Floristics - University Of Wisconsin-Madison

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FloristicsBanksia coccinea - Australia

Historical Biogeography“What lives where and why?” Historical biogeography is the flipside to ecologicalbiogeography Most of its practitioners are not geographers butsystematists specializing on specific groups of organismsThree phases of historical biogeography are usually seen:1. Descriptive — distributions and areas (floristicand faunistic biogeography)

Historical Biogeography“What lives where and why?” Historical biogeography is the flipside to ecologicalbiogeography Most of its practitioners are not geographers butsystematists specializing on specific groups of organismsThree phases of historical biogeography are usually seen:2. Narrative — using historical (geological andevolutionary) events and ad hoc assumptions as a basisfor explaining a given distribution pattern (includingdispersal biogeography)

Historical Biogeography“What lives where and why?” Historical biogeography is the flipside to ecologicalbiogeography Most of its practitioners are not geographers butsystematists specializing on specific groups of organismsThree phases of historical biogeography are usually seen:3. Analytical — comparison of the patterns of relationships ofdifferent groups of organisms occupying similar areas to findcommon biogeographic patterns vicariance biogeography cladistic biogeography phylogenetic biogeography

Floristic BiogeographyBasic to the study of floristics is knowledgeof the geographic distribution of organisms Geographic distributions are limited No species completely cosmopolitan Most species and genera, and even familiesand orders are restricted in distributionAmborella trichopodais endemic to NewCaledonia

Floristic BiogeographyBasic to the study of floristics is knowledgeof the geographic distribution of organisms Geographic distributions are limited No species completely cosmopolitan Most species and genera, and even familiesand orders are restricted in distributionWhite spruce is endemic to borealforest of North America; Picea(spruce genus) is restricted to theNorth Hemisphere

Floristic BiogeographyBasic to the study of floristics is knowledgeof the geographic distribution of organisms Geographic distributions are limited No species completely cosmopolitan Most species and genera, and even familiesand orders are restricted in distributionGunnera (Gunneraceae)has a wider but stillpatchy distribution(Hawaiian species here)

Limits to Distributions climatic: temperature, precipitation, seasonality topographic: mountains, oceans habitat: soil, pH, water availability, sun vs. shade biotic: competition, predation, coevolution history: age, dispersal, sundering, speciation

Limits to Distributions climatic: temperature, precipitation, seasonality topographic: mountains, oceans habitat: soil, pH, water availability, sun vs. shade biotic: competition, predation, coevolution history: age, dispersal, sundering, speciationPalm family is limited by severe cold temps due to their singleterminal bud at end of the stem

Limits to Distributions climatic: temperature, precipitation, seasonality topographic: mountains, oceans habitat: soil, pH, water availability, sun vs. shade biotic: competition, predation, coevolution history: age, dispersal, sundering, speciationClimate but also topography limits the extent of coniferousboreal forest species

Limits to Distributions climatic: temperature, precipitation, seasonality topographic: mountains, oceans habitat: soil, pH, water availability, sun vs. shade biotic: competition, predation, coevolution history: age, dispersal, sundering, speciationHabitat requirements force the distributions of kinglets andredback voles to match that of the coniferous boreal forests

Limits to Distributions climatic: temperature, precipitation, seasonality topographic: mountains, oceans habitat: soil, pH, water availability, sun vs. shade biotic: competition, predation, coevolution history: age, dispersal, sundering, speciationEpifagusvirginianaBeech drops(root parasite onlyon Americanbeech)American beech – Fagus grandifolia

Limits to Distributions climatic: temperature, precipitation, seasonality topographic: mountains, oceans habitat: soil, pH, water availability, sun vs. shade biotic: competition, predation, coevolution history: age, dispersal, sundering, speciationArgyroxiphium - silversword

Distribution PatternsEvery species (or higher taxa) has a particular distributionthat varies in three important features: range: entire region or area ofoccurrenceClematis fremontii (leatherflower) isrestricted to three midwestern statesleatherflower

Distribution PatternsEvery species (or higher taxa) has a particular distributionthat varies in three important features: degree of geographicalcontinuityClematis fremontii (leatherflower) isdiscontinuous in distribution across its rangeas it is restricted to limestone glades

Distribution PatternsEvery species (or higher taxa) has a particular distributionthat varies in three important features: frequency of occurrenceClematis fremontii (leatherflower) is frequentbut aggregrated in individual limestoneglades

Distribution PatternsWhat kinds of distribution patterns? Continuous Endemic Disjuncts (discontinuous)aquatic Ceratophyllum demersum (coons-tail, hornwort) iswidespread, continuous, nearly cosmopolitan

Distribution PatternsWhat kinds of distribution patterns? Continuous Endemic Disjuncts (discontinuous)Nothofagus (southern beech)is endemic to severaltemperate southernhemisphere areas, thusdisjunct

Distribution PatternsWhat kinds of distribution patterns? Continuous Endemic Disjuncts (discontinuous)Osmorhiza chilensis (sweet cicely) shows an amphi-tropical disjunction anda western N. American - Great Lakes – eastern N. American disjunction

Distribution PatternsWhat kinds of distribution patterns? Continuous Endemic Disjuncts (discontinuous)Empetrum (crowberries) of boreal and tundra and Larrea(creosote) of deserts also show amphi-tropical disjunctions

Distribution PatternsWhat kinds of distribution patterns? Continuous Endemic Disjuncts (discontinuous)1 species of Pitcairnia in westAfricaFamily Bromeliaceae (pineapples) shows continuous distribution throughoutAmericas, endemic to this region, except for peculiar disjunct in West Africa

Distribution PatternsWhat kinds of distribution patterns? Continuous Endemic Disjuncts (discontinuous) S. amazonica water dispersedFamily Humiraceae shows same peculiar disjunct in West Africa- why? (not a floristic question)

Distribution PatternsWhat kinds of distribution patterns? Continuous Endemic Disjuncts (discontinuous) all nine taxa described survive andreproduce in accordance to specificenvironmental requirements each occupies a precise area or rangefirst determined by history (area/life) actual ranges are limited byecological or biological features for invasive weeds, perhaps theoppositeLythrum salicaria purple loosestrife

Distribution PatternsTypes of continuous patterns? Cosmopolitan: distributed all over the globe indifferent to many environmental conditionsCeratophyllum in waterTaraxacum (dandelion) on land

Distribution PatternsTypes of continuous patterns?— can be taxa above species Cosmopolitan: distributed all over the globe indifferent to many environmental conditionsChiroptera - bats

Distribution PatternsTypes of continuous patterns? Circum-boreal: [circum-austral rare!]Rhododendron lapponicum lapland rosebay (Ericaceae)

Distribution PatternsTypes of continuous patterns? Pantropic: distributionlimited by oceans intropical & subtropicallatitudesPalmae - palm family

Distribution PatternsTypes of endemic patterns? Taxonomic (evolutionary) relicts: sole survivors ofonce diverse taxonomic groupsThe primitive angiosperm Degeneria belongs to a lineagethat was more species-rich as seen in the fossil record

Distribution PatternsTypes of endemic patterns? Taxonomic (evolutionary) relicts: sole survivors ofonce diverse taxonomic groupsThe gymnosperm Ginkgo biloba belongs to an ancient fossillineage going at least to the Mesozoic Era

Distribution PatternsTypes of endemic patterns? Taxonomic (evolutionary) relicts: sole survivors ofonce diverse taxonomic groupsGinkgoes are abundant in the fossil record, butonly one species is extant, all others are extinct

Distribution PatternsTypes of endemic patterns? Climatic (biogeographical) relicts: narrowlyendemic survivors of once widespread taxaGinkgo biloba is now restricted to a small area of China, but fossilevidence shows Ginkgo widespread in the temperate N Hemisphereas recently as the Pliocene (3 mya)

Distribution PatternsTypes of disjunct patterns? Many types! – we will look atseveral later Involve interplay betweenearth history and biologicalhistoryClassification of major distributions ofseed plants (Thorne 1972; Stott 1982)

Distribution PatternsTypes of disjunct patterns?Mesosaurus brasiliensis amphi-Atlantic distribution of thePermian reptile Mesosaurus wasused by Alfred Wegener asevidence for continental driftClassification of major distributions ofseed plants (Thorne 1972; Stott 1982)

Provincialism— one of the most important concepts inbiogeography, but what does “provincialism” mean?Definition by Webster’s Dictionary:2. [n] a lack of sophistication

Provincialism— one of the most important concepts inbiogeography, but what does “provincialism” mean?Definition by Webster’s Dictionary:1. [n] a partiality for some particular place2. [n] a lack of sophisticationWhen the ranges of organisms are examined closely, it is seenthat endemic forms are neither randomly nor uniformlydistributed across the earth but instead are clumped inparticular regions.

ProvincialismThree patterns are observed:1. the most closely related species tend to haveoverlapping or adjacent ranges withinrestricted parts of continents - parapatricRhea americanaRhea pennata(R. darwinii)Darwin noted this withrheas in 1833Law of Representative Species - repeatedbiogeographical observation

ProvincialismThree patterns are observed:1. the most closely related species tend to haveoverlapping or adjacent ranges withinrestricted parts of continents - parapatricExample: Banksia (Proteaceae) and Eucalyptus (Myrtaceae)

ProvincialismBanksia candelleanaBanksia coccineaBanksia marginataBanksia epica

ProvincialismThree patterns are observed:1. the most closely related species tend to haveoverlapping or adjacent ranges within restrictedparts of continents - parapatric2. a significant portion of orders or families and somegenera have markedly disjunct ranges, with taxaliving in widely separated regions of continentsor on different continents — allopatric,vicariantRoupalaProteaProteaceaeBanksia

ProvincialismThree patterns are observed:1. the most closely related species tend to haveoverlapping or adjacent ranges within restrictedparts of continents - parapatric2. a significant portion of orders or families and somegenera have markedly disjunct ranges, with taxaliving in widely separated regions of continents oron different continents — allopatric, vicariant3. completely unrelated taxa, both plants and animals,show similar patterns of endemism —they shareareas of endemism — sympatric

ProvincialismEucalyptus paucifloraEucalyptus apiculataEucalyptus manniferaEucalyptus curtisii

ProvincialismAllopatric (vicariant)disjunctions ofrelated nces ofunrelated taxaAreas of Endemism -GuavaShared areas bymany unrelatedplants, fungi, andanimalsSyzygiumMyrtaceaeEucalyptus

ProvincialismRe-examine the distribution pattern of Nothofagaceae . . .Nothofagaceae

Provincialism. . . a very similar distribution pattern is seen with Stylidiaceae . . .NothofagaceaeStylidiaceaeDonatia

Provincialism. . . and with Acaena (Rosaceae)NothofagaceaeStylidiaceae Many species of Acaena(Rosaceae) occur in the same areasas Nothofagus and Stylidiaceae Does this mean that these twotaxa and Acaena have a similarhistory that gives rise to thispattern?

ProvincialismThis same pattern of endemic distribution in the temperatesouthern hemisphere is repeated by many unrelatedgroups of organisms!

ProvincialismQuestion to ponder: What do areas of endemism mean? Why do southern beeches show distributions similar to chironomidmidges when neither is dependent upon the other? Is it that both have independently dispersed and become adapted tosimilar southern hemisphere habitats (ecology!), or does history of the biotas and areas in which they occur provide adifferent and perhaps better answer simultaneously addressing alltaxa?Allopatric (vicariant)disjunctions ofrelated taxaSympatricoccurrences ofunrelated taxaAreas of Endemism Shared areas bymany unrelatedplants, fungi, andanimals

ProvincialismQuestion to ponder: What do areas of endemism mean? These are questions not answered by floristics but require otherinformation about earth history and history of the organisms They are answered (or attempted to!) in the narrative andanalytical phases of biogeographyAllopatric (vicariant)disjunctions ofrelated taxaSympatricoccurrences ofunrelated taxaAreas of Endemism Shared areas bymany unrelatedplants, fungi, andanimals

Floristic SystemsProvincialism and Floristic Kingdoms Provincialism noted by early biogeographers: Schouw (1823),de Candolle (1855), Sclater (1858) and Wallace (1876). Impressed by the differences in the biotas on the variouslandmasses. Goal was to identify these units of different biota and thephysical or historical barriers that prevented the exchange ofspecies. Result was a division of the earth into a hierarchy of regionsreflecting patterns of faunal and floral similarities.1. Descriptive Historical Biogeography! — distributions andareas (floristic/faunistic geography)

Floristic SystemsProvincialism and Floristic KingdomsFirst map of botanical geography showing 25“kingdoms”, derived from the work of JoakimSchouw (and Humboldt)Joakim Frederik Schouw(1789 - 1852)Danish botanist and geographer,student of Humboldt. Producedfirst comprehensive textbook onplant geography.

Faunistic SystemsProvincialism and Faunistic KingdomsSix Faunal Provinces - Sclater 1858 Likewise, ornithologists andmammalogists defined faunisticprovinces Philip Lutley Sclater (1829–1913), Britishornithologist who described 1067 speciesand 135 genera of birds, published in 1858an important paper in which he divided theworld into biogeographic regions on thebasis of birds

Faunistic SystemsProvincialism and Faunistic KingdomsSix Faunal Provinces - Sclater 1858 Alfred Wallace later elaboratedon the Oriental and Australianprovinces in some detail basedon mammals and birdsAlfred Wallace’s Line (1876)

Floristic SystemsProvincialism and Floristic KingdomsFloristics generally uses the following hierarchical scheme:Kingdom (Realm) — distinctive floras; endemic familiesRegion — generic endemism highProvince (Domain) — species endemism highDistrict — subspecies endemism only

Floristic SystemsProvincialism and Floristic KingdomsRonald Good’s floristicsystem is the most wellknown with 6 kingdoms.Good, Ronald, 1947. The Geography ofFlowering Plants.

Floristic SystemsProvincialism and Floristic KingdomsFloraNote that floristic kingdomsinclude two additional relativeto the faunistic1. South African or Cape2. Antarctic— an indication of the moreremarkable levels of endemismseen in plants relative to animalsFauna

Floristic SystemsProvincialism and Floristic KingdomsFlora. . . but the merging of twofaunistic provinces into onefloristic kingdom —1. Ethiopian (African)2. Oriental Paleotropical floristic kingdom1. Paleoarctic2. Neoarctic Boreal floristic kingdomFauna

Floristic SystemsProvincialism and Floristic KingdomsThe six floristic kingdoms (colors, red lines) are here shown divided into 35regions (blue lines) based on Takhtajan (1978). Note the differentplacements of Boreal and Antarctic lines.

Floristic SystemsBoreal or Holarctic Kingdom (3subkingdoms, 9 regions) largest of the six kingdoms, 1/2 of surface Europe, N. Africa, temp. Asia, N. America 60 families raceae

Floristic SystemsPaleotropical Kingdom (5subkingdoms, 13 regions) tropical Old World, not Australia, Pacific 40 families endemicNepenthaceae

Floristic SystemsNeotropical Kingdom (5 regions) s. Florida, C. America, Antilles, most S.America 25 families endemicTovariaceaeAboboldaceaeCyclanthaceae

Floristic SystemsCape Kingdom (1 region) smallest kingdom, southern South Africa exceptionally diverse 8 families endemicPenaeaceaeRoridulaceaeGreyiaceae

Floristic SystemsAustralian Kingdom (3 regions) isolated island continent distinctive flora and high endemism seen in many biome types 18 families endemicAustrobaileyaceaeXanthorrhoeaceae s.s.Cephalotaceae

Floristic SystemsAntarctic or Holantarctic Kingdom temperate S. America, New Zealand,Antarctica 12 families endemicEucryphiaceaeNothofagaceae &Misodendraceae(parasite on Nothofagus)

dispersal biogeography) Three phases of historical biogeography are usually seen: Historical biogeography is the flipside to ecological biogeography Most of its practitioners are not geographers but systematists specializing on specific groups of organisms Historical Biogeography "What lives where and why? "

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