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Sys tematics of theOsteocephalus buckleyi species complex... 1

Systematics of the Osteocephalus buckleyi speciescomplex (Anura, Hylidae) from Ecuador and Peru

Santiago R. Ron1,, Pablo J. Venegas2,, Eduardo oral1,3,, Morley Read1,| ,Diego A. Ortiz1,, Andrea L. Manzano1,4,#

1 Museo de Zoologa, Escuela de Biologa, Ponticia Universidad Catlica del Ecuador, Av. 12 de Octubre yRoca, Aptdo. 17-01-2184, Quito, Ecuador2 Divisin de Herpetologa-Centro de Ornitologa y Biodiversidad

(CORBIDI), Santa Rita N105 Of. 202, Urb. Huertos de San Antonio, Surco, Lima, Per3 Current address:Facultad de Ciencias Ambientales, Universidad Internacional SEK, Quito, Ecuador4 Current address: BiologyDepartment, HH227, San Francisco State University, 1600 Holloway Avenue, San Francisco, CA 94132, USA

urn:lsid:zoobank.org:author:ACF9C463-F771-459C-B22B-AF6B9902DF57 urn:lsid:zoobank.org:author:15AD03E1-9ACF-4F38-AA96-09A5A56A3DC4 urn:lsid:zoobank.org:author:1A4F31AF-5629-4EE5-83DD-1EBB93139A5B | urn:lsid:zoobank.org:author:10D75453-75A0-49C7-B577-A7687398CEF0

urn:lsid:zoobank.org:author:51147772-F315-412E-BA89-B8990CA49544 # urn:lsid:zoobank.org:author:7C29ADA6-00FB-45DC-9A5B-7B9A217B71F6

Corresponding author: Santiago R. Ron ([email protected])

Academic editor: F. Andreone | Received 24 June 2012 | Accepted 5 October 2012 | Published 18 October 2012

urn:lsid:zoobank.org:pub:163872F0-BE66-436C-9663-EE2226C358AB

Citation: Ron SR, Venegas PJ, oral E, Read M, Ortiz DA, Manzano AL (2012) Systematics of theOsteocephalus buckleyispecies complex (Anura, Hylidae) from Ecuador and Peru. ZooKeys 229: 152.doi: 10.3897/zookeys.229.3580

Abstract We present a new phylogeny, based on DNA sequences of mitochondrial and nuclear genes, for frogsthe genusOsteocephalus with emphasis in theOsteocephalus buckleyi species complex. Genetic, morpho-logic, and advertisement call data are combined to dene species boundaries and describe new speciTe phylogeny shows strong support for: (1) a basal position ofO. taurinus + O. oophagus , (2) a cladecontaining phytotelmata breeding species, and (3) a clade that corresponds to theO. buckleyi species com-plex. Our results document a large proportion of hidden diversity within a set of populations that wepreviously treated as a single, widely distributed species,O. buckleyi . Individuals assignable toO. buckleyi formed a paraphyletic group relative toO. verruciger and O. cabrerai and contained four species, one ofwhich isO. buckleyi sensu stricto and three are new. wo of the new species are shared between Ecuador andPeru (O. vilmae sp. n. andO. cannatellai sp. n.) and one is distributed in the Amazon region of southernPeru (O. germani sp. n.). We discuss the difficulties of using morphological characters to dene speciesboundaries and propose a hypothesis to explain them.

ZooKeys 229: 152 (2012)

doi: 10.3897/zookeys.229.3580

www.zookeys.org

CopyrightSantiago R. Ron et al. This is an open access article distributed under the terms of theCreative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

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Santiago R. Ron et al. / ZooKeys 229: 152 (2012)2

ResumenPresentamos una nueva logenia, basada en secuencias de ADN de genes nucleares y mitocondrialpara ranas del gneroOsteocephalus con nfasis en el complejo de especiesOsteocephalus buckleyi . Datosgenticos, morfolgicos, y de cantos de anuncio se combinan para denir lmites de especies y descrnuevas especies. La logenia muestra un soporte fuerte para: (1) una posicin basal deO. taurinus +O.oophagus , (2) un clado que contiene especies con reproduccin en totelmatas, y (3) un clado que cor-responde el complejo de especiesO. buckleyi . Nuestros resultados documentan una gran proporcin dediversidad escondida dentro de un grupo de poblaciones que previamente haban sido tratadas comuna sola especie ampliamente distribuida,O. buckleyi . Los individuos asignables aO. buckleyi formaronun grupo paraltico en relacin aO. verruciger y O. cabrerai y contuvieron cuatro especies, una de lascuales esO. buckleyi sensu scricto y tres son nuevas. Dos de las nuevas especies estn compartidas entreEcuador y Per (O. vilmae sp. n. yO. cannatellai sp. n.) y una est en el sur de Per (O. germani sp. n.)Discutimos las dicultades de usar caracteres morfolgicos para denir lmites de especies y proponeuna hiptesis para explicarla.

Keywords Advertisement calls, Amazon, Anura, Cryptic species, Morphology,Osteocephalus buckleyi , Phylogeny

Introduction

Te Upper Amazon region has the highest alpha diversity of amphibians in the Worldwith several sites exceeding 100 species in less than 10 km2 (Bass et al. 2010). Re-markably, these gures may vastly underestimate the total diversity as shown by tdiscovery of large numbers of cryptic species with the use of genetic markers (eFouquet et al. 2007; Funk et al. 2011; Padial and De la Riva 2009; Ron et al. 2006)Tese preliminary efforts suggest that the use of genetic characters is crucial to attaia complete understanding of the diversity and evolutionary history of Amazoniaamphibians. Tis necessity is particularly pressing in widespread taxa with pervasitaxonomic problems.

One such group isOsteocephalus , a genus of hylid frogs widely distributed in the

Amazon Basin, Guianas and upper drainages of the Magdalena and Orinoco rive(Frost 2010).Osteocephalus are arboreal and nocturnal frogs with reproduction modesvarying from deposition of eggs in lentic water and exotrophic tadpoles to depositiof eggs in bromeliads and oophagus tadpoles and biparental care (Crump 1974; Jungfer and Weygoldt 1999). Tere are 24 described species and reports of undescribedspecies are frequent (e.g., Jungfer 2010; Moravec et al. 2009; Ron et al. 2010). Tere only one formally dened species group withinOsteocephalus , theO. buckleyi complex.It was rst proposed by Cochran and Goin (1970) to allocateO. buckleyi(Boulenger1882),O. pearsoni(Gaige 1929), andO. cabrerai (Cochran and Goin 1970). Its rst

large scale review was carried out by rueb and Duellman (1971) who examined tmorphology of specimens from seven countries and concluded that theO. buckleyi complex (excludingO. verruciger Werner 1901) consisted of a single, morphologi-


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Systematics of theOsteocephalus buckleyi species complex... 3

cally variable and widely distributed species. Tey synonymizedO. cabrerai , O. carri(Cochran and Goin 1970), andO. festae (Peracca 1904) underO. buckleyi.Te threespecies have been subsequently resurrected (Duellman and Mendelson 1995; Jungf2010; Lynch 2006). Recent reviews (Jungfer 2010; 2011; Moravec et al. 2009; Roet al. 2010) imply that theO. buckleyi species complex consists of nine species:O.buckleyi , O. cabrerai , O. carri , O. duellmani Jungfer 2011,O. festae , O. inframaculatus(Boulenger 1882),O. mutabor Jungfer and Hdl 2002,O. verruciger and an unde-scribed species sister toO. verruciger . A phylogeny based on mitochondrial DNA re-vealed strong support for theO. buckleyi complex as well as paraphyly inO. verruciger andO. buckleyi (Ron et al. 2010).

Despite recent contributions to the taxonomy of the group (e.g., Jungfer 2010;2011) theO. buckleyi species complex still contains undescribed species as well as alphtaxonomic problems (Jungfer 2010; Ron et al. 2010) which attest the difficulties ocorrectly identifying species boundaries on the basis of morphological evidence aloHerein we integrate genetic, morphological and advertisement call data to assess tphylogenetic relationships and species boundaries among populations of theO. buck-leyi complex from Ecuador and Peru. Te results demonstrate the existence of threenew species, which are formally described here.

Methods

For ease of comparison, we generally follow the format of rueb and Duellma(1971) for diagnosis and description. Morphological terminology and abbreviations follow Lynch and Duellman (1997). Notation for hand and foot webbing isbased on Myers and Duellman (1982). Sex was determined by the texture of dorsskin, the presence of nuptial pads or vocal sac folds, and by gonadal inspectioSpecimens were xed in 10% formalin and preserved in 70% ethanol. Snout-venlength is abbreviated as SVL. Examined specimens (listed in the type-series a Appendix I) are housed at the collection of the Divisin de Herpetologa, Centr

de Ornitologa y Biodiversidad (CORBIDI), Herpetology Collection at EscuelPolitcnica Nacional (EPN-H), Museo de Historia Natural at Universidad SanMarcos (MUSM), Museo de Zoologa at Ponticia Universidad Catlica del Ecuador (QCAZ), and Natural History Museum (BMNH). Te pencil drawing of theholotype ofO. cannatella sp. n. was made using a Wild Heerbrugg M3B 10/21stereo microscope equipped with a camera lucida.

Principal Components Analysis (PCA) and Discriminant Function Analysis (DFAwere used to assess the degree of morphometric differentiation between species. Owell preserved specimens (Simmons 2002) were measured for the following eight m

phological variables, following Duellman (1970): (1) SVL; (2) head length; (3) hewidth; (4) tympanum diameter; (5) femur length; (6) tibia length; (7) foot lengthand (8) eye diameter. All variables were log-transformed. o remove the effect of


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variation with SVL, the PCA was applied to the residuals from the linear regressiobetween the seven measured variables and SVL. We applied a multivariate analysisvariance (MANOVA) to tests for morphometric differences between sexes. Becausefound signicant differences inO. buckleyi , the PCA and DFA were applied on each sexseparately. For the PCA, only components with eigenvalues > 1 were retained. Te DFAwas applied to the measured variables without size correction because we wanted tosess discriminability among species based on all the variables, including SVL. Samsizes are:O. buckleyi 24 males, 3 females; O. cabrerai 7 males;O. cannatellai sp. n. 33males, 3 females;O. festae 7 males, 18 females;O. germani sp. n. 2 males, 5 females;O.verruciger 22 males, 5 females; andO. vilmaesp. n. 4 males. Both PCA and DFA wereconducted in JMP 8.01 (SAS Institute 2008). Measurements were made using digcalipers (to the nearest 0.01 mm).

Advertisement calls recordings were made with a Sennheiser ME-67 directiomicrophone with digital recorder Olympus LS10. Calls were analyzed using softwRaven 1.2.1 (Charif et al. 2004) at a sampling frequency of 22.1 kHz and a frequencresolution of 21.5 Hz. Calls consist of two components, the rst is a rattle note anthe second is a quack note. Measured call variables are: (1) call rate: number of caper second, (2) dominant frequency: frequency with the most energy, measured alonall the call, (3) duration of rst component note: time from the beginning to the endof note, (4) duration of second component: time from beginning of rst quack to thend of the last, (5) rst component interval: time from the end of last note of the rcomponent to the beginning of the rst note of the second component, (6) number opulses: number of pulses in a rst component note, (7) pulse rate: number of pulseduration of rst component note, (8) duration of second component note: durationfrom beginning to end of a single quack, (9) quack rate: number of quacks/duratioof second component. If available, several calls or notes were analyzed per individto calculate an individual average. Original recordings are deposited in the audarchive of the QCAZ and are available through the AmphibiaWebEcuador websi(http://zoologia.puce.edu.ec/vertebrados/anbios/).

DNA extraction, amplication, and sequencing

otal DNA was extracted from muscle or liver tissue preserved in 95% ethanol or tsue storage buffer using standard phenolchloroform extraction protocols (Sambroet al. 1989). Polymerase chain reaction (PCR) was used to amplify the mitochondrigenes 12S rRNA ,16S rRNA, ND1 (with anking tRNA genes), CO1, and controregion. We amplied one DNA fragment for 12S, CO1, and the control region andone or two overlapping fragments for the last ~320 bp of 16S and the adjacent ND

using primers listed in Goebel et al. (1999) and Moen and Wiens (2009). We also amplied the nuclear genePOMC as a single fragment using primers listed by Wiens et al.(2005). PCR amplication was carried under standard protocols. Amplied producwere sequenced by the Macrogen Sequencing eam (Macrogen Inc., Seoul, Korea).
http://zoologia.puce.edu.ec/vertebrados/anfibioshttp://zoologia.puce.edu.ec/vertebrados/anfibios


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Phylogenetic analyses

We estimated phylogenetic relations between species ofOsteocephalus based on newly gen-erated sequence data for ve mitochondrial (12S RNA,CO1, 16S , ND1, control region)and one nuclear gene (POMC ) for a total of up to 4170 bp. o expand the species sam-pling, we also included sequences from GenBank. All samples are listed in able 1. the outgroup, we included one sample ofTrachycephalus jordaniand one ofT. typhonius(based on Faivovich et al. 2005 and Wiens et al. 2010). Te completeness of the sequences varied considerably among individuals (specially for samples from GenBank whtypically lacked three or more loci). Nevertheless, we included samples with missing because analyses of both empirical and simulated matrices have shown that taxa wmissing sequences can be accurately placed in model-based phylogenetic analyses ifnumber of characters is large, as in our matrix (for a review see Wiens and Morrill 201

Preliminary sequence alignment was done with MAFF 6.814b software with the INS-i algorithm (Katoh et al. 2002). Te sequence matrix was imported to Mesquite (version 2.72; Maddison and Maddison 2009) and the ambiguously aligned regions were a justed manually to produce a parsimonious alignment (i.e., informative sites minimizeIn protein coding loci, DNA sequences were translated to amino acids with Mesquite aid the manual alignment. Phylogenetic trees were obtained using Bayesian inference.

Because our dataset includes several loci, it is unlikely that it ts a single modelnucleotide substitution. Tus, we partitioned the data to analyze each partition undera separate model. Te best model for each partition was chosen with JModel est version 0.1.1 (Posada 2008) using the Akaike Information Criterion with sample sizcorrection as optimality measure. We also evaluated three different partition strategi(i) a single partition, (ii) six partitions (one per loci), and (iii) twelve partitions (one each codon position in protein coding loci plus one for each non protein coding lociTe best partition strategy was chosen by estimating Bayes factors using a threshold 10 as evidence in favor of the more complex partition (Brandley et al. 2005).

Each Bayesian analysis consisted of two parallel runs of the Metropolis couplMonte Carlo Markov chain for 5 106 generations. Each run had four chains with a

temperature of 0.05. Te prior for the rate matrix was a uniform dirichlet and all to-pologies were equally probable a priori. Convergence into a stationary distribution wdetermined by reaching average standard deviation split frequencies < 0.05 betweruns. We also used software racer ver. 1.5 (Rambaut and Drummond 2007) to visually inspect convergence and stationarity of the runs. Te rst 50% of the sampledgenerations were discarded as burn-in and the remaining were used to estimate thBayesian tree, posterior probabilities and other model parameters. Phylogenetic anases were carried out in MrBayes 3.2.1 (Ronquist et al. 2012).

Because the only nuclear gene analyzed had low variability and few informati

sites, it was concatenated to the mitochondrial genes into a single matrix. We reconize the advantages of species-tree methods (e.g., Edwards et al. 2007) but could nuse them given the insufficient number of nuclear genes sampled. We encourage tapplication of those methodologies in future phylogenetic inferences inOsteocephalus .


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nih.gov/nuccore/JX847063http://www.ncbi.nlm.nih.gov/nuccore/JX875599http://www.ncbi.nlm.nih.gov/nuccore/JX875725http://www.ncbi.nlm.nih.gov/nuccore/JX875650http://www.ncbi.nlm.nih.gov/nuccore/JX875801http://www.ncbi.nlm.nih.gov/nuccore/JX847062http://www.ncbi.nlm.nih.gov/nuccore/JX875598http://www.ncbi.nlm.nih.gov/nuccore/JX875799http://www.ncbi.nlm.nih.gov/nuccore/JX875703http://www.ncbi.nlm.nih.gov/nuccore/JX875865http://www.ncbi.nlm.nih.gov/nuccore/JX847106http://www.ncbi.nlm.nih.gov/nuccore/JX875640http://www.ncbi.nlm.nih.gov/nuccore/JX875798http://www.ncbi.nlm.nih.gov/nuccore/JX875702http://www.ncbi.nlm.nih.gov/nuccore/JX875864http://www.ncbi.nlm.nih.gov/nuccore/JX847105http://www.ncbi.nlm.nih.gov/nuccore/JX875639http://www.ncbi.nlm.nih.gov/nuccore/JX875793http://www.ncbi.nlm.nih.gov/nuccore/JX875724http://www.ncbi.nlm.nih.gov/nuccore/JX875858http://www.ncbi.nlm.nih.gov/nuccore/JX847099http://www.ncbi.nlm.nih.gov/nuccore/JX875633http://www.ncbi.nlm.nih.gov/nuccore/JX875784http://www.ncbi.nlm.nih.gov/nuccore/JX875694http://www.ncbi.nlm.nih.gov/nuccore/JX875850http://www.ncbi.nlm.nih.gov/nuccore/JX875629http://www.ncbi.nlm.nih.gov/nuccore/JX875783http://www.ncbi.nlm.nih.gov/nuccore/JX875672http://www.ncbi.nlm.nih.gov/nuccore/JX875849http://www.ncbi.nlm.nih.gov/nuccore/JX847095http://www.ncbi.nlm.nih.gov/nuccore/JX875628http://www.ncbi.nlm.nih.gov/nuccore/JX875782http://www.ncbi.nlm.nih.gov/nuccore/JX875714http://www.ncbi.nlm.nih.gov/nuccore/JX875848http://www.ncbi.nlm.nih.gov/nuccore/JX847094http://www.ncbi.nlm.nih.gov/nuccore/JX875627http://www.ncbi.nlm.nih.gov/nuccore/JX875778http://www.ncbi.nlm.nih.gov/nuccore/JX875722http://www.ncbi.nlm.nih.gov/nuccore/JX875845http://www.ncbi.nlm.nih.gov/nuccore/JX847092http://www.ncbi.nlm.nih.gov/nuccore/JX875625http://www.ncbi.nlm.nih.gov/nuccore/JX875765http://www.ncbi.nlm.nih.gov/nuccore/JX875689http://www.ncbi.nlm.nih.gov/nuccore/JX875833http://www.ncbi.nlm.nih.gov/nuccore/JX847087http://www.ncbi.nlm.nih.gov/nuccore/JX875618http://www.ncbi.nlm.nih.gov/nuccore/JX875764http://www.ncbi.nlm.nih.gov/nuccore/JX875677http://www.ncbi.nlm.nih.gov/nuccore/JX875832http://www.ncbi.nlm.nih.gov/nuccore/HQ600640http://www.ncbi.nlm.nih.gov/nuccore/HQ600607http://www.ncbi.nlm.nih.gov/nuccore/JX875763http://www.ncbi.nlm.nih.gov/nuccore/JX875720http://www.ncbi.nlm.nih.gov/nuccore/JX875831http://www.ncbi.nlm.nih.gov/nuccore/HQ600639http://www.ncbi.nlm.nih.gov/nuccore/HQ600606http://www.ncbi.nlm.nih.gov/nuccore/JX875754http://www.ncbi.nlm.nih.gov/nuccore/JX875686http://www.ncbi.nlm.nih.gov/nuccore/JX875822http://www.ncbi.nlm.nih.gov/nuccore/HQ600634http://www.ncbi.nlm.nih.gov/nuccore/HQ600601http://www.ncbi.nlm.nih.gov/nuccore/JX875753http://www.ncbi.nlm.nih.gov/nuccore/JX875708http://www.ncbi.nlm.nih.gov/nuccore/JX875821http://www.ncbi.nlm.nih.gov/nuccore/HQ600633http://www.ncbi.nlm.nih.gov/nuccore/HQ600600http://www.ncbi.nlm.nih.gov/nuccore/JX875742http://www.ncbi.nlm.nih.gov/nuccore/JX875718http://www.ncbi.nlm.nih.gov/nuccore/JX875812http://www.ncbi.nlm.nih.gov/nuccore/JX847081http://www.ncbi.nlm.nih.gov/nuccore/JX875611http://www.ncbi.nlm.nih.gov/nuccore/DQ380378http://www.ncbi.nlm.nih.gov/nuccore/EU034082http://www.ncbi.nlm.nih.gov/nuccore/JX875737http://www.ncbi.nlm.nih.gov/nuccore/JX847070http://www.ncbi.nlm.nih.gov/nuccore/JX875736http://www.ncbi.nlm.nih.gov/nuccore/JX875657http://www.ncbi.nlm.nih.gov/nuccore/JX875808http://www.ncbi.nlm.nih.gov/nuccore/JX847069http://www.ncbi.nlm.nih.gov/nuccore/JX875608http://www.ncbi.nlm.nih.gov/nuccore/JX875735http://www.ncbi.nlm.nih.gov/nuccore/JX875807http://www.ncbi.nlm.nih.gov/nuccore/JX847068http://www.ncbi.nlm.nih.gov/nuccore/JX875607http://www.ncbi.nlm.nih.gov/nuccore/JX875734http://www.ncbi.nlm.nih.gov/nuccore/JX875806http://www.ncbi.nlm.nih.gov/nuccore/JX847067http://www.ncbi.nlm.nih.gov/nuccore/JX875606http://www.ncbi.nlm.nih.gov/nuccore/EU034116http://www.ncbi.nlm.nih.gov/nuccore/DQ380378http://www.ncbi.nlm.nih.gov/nuccore/EU034082http://www.ncbi.nlm.nih.gov/nuccore/JX875744http://www.ncbi.nlm.nih.gov/nuccore/JX875680http://www.ncbi.nlm.nih.gov/nuccore/HQ600629http://www.ncbi.nlm.nih.gov/nuccore/HQ600596http://www.ncbi.nlm.nih.gov/nuccore/EU034081


7/52


M u s e u m

N o .

S p e c i e s

G e n b a n k A c c e s s i o n N o .

R e f e r e n c e

1 6 S - N D 1

1 2 S

C O 1

C o n t r o l R e g i o n

P O M C

C O R B I D I 1 2 0

O . c

a b r e r a

i

J X 8 7 5 6 0 0

- -

- -

J X 8 7 5 6 5 1

J X 8 7 5 7 2 7 T i s s t u d y

C O R B I D I 5 8 1 9

O . c

a b r e r a

i

J X 8 7 5 6 0 4

J X 8 4 7 0 6 6

J X 8 7 5 8 0 4

J X 8 7 5 6 5 5

J X 8 7 5 7 3 1 T i s s t u d y

C O R B I D I 5 8 2 1

O . c

a b r e r a

i

J X 8 7 5 6 0 5

- -

J X 8 7 5 8 0 5

J X 8 7 5 6 5 6

J X 8 7 5 7 3 2 T i s s t u d y

L S U M Z H - 1

3 7 2 0

O . c

a b r e r a

i

- -

A Y 8 4 3 7 0 5

- -

- -

- -

F a i v o v i c h e t a l . 2 0 0 5

Q C A Z 2 7 9 2 3

O . c

a b r e r a

i

J X 8 7 5 6 1 7

J X 8 4 7 0 8 6

J X 8 7 5 8 2 7

J X 8 7 5 7 0 9

J X 8 7 5 7 6 0 T i s s t u d y

Q C A Z 2 8 2 3 1

O . c

a b r e r a

i

H Q 6 0 0 6 2 1 H Q 6 0 0 6 5 4 J X 8 7 5 8 3 0

J X 8 7 5 7 1 0


C O R B I D I 9 3 6 8

O . c

a n n a t e

l l a i

- -

J X 8 4 7 0 7 2

- -

J X 8 7 5 6 5 8

- -

T i s s t u d y

C O R B I D I 9 3 7 0

O . c

a n n a t e

l l a i

J X 8 7 5 6 4 3

J X 8 4 7 0 7 4

- -

J X 8 7 5 6 6 0

- -

T i s s t u d y

C O R B I D I 9 3 9 4

O . c

a n n a t e

l l a i

J X 8 7 5 6 4 4

J X 8 4 7 0 7 5

- -

J X 8 7 5 6 6 1

- -

T i s s t u d y

C O R B I D I 9 5 0 7

O . c

a n n a t e

l l a i

J X 8 7 5 6 4 5

J X 8 4 7 0 7 7

- -

J X 8 7 5 6 6 2

- -

T i s s t u d y

Q C A Z 2 5 4 6 9

O . c

a n n a t e

l l a i

H Q 6 0 0 6 1 7 H Q 6 0 0 6 5 0 J X 8 7 5 8 2 3

J X 8 7 5 6 8 7


Q C A Z 3 1 0 1 6

O . c

a n n a t e

l l a i

J X 8 7 5 6 2 1

J X 8 4 7 0 8 9

J X 8 7 5 8 3 9

J X 8 7 5 7 1 2

J X 8 7 5 7 7 1 T i s s t u d y

Q C A Z 3 1 0 3 2

O . c

a n n a t e

l l a i

J X 8 7 5 6 2 2

J X 8 4 7 0 9 0

J X 8 7 5 8 4 0

J X 8 7 5 6 9 1

J X 8 7 5 7 7 2 T i s s t u d y

Q C A Z 3 1 0 3 3

O . c

a n n a t e

l l a i

J X 8 7 5 6 2 3

- -

J X 8 7 5 8 4 1

J X 8 7 5 6 6 8

J X 8 7 5 7 7 3 T i s s t u d y

Q C A Z 3 2 5 0 6

O . c

a n n a t e

l l a i

H Q 6 0 0 6 1 8 H Q 6 0 0 6 5 1 J X 8 7 5 8 4 3

J X 8 7 5 6 9 2


Q C A Z 3 2 5 0 8

O . c

a n n a t e

l l a i

H Q 6 0 0 6 1 9 H Q 6 0 0 6 5 2 J X 8 7 5 8 4 4

J X 8 7 5 6 9 3


Q C A Z 3 7 1 7 5

O . c

a n n a t e

l l a i

H Q 6 0 0 6 2 0 H Q 6 0 0 6 5 3 J X 8 7 5 8 4 6

J X 8 7 5 7 1 3


Q C A Z 3 9 6 3 3

O . c

a n n a t e

l l a i

J X 8 7 5 6 3 0

J X 8 4 7 0 9 6

J X 8 7 5 8 5 2

J X 8 7 5 6 7 8

J X 8 7 5 7 8 6 T i s s t u d y

Q C A Z 4 0 2 5 8

O . c

a n n a t e

l l a i

J X 8 7 5 6 3 1

J X 8 4 7 0 9 7

J X 8 7 5 8 5 4

J X 8 7 5 6 9 6

J X 8 7 5 7 8 8 T i s s t u d y

Q C A Z 4 5 9 0 9

O . c

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l l a i

J X 8 7 5 6 3 5

J X 8 4 7 1 0 1

J X 8 7 5 8 6 0

J X 8 7 5 7 0 1

J X 8 7 5 7 9 5 T i s s t u d y

Q C A Z 4 6 4 7 2

O . c

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J X 8 7 5 6 3 8

J X 8 4 7 1 0 4

J X 8 7 5 8 6 3

J X 8 7 5 7 1 7

J X 8 7 5 7 9 7 T i s s t u d y

Q C A Z 4 9 5 7 2

O . c

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J X 8 7 5 6 4 1

J X 8 4 7 1 0 7

J X 8 7 5 8 6 6

J X 8 7 5 6 7 4

J X 8 7 5 8 0 0 T i s s t u d y

C B F 6 0 5 1

O . c

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F J 9 6 5 3 0 0

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M o r a v e c e t a l . 2 0 0 9
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