YMPEV 3988 No.

of Pages 11, Model 5G

22 August 2011

Molecular Phylogenetics and Evolution xxx (2011) xxx–xxx

1

Contents lists available at SciVerse ScienceDirect

Molecular Phylogenetics and Evolution

journal homepage: www.elsevier.com/locate/ympev

2 Phylogenetic relationships of the mockingbirds and thrashers (Aves: Mimidae)

3 Irby J. Lovette a,⇑, Brian S. Arbogast b, Robert L. Curry c, Robert M. Zink d, Carlos A. Botero e,
4 John P. Sullivan a, Amanda L. Talaba a, Rebecca B. Harris a, Dustin R. Rubenstein f, Robert E. Ricklefs g,
5 Q1 Eldredge Bermingham h
6 a
Fuller Evolutionary Biology Program, Cornell Lab of Ornithology, Cornell University, 159 Sapsucker Woods Rd., Ithaca, NY 14950, USA
7 b
Department of Biology and Marine Biology, University of North Carolina Wilmington, 601 S. College Rd., Wilmington, NC 28403, USA
8 c
Department of Biology, Villanova University, 800 Lancaster Ave., Villanova, PA 19085, USA
9 d
Bell Museum, 1987 Upper Buford Circle, University of Minnesota, St. Paul, MN 55108, USA
10 e
National Evolutionary Synthesis Center, 2024 W Main St., Suite A200, Durham, NC 27707, USA
11 f
Department of Ecology, Evolution and Environmental Biology, Columbia University, 1200 Amsterdam Ave., New York, NY 10027, USA
12 g
Department of Biology, University of Missouri at St. Louis, 8001 Natural Bridge Rd., St. Louis, MO 63121, USA
13 h
Smithsonian Tropical Research Institute, Balboa, Panama

14
15
a r t i c l e i n f o a b s t r a c t
1
3 7
1
18 Article history: The mockingbirds, thrashers and allied birds in the family Mimidae are broadly distributed across the 32
19 Received 1 January 2011 Americas. Many aspects of their phylogenetic history are well established, but there has been no previous 33
20 Revised 3 June 2011 phylogenetic study that included all species in this radiation. Our reconstructions based on mitochondrial 34
21 Accepted 11 July 2011
and nuclear DNA sequence markers show that an early bifurcation separated the Mimidae into two clades, 35
22 Available online xxxx
the first of which includes North and Middle American taxa (Melanotis, Melanoptila, Dumetella) plus a small 36
radiation that likely occurred largely within the West Indies (Ramphocinclus, Allenia, Margarops, Cinclocer- 37
23 Keywords:
thia). The second and larger radiation includes the Toxostoma thrasher clade, along with the monotypic 38
24 Mimidae
25 Mockingbird
Sage Thrasher (Oreoscoptes) and the phenotypically diverse and broadly distributed Mimus mockingbirds. 39
26 Thrasher This mockingbird group is biogeographically notable for including several lineages that colonized and 40
27 Galapagos diverged on isolated islands, including the Socorro Mockingbird (Mimus graysoni, formerly Mimodes) 41
28 Phylogeny and the diverse and historically important Galapagos mockingbirds (formerly Nesomimus). Our recon- 42
29 Classification structions support a sister relationship between the Galapagos mockingbird lineage and the Bahama 43
30
Mockingbird (M. gundlachi) of the West Indies, rather than the Long-tailed Mockingbird (M. longicaudatus) 44
or other species presently found on the South American mainland. Relationships within the genus Toxos- 45
toma conflict with traditional arrangements but support a tree based on a preivous mtDNA study. For 46
instance, the southern Mexican endemic Ocellated Thrasher (T. ocellatum) is not an isolated sister species 47
of the Curve-billed thrasher (T. curvirostre). 48
Ó 2011 Published by Elsevier Inc. 49

50
51

52 1. Introduction et al., 1997; Derrickson, 1988; Botero et al., 2009a); the group as a 64
whole shows extensive variation in the extent and use of acoustic 65
53 The mockingbirds, thrashers and allied birds that comprise the signals, and this behavioral diversity has informed comparative 66
54 avian family Mimidae are well-known models in evolutionary biol- studies of how complex song traits evolve (Botero et al., 2009b). 67
55 ogy and behavioral ecology. The Galapagos mockingbirds inspired The group is likewise notable for its high interspecific variation in 68
56 Charles Darwin when he noted the morphological differences social and mating systems (Curry, 1989; Curry and Grant, 1990), 69
57 among forms from several islands and their differentiation from re- as even within the mockingbirds breeding groups vary across a 70
58 lated species that occur on the South American mainland (Darwin, sociality spectrum from socially monogamous pairs, to parents 71
59 1845, 1859). Other species of mockingbirds and thrashers have with helpers who are previous offspring, to plural breeding groups 72
60 some of the most extensive song repertoires known in all birds that contain multiple reproducing pairs (Curry, 1988). 73
61 (e.g., Kroodsma and Parker, 1977; Derrickson, 1987; Read and No previous phylogenetic survey has included a full sample of 74
62 Weary, 1992), and many functional and behavioral aspects of their mimid species, and this lack of a completely inclusive phylogenetic 75
63 vocal and learning abilities have been characterized (e.g., Kroodsma hypothesis has hindered some comparative analyses of trait varia- 76
tion in this group. Nonetheless, many aspects of the phylogenetic 77
⇑ Corresponding author. Fax: +1 607 254 2486. history of the Mimidae radiation are well established. The mimid 78
E-mail address: IJL2@cornell.edu (I.J. Lovette). clade is sister to the more diverse Old World starlings (Sturnidae; 79

1055-7903/$ - see front matter Ó 2011 Published by Elsevier Inc.

doi:10.1016/j.ympev.2011.07.009

Please cite this article in press as: Lovette, I.J., et al. Phylogenetic relationships of the mockingbirds and thrashers (Aves: Mimidae). Mol. Phylogenet. Evol.
(2011), doi:10.1016/j.ympev.2011.07.009
 YMPEV 3988 No. of Pages 11, Model 5G
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2 I.J. Lovette et al. / Molecular Phylogenetics and Evolution xxx (2011) xxx–xxx

80 Beecher, 1953; Stallcup, 1961; Sibley and Ahlquist, 1984, 1990; shown that the historical population structure within the Nesomi- 146
81 Malcarney et al., 1994; Voelker and Spellman, 2004; Ericson and mus group does not fully track the traditional species boundaries 147
82 Johansson, 2003; Cibois and Cracraft, 2004; Barker et al., 2004; that were defined based on phenotypic traits, suggesting instead 148
83 Zuccon et al., 2006; Lovette and Rubenstein, 2007). The Mimi- that this radiation has a complex within-archipelago history of dis- 149
84 dae + Sturnidae are in turn sister to the lineage comprising the persal, gene flow, and phenotypic differentiation (Arbogast et al., 150
85 two species of Oxpeckers (Buphagidae), and fall within a broader 2006; Hoeck et al., 2010). Several studies have found the Nesomi- 151
86 songbird radiation that includes the thrushes and dippers (Cibois mus group to be embedded within the broader Mimus clade 152
87 and Cracraft, 2004; Zuccon et al., 2006; Lovette and Rubenstein, (Arbogast et al., 2006; Lovette and Rubenstein, 2007) and taxo- 153
88 2007). Several phenotypically odd New World taxa (e.g., Donacobi- nomically this group is now merged into Mimus (Remsen et al., 154
89 us, Rhodinocichla) have at times been suggested to be allied to the 2010), but precisely where the Nesomimus lineage falls in relation 155
90 Mimidae, but phylogenetic evidence has since shown that they fall to these other mockingbirds remains uncertain. 156
91 elsewhere within the Passeriformes (Seutin and Bermingham, A somewhat analogous situation involves the Socorro Mocking- 157
92 1997; Alström et al., 2006; Johansson et al., 2008; Gelang et al., bird (Mimus graysoni), a morphologically distinct endangered spe- 158
93 2009). In reconstructions based on molecular data, a long and cies endemic to its remote namesake island. This species was 159
94 highly supported branch separates the Sturnidae from the earliest separated into the monotypic genus Mimodes until analyses of 160
95 node within the Mimidae (Zuccon et al., 2006; Lovette and mitochondrial DNA characters revealed that it falls well within 161
96 Rubenstein, 2007), thereby strongly supporting the monophyly of the Mimus mockingbird group (Barber et al., 2004). 162
97 the Mimidae as it is currently classified (e.g., A.O.U., 1998). Relationships within the Tropical (M. gilvus) and Northern (M. 163
98 The broad outline of phylogenetic relationships within the polyglottos) Mockingbird complex are complex and not fully re- 164
99 Mimidae is also well established (Hunt et al., 2001; Lovette and solved. These taxa hybridize in their zone of contact in central 165
100 Rubenstein, 2007). Species in the family fall into two clades, the Mexico (Wetmore, 1943), and they have sometimes been consid- 166
101 first of which comprises three species found in Mexico and north- ered conspecific. They are generally classified as separate species 167
102 ern Central America (Melanoptila and Melanotis), plus a group that (e.g., Davis and Miller, 1960; Mayr and Short, 1970; A.O.U., 1998) 168
103 has radiated mostly within the West Indian archipelago (Cinclocer- and mitochondrial DNA-based phylogenetic reconstructions sug- 169
104 thia, Margarops, Allenia, Ramphocinclus). The migratory and mono- gest that some populations of these taxa are well differentiated 170
105 typic Gray Catbird (Dumatella carolinensis) which breeds in North (Hunt et al., 2001). Morphological traits suggest this complex fur- 171
106 America is embedded within this otherwise West Indian clade ther includes the San Andrés Mockingbird, a distinctively large- 172
107 (Hunt et al., 2001; Lovette and Rubenstein, 2007). At the intraspe- billed population found only on its namesake islands in the wes- 173
108 cific level, the phylogeography of all endemic West Indian species tern Caribbean that has most often been considered a subspecies 174
109 found on more than one island (Cinclocerthia ruficada, C. gutturalis, of gilvus (e.g., Davis and Miller, 1960; A.O.U., 1998; Dickinson, 175
110 Margarops fuscatus, Allenia fusca) has been explored using mito- 2003), but which is sometimes treated as the full species M. mag- 176
111 chondrial markers (Hunt et al., 2001). nirostris (e.g., Bond, 1956). 177
112 The second and larger of the two subclades within the Mimidae Here we use mitochondrial and nuclear intron DNA sequence 178
113 comprises the continental thrashers (Toxostoma and Oreoscoptes) markers to reconstruct the phylogenetic relationships among all 179
114 and the mockingbirds (Mimus, plus the species formerly assigned Mimidae species, which we hope will aid in future comparative 180
115 to Nesomimus and Mimodes). Relationships among Toxostoma pop- analyses of morphological, ecological, and behavioral trait varia- 181
116 ulations and species have been reconstructed in an extensive series tion in this group. 182
117 of studies using allozyme, mitochondrial DNA, morphological, and
118 behavioral characters (Zink et al., 1997, 1999; Zink and Blackwell-
119 Rago, 2000; Zink et al., 2001; Rojas-Soto, 2003; Sgariglia and Burns, 2. Materials and methods 183
120 Q2 2003; Rojas-Soto et al., 2007). Collectively, these studies have sup-
121 ported the monophyly of the species assigned to Toxostoma, clari- 2.1. Taxonomic and molecular marker sampling 184
122 fied the relationships among many of their populations, and
123 identified some populations that might warrant future recognition We included in our analyses sequences from all 34 currently 185
124 as full species (e.g., Rojas-Soto et al., 2007). At a slightly deeper recognized species of Mimidae (A.O.U., 1998; Dickinson, 2003), 186
125 phylogenetic level, the Toxostoma clade is sister to the remaining including multiple representatives of most species (Appendix 187
126 taxa in this thrasher/mockingbird group. The monotypic Sage A). The majority of the samples obtained for this study allowed 188
127 Thrasher (Oreoscoptes montanus) is in turn sister to a group that in- the extraction of high-quality DNA, and from these we generally 189
128 cludes all of the Mimus mockingbirds (Lovette and Rubenstein, obtained sequences of five protein-coding mtDNA genes (ND2, 190
129 2007). CO1, CO2, ATPase8, and ATPase6) with a combined length of 191
130 The Mimus mockingbird radiation is relatively recent and pre- 4118 nucleotides, and sequences from three autosomal nuclear 192
131 sents a number of phylogenetic and systematic challenges. Mock- intron loci, beta-fibrinogen introns 5 (FGB-5; Kimball et al., 193
132 ingbird species are distributed throughout much of North, 2009) and 7 (FGB-7; Prychitko and Moore, 1997), transforming 194
133 Central, and South America, and different lineages have colonized growth factor beta-2 intron 5 (TGFB2–5; Primmer et al., 2002), 195
134 both archipelagos (the West Indies and the Galapagos) and conti- and rhodopsin intron 1 (RHO-1; Primmer et al., 2002). The com- 196
135 nental satellite islands (Isla Socorro in the Pacific Ocean off of Mex- bined length of these nuclear loci is 2963 aligned nucleotides 197
136 ico; the Islas San Andrés in the Caribbean off of southern Central (not including regions of ambiguous alignment). Our laboratory 198
137 America). As is often the case for insular birds, many of the island methods for the generation of these sequences have been de- 199
138 mockingbird lineages have differentiated substantially in morpho- tailed elsewhere (Lovette and Rubenstein, 2007; Lovette et al., 200
139 logical and behavioral traits. This differentiation is pronounced 2010). In some cases (see Appendix A) we did not obtain the full 201
140 within the Galapagos, where the four generally recognized species set of loci from a sample, as was also true in the few cases when 202
141 have evolved differences in bill shape and size, plumage pattern, the sequences were obtained from studies conducted in other 203
142 body size, song, and mating systems, among other traits (Gulledge, laboratories (Appendix A; Barber et al., 2004; Arbogast et al., 204
143 1975; Abbott and Abbott, 1978; Curry, 1989). Phylogeographic sur- 2006). 205
144 veys of the Galapagos populations using mitochondrial (Arbogast We generated sequences from three species (Mimus graysoni, 206
145 et al., 2006) and microsatellite (Hoeck et al., 2010) markers have Toxostoma ocellatum, Toxostoma guttatum) and four Mexican Mimus 207

Please cite this article in press as: Lovette, I.J., et al. Phylogenetic relationships of the mockingbirds and thrashers (Aves: Mimidae). Mol. Phylogenet. Evol.
(2011), doi:10.1016/j.ympev.2011.07.009
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208 gilvus individuals (a species from which we also included to each of the three independent nuclear loci (FGB-5 and FGB-7 270
209 high-quality samples from other locations) from toe-pads taken are linked and were grouped into a single partition) and separate 271
210 from museum skin samples. Toe-pad samples were extracted and partitions for each codon position in the mtDNA sequences. Like- 272
211 amplified in a dedicated ancient-DNA laboratory facility using wise, the four mtDNA genes are a single linkage group and were 273
212 stringent protocols as described in Lovette and Rubenstein analyzed together. For each partition (mtDNA1stcodon, 274
213 (2007). From the M. graysoni and T. guttatum samples we se- mtDNA2ndcodon, mtDNA3rdcodon, FGB-5 + FGB-7, TGFB2–5, RHO- 275
214 quenced nearly the entire set of mtDNA coding genes; from the 1), we used the program MrModeltest v2 (Nylander, 2004) to iden- 276
215 T. occellatum and M. gilvus samples we sequenced only the mtDNA tify the best-fitting model under the Bayesian information criterion 277
216 ND2 gene. (BIC; Akaike, 1973; Kass and Raftery, 1995; Posada and Buckley, 278
217 Preliminary analyses showed that in all cases, sequences from 2004). The preferred model for all mtDNA partitions was 279
218 conspecific samples had very high similarity. Because we are pri- GTR + G + I and the preferred model for the Fib 5&7, TGFB5 and 280
219 marily concerned here with interspecific relationships, and to re- Rho1 partitions was HKY + G. These two models were applied 281
220 duce computation times, we selected a single representative of respectively to all analyses involving these data partitions. 282
221 each species to include in some phylogenetic analyses (Appendix
222 A), although we report additional analyses (see below) with all 2.4. Gene tree analyses 283
223 samples included.
224 Previous studies have provided robust evidence that the Mimi- To assess congruence among independent gene trees, we ran 284
225 dae species included here form a monophyletic group that is the separate Bayesian MCMC phylogenetic analyses on each of the 285
226 sister-group of the starlings (Sturnidae; e.g., Sibley and Ahlquist, four unlinked data partitions (mtDNA, FGB-5 + FGB-7, TGFB2–5, 286
227 1990; Barker et al., 2004; Cibois and Cracraft, 2004; Zuccon et al., RHO-1). To facilitate comparisons across datasets, these analyses 287
228 2006; Voelker and Spellman, 2004; Lovette and Rubenstein, included only single representatives of 30 Mimidae species from 288
229 2007). We included single representatives of five Sturnidae subc- which we sampled all loci (Mimidae species entirely missing from 289
230 lades (see Lovette and Rubenstein, 2007; Lamprotornis superbus, these nuclear gene trees are M. graysoni, T. guttatum, T. ocellatum, 290
231 Onychognathus morio, Aplonis panayensis, Sturnus vulgaris, Rhabdor- and T. curvirostre). Each gene-tree analysis was run in MrBayes for 291
232 nis inornatus) as outgroups, along with one representative (Bupha- 1 x 107 generations sampled every 1 x 104 generation, with five 292
233 gus africanus) of the sister group to the Mimidae + Sturnidae, and replicated runs of four chains with default heating. Plots of –ln 293
234 one more distant outgroup taxon (Catharus guttatus). likelihood scores, parameter values, and posterior probability 294
scores for individual nodes suggested that each analysis had con- 295
235 2.2. Bayesian phylogenetic methods verged before (usually well before) the 5  106 generation. The 296
topologies of all well-supported nodes of the two independent 297
236 We based all phylogenetic reconstructions on Markov chain runs also converged by the 5  106 generation, as did the poster- 298
237 Monte Carlo (MCMC) analyses conducted using MrBayes 3.1.2. ior probability scores of individual nodes as assessed graphically 299
238 (Huelsenbeck and Ronquist, 2001; Ronquist and Huelsenbeck, using AWTY. We discarded the first 9  10 generations as ‘burn- 300
239 2003). The number of MCMC generations, number of concurrent in’ and combined the remaining 100 samples from each run to 301
240 chains, and number of separate runs varied among analyses, as de- calculate 90% majority-rule consensus topologies for each gene 302
241 scribed below. All MCMC runs were sampled every 1  104 gener- tree. The .con file created by MrBayes was visualized in FigTree 303
242 ations and were run using the parallel computing resources of the v.1.2.2. This consensus tree was confirmed in PAUP by combining 304
243 Computational Biology Service Unit at the Cornell University The- the last 90% of each .t file into one .t file. A majority-rule consen- 305
244 ory Center. sus tree was then visualized. 306
245 For each set of MCMC analyses, we evaluated the ‘burn-in’ per- We used a similar approach to analyze the mitochondrial se- 307
246 iod by plotting variation in log likelihood scores and individual quences alone. 308
247 parameter values across the duration of the run, and by evaluat-
248 ing variation in the posterior probability scores of the most vari-
2.5. Combined data analyses 309
249 able nodes using the web-based program AWTY (Wilgenbusch
250 et al., 2004). We then discarded the pre-stationarity output, and
We conducted further MCMC analyses that included all loci 310
251 to be conservative at least the initial 25% of the apparently sta-
sampled from all samples listed in Appendix A. These more com- 311
252 tionary samples, based on the slowest-converging metric. We
prehensive analyses were run with six partitions, three for the 312
253 evaluated convergence among runs by comparing the resulting
mitochondrial codon positions and one each for the three unlinked 313
254 consensus topologies, and by plotting the posterior probability
nuclear intron loci. Five independent runs, each with four concur- 314
255 scores of individual nodes using AWTY. The retained post-sta-
rent chains, were run for 1.3  107 generations. The first 1  106 315
256 tionarity samples from separate but convergent runs were then
generations were discarded as burn-in; the remaining samples 316
257 combined to calculate majority-rule consensus trees, posterior
from the independent runs were highly congruent, and the com- 317
258 probability values for each node, mean branch lengths, and
bined post-burn-in samples from all five runs were used to derive 318
259 parameter estimates.
a 50% majority rule consensus tree and to calculate mean branch 319
260 In addition to the phylogenetic analyses covered in detail be-
lengths. 320
261 low, we also ran a variety of preliminary analyses using other
262 phylogenetic methods (maximum likelihood and maximum parsi-
263 mony), alternative data partitioning schemes, and subsets of the 3. Results 321
264 overall dataset. Those results were highly consistent with the re-
265 sults of the comprehensive Bayesian analyses reported here. All sequences new to this study have been deposited in Gen- 322
Bank; Appendix A gives accession numbers for both these and all 323
266 2.3. Data partitions and model selection previously reported sequences. 324
The Bayesian analyses of all concatenated loci from all samples 325
267 Our dataset comprised a heterogeneous mix of mtDNA protein- produced a topology (Fig. 1) in which nearly all nodes are highly 326
268 coding and nuclear non-coding loci, and hence we used six data supported, and which is largely consistent with the relationships 327
269 partitions in our likelihood analyses, assigning single partitions found in previous studies that included subsets of these taxa as de- 328

Please cite this article in press as: Lovette, I.J., et al. Phylogenetic relationships of the mockingbirds and thrashers (Aves: Mimidae). Mol. Phylogenet. Evol.
(2011), doi:10.1016/j.ympev.2011.07.009
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4 I.J. Lovette et al. / Molecular Phylogenetics and Evolution xxx (2011) xxx–xxx

Fig. 1. Relationships among species of Mimidae estimated by Bayesian MCMC analyses of concatenated mitochondrial and nuclear intron sequences subdivided into six
partitions (three mitochondrial partitions by codon position; nuclear locus partitions FGB-5 + FGB-7, TGFB2-5, RHO-1). The topology shown here is a 50% majority rule
consensus of post-burn-in trees. Numbers adjacent to nodes indicate posterior probability values >75. Topology is rooted to outgroup taxa (not shown) in the sister family
Sturnidae as described in the text. Geographic descriptors at right indicate primary area of breeding distribution.

Please cite this article in press as: Lovette, I.J., et al. Phylogenetic relationships of the mockingbirds and thrashers (Aves: Mimidae). Mol. Phylogenet. Evol.
(2011), doi:10.1016/j.ympev.2011.07.009
 YMPEV 3988 No. of Pages 11, Model 5G
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329 scribed above. Novel or potentially important aspects of this tree 4. Discussion 354
330 include the placement of the Galapagos mockingbird lineage as sis-
331 ter to the Bahama mockingbird (Mimus gundlachi) with a posterior 4.1. Phylogenetic relationships within the Mimidae 355
332 probability score of 100; this relationship was found with lower
333 support in previous studies (Arbogast et al., 2006). As has been Fig. 1 presents our most comprehensive summary of the phylo- 356
334 found previously, both the Galapagos Mockingbird group (previ- genetic relationships within the Mimdae based on our analyses of 357
335 ously separated as Nesomimus) and the Socorro Mockingbird (pre- mitochondrial and nuclear DNA sequence markers. Under this 358
336 viously in Mimodes) are embedded within a shallow clade that also phylogenenetic hypothesis, the Mimidae can be subdivided by a 359
337 includes all Mimus mockingbird taxa. Taxa that have not been in- basal split into two daughter clades, one of which comprises the 360
338 cluded in earlier molecular phylogenetic reconstructions include ‘blue’ mockingbirds, New World catbirds, and Antillean thrashers, 361
339 the Brown-backed Mockingbird (Mimus dorsalis) of the southwest- the other containing the Mimus mockingbirds and the continental 362
340 ern Andes which falls within a clade of other continental South thrashers. 363
341 American mockingbirds, and the San Andrés Mockingbird (M. [gil-
342 vus] magnirostris) which falls within the Northern/Tropical (M. 4.1.1. Blue mockingbirds, new world catbirds, and antillean thrashers 364
343 polyglottos/gilvus) mockingbird complex. The blue mockingbird/catbird/antillean thrasher clade contains 365
344 As expected at this level of divergence, all phylogenetic analyses nine species that are currently assigned to seven genera (Melanotis, 366
345 based on single nuclear loci had low resolution (Fig. 2), although Melanoptila, Dumetella, Ramphocinclus, Allenia, Margarops, Cinclo- 367
346 each provided some support for relationships that receive higher certhia; A.O.U., 1998). Relationships within this group were ex- 368
347 support in the in the concatenated analysis. In contrast, the analy- plored by Hunt et al. (2001) using mitochondrial markers and by 369
348 ses based on mitochondrial sequences alone (Fig. 3) produced a Lovette and Rubenstein (2007) using both mitochondrial and nu- 370
349 strongly supported topology that was identical at the species and clear DNA data; our topologies here are consistent with those ear- 371
350 higher level to that of the concatenated analysis (Fig. 1). Consid- lier results and the historical interpretations based on them. An 372
351 ered together, these results indicate that the topology in the con- early split within this clade separates the blue mockingbird lineage 373
352 catenated analysis is driven overwhelmingly by the signal in the from a clade containing the remaining catbirds and Antillean 374
353 mitochondrial data partition. thrashers. The two Melanotis blue mockingbirds have allopatric 375

Fig. 2. Three single-locus gene tree hypotheses based on nuclear intron sequences. Topologies represent 50% majority rule consensus of post-burn-in trees. Numbers adjacent
to nodes indicate posterior probability values >75%. All reconstructions were rooted using Sturnidae outgroup taxa (not shown).

Please cite this article in press as: Lovette, I.J., et al. Phylogenetic relationships of the mockingbirds and thrashers (Aves: Mimidae). Mol. Phylogenet. Evol.
(2011), doi:10.1016/j.ympev.2011.07.009
 YMPEV 3988 No. of Pages 11, Model 5G
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6 I.J. Lovette et al. / Molecular Phylogenetics and Evolution xxx (2011) xxx–xxx

100 Cinclocerthia ruficauda

100 Cinclocerthia gutturalis
100 Margarops fuscatus
Allenia fusca
100

Dumetella carolinensis
100 98

Ramphocinclus brachyurus
100
Melanoptila glabrirostris

100 Melanotis caerulescens

Melanotis hypoleucus
100
Mimus dorsalis
100 Mimus triurus
100 Mimus saturninus

100 Mimus patagonicus

Mimus thenca
Mimus gilvus (Trinidad)
Mimus gilvus (Colombia)
100
Mimus polyglottos
100
Mimus gilvus (Mexico)
100
Mimus gilvus (Isla San Andres)
100

Mimus gilvus (Mexico)

98
Mimus graysoni
100
88 Mimus longicaudatus
Mimus gundlachii
100 Mimus parvulus
100
100 Mimus trifasciatus
100 Mimus macdonaldi
Mimus melanotis
Oreoscoptes montanus
100 Toxostoma cinereum
100
Toxostoma bendirei
100
Toxostoma crissale
100

Toxostoma lecontei
91

Toxostoma redivivum
100

Toxostoma rufum
100

97
Toxostoma longirostre
91
Toxostoma guttatum
99
Toxostoma ocellatum
Toxostoma curvirostre
Fig. 3. Bayesian likelihood reconstruction of relationships among Mimidae species based on mitochondrial sequences alone. Numbers adjacent to nodes indicate posterior
probability values >75%. All reconstructions were rooted using Sturnidae outgroup taxa (not shown).

Please cite this article in press as: Lovette, I.J., et al. Phylogenetic relationships of the mockingbirds and thrashers (Aves: Mimidae). Mol. Phylogenet. Evol.
(2011), doi:10.1016/j.ympev.2011.07.009
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I.J. Lovette et al. / Molecular Phylogenetics and Evolution xxx (2011) xxx–xxx 7

376 distributions in Mexico and northern Central America, and they are (A.O.U., 1998, and later supplements). Within the first of these 441
377 clearly sister taxa although they are separated by substantial ge- groups, the Gray (T. cinereum) and Bendire’s (T. bendirei) thrashers 442
378 netic divergence in comparison to many other sister species pairs are sister species separated by relatively low genetic divergence, 443
379 within the Mimidae (Fig. 3). consistent with the expectation that these taxa are closely allied 444
380 All but one of the remaining taxa in the catbird/antillean (e.g., Mayr and Short, 1970; A.O.U., 1998; Zink et al., 1999). This 445
381 thrasher group have distributions within the Caribbean basin. group also contains the more divergent sister-species pair formed 446
382 The basal split within this clade separates the Black Catbird (Mela- by the Crissal (T. crissale) and Le Conte’s (T. lecontei) thrashers, 447
383 noptila glabrirostris) of the Yucatan peninsula from the remaining which together are in turn sister to the California Thrasher 448
384 species. This general topology, with two successive early-splitting (T. redivivium). 449
385 lineages now found on the Central American mainland, may hint The second Toxostoma clade includes the Curve-billed Thrasher 450
386 at the route of colonization into the West Indies taken by the (T. curvirostre) as sister to the remaining four species; this topology 451
387 ancestor of the radiation now found in that island system. The sis- is not consistent with the suggestion (Sibley and Monroe, 1990) 452
388 ter-clade to Melanotis comprises five species of thrashers resident that T. curvirostre is most closely allied to the Ocellated Thrasher 453
389 within the West Indies, plus the Gray Catbird (Dumetella carolinen- (T. ocellatum). However, the clade formed by the Brown (T. rufum), 454
390 sis), a migratory species with a broad breeding range that spans Long-billed (T. longirostre), and Cozumel (T. guttatum) thrashers 455
391 much of the United States and southern Canada, and which winters matches previous predictions about their close affinities (Mayr 456
392 in the continental regions surrounding the Gulf of Mexico and on and Short, 1970; A.O.U., 1998; Zink et al., 1999; Sibley and Monroe, 457
393 the western islands in the Greater Antilles. As found previously 1990). 458
394 (Hunt et al., 2001; Lovette and Rubenstein, 2007), Dumetella is
395 embedded within the Antillean thrasher radiation as the sister tax- 4.1.3. Mimus mockingbirds 459
396 on of Ramphocinclus brachyurus, the White-breasted Thrasher, a The Mimus mockingbirds form a shallow clade of 14 or 15 spe- 460
397 highly endangered species found only on the Lesser Antillean is- cies that collectively are sister to the Sage Thrasher (Oreoscoptes 461
398 lands of Martinique and St. Lucia. Based on the phylogenetic evi- montanus) lineage (Fig. 1). Subgroups within this relatively recent 462
399 dence, it seems most likely that the ancestral Dumetella lineage radiation include a clade of five species endemic to South America. 463
400 recolonized North America from the West Indies, adding to the Within this South American group, the two southernmost species 464
401 growing list of avian examples of such ‘reverse colonization’ of (the Chilean Mockingbird M. thenca and Patagonian Mockingbird 465
402 continental areas from island source populations that have been M. patagonicus) are allopatric sister taxa separated by the heights 466
403 identified from phylogenetic evidence (Bellemain and Ricklefs, of the southern Andes. The range-restricted Brown-backed Mock- 467
404 2008; Sheldon et al., 2009; Sturge et al., 2009). The Dumetella line- ingbird (M. dorsalis), which has not been included in previous phy- 468
405 age likewise probably transitioned to long-distance migration from logenetic surveys of this genus, is sister to the White-banded 469
406 a sedentary ancestor, a change consistent with predictions that Mockingbird (M. triurus), and these two species are in turn sister 470
407 avian migratory behaviors have high evolutionary lability despite to the broadly distributed Chalk-browed Mockingbird (M. 471
408 the complex suite of traits necessary to sustain successful migra- saturninus). 472
409 tion (Berthold, 1999; Zink, 2002). A second group within Mimus comprises the Northern (M. 473
410 The five species of resident Antillean thrashers are all found in polyglottos) and Tropical (M. gilvus) Mockingbirds along with 474
411 the Lesser Antilles, with one (Margarops fuscatus) extending its two island forms, the Socorro Mockingbird (Mimus [Mimodes] 475
412 range to Puerto Rico and the Bahamian archipelago. These birds graysoni) and the San Andrés Mockingbird (M. [gilvus] magniros- 476
413 have a complex pattern of island occupancy, with some Lesser tris). As found previously (Barber et al., 2004), M. graysoni is 477
414 Antillean islands supporting all five species. The morphological embedded within the larger Mimus clade, where it sister to the 478
415 and behavioral variation within this shallow clade of thrashers is remaining members of the M. polyglottos/gilvus group. Relation- 479
416 correspondingly high, and this group may represent the sole avian ships among the polyglottos/gilvus group are complicated, as both 480
417 example of a Lesser Antillean adaptive radiation (Ricklefs and Ber- M. magnirostris and M. polyglottos fall within the M. gilvus cluster, 481
418 mingham, 2008). thereby rendering M. gilvus paraphyletic. At this level of diver- 482
419 The high morphological distinctiveness of the Antillean gence, nearly all of the phylogenetic signal in our analyses derives 483
420 thrasher species has resulted in all but Cinclocerthia being currently from the mitochondrial loci (see Fig. 2), so this paraphyly reflects 484
421 classified into monotypic genera (A.O.U., 1998; Banks et al., 2002). the history of the associated mitochondrial gene tree. In the ab- 485
422 Although many of these taxa are morphologically divergent from sence of new behavioral and genetic information from the M. 486
423 the other species in this clade, it could be argued that this group polyglottos/gilvus hybrid zone in Central Mexico, we recommend 487
424 is generically over-split, as this Antillean thrasher clade is likely retaining the current species-level taxonomy for these groups, 488
425 no older than the Mimus or Toxostoma clades which similarly con- recognizing both M. gilvus and M. polyglottos but considering M. 489
426 tain a range of species with high phenotypic variation, including magnirostris a subspecies of M. gilvus. 490
427 some island forms that have differentiated notably from their con- A final group within Mimus comprises the Galapagos mocking- 491
428 tinental allies. There is no intermediate generic reassignment that bird clade, and their apparent sister taxon, the Bahama Mocking- 492
429 recognizes monophyletic subgroups within this clade while also bird (M. gundlachii). 493
430 reflecting the major phenotypic differences among these taxa. If
431 the genera we currently recognize as Melanoptila, Dumetella, Ram- 4.2. Historical biogeography of oceanic island mockingbirds 494
432 phocinclus, Allenia, Margarops, and Cinclocerthia were grouped into
433 a single genus, Dumetella Wood, 1837 would have nomenclatural The current distribution of Mimus mockingbirds suggests that 495
434 priority. these birds (or their recent ancestors) are accomplished over- 496
water dispersers, as at least five Mimus lineages have indepen- 497
435 4.1.2. Toxostoma thrashers dently colonized one or more oceanic islands. In addition to being 498
436 Our reconstructions of species-level relationships of the Toxos- found across much of North America, the Northern Mockingbird 499
437 toma thrashers are highly consistent with previous phylogenetic (M. polyglottos) is resident through the Bahamas, the large islands 500
438 hypotheses for this group (Zink et al., 1999; Lovette and Ruben- of the Greater Antilles except for Jamaica, and the Cayman Is- 501
439 stein, 2007). We found these birds to be subdivided into two clades lands; phylogeographic studies have shown that these West In- 502
440 (Fig. 1), each comprised of five currently recognized species dian populations are closely allied to the more northerly 503

Please cite this article in press as: Lovette, I.J., et al. Phylogenetic relationships of the mockingbirds and thrashers (Aves: Mimidae). Mol. Phylogenet. Evol.
(2011), doi:10.1016/j.ympev.2011.07.009
 YMPEV 3988 No. of Pages 11, Model 5G
22 August 2011

8 I.J. Lovette et al. / Molecular Phylogenetics and Evolution xxx (2011) xxx–xxx

504 continental populations, implying a recent set of island coloniza-

EF484115

EF484120
505 tions (Hunt et al., 2001). The Tropical Mockingbird (M. gilvus) has

TGFB2-4

Pending

Pending

Pending

EF484125

EF484119
similarly colonized the most of the Lesser Antillean island chain

Pending

Pending
506
507 from South America (Hunt et al., 2001). In the western Caribbean,

—
—
508 a separate colonization by a lineage in the M. polyglottos/gilvus
509 group gave rise to the unusually large-billed population (spe-

EF472856

EF472861
Pending

Pending

Pending

EF472866

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510 cies/subspecies M. magnirostris) on San Andrés Island; the mito-

Pending

Pending
Pending
Rho-1
511 chondrial evidence suggests that the San Andrés population is
512 allied to M. gilvus lineages from southern Mexico (Fig. 1), thereby

—
513 suggesting its likely colonization source. The more divergent

EF471844

EF471849
514 mockingbird population (M. graysonii) endemic to Isla Socorro–a

Pending

Pending

Pending

EF471854

EF471848
Nuclear intron loci

Pending

Pending
Pending
515 small, arid island in the Pacific Ocean located about 600 km west

Fib-7
516 of the central Mexican coast – is apparently derived from the

—
517 lineage that is also ancestral to the entire M. polyglottos/gilvus

EF468323

EF468328
518 group, implying a substantially earlier colonization of that remote

Pending

Pending

Pending

EF468333

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519 island along with a substantial period of isolation of M. graysonii.

Pending

Pending
Fib-5
520 (Fig. 1).
The mockingbird dispersal event with the most notable evolu-

—
—
521
522 tionary consequences was the colonization of the Galapagos by

EF486776

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e
the ancestor of the lineages that then radiated in that archipelago.

Pending

Pending

Pending
ATPases

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523

Pending

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Pending
524 Darwin noted the affinities of the Galapagos mockingbirds to spe-
525 cies of Mimus from the South American continent (Darwin, 1859),

—
and it was long assumed that the Galapagos lineage was most

Taxa included in this study, tissue types, collecting localities, institutional sources, and GenBank accession numbers.
526

EF484317

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527 closely related to species found on that relatively nearby source.

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Pending

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528 Mitochondrial sequence-based analyses by Arbogast et al.

CO2
Mitochondrial coding genes
529 (2006) instead identified the Bahama Mockingbird (M. gundlachii)

—
530 as the sister lineage to the Galapagos clade, a finding mirrored in
531 our phylogenetic results presented here (Fig. 1) based on some-

EF484216

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Pending

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EF484226

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532 what more extensive sampling of DNA markers. In addition to

Pending

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Pending
533 being recovered in our phylogenetic analyses of mitochondrial se- CO1
534 quences (Fig. 3), the sister relationship between the Bahama

—
535 Mockingbird and the Galapagos Mockingbird group was notably
EF468192

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536 recovered in the independent topologies from two of our three Pending

Pending

Pending

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537 nuclear loci (Fig. 2). Several other Galapagos vertebrates are allied
d
ND2

538 to West Indian taxa (reviewed in Parent et al., 2008), suggesting

539 that in each case the common ancestor of these island forms
Mexico, Quintana Roo, Isla
Bahamas, Grand Bahama I.

USA, New York, Tompkins

USA, New York, Tompkins

540 was prone to over-water dispersal.

USA, California, San Bernardino

USA, New York, Tompkins Co.
Trinidad, Chacachacare I.
5. Uncited reference

Mexico, Quintana Roo

541

542 Q3 Posada and Crandal (1998).

Cozumel
Locality

Colombia
Colombia
Mexico

543 Acknowledgments
Co.

Co.

544 We thank B. McCleery and D. Rabosky for advice and assis-

Typeb

545 tance, and two reviewers for the comments on the manuscript.
T

B
B
T
T
T

Part of this work was carried out using the resources of the Com-
S

546
547 putational Biology Service Unit at Cornell University which is par-
548 tially funded by Microsoft Corporation. We thank the following
Museum sourcea &

549 institutions, their field collaborators, and their collections staff

LSUMNS-B0081

550 for the loan of genetic materials: Academy of Natural Sciences,

STRI-BHDCA4

LSUMNS-B21369
CUMV-50343

CUMV-50532

CUMV-50356

551 Philadelphia; Bell Museum of Natural History, University of Min-

sample No.

CUMV-33468
CUMV-50469
STRI-CCMGI1

552 nesota; Cornell University Museum of Vertebrates; Louisiana

CAB111c
CAB110c

553 State University Museum of Natural Science; Museo Nacional de

554 Historia Natural, Santiago, Chile; San Diego State University
555 Museum of Biodiversity; Wake Forest University. We thank
556 B. Arbogast and D. Cadeña for providing previously unpublished
557 sequences generated in their laboratories. We are grateful to
polyglottos
polyglottos
glabrirostris

glabrirostris
carolinensis

carolinensis

carolinensis

558 P. Tubaro of the Museo Argentino de Ciencias Naturales, Buenos

Melanoptila

Melanoptila

gilvus
gilvus
gilvus
gilvus

Aires for his guidance in obtaining specimens of South American

Dumetella

Dumetella

Dumetella

559
Appendix A

560 mockingbird species. CAB is funded by the National Evolutionary

Taxon

Mimus
Mimus
Mimus
Mimus
Mimus
Mimus

561 Synthesis Center (NESCent), NSF Grant EF-0905606. This Project

562 was additionally supported by awards NSF-DEB-0924741 and
563 NSF-DEB-0515981.

Please cite this article in press as: Lovette, I.J., et al. Phylogenetic relationships of the mockingbirds and thrashers (Aves: Mimidae). Mol. Phylogenet. Evol.
(2011), doi:10.1016/j.ympev.2011.07.009
 Appendix A (continued)
(2011), doi:10.1016/j.ympev.2011.07.009
Please cite this article in press as: Lovette, I.J., et al. Phylogenetic relationships of the mockingbirds and thrashers (Aves: Mimidae). Mol. Phylogenet. Evol.

22 August 2011
YMPEV 3988
Taxon Museum sourcea & Typeb Locality Mitochondrial coding genes Nuclear intron loci
sample No.
ND2d CO1 CO2 ATPasese Fib-5 Fib-7 Rho-1 TGFB2-4
Mimus gilvus CUMV-33466 S Mexico, Campeche Pending — — — — — — —
Mimus gilvus CUMV-33464 S Mexico, Yucatan Pending — — — — — — —
Mimus gilvus CUMV-33465 S Mexico, Yucatan Pending — — — — — —
Mimus magnirostris CAB-MMAG1c B Colombia, San Andres I. Pending Pending Pending Pending Pending Pending Pending Pending
Mimus magnirostris CAB-MMAG2c B Colombia, San Andres I. Pending Pending Pending Pending Pending Pending Pending Pending
Mimus gundlachi STRI-JAMGU1 T Jamaica, Portland Ridge EF468198 EF484222 EF484323 EF486782 EF468329 EF471850 EF472862 EF484121
Mimus thenca MNHN-2676 T Chile EF468204 EF484228 EF484329 EF486788 EF468335 EF471856 EF472868 EF484127
Mimus longicaudatus LSUMNS-B5229 T Peru, Lambayeque EF468200 EF484224 EF484325 EF486784 EF468331 EF471852 EF472864 EF484123
Mimus longicaudatus LSUMNS-B3810 T Peru, Ica Pending Pending Pending Pending Pending Pending Pending Pending
Mimus saturninus CUMV-50582[009] T Argentina, Buenos Aires Prov. EF468203 EF484227 EF484328 EF486787 EF468334 EF471855 EF472867 EF484126
Mimus saturninus CUMV-50581[008] T Argentina, Buenos Aires Prov. Pending Pending Pending Pending Pending Pending Pending Pending
Mimus patagonicus CUMV-50577[004] T Argentina, Jujuy Prov. EF468201 EF484225 EF484326 EF486785 EF468332 EF471853 EF472865 EF484124
Mimus patagonicus CUMV-50578[001] T Argentina, Jujuy Prov. Pending Pending Pending Pending Pending Pending Pending Pending

I.J. Lovette et al. / Molecular Phylogenetics and Evolution xxx (2011) xxx–xxx
Mimus triurus CUMV-MACH14 T Argentina, Buenos Aires Prov. EF468205 EF484229 EF484330 EF486789 EF468336 EF471857 EF472869 EF484128
Mimus dorsalis CUMV-50579[002] T Argentina, Jujuy Prov. Pending Pending Pending Pending Pending Pending Pending Pending
Mimus dorsalis CUMV-50580[006] T Argentina, Jujuy Prov. Pending Pending Pending Pending Pending Pending Pending Pending
Mimus parvulus WFU-Isab02d B Ecuador, Galapagos, Isabella I. AY311553 DQ083935 — DQ087201 Pending Pending Pending Pending
Mimus parvulus WFU-Scz2d B Ecuador, Galapagos, Santa Cruz I. AY311553 DQ083934 — DQ087200 Pending Pending Pending Pending
Mimus trifasciatus WFU-Ntr03d B Ecuador, Galapagos, Champion I. AY311544 DQ083929 — DQ087195 Pending Pending Pending Pending
Mimus macdonaldi WFU-Nma002d B Ecuador, Galapagos, Española I. AY311555 DQ083930 — DQ087196 Pending Pending Pending Pending
Nesomimus melanotis WFU-Scr01d B Ecuador, Galapagos, San AY311576 DQ083931 — DQ087197 Pending Pending Pending Pending
Cristóbal I.
Oreoscoptes montanus LSUMNS-B19513 T USA, California, San Bernardino EF468206 EF484230 EF484331 EF486790 EF468337 EF471858 EF472870 EF484129
Co.
Mimus graysoni Barber et al., 2004 Mexico, Isla Socorro AY758199 — — — — — — —
Mimus graysoni CUMV-24114 S Mexico, Isla Socorro Pending Pending Pending Pending — — — —
Toxostoma rufum LSUMNS-B0490 T USA, Louisiana, Cameron Parish EF468209 EF484233 EF484334 EF486793 EF468340 EF471861 EF472873 EF484132
Toxostoma rufum CUMV-50815 T USA, New York, Tompkins Co. Pending Pending Pending Pending Pending Pending Pending Pending
Toxostoma cinereum LSUMNS-B6745 T Mexico, Baja California EF468208 EF484232 EF484333 EF486792 EF468339 EF471860 EF472872 EF484131
Toxostoma curvirostre BMNH-CBTH024 T Mexico, San Lius Potosi Pending Pending Pending Pending — — — —
Toxostoma curvirostre BMNH-CBTH051 T Mexico, Chihuahua Pending Pending Pending Pending — — — —
Toxostoma longirostre BMNH-BTH2TX T USA, Texas, Atascosa Co. Pending Pending Pending Pending Pending Pending Pending Pending
Toxostoma longirostre BMNH-BTH4TX T USA, Texas, Atascosa Co. Pending Pending Pending Pending Pending Pending Pending Pending
Toxostoma guttatum CUMV-30516 S Mexico, Quintana Roo, Cozumel I. Pending Pending Pending Pending — — — —
Toxostoma ocellatum CUMV-29035 S Mexico, Quintana Roo, Cozumel I. Pending — — — — — — —
Toxostoma bendirei BMNH-BETH4 T USA, Arizona, Pima Co. Pending Pending Pending Pending Pending — Pending Pending
Toxostoma bendirei BMNH-BETH5 T USA, Arizona, Pima Co. Pending Pending Pending Pending Pending Pending Pending Pending
Toxostoma crissale BMNH-CRTH2AZ T USA, Arizona, Cochise Co. Pending Pending Pending Pending Pending Pending Pending Pending
Toxostoma crissale BMNH-CRTH6CA T USA, California, San Bernardino Pending Pending Pending Pending Pending Pending Pending Pending
Co.
Toxostoma lecontei BMNH-LETH7AZ T USA, Arizona, La Paz Co. Pending Pending Pending Pending Pending Pending Pending Pending
Toxostoma lecontei BMNH-LETH8AZ T USA, Arizona, La Paz Co. Pending Pending Pending Pending Pending — Pending Pending
Toxostoma redivivium SDSU-2311 T USA, California, Lake Co. Pending Pending Pending Pending Pending Pending Pending Pending

No. of Pages 11, Model 5G

Toxostoma redivivium SDSU-2318 T USA, California, Tuolumne Co. Pending Pending Pending Pending Pending Pending Pending Pending
Toxostoma redivivium SDSU-2375 T USA, California, San Diego Co. Pending Pending Pending Pending Pending Pending Pending Pending
Toxostoma redivivium SDSU-2378 T USA, California, San Diego Co. Pending Pending Pending Pending Pending Pending Pending Pending
Ramphocinclus STRI-SLRBR2 T St. Lucia EF468207 EF484231 EF484332 EF486791 EF468338 EF471859 EF472871 EF484130
brachyurus
Ramphocinclus STRI-SLRBR1 T St. Lucia Pending — — — — — — —
brachyurus
Melanotis LSUMNS-B0022 T Mexico, Puebla EF468193 EF484217 EF484318 EF486777 EF468324 EF471845 EF472857 EF484116
caerulescens
Melanotis LSUMNS-B0048[18] T Mexico, Puebla Pending Pending Pending Pending Pending Pending Pending Pending

(continued on next page)

9
 YMPEV 3988 No. of Pages 11, Model 5G
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10 I.J. Lovette et al. / Molecular Phylogenetics and Evolution xxx (2011) xxx–xxx

Institutional sources of samples: ANSP: Academy of Natural Sciences, Philadelphia, Pennsylvania, USA; BMNH: Bell Museum of Natural History, University of Minnesota, Minnneapolis, Minnesota, USA; CUMV: Cornell
University Museum of Vertebrates, Ithaca, New York, USA; LSUMNS: Louisiana State University Museum of Natural Science, Baton Rouge, Louisiana, USA; MNHN: Museo Nacional de Historia Natural, Santiago, Chile; SDSU: San
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Please cite this article in press as: Lovette, I.J., et al. Phylogenetic relationships of the mockingbirds and thrashers (Aves: Mimidae). Mol. Phylogenet. Evol.
(2011), doi:10.1016/j.ympev.2011.07.009