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Vipera aspis meeting- Leysin, Switzerland - 2015
Fernando Martínez-‐Freiría
CIBIO/InBio, Centro de Investigação em Biodiversidade e Recursos Genéticos. Universidade do Porto. Portugal.
European vipers à genus Vipera
ammodytes
3 clades aspis
pelias Miocene -‐ Pliocene Allopatric speciation processes Reproductive isolation
pre-‐mating barriers post-‐mating barriers
13 species in Europe (???) Parapatric distribution Contact zones:
-‐ sympatry -‐ interspecific competition -‐ gene flow
3 species Vipera seoanei Lataste, 1879 Vipera aspis (Linnaeus, 1758) Vipera latastei Boscá, 1878
Iberian vipers
Euro-‐Siberian
Mediterranean
Different morphological traits
Iberian vipers
Similar ecological traits
Ambush predators Diet
Iberian vipers
Activity
Hibernation -‐ activity periods Moulting periods
Reproduction
Feeding
Mating spring autumn
Viviparous
Parapatric distribution
Different habitat requirements
Different thermoregulatory capacities and abilities
Iberian vipers
V. seoanei – V. aspis
Contact among the 3 species
High course of the
Ebro river
V. aspis – V. latastei
contact zones among Iberian vipers
Morphological convergence
V. seoanei – V. latastei
Local scale latastei-‐seoanei à allopatry aspis-‐seoanei à allopatry aspis-‐latastei à ¿?
1. Distribution 2. Potential distribution
a) environmental factors b) sympatry areas
3. Morphology
a) geographic variation b) environmental factors
4. Ecological traits 5. Gene flow and environmental correlates
a) genetic structure b) hybridization c) ecological divergence d) environmental transition – species traits
V. aspis, V. latastei and V. seoanei
V. aspis and V. latastei
V. aspis, V. latastei and hybrids VA x VL
V. aspis, V. latastei and V. seoanei
Intermediate forms and syntopy among vipers (V. aspis and V. latastei) in Northern Iberian Peninsula
F. Martínez-‐Freiría, J.C. Brito & M. Lizana (2006). The Herpetological Bulletin 97.
Aim # 1
Distribution
intermediate vipers
V. aspis V. latastei
GIS-‐based niche models identify environmental correlates sustaining a contact zone between three species of European vipers
F. Martínez-‐Freiría, N. Sillero, M. Lizana & J.C. Brito (2008). Diversity & Distributions 14.
Aim # 2
Potential distribution
Morphological criterium Gps locationsà UTM 1x1 km squares:
-‐ 48 V. aspis -‐ 54 V. latastei -‐ 19 V. seoanei
Intermediate forms were not used ENMs Maxent 3.0.4 beta ArcMap 9.2 (GIS) – “ensemble forecasting”
7 EGVs
Material and methods
V. aspis V. latastei V. seoanei
Altitude 5.76 (3.0 - 8.5) 2.56 (1.1 - 4.4) 7.67 (4.0 - 14.5)
Precipitation 12.52 (10.0 - 19.7) 54.80 (50.9 - 57.5) 20.28 (13.4 - 27.3)
Evapotransp. 44.71 (40.5 - 48.9) 16.53 (14.8 - 18.8) 17.95 (12.5 - 24.7)
Subst. cover 14.55 (10.9 - 17.6) 0.89 (0.3 - 1.6) 2.15 (0.4 - 5.4)
Slope 9.58 (4.3 - 16.1) 9.65 (5.7 - 16.0) 16.62 (11.2 - 20.0)
Max. Temp. 4.85 (2.7 - 8.9) 7.97 (1.4 - 14.2) 6.25 (3.4 - 10.0)
Min. Temp. 7.99 (5.0 - 10.5) 7.60 (5.7 - 9.9) 29.10 (21.8 - 41.7)
AUC training 0.915 (0.903 - 0.927) 0.951 (0.943 - 0.960) 0.961 (0.946 - 0.970)
AUC test 0.782 (0.707 - 0.865) 0.873 (0.809 - 0.935) 0.780 (0.708 - 0.923)
EGVs importance
Response curves
Results
V. aspis —— V. latastei – · – V. seoanei -------
Precipitation (mm/year)
Slope (º)
areas of potential
distribution
areas of potential sympatry
V. aspis -‐ V. latastei 76 km2
V. aspis -‐ V. seoanei 23 km2 V. latastei -‐ V. seoanei 2 km2
Results
V. seoanei V. aspis
V. latastei
VA -‐VL
VA -‐VS
VL -‐VS
Geographical patterns of morphological variation and environmental correlates in contact zones: a multi-‐scale approach using two Mediterranean vipers
(Serpentes) F. Martínez-‐Freiría, X. Santos, J.M. Pleguezuelos, M. Lizana & J.C. Brito (2009).
Journal of Zoological Systematics and Evolutionary Research 47.
Aim # 3
Morphology
2 scales
regional: NE Iberia 64500 km2
local: High Ebro 300 km2
Study areas
728 vipers à 276 museum collections and 452 field work in the High Ebro
V. aspis V. latastei Intermediate
vipers analyses
NE Iberia
specimens 369 261 98 630
UTM 10x10 99 41 5 135
High Ebro
specimens 183 179 92 362
Material and methods
Morphological criterium and genetic analyses intermediate vipers à mostly hybrids
Idrisi Kilimanjaro 14.01
ArcGis 9.2
9 meristic traits apical scales à APICA supralabials à SUPRAR
infralabials à INFRAR perioculars à PERIR
loreals à LORER intercantals+intersupraoculars à INTER
ventralsà VENT subcaudalsà SUBCA
dorsal marksà DMARK
Models
NE Iberia
Clines, N-‐S oriented
Results
High Ebro
Clines, N-‐S oriented 3 areas of morphological
variability
Results
Environmental correlates
SCALE TRAIT ALT PREC PRECS TEMP TEMPS MINT MAXT
Regional: NE Iberia
APICA -0.176 -0.722 0.486 0.572 0.555 0.151 0.516
DMARK 0.358 0.734 -0.698 -0.441 -0.528 -0.386 -0.629
INTER -0.136 -0.655 0.493 0.535 0.718 0.144 0.484
VENTF 0.363 0.708 -0.595 -0.336 -0.225 -0.437 -0.602
VENTM 0.501 0.757* -0.766* -0.368 -0.459 -0.544 -0.716
PC1 0.329 0.789* -0.667 -0.509 -0.568 -0.354 -0.647
Local: High Ebro
APICA 0.195 -0.734 -0.172 0.709 0.762* -0.307 0.487
DMARK -0.219 0.762* 0.187 -0.753* -0.803* 0.337 -0.499
INTER 0.097 -0.676 -0.096 0.573 0.631 -0.196 0.483
VENTF -0.145 0.605 0.125 -0.552 -0.649 0.227 -0.400
VENTM -0.111 0.795* 0.091 -0.689 -0.700 0.239 -0.576
PC1 -0.173 0.775* 0.150 -0.720 -0.770* 0.291 -0.529
Results
Spatial and temporal segregation allows coexistence in a hybrid zone among two Mediterranean vipers (Vipera aspis and V. latastei)
F. Martínez-‐Freiría, M. Lizana, J.P. do Amaral & J.C. Brito (2010). Amphibia-‐Reptilia 31.
Aim # 4
Ecological traits
Diet V. aspis: 37 (181) V. latastei: 48 (172) Hybrids: 28 (85) 4 categories: invertebrates, reptiles, Soricomorpha and Rodentia. χ2 tests Indexes: Levin, Pianka
Micro-‐habitat – Activity
-‐ radio-‐tracking of 10 males (4 V. aspis, 3 V. latastei and 3 hybrids)
Micro-‐habitat: 297 presences/absences area of 9m2 Logistic regression
Activity, 3 indicators: 1) % of active males 2) ratio moved length /days with movement (AMR) 3) home range size, using KERNEL
ArcGis 9.2 χ2 tests
Material and methods
Material and methods Reproduction
-‐ captured and road-‐kills -‐ 13 pregnant females (4 V. aspis, 7 V. latastei and 2 hybrids)
Potential fecundity, effective fecundity and reproductive effort.
New-‐borns:
Biometry (SVL and Weight) ANOVA tests Mortality (NSB) Moulting traits (NMAP and NWFS)
Frequency of reproduction Test de χ2 Demography
Relative abundance: standardized transects (18.85km) Mortality on roads (3.62 km around transects) χ2 tests KIA’s
Ad-‐hoc observations
Results – diet
similar niche breadth Levin’s index = 0.22 – 0.44
high overlap of trophic niche Pianka’s index > 0.93
temporal differences: feeding frequency (VA < VL < HY) feeding period (HY < VL < VA)
competition for preys
V. aspis V. latastei Hybrids Mollusca 0 2.17 3.57 Coleoptera 0 4.35 0
Invertebrates 0 4.17 3.57
Chalcides 5.41 6.52 14.29 Podarcis 0 2.17 0
Coronella 0 2.17 0 Vipera 2.7 2.17 0
Reptiles 8.11 14.58 10.71
Sorex 8.11 13.04 10.71 Crocidura 8.11 15.22 14.29
Soricomorpha 16.22 20.83 17.86
Microtus 43.24 32.61 42.86 Apodemus 32.43 17.39 25
Rodentia 75.68 60.42 67.86 Mammals 91.89 81.25 85.71
similar preys
V. aspis V. latastei Hybrids Spring
Bushes (β) 0.06** 0.17* - Rocks (β) 0.07*** 0.27* 0.11* Slope (β) - -2.87 - Trees (β) - 0.14 0.20* Constant -2.28 -9.87 -5.41 AUC 0.89 0.99 0.99 S.D. 0.04 0.01 0.01 CCR % train. pres. 82.86 95 94.74 % train. abs. 88.57 95 89.47 % test pres. 100 71.43 100 % test abs. 66.67 100 84.62 Total presences 47 27 51
Summer Bushes (β) 0.12* 0.1 0.07* Musgo (β) - 0.17 - Rocks (β) 0.08* - - Soil (β) - - 0.24* Trees (β) 0.13* - 0.05* Constant -7.46 -4.93 -4.45 AUC 0.94 0.98 0.94 S.D. 0.03 0.02 0.04 CCR % train. pres. 86.36 91.7 89.47 % train. abs. 81.82 91.7 84.21 % test pres. 57.14 59 100 % test abs. 71.43 100 66.67 Total presences 29 16 25
V. aspis V. latastei Hybrids Autumn Bushes (β) 0.22* 0.12* 0.18 Rocks (β) - - 0.13 Slope (β) 3.46* 4.81** - Trees (β) 0.08** - - Constant -15.65 -14.32 -7.9 AUC 0.97 0.97 0.99 S.D. 0.02 0.02 0.01 CCR % train. pres. 92 100 92.9 % train. abs. 88 90.9 92.9 % test pres. 62.5 91.67 100 % test abs. 87.5 75 100 Total presences 33 45 19
Similar elements during most part of the annual cycle: bushes and rocks
Different elements in some seasons: summer Field observations à different locals for
hibernation and gestation.
Results – micro-‐habitat selection
competition – segregation for space
Significant differences
Home range size
Movement rate (AMR)
Results – activity patterns
segregation in temporal axis
V. aspis V. latastei Hybrids
Detected mating period APRIL APRIL MARCH
Minimum size of sexual maturation (mm) Females
Embryos or developed follicles 341 300 414
Mating or with reproductive behaviour 390 470 470
Males Mating or guarding females 395 400 440
Date of parturition AUG-SEP AUG-SEP AUG
V. aspis V. latastei Hybrids Fecundity
Avg nº follicles 7.39 (133 foll) 8.80 (88 foll) 8.80 (88 foll) S.D. 2.56 1.52 1.44 Avg nº embryos 5.75 (46 emb) 7.7 (77 emb) 5.6 (28 emb) S.D. 2.55 2.45 1.67 Avg reprod effort 0.59 0.74 0.79
S.D. 0.24 0.13 0.28
V. aspis V. latastei Hybrids Newborns traits
Avg SVL (mm) 165.86*** 152.42*** 157.58*** S.D. 11.09 7.85 19.36 Avg weight (g) 5.28 4.8 4.75 S.D. 0.88 0.87 1.28 Avg NSB 0.75* 0.94* 0.83* S.D. 0.44 0.23 0.39 Avg NMAP 0.61* 0.81* 0.50* S.D. 0.5 0.39 0.52 Avg NWFS 0.14** 0.00** 0.00** S.D. 0.36 0 0
V. aspis V. latastei Hybrids Frequency of reproduction (%) 2004* 63.2 65 80
2005* 11.1 20 14.3 2006* 18.2 35.3 26.7 Average 30.8 40.1 40.3
S.D. 28.23 22.88 34.92
Triennial reproductive cycle Similar traits
Fecundity: VL > VA >HY Reproductive effort: HY > VL > VA
SVL: VA > HY > VL NSB: VL> HY > VA NMAP: VL > VA> HY NWFS: VA > VL= HY
Results – reproductive traits
Hybrids had lower fitness
Significant differences -‐ Hybrids were more abundant
-‐ Males of V. aspis were less abundant
-‐ Hybrids had low mortality
-‐ Females of V. latastei had high mortality
Results – demographic traits
Parental species had disadvantages Hybrids had a good performance
Hybridization at an ecotone: ecological and genetic barriers between three Iberian vipers
P. Tarroso, R. Pereira, F. Martínez-‐Freiría, R. Godinho & J.C. Brito (2014) Mol Ecol 23
Aim # 5
Gene flow and environmental correlates
DNA of 218 vipers: 210 nuDNA, 8 microsatellites 211 mitDNA, 2 RFLPs
Structure 2.3.3
cluster membership probabilities (CMPs) NewHybrids à hybrids and generation? HybridLab à simulations and thresholds ENMs: Artificial Neural Networks àSimapse 9 EGVs
CMPs of parental taxa SPCA of the EGVs
Material and methods
Suitability maps were contrasted against nuDNA, mtDNA and morphological traits
Cluster membership probabilities (CMPs)
3 populations Pop 1 à V. latastei 68 91 % Pop 2 à V. aspis 87 87 % Pop 3 à V. seoanei 13 100 %
Results – population structure
Interpolation of genetic affinity (IDW)
42 ind with admixed ancestry; 41 VAxVL, 1 VLxVS; none F1
mtDNA à mitochondrial introgression Pop 1 (VL) à 4 ind VA mtDNA Pop 2 (VA) à 8 ind VL mtDNA
Results – environmental analyses
SPCA of the EGVs
strong ecological gradient à ECOTONE
Suitable areas
EGVs importance Response curves
V. seoanei
V. aspis
V. latastei
species are ecologically divergent
hybrids mostly occur in sub-‐optimal areas to parentals
Results – environmental analyses
Results – population structure + environmental analyses mtDNA nuDNA morphology
high spatial concordance among traits and ENMs most hybrids occur in unsuitable areas for parentals
Vipera seoanei Vipera aspis
Vipera latastei
ALLOPATRY
DIFFERENT HABITAT à pre-mating BARRIER
SIMPATRY coexistance
Intermediate vipers
HYBRIDS
Vipera aspis
Vipera latastei
ADAPTATION Morphological convergence
HIBRIDIZATION
Eurosiberian Mediterranean
HYBRIDS
Vipera aspis
Vipera latastei Interspecific competition
(habitat, micro-habitat and diet)
Niche segregation (spatial and temporal)
PARENTAL à disadvantages
HYBRIDS à dis- // advantages
BALANCE population dynamics
ECOTONE
limitation to spread
suboptimal for parental taxa
SIMPATRY
coexistence
Ecological segregation as barrier to gene flow
endogenous / exogenous selection??
§ Co-‐authors of the works: J.C. Brito, P. Tarroso, X. Santos, J.M. Pleguezuelos, N. Sillero, J.P. de Amaral, M. Lizana, R. Pereira, R. Godinho.
§ Family and friends.
§ Asociación Socio-‐cultural Hoces del Alto Ebro y Rudrón (Burgos, Spain).
§ Pre-‐doctoral grant AP2003-‐2633 from Ministerio de Educación, Cultura y Deporte (Spain).
§ Post-‐doctoral grant SFRH/BPD/109119/2015 from FCT (Portugal)
§ Project POCTI⁄∕BIA-‐BDE⁄∕55596⁄∕2004 from Fundacâo para a Ciência e Tecnologia (FCT, Portugal).
§ Project 18.JCY4 463A.C.03 from Junta de Castilla y León (Spain).
