Biodiversity of Cacti

The Cactaceae with c.
11435 species are the mosst important

plant familyy of the arid regions of th
he Americas. Recent revisions and molecular studies resulted in an
n improved
knowledge of the phyloggeny and taxo
onomy of thiss group. Due
to their high
h value as ornnamental plan
nts, countless publications
with data on ecological ppreferences an
nd geographicc occurrence
of the speciies are availabble. In this volume, the disttribution areas of all ca
actus species are mapped. On this basis, we identified and ch
haracterized sseven geogra
aphical centers of cactus
diversity. Overall
O
diversitty patterns off the family, as well as, diversity patterns of all taxxonomic subgroups, growth
h forms, and
pollination syndromes aare presented and mapped on the phylogeny of the Cactaceaee. More than 50% of the species
s
have
extremely small distribbution rangess, resulting in
i potential
threat and insufficient ccoverage by existing prote
ected areas.
This volume
e presents thee most comprrehensive biog
geographical
analysis of one
o of the largger plant families.
Map 26
1-2
3-4
5-6
7-8
9 - 12
13 - 15
16 - 18
19 - 22
Cactaceae:
g
Generic diversity (128 gen.)
Genera per 2500 km
1500 km
' Barthlott et al. 2013
Text zweisprachig Engl ish und Deutssch

c. 205 pagges, c. 333 co lored maps sh
howing the distribution of
all Cactus species
c. 60 colo
or figures inccl. 44 fotos of
o cacti specie
es and their
habitats
Analyses and
a diversity maps for all tribes and ge
enera, major
growth forms, and polliination syndro
omes
Based on the most receent phylogene
etic studies
BARTHLOTT, W., BURSTEEDDE, K., GEFFERT
E
, J.L., IBISCH, P.L.,
KOROTKOVA, N., MIEBACH, A., RAFIQPOORR, M.D., STEIN, A. & MUTKE,
J. (2015): Biogeography and Biodiversity of Cacti. Schumannia
7. 205 page
es, 333 maps,, ca. 60 photo
os and other figures. 21.0
x 29.7 cm (A4), Hardcoover Price: 39,- + tran
nsport, ISSN
1437-2517. Bezugsadressse (to order by): gs@dkg.eu
u
Wilhelm Barthlott, Kerstin Burstedde, Jan Laurens Geffert, Pierre L. Ibisch, Nadja Korotkova,
Andrea Miebach, M. Daud Rafiqpoor, Anke Stein & Jens Mutke
Biogeography and biodiversity of cacti

Biogeographie und Biodiversitt der Kakteen
Schumannia
Ein Sonderheft der

Deutschen Kakteen-Gesellschaft e.V.
und der Gesellschaft sterreichischer Kakteenfreunde
Schumannia 7
2015
herausgegeben von
Detlev Metzing
Abstract / Kurzfassung / Resumen
Abstract
The Cactaceae with c. 1,435 species are the most important plant family of the arid regions of the Americas. Recent revisions and molecular studies resulted in an improved knowledge of their phylogeny and taxonomy. Due to their high value as ornamental plants, countless
publications with data on ecological preferences and geographic occurrence of the species are available. In this volume, the distribution areas of all cactus species are mapped. On this basis, we identified and characterized seven geographical diversity centers. Overall diversity
patterns of the family, as well as, diversity patterns of all taxonomic subgroups, growth forms, and pollination syndromes are presented and
mapped on the phylogeny of the Cactaceae. More than 50% of the species have extremely small distribution ranges, resulting in potential
threat and insufficient coverage by existing protected areas. This volume presents the most comprehensive biogeographical analysis of one
of the larger plant families, illustrated by 333 colored maps and 55 color figures on c. 200 pages.
Kurzfassung
Die Cactaceae sind mit rund 1.435 Arten die bedeutendste Pflanzenfamilie der amerikanischen Trockengebiete. Aktuelle Revisionen und
molekulare Analysen haben das Verstndnis von Taxonomie und Phylogenie erheblich verbessert. Aufgrund des weltweiten Interesses an
dieser Zierpflanzengruppe liegen unzhlige Publikationen zu den kologischen Ansprchen und Standorten vor. In diesem Band wurden
die Verbreitungsmuster aller Arten erfasst und auf farbigen Karten dargestellt.Auf dieser Grundlage wurden sieben Diversittszentren identifiziert und nher charakterisiert. Diversittsmuster der Gesamtfamilie wie auch aller taxonomischen Untergruppen sowie z. B. Wuchsformen und Bestubungssyndromen werden prsentiert und unter Einbeziehung phylogenetischer Analysen diskutiert. ber 50 % der Kakteen haben sehr kleine Verbreitungsareale, womit in vielen Fllen eine potentielle Gefhrdung und unzureichende Abdeckung durch existierende Schutzgebiete einhergeht. Illustriert durch insgesamt 333 farbige Karten und 55 farbige Abbildungen auf ca. 200 Seiten wird die
bisher umfangreichste biogeographische Analyse einer greren Pflanzenfamilie vorgelegt.
Resumen
Biogeografa y Biodiversidad de las Cactceas. Cactaceae con aprox. 1435 especies, es la familia ms importante de plantas en las zonas ridas de Amrica. Revisiones y estudios moleculares recientes han mejorado el conocimiento de la taxonoma y filogenia de este grupo. Debido a su importancia como plantas ornamentales, estn disponibles un sinnmero de publicaciones sobre las preferencias ecolgicas y las
distribuciones geogrficas de las especies. En este volumen estn representadas grficamente las reas de distribucin geogrfica de todas
las especies de cactos. Con base en esta informacin hemos identificado siete centros geogrficos de diversidaden el grupo. En la filogenia
de Cactaceae se representan los patrones de diversidad general de la familia, al igual que los patrones de diversidad de sus subgrupos taxonmicos, formas de crecimiento y sndromes de polinizacin. Ms del 50% de las especies presentan mbitos de distribucin geogrfica muy
restringidos, resultando en una amenaza potencial y presencia insuficiente en las reas protegidas en existencia. Este volumen presenta el
anlisis biogeogrfico ms comprensivo de una de las familias ms grandes de plantas, ilustrado con 55 figuras y 333 mapas a color en aprox.
200 pginas.
Corresponding Author / Korrespondenzautor:

Prof. Dr. Wilhelm Barthlott
Nees Institute for Biodiversity of Plants, University of Bonn
Venusbergwerg 22
53115 Bonn Germany
e-Mail: barthlott@uni-bonn.de
www.lotus-salvinia.de
Schumannia 7
2015
205
Contents / Inhalt
Foreword and acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Vorwort und Danksagungen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
6
7
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Einleitung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Cactus ecology and biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

kologie und Biogeographie der Kakteen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
2.1
Cacti and their habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

Kakteen und ihre Lebensrume . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
2.2
Biogeography and palaeo-history . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

Biogeographie und Florengeschichte . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Phylogeny, evolution and systematics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

Phylogenie, Evolution und Systematik . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
3.1
Finding the closest relatives of cacti . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

Die nchsten Verwandten der Kakteen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
3.2
Age estimates for the Cactaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

Schtzungen des Alters der Cactaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
3.3
Current understanding of phylogenetic relationships within Cactaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

Das aktuelle Verstndnis der phylogenetischen Beziehungen innerhalb der Cactaceae . . . . . . . . . . . . . . . . . . . . . . . . . 27
3.4
Basal cactus lineages and the origin of cacti . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

Frhe Kakteen-Linien und der Ursprung der Kakteen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
3.5
Relationships within Cactoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

Beziehungen innerhalb der Cactoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
3.6
Concluding remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Schlussbemerkungen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Mapping the diversity of cacti . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

Kartographische Darstellung der Kakteen-Diversitt . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
4.1
Main data sources . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

Wichtige Datenquellen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
4.2
Generating the distribution maps . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

Erstellung der Verbreitungskarten . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
4.3
Data analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Datenanalyse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
4.4
Quality and reliability of data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

Qualitt und Zuverlssigkeit der Daten . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Schumannia 7
2015
Patterns of diversity and endemism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Muster von Diversitt und Endemismus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
37
51
5.1
General overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Allgemeiner berblick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
37
51
5.2
Centres of diversity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Diversittszentren . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
37
51
5.3
Range-sizes of Cacti . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Arealgren von Kakteen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
41
56
5.4
Spatial distribution of growth forms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Rumliche Verbreitung der Wuchsformen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
44
62
5.5
Spatial distribution of pollination syndromes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Rumliche Verteilung der Bestubungssyndrome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
47
63
5.6
Diversity pattern of genera, tribes and subfamilies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Diversittsmuster der Gattungen, Triben und Unterfamilien . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
48
66
5.7
Diversity gradients . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Diversittsgradienten . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
50
67
Conservation and hotspots: cactus diversity in change . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Kakteen-Diversitt im Wandel und Naturschutz . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
68
73
6.1
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Einleitung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
68
73
6.2
Threats to Cactaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Gefhrdungen der Kakteen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
69
74
6.3
Cacti as threats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Kakteen als Bedrohung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
73
81
6.4
Country profiles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Lnderprofile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
73
81
Distribution maps of Cactaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Verbreitungskarten der Cactaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
82
89
7.1
Subfamily: Pereskioideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Unterfamilie: Pereskioideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
82
89
7.2
Subfamily: Maihuenioideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Unterfamilie: Maihuenioideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
82
89
7.3
Subfamily: Opuntioideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Unterfamilie: Opuntioideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
82
89
7.4
Subfamily: Cactoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Unterfamilie: Cactoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
83
90
References/Literatur . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 202
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205
Kurzfassung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205
Resumen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205
Schumannia 7
2015
Foreword and acknowledgements

Wilhelm Barthlott
Cacti are, just like bromeliads or hummingbirds, creatures of the New World.About 1440
species populate the Americas; some of them
are globular and about two centimetres small
(Blossfeldia), while others are columnar and
up to 14 metres tall (Carnegiea). Their distribution ranges from the boreal coniferous
forests in the North (south-west Canada) up to
the grasslands of the Pampa in the south. Cacti
inhabit diverse habitats, from the coast to the
high mountains, from deserts up to the rainforest canopy.
The often densely spined succulents (called
herbal crystals by the poet Adalbert Stifter)
are characterized by magnificent flowers. Not
only scientists but also plant enthusiasts are fascinated by this plant family.They are covered by
important collections, national and international cactus and succulent societies, about a
dozen journals and an enormous amount of literature. Hence, cacti are one of the most wellstudied major plant families.
The first modern monographs in the new millennium have been presented by Edward F.Anderson as The cactus fanily (ANDERSON 2001,
German editions as Das groe KakteenLexikon, ANDERSON 2005, 2011) and David
HUNT (2006). The amount of published data is
overwhelming, but a modern biogeographic
overview for the family is still missing. There
are early biogeographic studies based on the
data available at that time. Karl SCHUMANN
(1899a) published Die Verbreitung der Cactaceae im Verhltnis zu ihrer systematischen
Gliederung [Distribution of Cactaceae in relation to their systematic classification] in
Berlin, and Curt Backeberg published his Zur
Geschichte der Kakteen [History of the cacti]
(BACKEBERG 1942b) and Verbreitung und
Vorkommen der Cactaceae [Distribution and
occurrence of Cactaceae] (BACKEBERG 1944).
Today, however, these pioneering treatments
are outdated, especially with regard to the present state of the knowledge of phylogeny, systematics, and molecular analyses.
This analysis of biogeography and biodiversity
of the cacti combines current knowledge with a
new objective in the context of modern biodiversity research. The family is analysed based
on systematic-taxonomic knowledge incorporating geographic and phylogenetic data. Initially, distribution maps were created for 1416 of
the 1438 accepted species. From these maps, diversity maps were generated for the relevant
taxonomic levels from genera to the entire fam-
ily, and the different centres of diversity were

circumscribed.At the same time, macroecological relations, distribution of growth forms, pollination syndromes, and other aspects were
recorded, spatially visualized, and correlated
with general biogeographical patterns and ecoregions. Issues of conservation were examined
under consideration of land use data and existing protected areas.This provides an important
foundation for the conservation of Cactaceae
and their habitats.
The plan for this publication emerged in the
early 1990s. First modern maps of phytodiversity had already been published by BARTHLOTT
(1983) for the CactaceaeRhipsalideae. As a
well-known and extensively studied group,
compared to other large families, the Cactaceae
will act as a role-model for the analysis of biodiversity and biogeography of a larger angiosperm group.
The monograph presented here is thus based
on three decades of preparatory work, as well as
the collaboration of many colleagues and students, who are either involved as authors or acknowledged for their contributions. Simone
Strecker (1992) submitted a diploma thesis
(Arealgeographie der Kakteen), which was
based on a manual compilation of diversity
maps of all Cactaceae genera. In subsequent
years, the data were complemented and edited
by Pierre Ibisch, utilizing vegetation maps and
his fieldwork in Bolivia. He also supervised preliminary studies and the digital processing of
distribution data by Georg Rauer, which was
supported by a research grant from the
Deutsche Kakteen-Gesellschaft (the German
Cactus Society)at that time still without true
geographic information systems (GIS).As part
of a large collaborative project (BIOTA)
funded by the German Federal Ministry for Education and Research, a working group (BIOMAPS) has been established in Bonn from
2001 to 2010 to map the global geographic diversity of angiosperms. Besides regional analyses, world maps of biodiversity of all land plant
groups were generated (e.g. BARTHLOTT & al.
1996, 2005, KIER & al. 2009, KREFT & al. 2010,
MUTKE & al. 2011). Therefore, the infrastructure and the aforementioned geographic information systems were available for digitization
and spatial analysis of the cactus data. This
started in 2007 and was carried out by two parallel diplom theses of Anke Stein and Kirsten
Burstedde (nee Hahne) that were finished in
spring 2009. In 2011 Andrea Miebach com-
pleted a bachelor thesis focusing on the evolution of functional traits in the Cactaceae. Jens
Mutke coordinated this work since 2007 with
great dedication and expertise.
In preparation for publication, all of the maps
were carefully revised and uniformly laid out
(among others by Laurens Geffert, Andrea
Miebach and Lina Samira Meiling (geb.
Mathar). Boris O. Schlumpberger provided data
on pollination ecology from his on-going research project. Nadja Korotkova analysed the
phylogenies based on molecular data, and geographer M. Daud Rafiqpoor edited the spatial aspects of the distribution maps.Almost all
of the above collaborators were, at least temporarily, funded by our long-term project Biodiversitt im Wandel of the Academy of Sciences and Literature in Mainz.
With the financial support of the Academy of Sciences and Literature in Mainz, we established a
small working group of the International Organization for Succulent Plant Study (IOS) at the
end of our long-term mapping work and met for
the first time in 2009 during the IOS Inter-Congress 2009 in Bonn. The colleagues carefully
checked and revised the distribution maps provided by us, and our special thanks go to David
Hunt, Nigel Taylor, Martin Lowry, Charles Graham, Paul Hoxey, and Ralf Bauer.
The list of people who contributed to the publication presented here is long.We are indebted
to all of them. However, this particularly applies to the Academy of Sciences and Literature in Mainz: Without a sustainable institutional framework of our long-term project
Biodiversitt im Wandel (19992014), the
complex project of mapping the cacti could not
have proceeded.
The most important historical work on biogeography of cacti was published in 1899 in
Berlin, written by Professor Karl Moritz Schumann, who founded the German Cactus Society (DKG) in 1892. This society has accompanied and supported our project from the beginning, and we are very much obliged to it.
We are especially grateful for the editor Detlev
Metzing, who worked with great competence
and intensity at the completion of this manuscript. It is a fortunate coincidence that this
work is published in the Schumannia, named
after the founder of the DKG.
Bonn, in January 2015
Wilhelm Barthlott
Schumannia 7
2015
1 Introduction
Wilhelm Barthlott
Presumably, cacti (Melocactus) were already

among the curiosities that Christopher Columbus presented as gifts to Queen Isabella of
Castile after his first voyage to the New World
in 1492. Carl Linnaeus (17071778) already had
a surprisingly clear picture of the delineation of
this family. In his Species Plantarum, published 1753, he listed 22 cactus species. Based
on his Sexual System of classification, he assumed a close relationship between morpho-
logically very different taxa, such as Melocactus and Opuntia, but also Pereskia, and he combined all these in the genus Cactus. With the
studies of Adrian Haworth (17681833), Joseph
Salm-Dyck Reifferscheidt (17731861), Augustin Pyrame de Candolle (17781841), and
Ludwig G. K. Pfeiffer (18051877), the systematic investigation in the middle of the 19th century was already far advanced. This was because these plants had stimulated the interest
of botanists and plant enthusiasts at an early

stage, caused by their bizarre and exotic
appearance, combined with modest cultivation
requirements of many species.
In this era, systematic recording of species was
at the heart of research. This did not only apply to cacti, but also to all other plant families.
Plant geography (and ecology) started early
with the exceptional singular Essai sur la Gographie des Plantes (Paris 1805) by the vi-
Fig. 1: The overlapping distribution ranges of ten of the Mexican globular cactus genera accepted by Curt BACKEBERG (1944), which he noticeably summarized under the name Boreoechinocacti. The lines are distribution boundaries of the genera. The Chihuahua centre of diversity (cf. Map11) is already clearly
visible, as already in the Karte der Verdichtung der Kakteenvorkommen [map of densification of cactus occurrences ](BACKEBERG 1958: Abb. 4) (from:
BACKEBERG 1944).
Abb. 1: Die berlagerten Areale von zehn der von Curt Backeberg akzeptierten Gattungen mexikanischer Kugelkakteen (BACKEBERG 1944), die er bezeichnenderweise unter dem Begriff Boreoechinocacti zusammenfasst. Die Linien sind die Arealgrenzen der Gattungen. Schon deutlich ist auf dieser Karte wie
auch auf der Karte der Verdichtung der Kakteenvorkommen (BACKEBERG 1958: Abb. 4) das Chihuahua-Diversittszentrum der Familie (vgl. Karte 11)
zu erkennen (aus: BACKEBERG 1944).
Schumannia 7
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2 Cactus ecology and biogeography

Jens Mutke
2.1 Cacti and their habitats

Cacti are one of the most characteristic plant
groups of the Americas, being almost endemic
to the New World. only one single epiphytic
species, Rhipsalis baccifera, has disjunct populations in the old World.
Latin America stands out for a high diversity
of different landscapes and vegetation zones
(Fig. 4). It may seem surprising to the non-specialist that habitats of cacti represent a large
range of environmental conditions, as well.
They grow from coastal sand dunes up to some
5000 metres elevation in the high Andes, and
from southern Canada to southern Argentina,
spanning a latitudinal range of more than 100
degree. Factors limiting the distribution range
of the family at high latitudes are extreme cold
and in most cases humid conditions with short
days during winter. In addition, more humid
parts of the mediterranean type winter rainfall
areas in northern California and south-central
Chile are avoided by most cacti. This seems
somewhat surprising, as especially some
species of Opuntia have been widely naturalized in the mediterranean Basin, including
large parts of southern Europe and northern
Africa. one important parameter limiting cactus growth in winter rainfall areas might be the
influence of frequent fires. Fire also limits the
establishment of cacti in many grass lands and
savannah ecosystems.
Even though cacti are an iconic element of
New World deserts, the family inhabits a wide
range of ecosystems (Figs. 521); cacti even
grow in humid tropical rainforests. However,
this is more of an exception and mainly true for
epiphytic species, some shrubby or arborescent
Pereskia in south-east Brazil and central America, and the specialists growing on isolated rock
outcrops (inselbergs) representing dry, azonal
habitat islands within rainforests. In general,
cacti are bad competitors under humid, fertile
conditions. Thus, most species are found in
ecosystems with limited water availability at
least during some months of the year, particular edaphic conditions such as the rock outcrops, saline, gypsum, serpentine substrates, or
sand fields. This includes open desert to semidesert vegetation types, as well as thorny shrub
and deciduous to semi-deciduous dry forests. It
Schumannia 7
2015
has been demonstrated by several authors that

many cacti species show higher rates of establishment and survival in the shadow of trees or
shrubs as nurse plants in semi-arid habitats
(e.g., LARREA-ALCZAR & al. 2008, mANDuJANo
& al. 2002).
on an ecoregional basis, the mexican pineoak forests of the Sierra madre oriental and
occidental as well as of the Trans-mexican
Volcanic belt belong to the areas with the
highest Cactus species richness, as well as the
meseta Central matorral and the Chihuahua
desert in mexico, the Central Andean Puna
Ecoregion sensu WWF (oLSoN & al. 2001) in
Peru and Bolivia, and the Bolivian montane
dry forests.
2.2 Biogeography and palaeo-history

Apart from the early studies by SCHumANN
(1899a) and BACKEBERG (1942) there are few
biogeographic studies for the cactus family
(but compare e.g. HERNNDEZ & GmEZ-HINoSTRoSA 2011, TAyLoR & ZAPPI 2004, THIEDE
1998). unfortunately, the analysis of biogeographic patterns of cacti is hampered by the
absence of a fossil record. Not even the age of
the family is known. Recent molecular clock
approaches using fossils of related taxa indicate a relatively short evolutionary history
compared to other flowering plant families
(compare chapter 3.2).
most cactus diversity today is linked to arid or
semi-arid conditions. The species rich arid regions of western North America (cf. Fig. 22)
and mexico are dominated by the almost exclusively North American cactus tribes Cacteae and Phyllocacteae, as well as many species of opuntioideae. These northern centres
of cactus diversity are separated from the dry
Caribbean Coast of Colombia and Venezuela
by the rainforests of southern Central America with some linkage through the Caribbean
Islands.
The humid rainforests of the Amazon Basin
and of the Choc region in western Colombia
separate these northern hemisphere cactus
habitats from suitable areas further south (cf.
Fig. 4). Dry valleys of the Andes in western
South America are to some degree connected
to the more southern regions of the Atacama

Desert at the Peruvian and Chilean Pacific
Coast. However, the geologically relative
young and somewhat wetter Northern Andes
especially in Colombia harbour only very few
cactus species.The Atacama Desert with many
endemic species especially of the tribe Cereeae
is separated from the species rich Bolivian Andean dry valleys and the western Chaco by the
high Andes reaching altitudes of more than
5000 m a.s.l. This Bolivian and northern Argentinian South Central Andes centre of cactus diversity has the highest diversity of different tribes and genera, as well as growth forms
of cacti. It is separated from the cactus diversity centre in the Caatinga in eastern Brazil by
the grasslands of the Cerrado, which are relatively poor in cacti, probably due to the frequent occurrence of fires. The high species diversity in the south-east Brazilian rainforest of
the mata Atlantica is a particular case owed
mainly to the presence of many epiphytic species of the tribe Rhipsalideae, as well as species
in azonal habitats like rock outcrops or coastal
sand dunes.A detailed description of the main
centres of Cactus diversity is given in chapter
5.2. For an overview on the biogeography and
ecology of cacti of Eastern Brazil see TAyLoR &
ZAPPI (2004).
The lack of data on the evolutionary history
of the Cactaceae makes it difficult to relate
the current distribution patterns to palaeogeographic conditions and events. Both BACKEBERG (1942b) as well as LEuENBERGER (1986)
conclude an origin of the Cactaceae and of
the most basal taxon in the phylogenetic tree,
the subfamily Pereskoideae with the only
genus Pereskia, in northern South America.
According to recent molecular studies this
genus is a grade with two clades at the basis
of the cactus phylogeny (compare NyFFELER &
EGGLI 2010b and Fig. 23). Today, the cladistically basalmost clade of Pereskia species occur in northern Venezuela, the Caribbean Islands, and Central Americaonly P. aureifolia occurs in south-east Brazil. In contrast, the
second clade of Pereskia species is almost restricted to south-east Brazil and the central
Andes of Bolivia and Peruwith P. aculeata
having a disjunct distribution area in the Carribean, as well.
13
3 Phylogeny, evolution, and systematics

Nadja Korotkova
3.1 Finding the closest relatives of cacti

The Cactaceae belong to the order Caryophyllales in which they are part of a group that
contains most of the succulent families of the
order: Cactaceae,Anacampserotaceae, Basellaceae, Didiereaceae, Halophytaceae, montiaceae, Portulacaceae, and Talinaceae (CuENouD
& al. 2002, SCHFERHoFF & al. 2009).The placement of Cactaceae within the Caryophyllales
(or in the former Centrospermae) dates back
to the 19th century, and molecular phylogenetic
studies have undoubtedly confirmed that
Cactaceae are part of the Caryophyllales. Nevertheless, the closest relatives of cacti have
been identified with confidence only recently.
A first hypothesis on the putative close relatives
of the cacti was suggested by THoRNE (1976),
who assumed a close relationship of Cactaceae
and Didiereaceae, Basellaceae and Portulacaceae and created the suborder Portulacinae
for this alliance. Recent studies have confirmed
that Cactaceae are closely associated with the
Portulacaceae, and even nested within them
(APPLEQuIST & WALLACE 2001, CuENouD & al.
2002, HERSHKoVITZ & ZImmER 1997). NyFFELER
(2007) could finally identify the Portulacaceae
tribe Anacampseroteae as the closest relatives
of cacti. The Portulacaceae were found to be
polyphyletic and several nomenclatural changes
became necessary. As one result, the Anacampserotae are now treated as the family
Anacampserotaceae and are the sister family of
the Cactaceae (NyFFELER & EGGLI 2010a).
3.2 Age estimates for the Cactaceae

Since Cactaceae are a New World family, absent from Africa despite of suitable habitats
(besides one Rhipsalis species and introduced
opuntias), it is generally assumed that they
evolved in South America after the break-up of
Gondwana during the late Jurassic and early
Cretaceous. This suggests a limiting age of
10090 million years when South America and
Africa were already separated (mAuSETH 1990).
An immediate age estimate for the Cactaceae
is not possible since no fossils are known.
The most comprehensive hypothesis on the
evolutionary history of the Cactaceae was
linked to the geological history of South America and was proposed by BACKEBERG (1942b).
24
He assumed an origin of the family in the tropical zone of mesoamerica during the Cretaceous, app. 130 million years, thus suggesting
that the Cactaceae are a rather ancient group.
much later, HERSHKoVITZ & ZImmER (1997) suggested the Cactaceae being a young group of
only 30 million years of age. Still, they did not
provide any plausible basis for this estimate.
The most recent evidence for the age of the
Cactaceae comes from molecular dating of the
Caryophyllales, yielding timeframes of 116104
million years (ANDERSoN & al. 2005, WIKSTRom
& al. 2001). No age estimates using a molecular clock particularly for the Cactaceae were
available until recently, due to the lack of fossils, which are used as calibration points in such
analyses. Age estimates for Cactaceae therefore still have to rely on known fossils of more
distantly related taxa.Thus, the molecular clock
calibrations only allow just setting an approximate timeframe when the family evolved.Two
studies using this approach yielded different
results: the age of the cacti was inferred as
19.13.1 million years (oCAmPo & CoLumBuS
2010) or 35 million years (ARAKAKI & al. 2011).
3.3 Current understanding of

phylogenetic relationships
within Cactaceae
The Cactaceae as a family were never seriously
questioned, but hypotheses on relationships
within the family were always troublesome.
Convergent evolution is frequent in this family (WALLACE & GIBSoN 2002), making interpretation of morphological features challenging. Columnar cacti, small globular cacti, and
epiphytes, as well as adaptations to major
pollinator groups (bats, birds) have evolved
several times independently and the flower
morphology is rather uniform, providing few
valuable characters. Furthermore, due to phenotypic plasticity as a response of the environmental conditions, individual taxa are morphologically variable, making interpretation of
morphological characters troublesome.
molecular phylogenetic methods contributed
much to the understanding of flowering plant
evolution and phylogenetics. But the relationships within the Cactaceae are still insufficiently understood, and few molecular phylogenetic studies have been conducted compared
to other popular plant families such as orchidaceae or Bromeliaceae.

The first DNA sequence data for Cactaceae
were published by WALLACE (1995). A more
comprehensive phylogenetic tree derived from
the chloroplast marker rbcL was provided by
WALLACE & GIBSoN (2002). The first comprehensive molecular phylogenetic study for the
Cactaceae was done by NyFFELER (2002) and
was based on datasets of the chloroplast regions trnK/matK and trnL-F. This study could
not entirely clarify relationships within the
family but identified major groups and furthermore revealed the para- or polyphyly of some
tribes and genera, e.g. Pereskia, Notocacteae,
Browningieae, Hylocereeae and Lepismium.
The most comprehensive phylogenetic hypotheses for the Cactaceae at present are based
on datasets of the plastid regions trnK/matK,
the rpl16 intron and trnL-F and the nuclear
gene ppc (HERNNDEZ-HERNNDEZ & al. 2011)
or on trnK/matK (BRCENAS & al. 2011). To
date, the study of BRCENAS & al. (2011) is the
Cactaceae study with the most species included
(666 species).The results of these studies are an
improvement compared to the previous ones,
but the results are still not satisfactory and the
relationships within Cactaceae remain partly
unresolved.
3.4 Basal cactus lineages and the

origin of cacti
In the following, the relationships are summarised as currently understood based on the
aforementioned studies and further studies of
single tribes and genera. The first branching is
the subfamily Pereskioideae (only Pereskia,
Fig. 26). As revealed by molecular data, Pereskia is not a monophyletic group but forms a
grade in the Cactaceae phylogeny (Figs. 24 &
25) with mesoamerican and Caribbean Pereskia species as the first branching group followed by the Andean Pereskia, which is sister
to the rest of the family (BuTTERWoRTH & WALLACE 2005, EDWARDS & al. 2005).
Pereskia has traditionally been interpreted as
the most ancestral cactus since the works of
SCHumANN (1899b) and many of the Pereskia
characters, which have been regarded as plesiomorhic within Cactaceae. These are the
woody stem, the presence of leaves (in contrast
Schumannia 7
2015
Fig. 25: Phylogenetic tree of the Cactaceae showing relationships between all Cactaceae genera based on the phylogenetic studies available
so far. The topology is based on HERNANDEZ-HERNANDEZ & al. (2011), NYFFELER
(2002), GRIFFITH & PORTER (2009) (Opuntioideae), BUTTERWORTH & al. (2002)
(Cacteae), KOROTKOVA & al. (2010, 2011) (Lymanbensonieae and Rhipsalideae), RITZ & al. (2012) (Tephrocacteae) and SCHLUMPBERGER & RENNER (2012)
for Echinopsis and allies. The suprageneric classification and the genera
follow NYFFELER & EGGLI (2012), with modifications based on KOROTKOVA & al.
(2010, 2011), RITZ & al. (2012) and SCHLUMPBERGER & RENNER (2012). Statistical
support for the main clades on tribal level is indicated with *** for maximum support and ** for low support. Support for the higher nodes is
not shown for better readability of the tree. Genera with (?) have so far
not been sampled in a phylogenetic analysis.
Abb. 25: Stammbaum der Cactaceae, gezeigt sind verwandtschaftliche
Beziehungen zwischen allen Cactaceae-Gattungen, basierend auf allen
bisher verfgbaren phylogenetischen Studien. Die Topologie basiert auf
HERNANDEZ-HERNANDEZ & al. (2011), NYFFELER (2002), GRIFFITH & PORTER (2009)
(Opuntioideae), BUTTERWORTH & al. (2002) (Cacteae), KOROTKOVA & al. (2010,
2011) (Lymanbensonieae und Rhipsalideae), RITZ & al. (2012) (Tephrocacteae) und SCHLUMPBERGER & RENNER (2012) fr Echinopsis und Verwandte.
Die supragenerische Klassifizierung und die Gattungen folgen NYFFELER &
EGGLI (2012), mit nderungen basierend auf KOROTKOVA & al. (2010), RITZ &
al. (2012) und SCHLUMPBERGER & RENNER (2012). Die statistische Untersttzung fr die wichtigsten Gruppen auf Tribus-Ebene wird mit *** fr
maximale Untersttzung und ** fr geringe Untersttzung angegeben.
Untersttzung fr die hheren Knoten wird zur besseren Lesbarkeit des
Baumes nicht gezeigt. Gattungen mit (?) wurden bisher in keiner phylogenetischen Analyse untersucht.
26
Schumannia 7
2015
4 Mapping the diversity of cacti

Jens Mutke, Kirsten Burstedde, Jan Laurens Geffert, Andrea Miebach, M. Daud Rafiqpoor,
Anke Stein & Wilhelm Barthlott
4.1 Main data sources

Given the fact that cacti have always fascinated
a large number of scientists, gardeners, and
plant enthusiasts, the taxonomic concepts used
vary a lot. Already Karl SCHumANN in (1899a)
starts his work on the Verbreitung der Cactaceae with a discussion on the problem of a
high degree of synonomy, nomina nuda, etc.
Around 14,000 species names of cacti had already been published in the 1970ies (BARTHLoTT 1977). These included a vast amount of
synonyms. Recent compilations of accepted

species for the family list between 1,431 species
in 124 genera (HuNT 2006) and 1,896 species in
127 genera (ANDERSoN 2001, 2005). For our
database, we used a slightly updated version of
the New Cactus Lexicon species list (Hunt,
pers. comm.).This list includes a few additional
names compared to the New Cactus Lexicon.
Distribution maps are presented in this volume
for only 1,416 species, since some species on
the original list are now regarded as synonyms
in our current database and since the natural
range of some widely naturalized taxa is insufficiently known (e.g., Opuntia ficus-indica). Infraspecific taxa are not considered and are all
included in the respective species distribution
maps.
Infrafamilial classification follows the tribes
and subfamilies of NyFFELER & EGGLI (2010b),
with minor modifications for the epiphytic Lymanbensonieae based on the results of KoRoTKoVA & al. 2010 (cf. Figs. 2425).
For our analyses, we used additional data regarding pollination syndromes (Schlumpber-
Fig. 32: Comparison of the diversity maps of Rhipsalinae from BARTHLOTT (1983) and Rhipsalideae from this work. The basic outline of the diversity
pattern was already published in 1983 based on a very limited database.
Abb. 32: Vergleich der Diversittskarten der Rhipsalinae aus BARTHLOTT (1983) und Rhipsalideae aus dieser Arbeit. Das Grundmuster der Diversittsverteilung wurde schon 1983 basierend auf relativ wenigen Daten erkannt.
30
Schumannia 7
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Chihuahua centre
PueblD-Oaxaca centre
367 species, 43 genera
Species rich genera:
Mammillaria
Echinocereus
Coryphantha
Opuntia
Cylindropuntia
Mammillaria
Opuntia
Ferocactus
Pachycereus
Selenicereus
94
44
40
27
17
6
5
4
3
3
3
Melocactus
Pilosocereus
Rhipsalis
Arrojadoa
Micranthocereus
Pereskia
Mammillaria
Stenocereus
Opuntia
Diversity centres of Cactaceae
Species per 2500 km
Jalisco centre
1-2
3-5
6 - 12
13 - 22
23 - 73
(defined as the top 5%

most species rich areas)
28
10
6
6
6
Caatinga centre
Sonora-Sinaloan centre
Mammillaria
Echinocereus
Ferocactus
Cylindropuntia
Opuntia
Stenocereus
Map 8
6
5
4
Mata Atlantica centre
Southern Central Andes centre

Echinopsis
Rebutia
Parodia
Gymnocalycium
Cleistocactus
16
8
7
5
5
5
59
28
23
19
14
Rhipsalis
Schlumbergera
Lepismium
Pilosocereus
24
5
3
3
1500 km
Barthlott et al. 2013
Map 8: Diversity centres of Cactaceae.

Karte 8: Diversittszentren der Cactaceae (definiert als die oberen 5 % artenreichsten Gebiete; Arten pro 2500 km; species rich genera = artenreiche Gattungen).
lie im Zentrum des Interesses einer ganzen Anzahl von Fachbotanikern, aber noch mehr von
Pflanzenliebhabern (siehe Kap. 1) steht: Es liegt
eine beinahe unberschaubare Flle von publizierten Daten zu Fundorten in Fachpublikationen vor, vorbildhaft sei mapping the Cacti
of mexico (HERNNDEZ & GmEZ-HINoSTRoSA
2011) genannt. Zudem liegen fr beinahe alle
greren Gattungen moderne Revisionen und
taxonomische Bearbeitungen bis hin zu modernen Gesamtbearbeitungen der Familie
(HuNT 2006) vor. Darber hinaus werden in den
zahlreichen nationalen Fachzeitschriften (z. B.
in der Bundesrepublik Deutschland Kakteen
und andere Sukkulenten und den uSA Cactus and Succulent Journal) beinahe monatlich ergnzende Fundortdaten publiziert. Es soll
aber angemerkt werden, dass die Datenlage im
36
Detail doch heterogen ist. Gattungen, die keinen hohen Sammlerwert besitzen (z. B. Opuntia), sind schlechter erfasst als ausgesprochene
Liebhaber-Gruppen (z. B. Mammillaria). Vielleicht eher politisch bedingt ist die Datenlage
fr unterschiedliche geographische Regionen
ebenfalls heterogen: Sdostbrasilien und mexiko sind hervorragend bearbeitet, fr Kuba ist
die Datenlage eher sprlich.
In noch hherem mae gilt dies fr die Erstellung der Diversittskarten, bei denen sich kleinere Fehler innerhalb der Arten-Arealkarten
der erfassten Arten aufheben. Zum Vergleich
haben wir in Abb. 32 unsere, auf einer relativ
engen Datenbasis beruhende Diversittskarte
der Rhipsalideen von 1983 der jetzt erarbeiteten Karte gegenbergestellt. Wenn man die
durch die unterschiedliche Projektion beding-
ten verschiedenen umrisslinien der Kontinente herausrechnet, sind die Karten beinahe
deckungsgleich. Insofern ist voraussehbar, dass
es z. B. an der Biodiversittskarte der gesamten
Familie (Karte 7) auch bei sich zunehmend verbesserter Datenlage in den nchsten Jahrzehnten kaum eine nderung mehr ergeben wird.
Zusammenfassend lsst sich feststellen, dass
die Familie der Cactaceae durch die vorliegenden publizierten Daten ein herausragendes
Beispiel fr die Analyse der Biogeographie
und Biodiversitt einer greren Pflanzenfamilie berhaupt darstellen. Es gibt wohl keine
zweite Familie entsprechender Gre innerhalb des ganzen Pflanzenreiches, die eine vergleichbare Analyse erlauben wrde.
Schumannia 7
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5 Patterns of diversity and endemism

Jens Mutke, Kirsten Burstedde, Jan Laurens Geffert, Andrea Miebach,
M. Daud Rafiqpoor, Anke Stein & Wilhelm Barthlott
5.1 General overview

Except for the high latitude boreal forests and
Tundra vegetation of North America and the
temperate rain forests of southern Chile, at
least some cactus species can be found anywhere in the Americas (map 7). Rhispsalis baccifera is the only species naturally found also in
the old World, ranging from tropical West
Africa via madagascar, mauritius, and Runion
to Sri Lanka.
Highest species richness of cacti ( 23 species
per 2,500 km, the 5% most species rich areas;
maps 8 & 9) can be found in seven distinct centres in mexico, Brazil, Argentina, and Bolivia.
At the genus level, additional centres are located
in Peru and in northern Venezuela (map 10)
these are covered more in detail in chapter 5.6.
The four North American centres of high
species richness in mexico and the southern
uSA are located in dry ecoregions (Table 1,
map 11): 1) the Chihuahua and the meseta
Central matorral, 2) the Puebla-oaxaca centre
with a diversity of different climates and habitats, as well as two minor areas 3) near the Pacific coast of Jalisco (Jalisco centre), and 4) at
the border of the southern Sonora desert to the
Sinaloan dry forests (Sonora-Sinaloan centre).
In South America, there are three main centres of cactus species richness in 1) the southern central Andes in Bolivia and northern Argentina, 2) the Caatinga, and 3) the coastal forest area of south-east Brazil. The latter centre
is mainly dominated by epiphytic species of the
Rhipsalideae (compare Table 2, map 12).
Intermediate species richness can be found all
over south-east Brazil, which is linked via a corridor of intermediate species richness in
uruguay and Paraguay to the Bolivian centre
of diversity. Coastal Peru and parts of the central Andean dry valleys show intermediate
species richness, but an especially high concentration of species with extremely restricted
distribution ranges. Northern Venezuela and
most parts of the Caribbean also show intermediate levels of species richness.
Low species richness ( 5 species per 2,500
km) occur in large parts of the Amazon Basin,
many fire-prone habitats, e.g., of the Brazilian
Schumannia 7
2015
Cerrado, the Llanos Savannah of Colombia

and Venezuela, mediterranean climate areas in
California and Chile, and in the temperate eastern uSA.
5.2 Centres of diversity

For the delineation of the centres of cactus
species richness (map 8,Tables 1 & 2), we used
the top 5% areas with highest species numbers
( 23 species per 2,500 km). To qualify as one
of the centres listed below, at least four adjacent grid cells of 50 x 50 km have to fulfil this
criterion.We distinguish seven diversity centres.
The four north/central American centres
Chihuahua centre: The mexican centre of diversity with the largest area and by far the highest species numbers is the large complex of
deserts and xeric shrublands of the Chihuahua
desert, including the mexican meseta Central
matorral (compare Table 1). It includes parts of
the north-eastern Sonora desert as well. Highest species richness can be found in the southeastern part of the centre with a diverse mosaic
of matorral and, e.g., the Sierra madre oriental Pine-oak Forests. map 11 shows the fine
scale patterns of species richness within the
mexican centres in smaller grid cells of 10 x 10
km. The centre is characterized especially by a
high diversity of globular species of the tribe
Cacteae and is the main centre of species richness of the opuntioideae. most other tribes are
absent or poorly represented. The growth
forms of shrubby and arborescent cacti also
have their centres of species richness in the
Chihuahua, whereas columnar cacti are almost
absent.
Puebla-Oaxaca centre: This mexican centre
mainly includes the Tehuacn-Cuicatln valley.
It is geographically isolated from the nearby
and much larger Chihuahua centre by the Transmexican volcanic mountain belt. Its topographic diversity (geodiversity in a wider sense)
results in a variety of vegetation types such as
tropical dry forests, xeric shrublands, thorn
shrubs, and deserts.The Puebla-oaxaca centre
has the highest concentration of columnar
species of the tribe Cacteae in mexico. In contrast to the Chihuahua centre, this centre is
home to a high number of cacti pollinated by
bats, most of them columnar. The mean range
size of the species occurring in this region is much
smaller compared to the Chihuahua centre. many
of the species are restricted to this region.
Sonora-Sinaloan centre: This centre comprises
a small area of high species richness at the
southern transition of the Sonora desert to the
Sinaloan dry forest dominated by deciduous
thorn forest. The species rich north-eastern
part of the Sonora desert, in contrast, is included in our larger Chihuahua centre (see
above). unlike the situation in those northern
parts of the Sonoran desert, frost is rare in the
southern Sonora-Sinaloan centre. The region
is characterised by a mixture of temperate and
tropical species.The growth forms of cacti present in the southern Sonora include globose
species (e.g. Mammillaria, Ferocactus) and columnar species (e.g. Stenocereus) of the Cacteae, columnar species of the Phyllocacteae subtribus Echinocereinae and shrubby Opuntia.
Jalisco centre: This small centre of high cactus
diversity at the pacific coast of the mexican state
Jalisco has a more humid climate compared to
the other northern diversity centres resulting in
tropical dry forest vegetation. As a result, this
centre is home not only to globular Mammilaria
species and abundant columnar and arborescent
cacti like Stenocereus, Cephalocereus (all Cacteae), and Opuntia, but also epiphytic species of
Epiphyllum and Selenicereus.
The three South American centres
Southern central Andes centre: The largest
South American centre of diversity also harbours the highest species number in South
America and is characterized by the highest
phylogenetic and morphological diversity. 10
out of the 11 tribes of cacti accepted here occur in the southern central Andes centre. With
eight tribes occurring per 2,500 km grid cell in
large parts of this centre, this is twice the number found in most parts of the Chihuahua centre. The centre is located in the Semiarid and
arid valleys of central and southern Bolivia and
northern Argentina. In the northern part of the
37
Map 10
1-2
3-5
6-8
9 - 12
13 - 22
Diversity centres of Cactaceae: genera

(defined as the top 5% most JHQXV rich areas)
Genera per 2500 km
1500 km
Map 10: Generic diversity centres of Cactaceae (highlighting the 5% most genus rich areas, compare map 26)
Karte 10: Diversittszentren der Cactaceae, Gattungen (definiert als die oberen 5 % gattungsreichsten Gebiete; Arten pro 2500 km).
Schumannia 7
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39
centre, original vegetation comprises deciduous forests with scattered naturally forest free
steep slopes with rock outcrops. In the south,
in the prepuna ecoregion, Andean shrublands
and semideserts are found. This centre harbours the highest number of globular cacti in
South America, but is also a centre of columnar cacti. It is the only South American region
with significant numbers of cushion forming
cacti. In addition to mainly bee-pollinated
species, it has a considerable number of sphingophilous species. Especially the western Andean part of the Bolivian province Tarija and
neighbouring areas of Salta and Jujuy in northern Argentina are extremely diverse when
looking at fine scale patterns of species richness
in 10 x 10 km grid cells (map 12).
Caatinga centre: This Brazilian centre is located in the southern part of the Caatinga
ecoregion and the Campos Rupestres linked to
mountain ranges of the Serra Espinhaco and
Chapada Diamantina.While the dominant vegetation types of the Caatinga region are xeric
shrubland and thorn forest, highest species diversity of cacti can be found in azonal habitat
islands such as rock outcrops (inselbergs) or
nutrient poor sandy soils.Although species diversity is much lower compared to the Chihuahua or the southern central Andes centre,
the Caatinga centre is home to a high number
of generaespecially of the tribe Cereeae.Accordingly, this centre is characterized by only
moderate species numbers of globose cacti, but
a high diversity of columnar species. The Caa-
tinga centre has the highest numbers of cacti

pollinated by bats and birds and only low numbers of bee-pollinated species.
Mata Atlntica centre: This centre in southeast Brazil is ecologically quite different from
all the others, as it is characterized by humid,
evergreen rain forest. Important taxa here are
the arborescent species Pereskia grandifolia,
and particularly a large number of epiphytic
species of Rhipsalis and related genera. Some
other species in this centre grow in isolated
azonal habitats such as coastal sand dunes or
rock outcrops (inselbergs, Figs. 5 & 6) (mostly
Cereeae, same genera as in the Caatinga centre). only small patches of the original vegetation of the south-east Brazil Atlantic rainforest
are left today.
Table 1: The North/Central American diversity centres
Centre
Chihuahua
Puebla-Oaxaca
Sonora-Sinaloan
Jalisco
area size (km)
1,020,000
47,500
12,500
12,500
vegetation
deserts and xeric

shrublands
tropical dry forests,

deserts and xeric
shrublands
transition of Sonoran
desert and Sinaloan
dry forests
tropical dry forest
main WWF
ecoregions
Chihuahuan desert (NA1303);

Central mexican matorral
(NA1302); Sierra madre
oriental pine-oak forests
(NA0303)
Tehuacn valley
matorral (NT1316),
Balsas dry forests
(NT0205)
Sonoran-Sinaloan
transition subtropical
dry forest (NA0201)
Jalisco dry forests

(NT0217)
dominating
taxa
Cacteae
Cacteae
Phyllocacteae
Echinocereinae, Cacteae,
Opuntia
Phyllocacteae
Echinocereinae &
Hylocereinae, Cacteae,
Opuntia
most species
rich growth form
globular
globular
globular
globular, epiphytic
dominating
pollination syndrome
bee pollination
bee pollination
bee pollination
bee pollination
Table 2: The South American diversity centres
Centre
Southern Central Andes
Caatinga
Mata Atlntica
area size (km)
297,500
157,500
40,000
vegetation
inter-Andean dry forests and

shrublands with steep rocky
slopes
xeric shrubland and thorn

forest with rock outcrops
atlantic rain forests, coastal

sand dunes, granit rock
outcrops
main WWF ecoregions
Bolivian montane dry forests

(NT0206), central Andean puna
(NT1002), southern Andean
yungas (NT0165)
Caatinga (NT1304), Campos

Rupestres montane savanna
(NT0703), Atlantic dry forests
(NT0202)
Serra do mar coastal forests

(NT0160); Bahia coastal
forests (NT0103)
dominating taxa
Trichocereinae
Cereinae
Rhipsalideae
most species rich growth form
globular
columnar
epiphytic
dominating pollination syndrome
bee pollination
bird pollination, bat pollination
bee pollination
40
Schumannia 7
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Map 11
Map 12
1 - 10
11 - 23
24 - 28
29 - 35
36 - 40
41 - 45
46 - 50
51 - 73
Diversity centres
1 - 10
11 - 23
24 - 28
29 - 35
36 - 40
41 - 50
51 - 60
61 - 73
Diversity centres
Diversity centres
in South America
(defined as the top
5% most species
rich areas)
Diversity centres in Central America

(defined as the top 5% most species rich areas)
Species per 100 km
250 km
500 km
Map 11: Diversity centres in Central America.

Karte 11: Diversittszentren in MIttelamerika (definiert als die
oberen 5 % artenreichsten Gebiete; Arten pro 100 km).
In addition to those seven centres presented

above, very small areas in Rio Grande do Sul
(south-east Brazil) and in south-east Paraguay
reach more than 23 species per 2,500 km. Especially south-east Paraguay is an important
area with high genus diversity (map 10, see
chapter 5.6 for a more detailed discussion of diversity patterns at higher taxonomic level).
Even though the Atacama Desert and the
western slopes of the Andes in Peru harbour
only moderate species diversity, they are home
to a) a high diversity of species with extremely
small distribution ranges (compare chapter 5.3)
and b) intermediate to high numbers of genera
(compare chapter 5.6). The Andes of northern
Peru are one of the few regions were at least 7
out of the 11 tribes of the cacti occur (map 27).
Additional centres of high taxonomic diversity
are the coastal mountains of northern Venezuela with high numbers of genera and tribes,
and parts of the Greater Antilles, especially
south-east Cuba and the islands of Hispaniola
and Puerto Rico (maps 26 & 27).
5.3 Range-sizes of Cacti

most species of cacti have extremely small distribution ranges. As a comparison: whereas
Schumannia 7
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Species per 100 km

Map 12: Diversity centres in South America.

Karte 12: Diversittszentren in Sdamerika (definiert als die
oberen 5 % artenreichsten Gebiete; Arten pro 100 km).
Birdlife International uses a threshold of

50,000 km range size for the definition of endemic bird areas, almost half of the cactus
species have range sizes smaller than 10,000
km (map 13). more than 300 species have
ranges even smaller than 2,000 km. Examples
of species with extremely small ranges, restricted to one or a few populations, are Discocactus horstii in minas Gerais (Brazil),
Aztekium ritteri and Geohintonia mexicana in
the Sierra madre oriental in Nuevo Leon
(mexico), and Cereus insularis on the islands of
Fernando de Noronha (Brazil) (compare Figs.
3537).
Looking at the distribution patterns of the
range-restricted species (narrow endemics
with range sizes smaller than 10,000 km, map
13), only some of them occur in one of the
seven diversity centres listed above. Narrow
endemic species are found only in the southern
central Andes centre (e.g. species of Cleistocactus, Parodia, Rebutia), the Puebla-oaxaca
centre (many Mammillaria species) as well as
in the (south-)eastern parts of the Chihuahua
centre (many species of Mammillaria, Thelocactus, Turbinicarpus, and others). Additional
centres of range-restricted cacti are parts of
the coastal Atacama Desert and the western
slopes of the Andes, especially in northern Peru
(e.g. Espostoa, Haageocereus, Matucana, and

many smaller genera), and the northern parts
of the Chilean matorral (mainly Copiapoa,
Eriosyce). Small numbers of range-restricted
species occur all over south-east Brazil, northern and central Argentina and Chile, and large
areas of mexico and Costa Rica.
At least some widespread species occur in almost all parts of the overall distribution range
of the Cactaceae.As a result, the median range
sizes for the species of a given place is higher
than 100,000 km in most grid cells (map 14)
even though most cactus species have distribution ranges of less than 10,000 km! However, those 50% species with small ranges are
scattered over many places and only few of
them co-occur in one place.
The species with the largest range size within
the Cactaceae is the tricontinental Rhipsalis
baccifera. Also within the Americas, Rhipsalis
baccifera has the largest distribution range of
all cacti species. many species with very large
range sizes are either epiphytic Rhipsalideae
or Hylocereinae (e.g., Hylocereus, Lepismium)
or shrubby opuntioideae. Nevertheless, even
within the epiphytic growth form, most species
of the genera Weberocereus in Central America or Schlumbergera in south-east Brazil have
very small known distribution ranges. In the
41
Map 13
1
2
3-4
51 - 6
72 - 8
93 -- 410
5-6
11
7 - 8- 15
16
- 25
9 - 10
11 - 15
Range-restricted
species
16 - 25
('endemics')
Range-restricted species
('endemics')
Species
per 2500 km
Species
per
km 10,000 km
with ranges2500
under
1500 km
with ranges under 10,000 km
Map 13: Range-restricted species (endemics).

Karte 13: Arten mit kleinem Areal (Endemiten; Arten pro 2500 km mit Arealen kleiner als 10.000 km).
42
Schumannia 7
2015
Map 15
1-2
3-5
6 - 10
11 - 15
16 - 25
26 - 35
36 - 45
46 - 56
Map 16
1-2
3-4
5-6
7-8
9 - 10
11 - 12
13 - 14
15 - 16
Growth form Globular:

Species diversity (692 sp.)
Growth form Columnar:

Species diversity (205 sp.)
1500 km
Species per 2500 km
1500 km
Species per 2500 km
Map 15: Growth form globular, species diversity (692 sp).

Karte 15: Wuchsform kugelig, Artendiversitt (692 Arten;
Arten pro 2500 km).
Fig. 33: An extraordinary life-form within the

Opuntieae: liana Tacinga braunii at its type
locality in Brazil (near Itaobim, state Minas
Gerais). Photo: W. Barthlott.
Abb. 33: Eine auergewhnliche Lebensform
innerhalb der Opuntieae: die Liane Tacinga
braunii an ihrem Typfundort in Brasilien
(bei Itaobim, Staat Minas Gerais).
44
Map 16: Growth form columnar, species diversity (205 sp).

Karte 16: Wuchsform sulig, Artendiversitt (205 Arten;
Arten pro 2500 km).
genus Rhipsalis, more than one third of the

species have distribution ranges smaller than
10,000 km. Looking at the largest genera of
the family, the range size distribution is very
uneven. In the large genus Mammillaria, which
is mostly restricted to mexico and southern
uSA, more than half of the species have distribution ranges smaller than 2,000 km (2/3
less than 10,000 km), whereas some taxa such
as M. heyderi, M. lasiacantha, or M. grahamii
are really widespread. A similar situation is
found in Parodia, where 70% of the species
mainly in Rio Grande do Sul (Brazil) and uruguay on the one hand and the southern central
Andes centre in Bolivia and Argentina on the
other handhave range sizes smaller than
10,000 km. In contrast, most species of the
genus Opuntia are widespread, and only a few
have restricted ranges of less than 10,000 km.
Echinopsis, which as a genus is restricted mainly to the (southern) central Andes, has species
of all range sizes, similar to Echinocereus in
mexico and the southern uSA. Gymnocalycium, which is mainly from Argentina, is an example for a genus where most species have
very small range sizessimilar, e.g., to Eriosyce
in Chile.
There are other genera, which as a genus have

large ranges, but include many restricted range
species. In the genus Pilosocereus, which occurs from south-east Brazil to northern mexico, almost half of the species have distribution
ranges smaller than 10,000 km, and only few
have ranges of several hundred thousand km.
A somewhat similar situation can be found in
the genus Melocactus, which has many restricted range species especially in south-east
Brazil but can be found even in mexico and
the Caribbean.
5.4 Spatial distribution of growth

forms
There is a strong difference in the diversity patterns of the two most species rich growth forms
within cacti: the globular and the columnar
species (maps 15 & 16). By far the highest
species numbers of globular cacti occur in the
Chihuahua centre (almost exclusively tribe
Cacteae). only moderate species numbers are
found in the southern central Andes centre
(mainly Cereeae subtribes Trichocereinae and
Notocacteae), and in Rio Grande do Sul
Schumannia 7
2015
Fig. 35: Discocactus horstii, extremely

range-restricted species (near Diamantina,
Minas Gerais, Brazil). Photo: W. Barthlott.
Abb. 35: Discocactus horstii, eine Art mit
extrem kleinem Verbreitungsgebiet
(bei Diamantina, Minas Gerais, Brasilien).
Fig. 36: Highly endemic Pelecyphora aselliformis in the Mexican state San Luis Potos. Photo:
W. Barthlott.
Abb. 36: Pelecyphora aselliformis ist im mexikanischen Staat San Luis Potos endemisch.
Fig. 38: Micranthocereus polyanthus in a field of white quartz sand near

Caetit (Bahia, Brazil). Photo: W. Barthlott.
Abb. 38: Micranthocereus polyanthus in weiem Quarzsand bei Caetit
(Bahia, Brasilien).
46
Fig. 37: Highly endemic Geohintonia mexicana

in Nuevo Len, Mexico. Photo: W. Barthlott.
Abb. 37: Geohintonia Mexicana, endemisch in
Nuevo Len, Mexiko.
Fig. 39: Cereus jamacaru, an example for an arborescent growth form

with columnar branches, with Werner Rauh (right) and Diedrich J. Supthut
(left) in the Brazilian state Bahia. Photo: W. Barthlott
Abb. 39: Ein Beispiel fr eine baumartige Wuchsform mit suligen sten:
Cereus jamacaru, mit Werner Rauh (rechts) und Diedrich J. Supthut (links)
im brasilianischen Staat Bahia.
Schumannia 7
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pischen Trockenwaldvegetation. In der Folge

ist dieses Zentrum nicht nur Heimat kugeliger
Mammillaria-Arten und hufiger suliger und
baumfrmiger Kakteen wie Stenocereus, Cephalocereus (alle Cacteae) und Opuntia, sondern auch von epiphytischen Arten der Gattungen Epiphyllum und Selenicereus.
Die drei sdamerikanischen Zentren
Sdliches Zentralanden-Zentrum: Das grte
sdamerikanische Diversittszentrum beherbergt auch die hchste Artenanzahl in Sdamerika und ist durch die hchste phylogenetische und morphologische Diversitt gekennzeichnet. Zehn der hier akzeptierten elf
Kakteen-Triben kommen in dem sdlichen
Zentralanden-Zentrum vor. mit in groen Teilen dieses Zentrums acht vorkommenden Triben pro 2.500-km-Rasterfeld ist die Anzahl
doppelt so gro wie in den meisten Gebieten
Fig. 43: Growth forms of cacti plotted on the phylogenetic tree of Fig. 25 (comp. the generic names).
Abb. 43: Wuchsformen der Kakteeen, in den phylogenetischen Stammbaum der Abb. 25 bertragen
(vgl. dort die Gattungsnamen). [Legende von oben nach unten: kugelig, sulig, sulig-baumfrmig,
strauchig, baumfrmig, epiphytisch, polsterfrmig.]
56
des Chihuahua-Zentrums. Das Zentrum befindet sich in den semiariden und ariden Tlern
des zentralen und sdlichen Boliviens sowie
des nrdlichen Argentiniens. Im nrdlichen
Teil des Zentrums umfasst die Vegetation laubwerfende Wlder mit dispersen, natrlich
waldfreien Steilhngen und felsigen Lichtungen. Im Sden, der Prpuna-koregion, dominieren Andenbuschlnder und Halbwsten.
Dieses Zentrum beherbergt die hchste Anzahl kugeliger Kakteen in Sdamerika, ist aber
auch ein Zentrum der Sulenkakteen. Es ist
die einzige sdamerikanische Region mit einer
signifikanten Anzahl polsterbildender Kakteen. Zustzlich zu den vorwiegend bienenbestubten Arten hat es eine beachtliche Anzahl schwrmerbltiger Arten. Besonders der
westliche Teil der bolivianischen Provinz Tarija und die benachbarten Gebiete Saltas und
Jujuys in Nordargentinien sind extrem vielfltig, wenn man den Artenreichtum in einer feinen Skala von 10 x 10 km-Rasterzellen betrachtet (Karte 12).
Caatinga-Zentrum: Dieses brasilianische Zentrum liegt im sdlichen Teil der Caatinga-koregion und den Campos Rupestres, die an die
Bergketten der Serra Espinhao und Chapada
Diamantina anknpfen. Whrend trockenes
Buschland und Dornwald die vorherrschenden
Vegetationstypen der Caatinga-Region sind, ist
die hchste Kakteenartendiversitt in azonalen Habitaten wie Felskuppen (sog. Inselberge)
oder auf nhrstoffarmen sandigen Bden zu
finden. obwohl die Artenvielfalt im Vergleich
zu dem Chihuahua- oder sdlichem Zentralanden-Zentrum viel niedriger ist, kommt hier
eine hohe Anzahl von Gattungen besonders
der Tribus Cereeae vor. Dementsprechend ist
dieses Zentrum durch eine mittlere Artenzahl
kugeliger Kakteen, aber eine hohe Diversitt
suliger Arten charakterisiert. Das CaatingaZentrum hat die hchsten Zahlen von durch
Fledermuse und Vgel bestubten Arten und
nur geringe Zahlen bienenbestubter Arten.
Mata Atlntica-Zentrum: Dieses Zentrum in
Sdostbrasilien ist von allen anderen Zentren
kologisch recht deutlich unterschieden, da es
durch feuchten, immergrnen Regenwald gekennzeichnet ist. Wichtige Taxa sind hier die
baumfrmige Art Pereskia grandifolia und besonders eine hohe Anzahl epiphytischer Arten
von Rhipsalis und verwandten Gattungen. Einige andere Arten dieses Zentrums wachsen
in isolierten azonalen Habitaten wie Kstendnen oder Felskuppen (Inselberge, Abb. 5 &
6) (meist Cereeae, die gleichen Gattungen wie
im Caatinga-Zentrum). Nur kleine Flecken der
ursprnglichen Vegetation des sdostbrasilianischen Regenwaldes sind heute noch brig.
Auer den sieben, oben vorgestellten Zentren
erreichen sehr kleine Gebiete in Rio Grande
Schumannia 7
2015
meter groe Verbreitungsgebiete. Eine etwas

hnliche Situation ist in der Gattung Melocactus zu finden, die viele kleinrumig verbreitete
Arten besonders in Sdostbrasilien hat, aber
sogar in mexiko und der Karibik vorkommt.
5.4 Rumliche Verbreitung der

Wuchsformen
Fig. 45: Convergent evolution of red hummingbird flowers in South and North America in unrelated
genera of small globular cacti: Mammillaria poselgeri from Mexico (Baja California) (left), Matucana
formosa from N-Peru (right). It is almost typical that a different flower syndrome within the predominantly entomophilous genus Mammillaria led to creation of a separate genus (Cochemiea poselgeri) in the past. Photos: W. Rauh.
Abb. 45: Konvergente Entstehung roter Kolibriblumen in Sd- und Nordamerika bei nicht verwandten Gattungen kleiner Kugelkakteen: links Mammillaria poselgeri aus Mexiko (Baja California),
rechts Matucana formosa aus N-Peru. Es ist beinahe typisch, dass ein abweichendes Bltensyndrom
innerhalb der vorwiegend entomophilen Gattung Mammillaria frher zur Abtrennung als eigene
Gattung (Cochemiea poselgeri) fhrte.
Fig. 46: Convergent evolution of night-blooming, white, hawkmoth pollinated flowers with extremely
long flower tubes in not related species: the dwarf columnar cactus Echinopsis mirabilis from Argentina (left), Epiphyllum phyllanthus (right), a widespread (Mexico to N-Argentina) rainforest epiphyte.
E. phyllanthus is one of the extremely spingophilous species with a flower tube of up to about 20 cm
length and can probably be pollinated only by the moth Cocytius cruentus. Photos: W. Barthlott.
Abb. 46: Konvergente Entstehung nachtblhender weier Schwrmerblumen mit extrem langen Bltenrhren bei nicht verwandten Arten: links der zwergige Sulenkaktus Echinopsis mirabilis aus Argentinien, rechts Epiphyllum phyllanthus, ein weitverbreiteter (Mexiko bis Nordargentinien) RegenwaldEpiphyt. E. phyllanthus ist einer der extrem spingophilen Arten mit einer Bltenrhre von bis ber
20 cm Lnge und kann mglicherweise nur von dem Schwrmer Cocytius cruentus bestubt werden.
62
Die Verbreitungsmuster der zwei artenreichsten Wuchsformen innerhalb der Kakteen (die
kugeligen und die suligen Arten) zeigen deutliche unterschiede (Karten 15 & 16). Bei weitem die hchsten Artenzahlen der Kugelkakteen kommen im Chihuahua-Zentrum vor
(fast ausschlielich der Tribus Cacteae). Nur
mittlere Artenzahlen findet man im sdlichen
Zentralanden-Zentrum (vorwiegend Cereeae
der Subtriben Trichocereinae und Notocacteae) und in Rio Grande do Sul (Sdost-Brasilien). Geringe Artenzahlen findet man in der
Caatinga (vorwiegend Melocactus). Im Gegensatz dazu gibt es fast keine Sulenkakteen im
zentralen Teil des Chihuahua-Zentrums; deren
hchste Diversitt ist in der Caatinga zu finden
(besonders Cereeae Subtribus Cereinae, Abb.
39), im sdlichen Zentralanden-Zentrum (besonders Cereeae Subtribus Trichocereinae)
und in der Atacama-Wste und den westlichen
Hngen der Anden in Peru (Cereeae Subtribus Trichocereinae). Das kleine Puebla-oaxaca-Zentrum in mexiko hat, im Gegensatz zu
dem benachbarten Chihuahua-Zentrum, ebenfalls eine signifikante Anzahl suliger Kakteen,
hauptschlich aus der Phyllocacteae Subtribus
Echinocereinae.
Die strauchigen und baumfrmigen Arten von
Pereskia und den opuntioideae (Karten 18 &
19,Abb. 40) weisen ihre hchsten Artenzahlen
im Chihuahua-Zentrum und dem Puebla-oaxaca-Zentrum auf, und nur eine geringe Artenvielfalt in Sdamerika.
Epiphytische Kakteen haben, verglichen mit
dem Rest der Familie, deutlich verschiedene
kologische Prferenzen (Karte 20). Die epiphytischen Rhipsalideae (Abb. 41, vorwiegend
sdamerikanisch, besonders Sdost-Brasilien)
und Hylocereinae (vorwiegend mittelamerika)
sind am artenreichsten in den Tieflandregenwldern der mata Atlntica und mittelamerikas. Jedoch kommen nur wenige Arten beider
Triben (z. B. Disocactus amazonicus und Strophocactus witii) in dem groen Regenwaldgebiet des Amazonasbeckens vor vorwiegend
im westlichen Amazonien.
Polsterbildende Kakteen (Abb. 42, hauptschlich Phyllocacteae-Echinocereinae und -Cylindropuntieae, sowie Maihuenia) sind an speziellen Standorten mit Frost oder stark trocknenden Winden, wie in den Hochanden Sd-
Schumannia 7
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6 Conservation and hotspots:

cactus diversity in change
Pierre L. Ibisch & Jens Mutke
6.1 Introduction
As cacti represent popular plants of great ornamental value, for a long time they have been
the subject of conservationist concerns (e.g.,
BENSON 1982, OLDFIELD 1984a, b). Due to a relatively better knowledge of many species, but
also as a consequence of early overexploitation for horticulutural uses, cacti, since the
1970s, were among the first plants to become
protected under international law. The whole
family Cactaceae is included in the Appendices
I and II of the Convention on international
trade in endangered species of wild fauna and
flora (CITES). This protection was especially
related to the international trade in endangered species and to high demands in the ornamental plant market, but it did not take
threats from other sources into account. With
increasing global conservation efforts aiming
at the establishment of a more effective protected area network, numerous species of Cactaceae have also benefitted from in-situ conservation measures. Still, the distribution
ranges of 19% of the species ranges do not
overlap with any protected area and 40% of
the species ranges are not covered by protected
areas of the IUCN categories I to IV. Most of
these have distribution ranges smaller than
50,000 km. Clearly, the representation within
a protected area does not automatically guarantee effective conservation, and the existence
of a species outside protected areas is not indicating imminent threat.
The geographical assessment of cacti presented here can be a starting point for a more
informed evaluation of their threat and conservation status. Clearly, many threats to biodiversity in general, and to cacti in particular,
have been dynamically unfolding and changing over the last decades. Still relevant treatments of cacti conservation have been provided by TAYLOR & al. (1997) and TAYLOR &
ZAPPI (2004), which, however can be complemented by integral and systematic assessments. The systematic discussion of threats to
cacti according to modern threat classification
approaches (SALAFSKY & al. 2008) is a first step,
68
Fig. 51: Shield inselberg in Bahia, Brazil, with Melocactus ferreophilus; agricultural not usable relict
habitat in a destroyed landscape. Photo: W. Barthlott.
Fig. 51: Schildinselberg in Bahia, Brasilien, mit Melocactus ferreophilus; ein landwirtschaftlich nicht
nutzbarer Relikt-Standort inmitten einer zerstrten Landschaft.
which has also been adopted in the course of

the elaboration of the IUCN Red List (IUCN
2012). Additionally, spatial analyses allow for
a certain differentiation of conservation problems in the different regions of the Cactaceae
range. In the future, these spatial analyses can
be refined further, e.g. including detailed gap
analyses.This is urgently required as the IUCN
Red List is far from being complete and up to
date.
The IUCN Red List still shows a strong bias
towards a few well investigated countries such
as Brazil and Mexico. About 48% of all cacti
species, which have been assigned to one of the
Red List categories Vulnerable, Endangered,
Critically endangered, Extinct in the wild or
Extinct, are Mexican taxa, and 49% are Brazilian. The analysis also might underestimate the
threat status of species living outside arid regions. According to the current Red List
(IUCN 2012), about 60% of the most endangered species (categories as above) would live
in deserts or rocky areas, and only about 12%
are forest taxa. However, in the case of many
epiphyte species from forest habitats (e.g., in
the very highly fragmented and deforested

Mata Atlantica region, Brazil) it seems likely
that the conservation status of many cactus
species is poor. IBISCH & al. (2000) indicated
that a few Bolivian epiphytic cacti would be at
least vulnerable to extinction, and the situation
is likely to have worsened.According to IUCN
(2012) however, the only epiphyte species that
are classified as vulnerable or more critical are
Rhipsalis cereoides, R. crispata, R. pilocarpa,
R. russellii, and Schlumbergera kautskyi (all
from Brazil). Intense monitoring efforts are required to detect significant population declines.
These are often affordable only for rich countries, where even single populations and varieties are carefully observed (e.g., compare isolated populations in the Sonoran desert of
south-central Arizona of Nichols turks head
cactus Echinocactus horizonthalonius var.
nicholii with a dramatic decline from 1995 to
2008 (MCINTOSH & al. 2011).
In the following we discuss the principal threats
to cacti adopting the standard classification by
SALAFSKY & al. (2007), which is also applied by
IUCN (2012).
Schumannia 7
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6.2 Threats to Cactaceae

Residential and commercial development, human settlements or other nonagricultural land
uses with a substantial footprint
Many fast growing cities exist in (sub)tropical
arid areas, where agriculture-related land use
change tends to be less important. With increasing human population growth, development, and urbanization, area consumption for
residential and commercial infrastructure becomes ever more relevant also for cacti. An
example is Cleistocactus acanthurus, which is
threatened by the development of Lima,
Chosica and satellite cities (MOTTRAM 1998). In
Latin America, a significant number of large
cities are located in more or less narrow arid
valleys with relevant occurrences of cacti. For
example, in Bolivia, the rapid growth of the city
of Cochabamba and satellite settlements drives
the conversion of habitats both of the bottom
and the slopes of the valley system affecting
cacti populations of Cleistocactus parviflorus,
Echinopsis obrepanda, Parodia schwebsiana
among others (previously even benefitted by
deforestation, IBISCH 2004b) (Fig. 55). In an assessment of the Cactaceae of the Quertano
semi-desert it was concluded that fewer species
were threatened by residential development
than other land-use changes; however, the corresponding impact was considered more severe
(CHVEZ & al. 2007). While agricultural landuse impacts mainly zonal ecosystems with
more or less fertile soils, residential development also affects azonal sites such as rock outcrops or steep slopes, especially in the case of
megacities suffering from decreasing space for
expansion (Figs. 5152). Non-agricultural land
consumption must not be neglected as a problem especially for specialist plants in azonal
habitats. Currently, 14% of the cacti recorded
by IUCN (2012), which have been assigned to
one of the Red List categories Vulnerable, Endangered, Critically endangered, Extinct in the
wild or Extinct suffer from threats belonging to
this category.
Agriculture and aquaculture threats from
farming and ranching as a result of agricultural
expansion and intensification, including silviculture, mariculture, and aquaculture
About 14% of all cacti species, which have
been assigned to one of the Red List categories
Vulnerable, Endangered, Critically endangered, Extinct in the wild or Extinct, are threatened by agriculture driven land-use change
(IBISCH & al. 2000, IUCN 2012, MLLER 2007).
For a long time, many cacti, in many regions
benefited from extensive land-use. Deforestation and the subsequent grazing in (semi-)arid
shrub-lands often led to the expansion of suit-
Schumannia 7
2015
able habitat for species that were stress-tolerant and resistant to herbivory, but poor competitors, including many terrestrial cacti (IBISCH
2004a). Indeed, grazing can benefit cactus populations (e.g., in the case of some Mammillaria
species in Tehuacn Valley, MARTORELL & PETERS 2009). However, with increasing intensity
of grazing the degradation of ground vegetation and soils leads to severe hydric erosion
triggering loss of habitat, and the intensification of agricultural land-use e.g. applying massive mechanical clearing, burning and irrigation the elimination of cactus habitat is promoted. With rapidly growing demand for
agricultural commodities and arable land, advancement of industrial agriculture even under rather marginal conditions, as well as progressing desertification, land conversion is becoming an ever more important threat to
species of Cactaceae. This is also increasingly
reflected in reports about principal threats: excessive pasturing and exploitation of the
desert, habitat fragmentation and land-use
changes, and anthropogenic fire are increasingly named as relevant conservation problems
(e.g., BRAILOVSKY SIGNORET & HERNNDEZ 2010,
CHVEZ & al. 2007, CLARK-TAPIA & al. 2005, EMMING 2005, GOLUBOV & al. 2009, LEON DE LA
LUZ 2005). In some cases, the compositional
change of plant communities is causing threats;
e.g., Opuntia humifusa, originally inhabiting
open Quercus savanna vegetation was reported
to suffer from shading by an encroaching
canopy of Quercus and Pinus, and smothering
by leaf litter (ABELLA & JAEGER 2004). Afforestation activities can threaten habitats of
Cactaceae, such as in the case of Neoporteria
aspillagae being affected by Pinus radiata plantations in Chile (NOVOA 2002).
Energy production and mining threats from
production of nonbiological resources
Mining and extraction of rocks can very specifically threaten species of Cactaceae at azonal
sites otherwise not affected by land-use changes.
For instance, this has been reported for Turbi-
Fig. 52: The highly mimetic Ariocarpus kotschoubeyanus (red circle) next to a flowering Proboscidea
in the Mexican state San Luis Potos. Photo: W. Barthlott.
Abb. 52: Der hoch mimetische Ariocarpus kotschoubeyanus (roter Kreis) neben einer blhenden
Proboscidea im mexikanischen Staat San Luis Potos.
69
Map 47
1
2
3
Protected areas
Canada: species diversity and protected areas

Species per 100 km
1000 km
Map 47: Canada. Species diversity and protected areas. Country area: 9.093.507 km; 3 species (0 endemic); 2 genera (0 endemic); 0 range-restricted
species; 0 species in CITES Appendix I; 0 species in the IUCN Red List.
Karte 47: Kanada. Artendiversitt und Schutzgebiete (Arten pro 100 km). Staatsgebiet: 9.093.507 km; 3 Arten (0 endemisch); 2 Gattungen (0 endemisch);
0 Arten mit begrenztem Areal; 0 Arten im CITES-Anhang I; 0 Arten in der IUCN-Roten-Liste.
Map 48
1-5
6 - 10
11 - 15
16 - 20
21 - 25
26 - 30
31 - 40
41 - 52
Protected areas
USA: species diversity and protected areas

Species per 100 km
1000 km
Map 48: USA. Species diversity and protected areas. Country area: 9,476,600 km; 173 species (32 endemic); 33 genera (1 endemic); 24 range-restricted
species; 18 species in CITES Appendix I; 6 species in the IUCN Red List.
Karte 48: USA. Artendiversitt und Schutzgebiete (Arten pro 100 km). Staatsgebiet: 9.476.600 km; 173 Arten (32 endemisch); 33 Gattungen
(1 endemisch); 24 Arten mit begrenztem Areal; 18 Arten im CITES-Anhang I; 6 Arten in der IUCN-Roten-Liste.
Schumannia 7
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71
7 Distribution maps of Cactaceae

Wilhelm Barthlott, Kerstin Burstedde, Jan Laurens Geffert, Pierre L. Ibisch, Nadja Korotkova,
Andrea Miebach, M. Daud Rafiqpoor, Anke Stein & Jens Mutke
Additional data and revisions contributed by Ralf Bauer, Graham Charles, Paul Hoxey,
David Hunt, Martin Lowry & Nigel Taylor
7.1. Subfamily: Pereskioideae

Pereskia Mill. (Maps 6466)
17 species.Woody climbers, shrubs or trees, normally 45 m; stems not conspicuously succulent, spiny; with broad persistent leaves. Flowers rotate white, orange, red or purplish. Fruits
fleshy globose, red, yellow or black. Distribution: Caribbean, Central America, northern
Colombia,Venezuela (Guyanas, coastal regions),
northern Peru (Andes), Bolivia, Paraguay,
northern Argentina (Andes and lowlands),
Brazil to northern Uruguay (Atlantic region).
Centres of diversity: Caribbean (Haiti), Bolivia
(eastern slopes of the Andes), eastern Brazil.
Main references: LEUENBERGER (1986), ANDERSON (2001), TAYLOR & ZAPPI (2004), HUNT
(2006).
7.2. Subfamily: Maihuenioideae

Maihuenia Phil. (Map 67)
2 species. Low cushion-forming, with persisting leaves. Flowers rotate, white, yellow or red.
Fruits fleshy. Distribution: central Chile to central Argentina. Main references: KIESLING
(1999), ANDERSON (2001), HOFFMANN & WALTER
(2004), HUNT (2006).
7.3. Subfamily: Opuntioideae

Austrocylindropuntia Backeb. (Map 68)
6 species. Small to large shrubs, cylindric, with
more or less tuberous, finally deciduous leaves.
Flowers rotate, yellow, pink or red. Fruits juicy.
Distribution: Ecuador to northern Argentina
(Andes). Main references: JRGENSEN (1999),
ANDERSON (2001), TAYLOR & ZAPPI (2004),
HUNT (2006).
Brasiliopuntia A. Berger (Map 69)
1 species. Lage tree up to 25 m, erect, with clusters of spines. Flowers near apex of leader or
terminal segment or borne by proliferation.
82
Flowers yellow. Fruits solitary or clustered, globose to ovoid. Distribution: Peru, Bolivia,
northern Argentina, southern Paraguay and in
coastal regions of Brazil. Main references: KIESLING (1999), ANDERSON (2001), TAYLOR & ZAPPI
(2004).
Consolea Lem. (Map 70)
3 species. Treelike with trunk and primary
branches cylindric, not articulated, ultimate;
segments flattened as in Opuntia. Flowers
small, diurnal. Fruits juicy, ovoid. Distribution:
Caribbean islands (mostly coastal rocks). Main
references: ANDERSON (2001), HUNT (2006).
Corynopuntia F. M. Knuth (Maps 7172)
14 species. Dwarf caespitose opuntioid shrubs;
branches cylindric-clavate. Flowers yellow or
rose to purple. Fruits ellipsoid fleshy at first,
later dry, yellow to brownish, smooth, sometimes spiny. Distribution: from central Mexico
to southern USA. Centre of diversity: Chihuahuan Desert. Main references: GUZMN &
al. (2003), HUNT (2006), USDA & NRCS
(2008).
Cumulopuntia F. Ritter (Map 73)
5 species. Low clumps. Flowers yellow, sometimes red; young shoots with small, cylindrical,
deciduous leaves. Fruits thick-walled and
fleshy. Distribution: southern Peru, south-western Bolivia, northern Chile and north-western
Argentina. Centre of diversity: south-eastern
Bolivia. Main references: BRAKO & ZARUCCHI
(1993), ANDERSON (2001), HUNT (2006).
Cylindropuntia (Engelm.) F. M. Knuth (Maps
7479)
33 species. Shrubby or treelike, branched. Flowers variously coloured. Fruits fleshy or dry. Distribution: south-western USA to Mexico,
Caribbean islands. Centre of diversity: northwestern Mexico to south-western USA (southern California to Arizona). Main references:
ANDERSON (2001), GUZMN & al. (2003), USDA
& NRCS (2008).
Grusonia Rchb. f. ex Britton & Rose (Map 80)

1 species. Low shrubs. Flowers variously
coloured. Fruits cylindric, various, fleshy or dry.
Distribution: northern Mexico (Coahuila).
Main references: GONZALEZ ELIZONDO & al.
(1991), VILLARREAL QUINTANILLA (2001), GUZMN & al. (2003).
Maihueniopsis Speg. (Map 81)
8 species. Low compact cushions; roots tuberous. Flowers yellow, orange, pink or red. Fruits
juicy, thick-walled, indehiscent. Distribution:
southern Bolivia,Argentina and eastern Chile.
Main references: KIESLING (1999), ANDERSON
(2001), HOFFMANN & WALTER (2004), HUNT
(2006).
Miqueliopuntia Fric ex F. Ritter (Map 82)
1 species. Shrubby, forming clumps up to 1.4 m,
Flowers almost white, fruits globose, pale green
to whitish, juicy. Distribution: central Chile
(coastal regions). Main references: ANDERSON
(2001), HOFFMANN & WALTER (2004).
Nopalea Salm-Dyck (Map 83)
3 species.Tree-like, with a cylindric trunk. Flowers purplish pink or red. Fruits ovoid. Distribution: north-eastern Mexico to Central America. Main references: KIESLING (1999), ANDERSON (2001), GUZMN & al. (2003), TAYLOR &
ZAPPI (2004), HUNT (2006).
Opuntia Mill. (Maps 8495)
64 species.Trees and shrubs, some low to cushion-forming; stem normally segmented; segments cylindric, clavate, subglobose, or more
or less flattened. Flowers rotate, yellow, pink,
red or whitish. Fruits umbilicate, fleshy or dry.
Distribution: southern Canada (temperate
forests), most parts of the USA, Central America, Mexico, Caribbean, north-eastern coast of
Colombia and Venezuela, Galpagos islands,
western Colombia, Ecuador, Peru, Bolivia,Argentina, Paraguay, Uruguay and south-eastern
Brazil. Centres of diversity: central Mexico
(southern Chihuahuan Desert); southern Mex-
Schumannia 7
2015
Map 67
Map 66
Pereskia
1000 km
aureiflora
nemorosa
sacharosa
stenantha
weberiana
Maihuenia
patagonica
poeppigii
Map/Karte 66: Pereskia.
500 km
Map/Karte 67: Maihuenia.
Map 68
Austrocylindropuntia
cylindrica
floccosa
pachypus
shaferi
subulata
vestita
500 km
Map/Karte 68: Austrocylindropuntia.
98
Schumannia 7
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Map 93
Opuntia
aciculata
atrispina
aureispina
bravoana
chlorotica
engelmannii
lagunae
littoralis
macrocentra
oricola
500 km
Map/Karte 93: Opuntia.
Map 95
Map 94
Opuntia
Opuntia
aurea
basilaris
huajuapensis
microdasys
pinkavae
pycnantha
rufida
streptacantha
500 km
fuliginosa
hyptiacantha
lasiacantha
leucotricha
pilifera
robusta
stenopetala
tehuacana
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250 km
109
Map 127
1
2
3
4
5
6
Cleistocactus: Diversity pattern
Species per 2500 km
1000 km
Map 127: Cleistocactus, diversity pattern.

Karte 127: Cleistocactus, Diversittsmuster (Arten pro 2500 km).
Map 128
Cleistocactus
baumannii
brookeae
buchtienii
candelilla
chrysocephalus
hyalacanthus
laniceps
luribayensis
parapetiensis
strausii
250 km
Map/Karte 128: Cleistocactus.
120
Schumannia 7
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Schumannia 7
2015
Index
The index covers genera, tribes, subfamilies, important synonyms, the country names of the country profiles and some key words.
Acanthocereus 83, 112

Acharagma 83, 112
Argentina 16, 36, 37, 80
Ariocarpus 69, 70, 72, 83, 112
Armatocereus 83, 113
Arrojadoa 63, 83, 113
Arthrocereus 83, 113
Astrophytum 72, 83, 114
Austrocactus 83, 114
Austrocylindropuntia 49, 82, 98
Aylostera = Rebutia
Aztekium 41, 83, 114
Bergerocactus 83, 114
biogeographic regions 22
biomes 14
Blossfeldia 15, 20, 25, 28, 83, 115
Blossfeldieae 23, 26
Bolivia 36, 3741, 47, 79
Brachycereus 20, 84, 115
Brasilicereus 84, 115
Brasiliopuntia 82, 99
Brazil 36, 3741, 78
Browningia 84, 115116
Caatinga centre 36, 40, 41, 65
Cacteae 23, 25, 26, 57, 58, 60
Cactoideae 26, 83, 55, 58
Calymmanthium 84, 116
Canada 71
Caribbean 72
Carnegiea 72, 73, 84, 116
Castellanosia 84, 116
Central America 35, 3741, 76
Cephalocereus 84, 117
Cereeae 23, 26, 66
Cereineae 26
Cereus 41, 46, 84, 117119
Chihuahua centre 36, 3740, 65
Chile 41, 81
Cipocereus 84, 119
Cleistocactus 41, 69, 84, 120121
climate factors 65
Coleocephalocereus 15, 84, 122
Colombia 13, 76
Conservation 6873
Consolea 82, 99
Copiapoa 17, 28, 41, 84, 122123
Corryocactineae 26
Corryocactus 84, 123123
Corynopuntia 82, 100
Coryphantha 84, 124126
Cuba 19, 49, 72
Cumarinia 84, 126
Cumulopuntia 16, 82, 101
202
Cylindropuntia 18, 82, 101103

Cylindropuntieae 23, 26, 59
Dendrocereus 19, 70, 84, 127
Denmoza 84, 127
Discocactus 41, 46, 63, 84, 127128
Disocactus 45, 63, 84, 128129
Echinocactus 20, 60, 84, 129
Echinocereinae 26
Echinocereus 21, 85, 130134
Echinopsis 16, 62, 69, 85, 135138
Ecuador 77
Epiphyllum 62, 85, 138139
Epithelantha 85, 139
Eriosyce 41, 85, 139140
Escobaria 85, 141
Escontria 85, 142
Espostoa 41, 85, 142
Espostoopsis 85, 142
Eulychnia 85, 143
Facheiroa 85, 143
Ferocactus 16, 85, 143145
Frailea 85, 145146
Geohintonia 41, 46, 85, 146
growth form 4445, 47, 49, 56
Grusonia 82, 104
Guyanas 77
Gymnocalycium 85, 147150
Haageocereus 41, 85, 150
Harrisia 85, 151
Hatiora 85, 151
Hylocereinae 25, 26
Hylocereus 86, 151152
Jalisco centre 36, 37, 40, 41, 65
Jasminocereus 86, 152
Lasiocereus 86, 152
latitudinal gradient 64
Leocereus 86, 153
Lepismium 86, 154
Leptocereus 86, 154
Leuchtenbergia 86, 155
Lobivia = Echinopsis
Lophophora 86, 155
Lymanbensonia 29, 86, 155
Lymanbensonieae 23, 25, 26
Maihuenia 25, 45, 82, 98
Maihuenieae 23
Maihuenioideae 25, 26, 82
Maihueniopsis 82, 104
Mammillaria 41, 62, 68, 70, 72, 86, 156162
Mammilloydia 86, 162
Mata Atlantica centre 36, 40, 41, 65
Matucana 41, 62, 86, 163
Melocactus 68, 70, 86, 163165
Mexico 36, 3740, 74

Micranthocereus 46, 86, 165
Mila 86, 166
Miqueliopuntia 82, 104
Myrtillocactus 86, 166
Neobuxbaumia 86, 167
Neolloydia 86, 167
Neoraimondia 86, 167
Neowerdermannia 86, 168
Nopalea 82, 104
Notocacteae 23, 25, 26, 61
Notocactus = Parodia
Obregonia 87, 168
Opuntia 19, 48, 69, 72, 82, 105109
Opuntieae 23, 26, 59
Opuntioideae 15, 20, 26, 3133, 55, 82
Oreocereus 87, 168
Oroya 87, 169
Ortegocactus 87, 169
Pachycereeae 25, 26
Pachycereus 18, 87, 169170
Paraguay 47, 49, 80
Parodia 41, 69, 87, 170173
Pediocactus 70, 72, 86, 174
Pelecyphora 46, 87, 174
Peniocereus 87, 174175
Pereskia 13, 20, 25, 82, 9798
Pereskieae 23
Pereskioideae 13, 20, 24, 26, 55, 82
Pereskiopsis 83, 110
Peru 41, 78
Pfeiffera 87, 176
Phyllocacteae 15, 23, 25, 26, 60
Pierrebraunia 87, 176
Pilosocereus 70, 87, 176179
Polaskia 87, 179
pollination 4750, 51, 57
Praecereus 87, 179
Pseudoacanthocereus 87, 180
Pseudorhipsalis 87, 180
Pterocactus 83, 110
Puebla-Oaxaca centre 36, 37, 40, 41, 65
Punotia 83, 110
Pygmaeocereus 87, 181
Quiabentia 83, 111
range size 4144
Rauhocereus 87, 181
Rebutia 41, 87, 181183
Rebutiinae 26
Rhipsalideae 10, 23, 26, 30, 61
Rhipsalidopsis 88, 184
Rhipsalinae 30
Rhipsalis 19, 20, 41, 48, 68, 88, 184187
Schumannia 7
2015

Biodiversity of Cacti

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The Cactaceae with c.

11435 species are the mosst important

Text zweisprachig Engl ish und Deutssch

Biogeography and biodiversity of cacti

Ein Sonderheft der

Abstract / Kurzfassung / Resumen

Corresponding Author / Korrespondenzautor:

Foreword and acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Cactus ecology and biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

Cacti and their habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

Biogeography and palaeo-history . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

Phylogeny, evolution and systematics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

Finding the closest relatives of cacti . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

Age estimates for the Cactaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

Current understanding of phylogenetic relationships within Cactaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

Basal cactus lineages and the origin of cacti . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

Relationships within Cactoideae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

Mapping the diversity of cacti . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

Main data sources . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

Generating the distribution maps . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

Quality and reliability of data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

Patterns of diversity and endemism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Spatial distribution of growth forms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Spatial distribution of pollination syndromes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Diversity pattern of genera, tribes and subfamilies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Conservation and hotspots: cactus diversity in change . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Distribution maps of Cactaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Foreword and acknowledgements

ily, and the different centres of diversity were

Presumably, cacti (Melocactus) were already

of botanists and plant enthusiasts at an early

2 Cactus ecology and biogeography

2.1 Cacti and their habitats

has been demonstrated by several authors that

2.2 Biogeography and palaeo-history

to the more southern regions of the Atacama

3 Phylogeny, evolution, and systematics

3.1 Finding the closest relatives of cacti

3.2 Age estimates for the Cactaceae

3.3 Current understanding of

to other popular plant families such as orchidaceae or Bromeliaceae.

3.4 Basal cactus lineages and the

4 Mapping the diversity of cacti

4.1 Main data sources

synonyms. Recent compilations of accepted

367 species, 43 genera

107 species, 27 genera

Species rich genera:

Species rich genera:

Diversity centres of Cactaceae

Species per 2500 km

Species rich genera:

(defined as the top 5%

Mata Atlantica centre

Southern Central Andes centre

Map 8: Diversity centres of Cactaceae.

5 Patterns of diversity and endemism

5.1 General overview

Cerrado, the Llanos Savannah of Colombia

5.2 Centres of diversity

Diversity centres of Cactaceae: genera

Genera per 2500 km

tinga centre has the highest numbers of cacti

Table 1: The North/Central American diversity centres