp

Scandinavia collection rules (2)

okay for species of LC in non protected areas for individual use

commerical collection requires permits

Germany collection rules (2)

okay for species of LC in non protected areas for individual use

commerical collection requires permits

Spain collection rules

forbidden, very strict

Portugal collection rules

okay for specified areas and species

France collection rules (2)

quote per person/day

forbidden in Pyrenees

Scandinavia species protection (3)

very strong,

habitat protection,

woodland key habitat

Germany species protection (4)

strong

red list

strictly protected areas

60 spp and also genus level

Spain species protection (2)

strong but slow laws

50 protected spp

Portugal species protection

medium/low

species protection France

strong

Scandinavia Laws (4)

country legislative,

forestry,

action plans,

ecosystem protection

Germany Laws (2)

BNatSchG

red lists

Spain Laws (3)

national lists

LIFE projects

Natura 2000

Portugal Law (2)

LIFE projects

Natura 2000

France Laws

National and regional system (ZNIEFF, based on red list)