Dr. Valerio Ketmaier [email protected]

Zoogeografia
Dr. Valerio Ketmaier
[email protected]
State of the Art:
Phylogeography
History, conceptual background and purview of
phylogeography
• 
The term phylogeography was introduced by Avise and co-workers in 1987
• 
Phylogeography is a field of study concerned with the principles and processes governing
the geographic distribution of genealogical lineages, especially those within and among
closely related species
• 
Phylogeography deals with the historical phylogenetic component of the spatial
distribution of lineages
• 
It is an integrative approach that combines diverse micro- and macro-evolutionary
disciplines
Microevolutionary disciplines
ethology
demography
Phylogeography
Paleogeography
Paleontology
Macroevolutionary disciplines
Pop genetics
Mol genetics
Phylogenetics
Dispersal
Vicariance
A
a
a1
B
b1 b2
A
b
a1
B
a2
C
B
C
a,b,c
B
a
C
a,b,c
c
B
a2
A
b1
B
C
c
Dispersal
A
a1
a2
b2
c
A
Area
Taxon
Vicariance
A
a1,b,
c1
c1
c2
A/C
C
a1
cc2
a2
A/B
b
B
Geographic distribution of genetic variants
Phylogeographic patterns
(aus Avise 2000)
Comparative phylogeography of Galapagos fauna
• 
• 
• 
• 
• 
The Galapagos Archipelago occupies a unique position in the history of evolutionary studies.
Oceanic islands are biologically simpler than continental regions.
The time of their geological formation is known.
Each island of the archipelago can be seen as a replicate natural experiment.
Relative limited number of lineages.
Geographical setting and geological history of the
archipelago
13 major islands; 6 smaller islands; 40 islets with official
names and many smaller unnamed islets ~ 1000 km from
Ecuador
• Islands are formed as the Nazca plate moves over an
hot spot (or mantle plume)
• Oldest islands located the southeast of the Archipelago
• Oldest emerged island 3-4 Ma old
• Sunken islands 14 Ma old
• Hot spot 80-90 Ma old
Ecology
• 
Dry climate with marked seasonality.
• 
A warm season (January to May) influenced by warm ocean currents (southward).
• 
A cool season (June to December) influenced by the Humboldt current (from the south to the
west).
• 
Higher islands have higher precipitation.
• 
Every 3-6 years changes in the direction of oceanic currents (El Nino) increase the amount of
rainfall dramatically.
• 
Vegetation on islands can be separated in six altitudinal zones, where plant composition is a
reflection of the humidity level.
Biogeography
• 
Three major track patterns connect the Galapagos with continental areas.
East Pacific track
-
Grehan Biol. J. Linn. Soc., 2001, 74: 267-287
Central AmericaCaribbean track
Dispersal capability
Pacific basin track
+
Study cases:
Land snails (genus Bulimulus)
• 
• 
About 80 among species and subspecies have been described for the Archipelago.
Molecular study based on two genes (COI and ITS1) to infer the spatial and temporal
patterns of speciation.
• Progression rule supported
• Sequence of species formation
approximates the pattern of
formation of the islands
• Older islands with monophyletic
and/or less diverse lineages
Parent & Crespi, Evolution 2006, 60: 2311-2328
Study cases:
Lava lizards (genus Microlophus)
• 
• 
At least 17 named and unnamed lineages recognized for the Archipelago.
Multiple mitochondrial genes sequenced (ND1, ND2, COI, tRNAs)
Peru
Peru + Ecuador
Kizirian et al., Mol. Phylog. Evol. 2004, 32: 761-769.
Study cases:
land and marine iguanas (genera Conolophus and Amblyrhynchus)
• 
• 
• 
Different ecology.
Are these two lineages each other’s closest relatives or not?
12s and 16s genes sequenced; all extant Iguanidae included in the study.
Rassmann, Mol. Phylog. Evol. 1997, 7: 158-172.
Study cases:
Darwin’s finches
• 
• 
• 
• 
• 
Darwin’s finches are one of a few classical examples of adaptive radiations.
Species show adaptive variation in beak size and shape and body size.
Variation at these traits is more typical of differences among families of bird.
Yet the radiation is believed to have occurred in less than three million years.
Three groups based on morphology and behavior: ground-finches (Geospiza); tree-finches
(Cactospiza, Camarhynchus, Platyspiza) and Warbler-finches (Certhidea, Pinaroloxias).
Open questions:
• Are Galapagos finches monophyletic?
• Is the Cocos finch directly related to the Galapagos finches?
• Did Warbler-finch descend from the same ancestor as the rest of
the group?
• Is the vegetarian finch really a member of the tree finch group?
• Is the distinction between ground and tree finches valid?
• MtDNA and microsatellite markers
Sato et al., PNAS 1999, 96: 5101-5106.
Freeland & Boag, The AUK 1999, 116: 577-588..
Petren et al., Proc. R. Soc. Lond. B., 266: 321-329
Study cases:
Darwin’s finches
Microsatellites
MtDNA
• Darwin’s finches constitute a monophyletic assemblage.
• Cocos finch are within Darwin’s finch assemblage.
• Warbler-finches are basal in the group’s phylogeny.
• The vegetarian finch is not a member of the tree finch group.
• Ground-finches are clearly a monophyletic group.
Sato et al., PNAS 1999, 96: 5101-5106.
Freeland & Boag, The AUK 1999, 116: 577-588.
Petren et al., Proc. R. Soc. Lond. B., 266: 321-329.
Study cases:
Giant Galapagos tortoises
• 
• 
The only extant giant tortoises along with those from Seychelles (but only a single population left).
15 described taxa, of which 11 extant in various stages of imperilment.
Questions:
• Are Galapagos tortoises a monophyletic lineage and where did they come from?
• Patterns and levels of diversification among as well as within islands. Systematics of the group.
Caccone et al., PNAS 1999, 9: 13223-13228.
• Colonization history of the Archipelago
Caccone et al., Evolution 2002, 56: 2052-2066.
• Restoration of endangered lineages. Conservation genetics.
Beheregaray et al., PNAS 2004, 101: 6514-6519.
• Multiple markers used.
Poulakakis et al., PNAS 2008, 105: 15464-15469.
Study cases:
Origin of giant Galapagos tortoises
6-12 Ma
Caccone et al., PNAS 1999, 9: 13223-13228.
Study cases:
Patterns and levels of diversification of giant Galapagos tortoises
Some of the haplotypes found at PBL, PBR, VD and RU differ
from haplotypes from the same location by 27-70 substitutions
while they differ by a few substitutions from haplotypes found in
geographically distant populations (PBL vs. VA/VD/SCR; PBR
vs. ESP; CPA vs PBL). Human induced translocations?
Caccone et al., Evolution 2002, 56: 2052-2066.
Study cases:
Patterns and levels of diversification of giant Galapagos tortoises
Pinta (Lonesome George)
Isabela North
Santiago
Isabela North
Isabela Central
S.Cruz
S.Cruz
Paz/S.Cruz
Isabela South
S.Cristobal
0.2-0.3 MA
Espanola
1.5-2.0 MA
Caccone et al., Evolution 2002, 56: 2052-2066.
Study cases:
Patterns and levels of diversification of giant Galapagos tortoises
Beheregaray et al., PNAS 2004, 101: 6514-6519.
Beheregaray et al., Science 2003, 302:75.
Considerable less genetic
diversity (Mt/ nucDNA)
Bootleneck
Age: 88.000-118.700 years
Vo l c a n i c e x p l o s i o n :
74.000-120.000 years
Study cases:
Conservation genetics of giant Galapagos tortoises
MtDNA
Microsatellites
Poulakakis et al., PNAS 2008, 105: 15464-15469.
• Isabela V.Wolf: 1669 individuals (about 1/5th of
the extant population size) screened at 12
microsatellite loci
• Their genetic profiles compared to the available
database of all extinct and extant Galapogos
tortoise species
•  The genotypes of 84 individuals from V.Wolf result
from hybridization events with a pure
C.elephantopus as one the parents
•  30 of these 84 individuals are less than 15 years
old
•  26 had a C.elephantopus mtDNA
•  There is a good chance that some of these
purebred C.elephantopus are still alive
Garrick et al., Genetics: in press.
•  The “extinct” species could be resurrected
through targeted breeding efforts
Outline
•  The peri-Mediterranean area: roots of its diversity
•  Four study cases
Cyprinids
Hewitt 1999
Stenasellus
Troglophilus/
Cyprinids
Murella
Current
diversity
• Stenasellidae is a group of exclusively freshwater stygobionts
• Limited ability to disperse through unsuitable ecological areas
• The case of the S. racovitzai / S. virei lineages
S.virei
S.racovitzai
.
‘50
‘99
‘99
‘64
‘23
‘95
>80%
MP
ML
NJ
?
9 Myr
Tuscany
Corsica
NW Sardinia
Stenasellus
racovitzai
SE Sardinia
29 Myr
Pyrenees
Stenasellus virei
Stenasellus assorgiai
Proasellus coxalis
Allozymes (15 loci); mtDNA (COI)
Ketmaier et al., 2003 Mol. Ecol.
N
c
Florence
Pisa
S
Siena
2-0.4 Myr
Elba Isl.
Grosseto
Viterbo
Pliocene coastline
Present coastline
Continental areas
Fossil islands
T.neglectus
T.ovuliformis
T.brevicauda
T.cavicola
DED
PEJ
DEL
T7
T.lazaropolensis
T38
T3
T39
T.tatyanae
E5D
T.bicakcii/BAL
T9
PIR
ADA2B
I5N
E1D
E4E
T.lagoi
T.marinae
T.gajaci
T.spinolosus
T.adamovici
T8
BEO
T6
T. andreinii andreinii
T. andreinii hydruntinus
T.ovuliformis
T.cavicola
A12
T22
T23
T.escalerai
Phylogeny
Bayesian Analysis
T.lagoi
T.spinolosus
T.marinae
T.adamovici
E1D
E4E
T.lazaropolensis
T.andreinii
T.neglectus
Greek Is.
Turkey
Balkans
S.Italy
T.ovuliformis
Combined data
(COI by cod+
12S/16S+
18S/28S)
T38
T39
T9
T7
BEO
DED
T8
T.cavicola
T6
T.brevicauda
T3
PEJ
ADA2B
I5N
PP ≥95%
PP = 100%
DEL
Balkans
PIR
T.bicakcii
BAL
A12
T22
T23
E5D T.gajaci
T.tatyanae
T.escalerai
Dolichopoda
Turkey
Formation of mid-Aegean Trench
Messinian
Quaternary
Climate Changes
Greek Is.
S.
Italy
Balkans
Turkey
Balkans
Turkey
15
12.5
10
Miocene
7.5
5
2.5
Pliocene
Chronology
0 Myr
Pleistocene
Lineages Through Time
Best Constant Rate Model (pure
birth)
AIC= -250.7077
Best Rate Variable Model (DDX)
AIC= -249.5808
∆AICrc= -1.126891
12
9.6
7.2
4.8
2.4
0
P= 0.647
Myr
The null hypothesis of constant speciation rate cannot be rejected
T.escalerai
T.tatyanae
T.gajaci
15
E5D
T22
T23
A12
BAL
T.bicakcii
I5N
ADA2B
T3
PEJ
T.brevicauda
T6
T.cavicola
T8
Myr
0
Turkey
PIR
DED
BEO
T7
T38
T39
T9
T.ovuliformis
T.neglectus
T.andreinii
T.lazaropolensis
T.adamovici
E1D
E4E
T.lagoi
T.spinolosusT.marinae
Dispersal within
inland Balkans
Dispersal from Anatolia
the Balkans
Balkans
DEL
Turkey
S.Italy
1.00
Balkans
Greek Is.
Bayesian Analysis of Geographic Distribution: Vicariance vs. Dispersal
Vicariance
Aegean / Anatolia
Dispersal back
to Anatolia
Dispersal to S. Italy
from Balkans
1.00
1.00
Dispersal to Med.
Anatolia from Aegean Anatolia
1.00
Dispersal to coastal Balkans
from inland Balkans
Dispersal to Caucasus
from Med. Anatolia
Biogeographic history
6
5
4
2
4
2
3
4
1
Ketmaier et al. 2000 Belg J Zool
Ketmaier et al. 2004 Subterranean Biology
Ketmaier et al. 2013 Subterranean Biology
Taylan et al. 2013 ZooKeys
Outline
•  Introduction to the model organisms (Cyprinids)
•  Current views and hypotheses on the diversification of
cyprinids in Southern Europe
•  Selected study cases: Scardinius (Rudds), Telestes
(Minnows), Rutilus (Roaches), and Alburnus (Bleaks)
•  Evolutionary and biogeographic inferences based on
molecular data
•  Effects of human interferences
Division of freshwater fishes according to
salinity tolerance
•  Primary: taxa that originated and spread in freshwaters strictly intolerant of salt water (Dipnoi, Polypteridae,
Cyprinidae etc.)
•  Primary-like: strictly freshwater species of ancient marine
origins (Gobiidae)
•  Secondary: mostly confined to freshwaters but relatively
salt-tolerant (Cichlidae, Cyprinodontidae, Poeciliidae etc.)
•  Peripheral: recent marine derivates and diadromous taxa
(Lota, Anguilla, Alosa, Salmo etc.)
Cyprinids
• 
The family belongs to the order Cypriniformes
• 
It’s the largest family of primary freshwater fish, with
about 220 genera and more than 2400 species
• 
Despite the diversity of the group little is known about
their relationships
Mayden et al. Mol Phylog Evol 2009 51: 500-514
Rüber et al. BMC Evol Biol 2007 7:38
20
18
16
14
12
10
N
Danubian
8
Non Danubian-Shared
6
Endemic
4
2
0
Tunisian
(1)
Maghrebo- Southern
Algerian
Iberian
(2)
(3)
Central
Iberian
(4)
EbroSouthern
Cantabric France (6)
(5)
PadanoVenetian
(7)
Tuscano- Dalmatian
Latium (8)
(9)
Albanian
(10)
Western
Greece
(11)
AegeoMacedoAnatolian
(12)
Danubian district
5
4
6
7
8
3
9
?
2
1
10
12
11
200 Km
Classical theory of early penetration and dispersal of freshwater fishes in western
Palearctic and in Southern Europe by river captures (Banarescu, 1960).
Pre Miocene in situ evolution?
England
Lower
Miocene
W Europe
Middle
Miocene
Central
Europe
Middle Central Asia
Miocene Siberia
Early
Miocene
E Asia
Quaternary glacial
maxima extinctions
Quaternary glacial
maxima survivals
Iberian Pen.
18 species
NW Africa
S France
Italy
W Balkans
Alternative theory of an early penetration of freshwater fishes in the periMediterranean area by a Messinian “sea” dispersal (Bianco, 1990).
Pre Miocene in situ evolution?
Central
Europe
Middle
Miocene
Para-Tethys
Early
Miocene
E Asia
Late
Messinian
Middle
Miocene
Middle
Miocene
W Europe
Mediterranean
“Lago Mare” phase
S France
&
N Spain
NW Africa
Iberian Pen.
S France
Italy
W Balkans
Tethys Ocean
Closure of the Tethys Ocean
Middle Miocene geography
Paratethys (dilution basin)
s
Alp
Dinarid
Balkan
Mediterranean (concentration basin)
Anatolian
•  The Mediterranean had a global negative balance (prevalence of evaporative losses on
meteoric and river inputs)
•  The Paratethys had a positive balance (prevalence of meteoric and river inputs on
evaporative losses)
Presalinity crisis event: closure of the AtlanticMediterranean seaway
Paratethys (freshwater basin)
Mediterranean (hypersaline basin)
The Mediterranean Messinian salinity crisis
Paratethys
(freshwater basin-”Lago Mare”)
Mediterranean (drying up)
The Messinian “Lago Mare” phase of the
Mediterranean
Paratethys
”Lago Mare” residual basins
”Lago Mare” phase of the Mediterranean
Late Messinian resettlement of marine conditions of the
Mediterranean
Mediterranean (oceanic salinity)
•  Black spots indicate ancient river drainages where Paratethys primary freshwater fishes
might have survived
Different timing: Quaternary vs. Messinian
Effects on phylogeny
Gradual hypothesis
Lago Mare hypothesis
N
14
5
4
6
7
8
3
10
9
11
13
12
2
1
200 Km
• Ketmaier et al. Ital J Zool 1998 65: 41-48
• Bianco & Ketmaier J Fish Biol 2001 59: 190-208
• Bianco, Busatto & Ketmaier Quad.ETP 2001 30:115-120
• Bianco & Ketmaier Environ Biol Fish 2003 68: 370
• Ketmaier et al. Zool Scr 2003: 13-22
• Ketmaier et al. Mol Phylog Evol 2004 32: 1061-1071
• Bianco et al. Biol Amb 2004 18: 7-14
• Bianco & Ketmaier Folia Zool 2005 42: 42-49
• Bianco, Zupo & Ketmaier J Fish Biol 2006 68: 150-155
• Ketmaier et al. Mol Phylog Evol 2008 49: 362-367
• Ketmaier et al. J. Fish Biol 2009 75: 997-1017
Scardinius acarnanicus (12)
Scardinius graecus (12)
Se1 (6)
100
Se14 (13)
+17
Se9 (14)
Se10 (14)
Se2 (7**)
Se3 (7**)
Scardinius erythrophthalmus
97 Se4 (7**)
+10 Se5 (8*)
75 Se6 (8*)
+6 Se7 (8*)
Se8 (8*)
Scardinius scardafa (8**)
100
Se12 (7)
+38
Se13 (11)
Scardinius erythrophthalmus
84
+4
*allochthnous anthropogenic populations
** autochthonous populations
98
+17
14
4
5
7
6
8
3
10
9
2
1
11
13
12
Se11 (14)
99
+19
98
+19 Ts2 (6)
100 Ts3 (14)
+13
Ts1 (5)
Telestes souffia
Ts4 (10)
96
+12
94
+23
98
+12
100
+16
100
+19
100
+26
Pachychilon pictus (11)
Lc2 (13)
96
+19
Tm1 (7)
Tm2 (7)
Tm3 (7)
Telestes
Tm4 (8)
muticellus
Tm5 (8)
94 Tm6 (9)
+23 Tm7 (9)
Telestes montenigrinus (11)
Leuciscus turskyi (10)
Leuciscus polylepis (7)
Telestes
Tp1 (12)
Tp2 (12)
pleurobipunctatus
Telestes beoticus (12)
Phoxinellus croaticus (10)
Phoxinellus metohiensis (10)
Lc1 (6)
Leuciscus cephalus
Tropidophoxinellus spartiaticus (12)
Leuciscus leuciscus (14)
Phoxinellus prespensis (12)
Cyprinus carpio (-)
0.01 substitutions/site
Ketmaier et al. Mol Phylog Evol 2004 32: 1061-1071
14
7
4
5
6
8
3
10
9
2
1
13
11
12
Ketmaier et al. Mol Phylog Evol 2008 49: 362-367
Effects on phylogeny
Gradual hypothesis
Lago Mare hypothesis
Semirara
Carabao
Boracay
Panay North
Panay South
North Gigante
Negros
Gallus g.
Pica p.
BO10
BO1
BO3
BO7
BO4
BO6
PN7
PN2
BO5
BO2
BO9
PN19
PN3
NE1
CA8
CA10
CA6
CA5
NE2
PN11
PN14
CA11
PN1
CA3
CA7
CA9
PN10
PN4
BO8
CA4
PS3
PN18
PN15
PN12
PN9
PS2
PN17
PS1
PN8
PN6
PN5
PN16
PN13
CA1
CA2
NG1
NE3
SE8
SE3
SE1
SE4
SE10
SE9
SE7
SE11
SE5
SE2
SE6
Plasmodium sp.
PN1B
PN15A
PN11C
NE3E
NE2M
CA7M
BO1C
CA1M
BO1E
CA5A
SE4D
SE5D
SE2I
SE9O
SE11A
SE7O
SE2A
SE3O
PN1A
NE2B
NE2J
PN15B
SE10E
CA8Q
CA5B
CA87
CA6M
CA8A
NE2L
NG1M
PN14U
PN13M
PN18M
PN1I
SE4F
SE6B
PN1F
NE3A
SE10C
SE2C
SE11K
SE6C
SE6A
SE8O
SE5A
SE1C
SE11T
SE10B
PS2M
CA8P
PN6M
SE11P
PN14Y
SE11L
PN11G
PN14A
SE1A
SE1I
• TreeMap superimposes the parasite tree (fish) on the host (area) tree to obtain reconciled trees
• Detect cospeciation, duplication and sorting events (19 cospeciation events, 39 duplications, 311 sorting
events in this particular case)
Silva-Iturriza et al. J Zool Syst Evol Res 2010 48: 269-278
14
There’s no statistical association
between the molecular tree and
the district area cladogram
(P=0.776)
7
4
5
6
8
3
10
9
2
1
13
11
12
District area cladogram
(Banarescu Hypothesis)
Telestes
Dalmatian (10)
Danubian (14)
W-Greece (12)
Padano-Venetian/
Tuscan-Latium/
Southern Italy (7-8-9)
Albanian (11)
Albanian (11)
Balkans
(12/13)
Dalmatian (10)
Padano-Venetian (7)
Ebro-Cantabric (5)
Padano-Venetian (7)
Southern France (6)
Southern France (6)
Danube (14)
Dalmatian (10)
Ebro-Cantabric (5)
(N/A)
Central Iberian (4)
Southern Iberian (3)
14
There’s no statistical association
between the molecular tree and
the district area cladogram
(P=0.596)
7
4
5
6
8
3
10
9
2
1
13
11
12
District area cladogram
(Banarescu Hypothesis)
Scardinius
W-Greece (12)
Danubian (14)
Danubian (14)
Albanian (11)
Aegeo-MacedoAnatolian (13)
Balkans
(12/13)
Southern France (6)
Padano-Venetian (7)
Padano-Venetian (7)
Southern France (6)
Tuscan-Latium (8) (N/A)
Ebro-Cantabric (5)
Padano-Venetian (7)
Albanian (11)
(N/A)
Central Iberian (4)
Southern Iberian (3)
Danubian (14)
14
7
4
5
6
8
3
9
2
Rutilus
1
There’s no statistical association
between the molecular tree and
the district area cladogram
(P=0.879)
10
13
11
12
Padano-Venetian (7)
District area cladogram
(Banarescu Hypothesis)
Albanian (11)
Danubian (14)
Padano-Venetian (7)
W-Greece (12)
Danubian (14)
Albanian (11)
Balkans
(12/13)
Tuscan-Latium (8) (N/A)
Padano-Venetian (7)
Danubian (14)
Southern France (6)
W-Greece (12)
Padano-Venetian (7)
Southern France (6)
Aegeo-MacedoAnatolian (13)
Ebro-Cantabric (5)
(N/A)
Padany-Venetian (7)
Central Iberian (4)
Southern Iberian (3)
Estimates of divergence times
Telestes
Scardinius
(6.3-5.1 Myr)
(5.7-4.8 Myr)
Rutilus
(5.6-4.5 Myr)
Messinian
(7.2-5.3 Myr)
Pliocene
(5.3-1.8 Myr)
Lago Mare phase
(5.3 Myr)
Pleistocene
(1.8-0.01 Myr)
BA
BI
• Diversification of leuciscine lineages has been
determined by intense vicariant events
• Vicariance was largely promoted by the paleoevolution of the Mediterranean region
• B anarescu and Bianco hypotheses are not
mutually exclusive
Take home message:
Banarescu hypothesis (BA) works at the amonggenera level
Bianco hypothesis (BI) targets divergence at the
within-genera level
Perea et al BMC Evol Biol 2010 10: 265
Padano-Venetian district north of the
Apennines. Four cyprinids and two
cobitids endemics.
Tuscano-Latium district: Central Italy
West of the Apennines. Two cyprinids
and one goby endemics.
Southern Italy district: south of Rivers
Vomano and Tiber. Two cyprinids and
one cobitid endemics.
Arrows indicate the directions of the
freshwater fish transfers.
•  The number of introduced species of European (in grey) and extra
European (in white) origin has been increasing steadily in the last five
centuries
•  All localities host species translocated from Northern Italy (in grey)
•  Eight localities (out of twelve) host species introduced from outside Italy (in black)
Bianco & Ketmaier J Fish Biol 2001 59: 190-208
Table 2. Illustrative examples of the interactions between non-native, translocated-native (*) and native freshwater fish species in Italy. Origin of native
species: D (Danubian), PV (Padano-Venetian), TL (Tuscano-Latium), AC (Apulo-Campania), ME (Mediterranean), IB (Iberian Peninsula).
Geographical areas are the same as in Fig. 1. Updated IUCN category (Bianco et al. 2013): CR (critically endangered), VU (vulnerable), EN
(endangered), NT (nearly threatened), LC (low concern). It is worth noting that P. genei and R. rubilio are threatened in their native ranges but pest
wherever they have been introduced.
Aliens & Translocated
Geographic origin
Rutilus rutilus
Chondrostoma nasus
D
D
Scardinius esperedicus*
Rutilus rubilio*
Rutilus rubilio*
Alburnus arborella*
Rutilus aula*
Chondrostoma genei*
Gambusia hoolbroki
Barbus graellsii
Barbus barbus
Telestes agassizi
Barbus balcanicus
Gobio gobio
Cobitis bilineata*
Esox Lucius
Silurus glanis
Padogobius bonelli*
PV
TL
TL
PV
PV
PV
USA
IB
D
D
D
D
PV
D
D
PV
Threatened natives
Rutilus pigus; Rutilus aula
Chondrostoma soetta;
Protochondrostoma genei
Scardinius scardafa
Rutilus aula
Alburnus albidus
Alburnus albidus
Rutilus rubilio
Squalius lucumonis
Aphanius fasciatus;
Gasterosteus gymnurus
Barbus tyberinus
Barbus plebejus
Telestes muticellus
Barbus caninus
Gobio benacensis
Cobitis zanandreai
Esox cisalpinus
All species
Padogobius nigricans
Geographic origin/
IUCN category
PV/EN; PV/LC
PV/EN
PV/EN
TL/CR
PV/LC
MER/VU
MER/VU
TL/NT
TL/CR
ME/LC
ME/LC
TL/VU
PV/VU
PV,TL/LC
PV/EN
PV/EN
AC/CR
PV/TL
-/TL/VU
Type of habitat where natives
and aliens species meet
Lacustrine
Lacustrine or Riverine
Lacustrine
Riverine
Riverine
Lacustrine
Lacustrine
Riverine
Wetlands
Lacustrine – Riverine
Lacustrine – Riverine
Riverine (Isonzo River)
Riverine (Isonzo River)
Riverine
Riverine (Volturno River)
Lacustrine – Riverine
Lacustrine
Riverine
Ketmaier & Bianco, in prep.
Alburnus albidus
Alburnus arborella
Ketmaier et al. J Fish Biol 2009 75: 997-1017
Ketmaier et al. J Fish Biol
2009 75: 997-1017
Ketmaier et al. J Fish Biol 2009 75: 997-1017
Ketmaier et al. J Fish Biol 2009 75: 997-1017
Ketmaier et al. J Fish Biol
2009 75: 997-1017
Fiorentino et al. 2008b. BJLS
Transect: 30 km
21 Areas
Fiorentino et al. 2013 MEC
Fiorentino et al. 2013.MEC
Fiorentino et al. 2013.MEC
Fiorentino et al. 2013.MEC
•  Southern Europe: a tale of tales
•  Current diversity is based on adding and mixing
different layers of complexity
•  History of Southern European lineages has been of
preservation rather than dramatic extinction/recolonization events