Per G P Ericson
email: per.ericson@nrm.se

Storrs L Olson

Martin Irestedt

Herculano Alvarenga

Jon Fjeldså

Circumscription of a monophyletic family for the tapaculos (Aves: Rhinocryptidae): Psiloramphus in and Melanopareia out

The tapaculos (Rhinocryptidae) are tracheophone, suboscine birds restricted to South and Central America. Most tapaculos share a number of internal and external characteristics that have been used to define the family taxonomically. The genera Melanopareia and Psiloramphus do not fully fit this pattern and have caused considerable dispute among taxonomists since they were first described. In this paper we delimit the systematic boundaries of the tapaculos and assess their generic relationships by an analysis of molecular sequence data. The results show that while Psiloramphus is nested well within the Rhinocryptidae, Melanopareia falls far outside that clade. A new family is erected for Melanopareia.

INTRODUCTION

The impudently named tapaculos (Rhinocryptidae) are a small group of tracheophone, suboscine passerines whose greatest generic diversity is in southern South America.

Most are large-footed, strong-legged ground birds reminiscent of some of the grounddwelling antthrushes (Formicariidae), with which they were often associated. The family is generally well defined by the presence of operculate nostrils, a tracheophone syrinx, a somewhat curved humerus and a four-notched sternum [START_REF] Ames | The morphology of the syrinx in passerine birds[END_REF][START_REF] Feduccia | Morphological similarities between the Menurae and the Rhinocryptidae, relict passerine birds of the Southern Hemisphere[END_REF][START_REF] Maurício | Hidden genetic diversity in Neotropical birds: molecular and anatomical data support a new genus for the "Scytalopus" indigoticus species-group (Aves: Rhinocryptidae)[END_REF]. Thus, Krabbe and Schulenberg (2003: 748) considered that the "tapaculos constitute a well-knit group the members of which are united by several derived characters. Only the genera Melanopareia and Psiloramphus differ to such a degree that their systematic position as tapaculos could be disputed." Furthermore, the phylogenetic relationships within the family Rhinocryptidae are poorly known, particularly regarding the placement of Liosceles and Acropternis, and the large austral species of Pteroptochos and Scelorchilus.

The early taxonomic history of the group was ably summarized by [START_REF] Sclater | On the Neotropical species of the family Pteroptochidae[END_REF]. d"Orbigny (1837) first erected a family "Rhinomyidaeae" (sic., p. 192) for Pteroptochos and his Rhinomya (= Rhinocrypta) using the operculate nostril to separate them from the Formicariidae. However, there was no coherent understanding of the suboscine groups until their distinction from the oscines was established by the pioneering work of [START_REF] Müller | Über die bisher unbekannten typischen Verschiedenheiten der Stimmorgane der Passerini[END_REF] on the syrinx, Müller showed that Scytalopus was a tracheophone suboscine and not a wren (Troglodytidae) and also that Scytalopus and Pteroptochus differed from other known passerines in having a 4-notched sternum. [START_REF] Sclater | On the Neotropical species of the family Pteroptochidae[END_REF] "Pteroptochidae" comprised Scytalopus (including the type species of what later became Myornis [START_REF] Chapman | The more northern species of the genus Scytalopus Gould[END_REF], Merulaxis, Liosceles, Pteroptochos (including the type species of what later became Teledromas Wetmore and [START_REF] Wetmore | A new genus and four new subspecies of American birds[END_REF]Scelorchilus Oberholser, 1923), Rhinocrypta, Hylactes (now included in Pteroptochos), Acropternis, and Triptorhinus (=Eugralla). This composition of the group was essentially maintained until [START_REF] Peters | Check-list of birds of the world[END_REF], except for the problem genera Psiloramphus and Melanopareia.

The three or four species of Melanopareia differ from other tapaculos by their rather slender build and boldly and attractively patterned plumage, and by sharing a semi-concealed white dorsal patch with various true antbirds (Thamnophilidae). They were originally described in the genus Synallaxis (Furnariidae), in which Sclater (1890) later submerged the genus. [START_REF] Salvin | On some new species of birds from western Ecuador[END_REF] described a new species from Ecuador as Formicivora speciosa, duly recognized in that combination by Sclater (1890:251) and others until Hellmayr (1906:334) showed that this was a synonym of Synallaxis elegans Lesson, in which genus Hellmayr continued to place it while regarding Salvin's allocation of it to Formicivora with incredulity. [START_REF] Ridgway | New genera, species, and subspecies of Formicariidae, Furnariidae, and Dendrocolaptidae[END_REF] appears to have overlooked this when he created a new genus Rhoporchilis for Formicivora speciosa. It was [START_REF] Hellmayr | Review of the birds collected by Alcide d'Orbigny[END_REF] who eventually established the modern concept of the genus in showing that Synallaxis elegans, S. torquata, and S. maximiliani were congeneric and would all fall under Reichenbach's earlier generic name Melanopareia and "find their natural place in the Formicariidae", where they stayed for only a few years (e.g. [START_REF] Cory | Catalogue of birds of the Americas and adjacent islands. Part III[END_REF]. Next came the observation of W.D.W. Miller that the sternum of Melanopareia was 4-notchedinformation that was conveyed to and presented by Wetmore (1926: 292). On this basis, [START_REF] Peters | Check-list of birds of the world[END_REF] included Melanopareia in the Rhinocryptidae, where it has resided since.

The other problem species, the Bamboo-wren Psilorhamphus guttatus, is a small bamboo specialist with a rather long, slender bill, a long tail and relatively weak feet, so that it bears little resemblance to large-footed terrestrial tapaculos (Fig. 1).

From the beginning [START_REF] Ménétriés | Monographie de la famille des Myiotherinae où sont décrites les espèces qui ornent le Musée de l'Académie Impériale des Sciences[END_REF], it was placed with the antbirds in the Myiotherinae (=Formicariidae). Sclater (1858:243) associated Psilorhamphus with Ramphocaenus (which is now in the oscine family Polioptilidae) in the Formicariidae with the comment that these genera "might perhaps be more naturally placed as a distinct subfamily of Pteroptochidae [=Rhinocryptidae]" despite the fact that "there is little external difference between the appearance of these birds and the true Wrens [Troglodytidae]". We are not aware, however, of any instance in which Psilorhamphus was placed in either the Troglodytidae or in a family with only sylviid-like genera, as might be inferred from [START_REF] Krabbe | Family Rhinocryptidae (Tapaculos)[END_REF].

Psilorhamphus continued to be associated with Ramphocaenus in the Formicariidaee.g. [START_REF] Sclater | Catalogue of the birds in the British Museum[END_REF] and Cory and Hellmayr (1924:205) although in the latter reference it was noted that W.D.W. Miller would show Psilorhamphus and Ramphocaenus to "constitute a separate family" in "a paper shortly to be published." Peters (1951: 213) later explained that Miller's death prevented publication of his results but that Wetmore (1943:306) had shown Ramphocaenus to have an oscine syrinx, and had told Peters that Microbates likewise was oscine and that he thought based on external morphology that Psilorhamphus was also probably oscine.

Therefore, Peters postponed his treatment of those genera for a future volume treating Sylviidae. [START_REF] Sick | Zur Kenntnis von Ramphocaenus (Sylviidae) und Psilorhamphus (Formicariidae)[END_REF] placed Ramphocaenus in the Sylviidae while provisionally referring Psilorhamphus to the Formicariidae. Then, [START_REF] Plótnick | Posicion [sic] sistemática del género "Psiloramphus" (Rhinocryptidae, Passeriformes)[END_REF] revealed that Psilorhamphus had a 4-notched sternum, a tracheophone syrinx, and had other characters, including an operculate nostril, indicating that it should be placed in the Rhinocryptidae. Thus, in Peters' Checklist Psilorhamphus appears as an addendum to the Rhinocryptidae that appeared in the volume on Sylviidae (Paynter 1964). [START_REF] Heimerdinger | Variation in the sternal notches of suboscine passeriform birds[END_REF] confirmed that the rhinocryptids they examined all had a 4-notched sternum but also showed that this condition obtained in at least two genera of grallarine Formicariidae, which was confirmed by [START_REF] Feduccia | Morphological similarities between the Menurae and the Rhinocryptidae, relict passerine birds of the Southern Hemisphere[END_REF]. [START_REF] Ames | The morphology of the syrinx in passerine birds[END_REF] made a thorough study of the anatomy of the syrinx in passerine birds and examined Melanopareia first-hand but had to rely on the description of [START_REF] Plótnick | Posicion [sic] sistemática del género "Psiloramphus" (Rhinocryptidae, Passeriformes)[END_REF] for Psilorhamphus. Although he noted that the cartilaginous elements of Melanopareia differed from all other tapaculos examined, he found no grounds for excluding either Melanopareia or Psilorhamphus from the Rhinocryptidae. [START_REF] Feduccia | Morphological similarities between the Menurae and the Rhinocryptidae, relict passerine birds of the Southern Hemisphere[END_REF] made the much unexpected discovery that the stapes in Melanopareia was of the primitive oscine type with a flattened footplate, rather than having an expanded, bulbous, fenestrate footplate as in all other suboscine birds. They went on to show other morphological similarities (which is all they ever claimed they were) between some of the Rhinocryptidae and the oscine Menurae (Menura and Atrichornis) of Australia. Although this observation was dismissed on the grounds that the characters involved are either primitive or convergent [START_REF] Krabbe | Family Rhinocryptidae (Tapaculos)[END_REF], the fact remains that the oscines and suboscines had to share a common ancestor and that the ancestor very likely looked like Atrichornis, Melanopareia, or one of the Rhinocryptidae such as Scelorchilus.

This paper aims to delimit the boundaries of the tapaculos and assess generic relationships within the group, supplementing the very detailed ongoing studies of relationships and speciation of small tapaculos (notably Scytalopus; Maurício et al. 2008, Cadena et al. unpublished) by providing a broader phylogenetic framework for the family.

MATERIALS AND METHODS

Taxon sampling, amplification and sequencing

This study includes representatives of all genera traditionally recognized in Rhinocryptidae (e.g. [START_REF] Ridgely | The Birds of South America[END_REF]Tudor 1994, Krabbe and[START_REF] Krabbe | Family Rhinocryptidae (Tapaculos)[END_REF], including representatives of the large genus Scytalopus, including one representative (indigoticus) of the "blue" species, which were recently placed in a separate genus Eleoscytalopus [START_REF] Maurício | Hidden genetic diversity in Neotropical birds: molecular and anatomical data support a new genus for the "Scytalopus" indigoticus species-group (Aves: Rhinocryptidae)[END_REF]. Three of the authors have significant field experience concerning the biology and vocalizations of tapaculos, and this was supplemented with comments and analyses of sound archives by Niels Krabbe (pers. comm.). Representatives of the main lineages within the tracheophone radiation serve as outgroups [START_REF] Ridgely | The Birds of South America[END_REF], Irestedt et al. 2002[START_REF] Krabbe | Family Rhinocryptidae (Tapaculos)[END_REF][START_REF] Chesser | Molecular systematics of New World Suboscine birds[END_REF]).

Three nuclear gene regions, myoglobin intron 2, ornithine decarboxylase (ODC) introns 6 to 7, and glyceraldehyde-3-phosphodehydrogenase (G3PDH) intron 11, were sequenced and used to estimate phylogenetic relationships. For each gene and taxon, multiple sequence fragments were obtained by sequencing with different primers. These sequences were assembled to complete sequences with SEQMAN II™ (DNASTAR Inc.). Positions where the nucleotide could not be determined with certainty were coded with the appropriate IUPAC code. GenBank accession numbers are given in Table 1. See [START_REF] Irestedt | Systematic relationships and biogeography of the tracheophone suboscines (Aves: Passeriformes)[END_REF], [START_REF] Allen | Novel intron phylogeny (ODC) supports plumage convergence in orioles (Icterus)[END_REF][START_REF] Allen | Novel intron phylogeny (ODC) supports plumage convergence in orioles (Icterus)[END_REF][START_REF] Fjeldså | Sapayoa aenigma: a New World representative of "Old World suboscines[END_REF] for extractions, amplifications, and sequencing procedures for fresh tissue/blood samples. Corresponding laboratory procedures for study skins are detailed in [START_REF] Irestedt | Nuclear DNA from old collections of avian study skins reveals the evolutionary history of the Old World suboscines (Aves, Passeriformes)[END_REF].

Phylogenetic inference and model selection

Due to the rather low number of insertions in the introns, the combined sequences could easily be aligned by eye. All gaps have been treated as missing data in the analyses. Bayesian inference (see, e.g., [START_REF] Holder | Phylogeny estimation: Traditional and Bayesian approaches[END_REF][START_REF] Huelsenbeck | Bayesian inference of phylogeny and its impact on evolutionary biology[END_REF]) was used to estimate the phylogenetic relationships. The models for nucleotide substitutions used in the analyses were selected for each gene individually by applying the Akaike Information Criterion (AIC, [START_REF] Akaike | Information theory as an extension of the maximum likelihood principle[END_REF]) and the program MrModeltest 2.2 [START_REF] Nylander | MrModeltest v.2.2. Program distributed by the author[END_REF] in conjunction with PAUP* [START_REF] Swofford | Phylogenetic analysis using parsimony (* and other methods)[END_REF].

Posterior probabilities of trees and parameters in the substitution models were approximated with MCMC and Metropolis coupling using the program MrBayes 3.1.1 [START_REF] Ronquist | MrBAYES 3: Bayesian phylogenetic inference under mixed models[END_REF]. Analyses were performed for both the individual gene partitions and the combined data set. In the analysis of the combined data set, the models selected for the individual gene partition were used. The chains for the individual gene partitions were run for 5 million generations while the chains for the combined data set were run for 10 million generations. Trees were sampled every 100th generations, and the trees sampled during the burn-in phase (i.e., before the chain had reached its apparent target distribution) were then discarded after checking for convergence, final inference was made from the concatenated outputs.

Sequence lengths and alignments

We were able to sequence all three gene regions almost completely for all included taxa (a few sequences miss some base pairs in the 3´ or 5´ends in the myoglobin or the ODC regions, and in the ODC region all sequences obtained from study skins lack a short fragment of 22 bp in the exon 7). Taking into account the missing base pairs, the sequences obtained varied in length between 667-701 bp for the myoglobin intron 2, and between 313-363 bp for the G3PDH intron 11, except for the two antthrushes Chamaeza and Formicarius which contains two large deletions in the G3PDH intron 11 which makes these sequences 251-252 bp long. In the ODC region all Rhinocryptidae and Furnariidae taxa have a large deletion in intron 7 and the sequences from these taxa range between 403 and 500 bp, while the sequences for all other taxa range between 586 and 624 bp.

Most indels observed in the introns were autapomorphic and mainly found in certain variable regions. Some indels vary in length between taxa, which makes it difficult to know if these indels are homologous or represent independent evolutionary events. Several apparently synapomorphic indels were also observed when mapping the data onto the tree topology obtained from the Bayesian analyses of the combined data set. A few indels were also found to be incongruent with the phylogenetic tree obtained from the analysis of the combined data set. These were generally found in the most variable regions and some of the single base pair insertions actually consist of different bases. For more details of indel length and positions see the alignments of the individual gene regions deposited at Genbank, accession numbers XX (myoglobin), XX (G3PDH), and XX (ODC).

Models for nucleotide substitutions

The priori selection of nucleotide substitution models suggested that the GTR + Γ model had the best fit for all three gene regions, but as the nucleotide state frequencies and gamma distribution differed between the partitions we applied a partitioned analysis of the combined data set. After discarding the burn-in phase the inference for myoglobin and G3PDH were based on a total of 45,000 samples from the posterior, the inference for ODC were based on a total of 40,000 samples, while the inference for the combined data set were based on a total of 95,000 samples. The posterior distribution of topologies is presented as a majority-rule consensus tree from the combined analysis in Figure 2.

RESULTS

The trees obtained from the Bayesian analyses of the individual gene partitions (Fig. 3) are overall topologically congruent. Melanopareia clusters with Thamnophilidae and Conopophagidae and we can therefore reject it as a member of the family Rhinocryptidae with high confidence. Apart from this, all traditional tapaculo genera form a monophyletic clade within a broader group which also contains Formicariidae sensu stricto and Furnariidae, in agreement with previous molecular studies of tracheophone suboscines [START_REF] Irestedt | Systematic relationships and biogeography of the tracheophone suboscines (Aves: Passeriformes)[END_REF][START_REF] Chesser | Molecular systematics of New World Suboscine birds[END_REF]).

Within the radiation of tapaculos there is also good support for two major clades. Clade 1 includes Teledromas, Acropternis, Rhinocrypta, Liosceles and Psilorhamphus and Clade 2 includes Scytalopus, Eugralla, Myornis, Merulaxis and Eleoscytalopus. The only conflict within the rhinocryptid radiation supported by posterior probabilities above 0.95 involves determining to which of the previous two clades Pteroptochos and Scelorchilus belong. In the ODC tree they group with Clade 1 (0.97), whereas the myoglobin tree indicates that these two taxa are sister to the Clade 2 (1.00). In the G3PDH tree this relationship is unresolved. Based on the overall congruence of the individual gene trees we believe that the tree obtained from the combined analysis (Fig. 2) represents the best estimate of the phylogenetic relationship of the tapaculos and in this Pteroptochos and Scelorchilus fall out as sister to Clade 2. This tree is fully congruent with the results of studies using other genetic markers, but with focus on detailed relationships within Clade 2 (Maurício et al. 2008 and Cadena et al., unpublished).

DISCUSSION

Melanopareia resembles members of the Rhinocryptidae in having the lacrimal bones partly fused with the ectethmoid (but the lacrimals are lacking in Conopophagidae, Thamnophilidae, Grallaridae and Formicariidae) and in having a 4-notched sternum.

The significance of these characters is uncertain because of the weak cranial ossification in these groups and a considerable flexibility (including varying degrees of developmental asymmetry) in the degree of ossification of the membranes serving as attachment of pectoral muscles [START_REF] Heimerdinger | Variation in the sternal notches of suboscine passeriform birds[END_REF]. The molecular data reject a close relationship of Melanopareia to the Rhinocryptidae. With four closely related extant species, Melanopareia represents a long, unbroken phylogenetic branch, and it may be difficult to tell with confidence whether this clade is nested within the Conopophagidae-Thamnophilidae complex or represents a relictual, basal tracheophone type of bird. [START_REF] Irestedt | Systematic relationships and biogeography of the tracheophone suboscines (Aves: Passeriformes)[END_REF] associated Teledromas with Melanopareia primarily because their vocalizations are confusingly similar, and Teledromas was considered to resemble a robust and pale version of Melanopareia. Both genera are reported to share a peculiarity of the pterylography of the flank region, and details of the nasal operculum and tarsal scutellation, and an X-ray photo suggested almost straight humeri (approaching those of Melanopareia; [START_REF] Irestedt | Systematic relationships and biogeography of the tracheophone suboscines (Aves: Passeriformes)[END_REF]. However, our DNA data and further examination of skeletal characters reject a closer relationship between them and places Teledromas centrally in Clade 1 of the Rhinocryptidae.

Psilorhamphus was placed with Polioptiline oscines based on the acutiplantar tarsus, but a similar tarsal scalation is also found in some antbirds, and osteology and syringeal morphology suggested placement with Rhinocryptidae. We were able to confirm from examination of skeletal specimens that Psilorhamphusas well as Teledromashave the expanded footplate of the stapes typical of other suboscines (except Melanopareia).

The distinctive appearance of Psilorhamphus may result from its divergent habits (albeit shared with Myornis, N. Krabbe pers. comm.), as it generally feeds by climbing in tangles of vine-like bamboo, occasionally up to 7 m, and it rarely feeds on the ground. This species is also known for its unbelievably loud and low-pitched vocalizations (for its small size, 13.5 cm): A fast series of hollow whistles at 0.9-1 kHz. Its sister taxon in Figure 2, Liosceles, also gives hollow whistles, but they are higher pitched (1.3 kHz) and are given at a slower pace; the pace and quality of the song notes of Psilorhamphus is most like the songs of larger species of Pteroptochos, which are even lower pitched (0.5-0.6 kHz). Interestingly, Psilorhamphus shares with Liosceles barred posterior underparts and distinctive whitish subterminal spots with a black outline on the middle and greater wing-coverts. Apart from this, it is difficult to see any external features supporting the suggested relationships within Clade 1.

The possible association of the large Chilean tapaculos (Pteroptochos, Scelorchilus) with Clade 2 receives some morphological support, as Pteroptochos has 14 rectrices, something that is also found in some species or individuals of Scytalopus, although other representatives of this genus have a reduced number of rectrices or asymmetrical tails [START_REF] Krabbe | Family Rhinocryptidae (Tapaculos)[END_REF]. Other suboscine birds typically have 12 rectrices, although there are many cases of reduction. Molecular relationships within Pteroptochos have been analyzed by [START_REF] Chesser | Molecular systematics of the rhinocryptid genus Pteroptochus[END_REF].

Within Clade 2, the Myornis-Eugralla-Scytalopus group is particularly well defined morphologically by small size and sooty-grey to blackish plumage and atrophied clavicles, which do not form a fused furcula [START_REF] Maurício | Hidden genetic diversity in Neotropical birds: molecular and anatomical data support a new genus for the "Scytalopus" indigoticus species-group (Aves: Rhinocryptidae)[END_REF]. This may reflect reduced flying ability of these specialized birds, which tunnel through the densest parts of the forest understorey. Myornis bears some resemblance to Merulaxis, in shape and juvenile plumage [START_REF] Krabbe | Family Rhinocryptidae (Tapaculos)[END_REF], but our result does not support such an association. On the other hand, Myiornis and Eugralla clearly fall outside the group of species of Scytalopus that we studied (except with G3PDH) and we therefore support keeping them in separate monotypic genera.

Under "Incertae sedis", [START_REF] Irestedt | Systematic relationships and biogeography of the tracheophone suboscines (Aves: Passeriformes)[END_REF] introduced a family "Melanopareiidae (new family, incl. Melanopareia and Teledromas)" and this family name has been accepted by SACC (Proposal #239, 2006). These sources, however, did not meet the requirements of the International Code of Zoological Nomenclature (ICZN 1999) as it lacked any description purporting to differentiate the taxon from other taxa, so the name was invalid at that point. The need still remains for separate family status for the genus, which we re-propose here:

  Melanopareiidae, new family Type and only included genus: Melanopareia Reichenbach, 1853. Diagnosis: Tracheophone suboscine passeriformes differing from other Tracheophonae except Rhinocryptidae (and a few other taxa: Myrmothera, Hylopezus, Pittasoma, Conopophaga) in having a 4-notched sternum and differing from the Rhinocryptidae and all other suboscines in retaining the primitive morphology of the stapes, with a flat, rather than inflated footplate. Zusammenfassung Umschreibung einer monophyletischen Familie für die Bürzelstelzer (Aves: Rhinocryptidae): Psiloramphus rein und Melanopareia raus Die Bürzelstelzer (Rhinocryptidae) sind tracheophone, suboscine Vögel, deren Vorkommen auf Süd-und Mittelamerika beschränkt ist. Die meisten Bürzelstelzer haben eine Reihe innerer und äußerer Merkmale gemeinsam, die benutzt worden sind, um die Familie taxonomisch zu definieren. Die Gattungen Melanopareia and Psiloramphus passen nicht vollständig in dieses Schema und haben zu beträchtlichen Disputen unter Taxonomen geführt, seit sie erstmals beschrieben worden sind. In diesem Artikel stecken wir die systematischen Grenzen der Bürzelstelzer ab und bewerten ihre Gattungsbeziehungen mit Hilfe einer Analyse molekularer Sequenzdaten. Die Ergebnisse zeigen, dass Melanopareia weit aus dieser Klade herausfällt, während Psiloramphus gut in die Rhinocryptidae hineinpasst. Eine neue Familie wird für Melanopareia eingerichtet.

Figure 1 .

 1 Figure 1. The bamboo-wren Psilorhamphus guttatus bears little external resemblance to typical members of the tapaculo family. With its grey iris, facial expression, bill shape, and wing-coverts with white dots Psilorhamphus instead resembles some antbirds (Dysithamnus, Myrmotherula) with which early ornithologists consequently placed it. Unlike other tapaculos Psilorhamphus spends most of the time above the ground. Photo: Edson Endrigo.

Figure 2 .

 2 Figure 2. Majority rule consensus tree obtained from the Bayesian analyses of the combined data set (myoglobin intron 2, ODC introns 6 and 7, and G3PDH intron 11).Posterior probability values are indicated at the node.

Figure 3 .

 3 Figure 3. The majority rule consensus trees obtained from the Bayesian analyses of the individual genes. A) myoglobin intron 2; B) ornithine decarboxylase introns 6 to 7; and C) glyceraldehyde-3-phosphodehydrogenase intron 11. Posterior probability values are indicated at the node.

Table 1 :

 1 Specimen data and GenBank accession numbers for samples used in the study. Samples vouchered with a study skin are indicated by an asterisk. "TP" indicates that the sample is obtained from toe-pads of an old study skin. Acronyms are AMNH, American Museum of Natural History, FMNH, Field Museum of Natural History, Chigaco, USA; NRM, Swedish Museum of Natural History; USNM, University States National Museum, Washington, USA; ZMUC Zoological Museum of Copenhagen. References: A, Irestedt et al. (2002); B: Ericson et al. (2002); C: Irestedt et al. (2004); D: Ericson & Johansson (2003); E: Fjeldså et al. (2005); F: Ericson et al. (2006); G: Fjeldså et al. (2003); H: Fjeldså et al. (2007).

Table 1

 1

	Species	Family: Subfamily	Voucher / Sample No.	Myoglobin	G3PDH	ODC
	Acropternis orthonyx	Rhinocryptidae	ZMUC 125695 *	new	new	new
	Merulaxis ater	Rhinocryptidae	ZMUC 128820	new	new	new
	Pteroptochos tarnii	Rhinocryptidae	AMNH RTC467 *	AY065774 A) AY590096 E)	new
	Rhinocrypta lanceolata	Rhinocryptidae	NRM 966793	AY065775 A) DQ438953 F) DQ435499 F)
	Teledromas fuscus	Rhinocryptidae	USNM BKS3703 *	new	new	new
	Psilorhamphus guttatus	Rhinocryptidae	ZMCU? Olson 2005	new	new	new
	Myornis senilis	Rhinocryptidae	ZMUC 134967 *	new	new	new
	Scytalopus parvirostris	Rhinocryptidae	ZMCU 128441	new	new	new
	Scytalopus speluancae	Rhinocryptidae	ZMCU 128818	new	new	new
	Scytalopus spillmannii	Rhinocryptidae	ZMUC 125091 *	AY065773 A) AY590097 E)	new
	Scytalopus zimmeri	Rhinocryptidae	ZMUC 126278 *	new	new	new
	Scytalopus superciliaris	Rhinocryptidae	USNM BKS 3592 *	new	new	new
	Eugralla paradoxa	Rhinocryptidae	NRM 570026 *, TP	new	new	new
	Scelorchilus rubecula	Rhinocryptidae	NRM 570029 *, TP	new	new	new
	Liosceles thoracicus	Rhinocryptidae	NRM 570027 *, TP	new	new	new
	Eleoscytalopus indigoticus	Rhinocryptidae	NRM 570028 *, TP	new	new	new
	Melanopareia maximiliani	Rhinocryptidae	ZMUC 125045 *	AY065785 A)	new	new
	Furnarius cristatus	Furnariidae: Furnariinae	NRM 966772 *	AY064255 B) AY590066 E) DQ435482 F)
	Philydor atricapillus	Furnariidae: Furnariinae	NRM 937334 *	AY065758 A) AY590076 E) EF212110 H)
	Synallaxis ruficapilla	Furnariidae: Furnariinae	NRM 956643 *	AY065763 A) AY590068 E) EF212119 H)
	Lepidocolaptes angoustirostris Furnariidae: Dendrocolaptinae	NRM 937184 *	AY065767 A) AY336576 G) DQ435486 F)
	Dendrocincla tyrannina	Furnariidae: Dendrocolaptinae	ZMUC 125661 *	AY442959 C) AY590087 E) EF212098 H)
	Chamaeza meruloides	Formicariidae	ZMUC 126604 *	AY065776 A) AY590095 E)	new
	Formicarius nigricapillus	Formicariidae	ZMUC 125987 *	AY065777 A)	new	new
	Grallaria squamigera	Grallariidae	ZMUC 124629 *	AY065778 A) AY677078 C)	new
	Dysithamnus mentalis	Thamnophilidae	NRM 956629 *	AY676995 C) AY677042 C)	new
	Terenura humeralis	Thamnophilidae	FMNH 389941	AY677004 C) AY677051 C)	new
	Thamnophilus caerulescens	Thamnophilidae	NRM 967007 *	AY065783 A) AY336587 G) DQ435504 F)
	Conopophaga aurita	Conopophagidae	ZMUC 125796 *	AY065784 A)		DQ435478 F)
	Conopophaga lineata	Conopophagidae	NRM 956653 *		AY336577 G)
	Pipra fasciicauda	Pipridae	NRM 947271 *	AY065787 A) AY336583 G ) DQ435495 F)
	Corythopsis delalandi	Tyrannidae	NRM 937282 *	AY065788 A) DQ435463 F) DQ435479 F)
	Elaenia flavogaster	Tyrannidae	NRM 966970 *	AY228295 D)	new	new

Acknowledgements

Our thanks go especially to those researchers who, for years, and often with very small economic means, have collected the specimens and tissue samples without which studies like this could not be made. N. Krabbe is thanked for comments to the manuscript. E. Endrigo kindly provided the photo of Psilorhamphus. The following institutions provided samples that have been included in the study: Zoological Museum of Copenhagen University, Swedish Museum of Natural History, National Museum of Natural History, American Museum of Natural History and Field Museum of Natural History. The Swedish Research Council provided financial support for the laboratory work (grant no. 621-2007-5280 to PE).


Thorsten J S Balsby
email: tjsbalsby@bio.ku.dk.

Poul Hansen

Element repertoire: change and development with age in Whitethroat Sylvia communis song

Keywords: song repertoire, song development, age, delayed maturation, whitethroat, Sylvia communis

Song repertoires are often important in sexual selection. In several species older males have larger repertoires than one-year old males. The development of large song repertoires within an individual is, however, poorly understood. We studied song element repertoire changes in five individual male whitethroats Sylvia communis sampled as one and two years old. Males increased the size of their element repertoire between first and second year. Song length and number of different elements per song did not change between years. Song element repertoire size, therefore, could be a reliable signal of male age. On average 44.3 % of the song elements in the first year repertoire were also found in the second year repertoire. Elements, shared between years, were found earlier in the songs and tended to occur in sequences. Sequences of shared elements also seemed to be conserved between years. The study suggests that the song element repertoire in the second year is partly based on the first year repertoire, which may explain why large song repertoires mainly are expressed by males at least two-year old. Song element repertoire can thus be a reliable indication of male age.

Introduction

Several oscine passerine species have very elaborate song repertoires, which may serve both inter-and intrasexual functions [START_REF] Catchpole | Bird song, biological themes and variation[END_REF]. Development of large repertoires has been hypothesized to be costly [START_REF] Catchpole | Song and female choice: good genes and big brains?[END_REF][START_REF] Buchanan | Developmental stress selectively affects the song control nucleus HVC in the zebra finch[END_REF]), but see [START_REF] Gil | The honesty of bird song: multiple constraints for multiple traits[END_REF], and large repertoire size has been shown to reflect aspects of male quality such as age, condition and parasite [START_REF] Catchpole | Bird song, biological themes and variation[END_REF][START_REF] Gil | Song characteristics are age dependent in the willow warbler Phylloscopus trochilus[END_REF][START_REF] Spencer | Parasites affect song complexity in a songbird[END_REF] or nutritional stress early in life [START_REF] Nowicki | Song learning, early nutrition and sexual selection in songbirds[END_REF].

Cross-sectional studies have demonstrated that old males have larger song repertoires in e.g. sedge warbler Acrocephalus schoenobaenus [START_REF] Birkhead | Song, sperm quality and testes asymmetry in the Sedge Warbler[END_REF], pied flycatcher Ficedula hypoleuca [START_REF] Lampe | Song structure reflects male quality in pied flycatchers (Ficedula hypoleuca)[END_REF], great tit Parus major [START_REF] Mcgregor | Song repertoires and lifetime reproductive success in the great tit (Parus major)[END_REF], European starling Sturnus vulgaris [START_REF] Eens | No overlap in song repertoire size between yearling and older starlings (Sturnus vulgaris)[END_REF], canary Serinus canaria [START_REF] Nottebohm | Developmental and seasonal changes in canary song and their relation to changes in the anatomy of song-control nuclei[END_REF], whitethroat Sylvia communis (Balsby 2000a) and red-winged blackbird Agelaius phoeniceus [START_REF] Yasukawa | Song repertoires and sexual selection in the red-winged blackbird[END_REF]). An age difference in song repertoire size could arise if older males include new elements in their repertoire or if males with larger repertoires have higher survival rates [START_REF] Espmark | Variations in the song of the pied flycatcher within and between breeding seasons[END_REF]. However, only in great tits a relation between song repertoire size and survival rate has been documented [START_REF] Lambrechts | Male quality, reproduction, and survival in the great tit (Parus major)[END_REF]. Longitudinal studies have shown that individual males increase their repertoire size with age, especially between first and second year, in pied flycatcher [START_REF] Espmark | Variations in the song of the pied flycatcher within and between breeding seasons[END_REF], willow warbler Phylloscopus trochilus [START_REF] Gil | Song characteristics are age dependent in the willow warbler Phylloscopus trochilus[END_REF], starling [START_REF] Mountjoy | Extended song learning in wild European starling[END_REF], sedge warbler [START_REF] Nicholson | Song sharing and repertoire size in the sedge warbler, Acrocephalus schoenobaenus: changes within and between years[END_REF]) and great reed warbler Acrocephalus arundinaceus, although the increase in repertoire size only applied to one of the two populations of great reed warblers [START_REF] Forstmeier | Does song reflect age and viability? A comparison between two populations of the great reed warbler Acrocephalus arundinaceus[END_REF]). These longitudinal studies indicate that the repertoires only develop fully in older males in a number of species.

Longitudinal changes are interesting, because they offer insight to the development of the song repertoire and may explain why mainly older males display large repertoires. Although a large repertoire would be advantageous in both interand intra-sexual interactions, the full repertoire is often delayed to older age, even though the songs in the repertoire could have been learned early in life. One explanation may be that males can not learn a large repertoire in one season, but need to build on the repertoire from one or more previous years. This could be tested by comparing the song element repertoire between years. In this case we should expect to find that many elements and the most frequently used elements are shared between years. Shared elements should also occur in identical sequences between years. The alternative possibility is that older males develop their repertoire independently of the repertoire of previous years. In that case, elements, shared between years, should not be used more frequently than non-shared elements and sequences of shared elements should not be conserved between years.

In this paper we present data on the longitudinal change of song element repertoires and composition in the whitethroat in order to investigate how males develop their song repertoire from first to second year. Male whitethroats use their song for mate attraction and territory defence and have a substantially higher song activity before they attract a female to their territory (Balsby 2000b). Cross-sectional studies of whitethroats have shown that one-year old males had smaller song repertoires than males two-years old or older. The song element repertoire seems to be a sexually selected trait in whitethroats. Females have been found to show a preference for males with a large song repertoire (Balsby 2000a), while males did not respond to differences in song repertoire size in a playback experiment [START_REF] Balsby | The meaning of song repertoire size and song length to male whitethroats Sylvia communis[END_REF]. Given the importance of repertoire size in sexual selection, we wanted to study the development of the repertoire size by comparing the song element repertoire in individual males as one-and two-year old

Methods

The birds in this study lived in hedge rows between agricultural fields on the south side of Fussing Lake, Denmark (5628N, 954E) in May and June 1995[START_REF] Catchpole | Song and female choice: good genes and big brains?[END_REF][START_REF] Birkhead | Song, sperm quality and testes asymmetry in the Sedge Warbler[END_REF] To colour band and age each male, we captured the birds in mist nets by means of playback. Each male was marked with a unique combination of colour rings and one metal ring for individual identification. One-year old males could be distinguished from older males on the basis of iris colour and moulting patterns [START_REF] Karlson | Fotografisk dokumentation av ålders-och könsskillnader hos fåglan -målsättning, arbetssätt och exempel på resultat (Photografic documentation of age and sex differences in birds -aims, methods and examples of results)[END_REF][START_REF] Svensson | Identification Guide to European Passerines[END_REF]. In total we colour banded 9 one-year old males and recorded their song. The following year we identified 5 of these males by their colour bands and recorded their song. We recorded and analysed a total of 7021 songs from the 5 males. We analysed an average of 813 songs (range 196 to 1162 songs) from one-year old males and an average of 590 songs (range 440 to 729 songs) from the same males in their second year. We recorded all males with a Sennheiser MKT 816T directional microphone and a Sony 5000 EMV tape recorder. All recordings took place between 0500 and 1200 h. Males were only recorded prior to attraction of a female, as song activity post female attraction is very scarce. For the spectrographic analysis we used Avisoft-SASLab Pro (www.avisoft.de). We sampled sounds at 22.05 kHz. Spectrograms were made with the hamming window, a time resolution of 5.8 ms and a bandwidth of 111 Hz.

We classified all song elements by visual comparison between spectrograms on the computer screen and a printed catalogue with all known song elements for each of the males (Fig. 1). Each song element was classified either as an already existing type or as a new type. As the analysis progressed we assigned each new element a unique number in the catalogue and printed them for the catalogue. Elements were categorized on basis of shape, frequency and length; elements were only categorized as different if the changes were distinctly different and not gradual. All elements could be categorized. The recordings of one-year and two-year old males were analysed separately.

We measured song length (as number of elements), number of different elements per song, rep1000 and max1000. Rep1000 measured the average number of different elements presented within 1000 elements calculated over the whole sequence at steps of 10 elements. Max1000 was the maximum number of different elements within 1000 elements found while calculating the rep1000 (Balsby 2000a). An estimate of the full repertoire may be obtained from a cumulative graph such as Fig. 2 if the graph approximated an asymptote [START_REF] Kroodsma | Song repertoires: problems in their definition and use[END_REF]. However, in the whitethroat most males continued to present new song elements even after more than 10,000 elements had been analysed (Balsby 2000a). Contrary to rep1000 and max1000, an estimate of the song element repertoire based on the cumulative graph would, therefore, depend on the number of elements analysed. Rep1000 and max1000 were, therefore, assumed to be a better measure, also for comparison between males and years.

We then compared the catalogue for the two years for each male to identify elements used in both years (shared elements) and elements that were only found in either first or second year (non-shared elements). We then translated each sequence so the element numbers as one-year old and two-year old referred to the same element for shared elements. From the translated sequence we quantified the sequential organization of the song (i.e. the number of times a combination of elements occurred in a particular sequence). For shared and non-shared elements we counted their frequency, the average position in the song of shared and non-shared elements measured as the number of elements counted from the start of each song. We also counted the number of times a sequence of shared elements occurred. This enabled us to compare the sequences and the use of shared elements between years within an individual.

Statistics

We used StatXact (Cytel Software Cooperation) for making the randomisation tests for comparison of two paired groups. Each test was based on 1000 resamplings and we report the exact p-value. For other tests, we used non-parametric statistics in accordance with [START_REF] Siegel | Nonparametric statistics for the behavioral sciences[END_REF]. In the sequence analysis, and element sharing we chose to test each test bird separately, because we were concerned about the individual pattern. This could give rise to issues with multiple testing, but most tests would still be significant after sequential bonferoni corrections [START_REF] Rice | Analysing tables of statistical tests[END_REF]. For comparison of sequence matrices between years we used Mantel test for skew symmetric case [START_REF] Mantel | The detection of disease clustering and a generalized regression approach[END_REF]. All tests are two-tailed.

Results

Song element repertoire size

All 5 males had larger song element repertoire as two-year old (Fig. 2), despite more elements had been analysed for the one-year old males except in one. The cumulative graph for one-year old males almost reached an asymptote, providing a good estimate of the song element repertoire size, while the cumulative graphs for two-year old males did not reach an asymptote, and thus underestimated the element repertoire size.

Despite the underestimate of the repertoire for 4 of the two-year old males, from which we analysed most elements as one year old, the total number of different elements increased at least 1.6-4.3 times from the first to the second year. Rep1000 and max1000 supported these observations, as two-year old males had significantly larger rep1000 and max1000 than one-year old males (Fig. 3a, b, respectively.

Randomization test accounting for paired design: rep100: p<0.01; max1000: p<0.01).

However, song length and number of different elements per song did not differ between years (Fig. 3c,d, resp. Randomization test accounting for paired design: song length: p=0.13; number of different elements per song: p=0.13).

Comparison of element repertoires in one and two years old males

Elements shared between years on average constituted 44 % (range 30 to 61 %) of the repertoire in one-year old males (Fig. 4), but on average only 20 % (range 7 to 27 %) of the repertoire in two-year old males. Four of the one-year old males used shared elements significantly more frequently than non-shared elements (Mann-Whitney U test for each male: U≥3216, n shared ≥97, n non-shared ≥118 p<0.001), while this was not the case for the last male, #5 (Mann-Whitney U test: U=1353, n shared =38, n non-shared =88, p=0.09). As two-year old, four of the males (#1 to 4) also used shared elements more frequently than non-shared elements (Mann-Whitney U test: U≥36662, n shared ≥97, n non-shared ≥431 p<0.001), whereas one male (#5) used non-shared elements more often than shared elements (Mann-Whitney U test: U=12355,n shared ≥38, n non-shared ≥510 p=0.04). Shared elements on average occurred earlier in the sequence of the song elements than non-shared elements in males #1-4 in both years (Mann-Whitney U test, one-year old: U≥5.02x10 5 , n shared ≥1844, n non-shared ≥422, p<0.0001; Mann-Whitney U test, two-year old: U≥ 1.27x10 5 , n shared ≥2249, n non-shared ≥1436, p<0.002). In male #5 shared elements occurred later in the song than non-shared elements in both years (Mann -Whitney U test, one-year old: U=1.59x10 6 , n shared =1079, n 2 =9983, p<0.0001; Mann-Whitney U test, two-year old: U=1.77x10 6 , n shared =453, n non-shared =8413, p=0.018).

Songs generally started with a shared element more frequently than one would expect based on the frequency of shared elements in each males' repertoire. The expected frequencies were thus calculated on basis of the proportion of shared and non-shared elements. All two-year old males used shared elements as the first element in the song more frequently than expected from the proportion of shared elements in their repertoire (Fig. 5, χ2=3284.6, df=4, p<0.001). In one-year old males the pattern was less clear as four of the males (#1-4) used shared elements as the first element in the song more often than expected from the proportion of shared elements in their repertoire and one male (#5) used shared elements as first song element less often than expected from the proportion of shared elements in its repertoire (Fig. 5, χ2=3199, df=4, p<0.001).

First-order transition of shared elements showed some stability between years.

Overall, shared elements tended to be followed by shared elements and non-shared elements tended to be followed by non-shared elements in both years for all the males (Chi-square test, one-year old: χ2>1180, df=1, p<0.001; two-years old: χ2>213, df=1, p<0.001). Dyads of shared elements tended to occur in identical sequences between years, as shown by the correlation between the frequency of the combinations of the first-order transitions in the first and second year for each male (Mantel test for asymmetric matrices, male1: t=50 r=0.29, male 2 t=59 r=0.19, male 3: t=114 r=0.35, male 4: t=96 r=0.33, male 5: t=3.9 r=0.17, df for all ∞, all p<0.001). Thus, sequences of pairs of elements were conserved between years. Furthermore, the longest sequence of shared elements that were identical between years were 4, 13, 16, 11 and 3 elements for male # 1 to 5, respectively.

Discussion

Male whitethroats increased their song element repertoire between the first and the second year of life, as also shown by the increased rep1000 and max1000. We have no data to quantify the change in repertoire size after the second year. However, the repertoire size of the two-year old males in this study were larger than the measured repertoire sizes in an earlier study (Balsby 2000a), in which the songs from 10 males, two years old or older, were analysed. We, therefore, believe that the major change in repertoire size in whitethroats occurs between first and second year, which would be similar to willow warblers [START_REF] Gil | Song characteristics are age dependent in the willow warbler Phylloscopus trochilus[END_REF]. The increase in repertoire size with age fits with other studies [START_REF] Gil | Song characteristics are age dependent in the willow warbler Phylloscopus trochilus[END_REF][START_REF] Nicholson | Song sharing and repertoire size in the sedge warbler, Acrocephalus schoenobaenus: changes within and between years[END_REF][START_REF] Forstmeier | Does song reflect age and viability? A comparison between two populations of the great reed warbler Acrocephalus arundinaceus[END_REF][START_REF] Mountjoy | Extended song learning in wild European starling[END_REF]. However, repertoire change with age did not occur in some populations of the great reed warbler [START_REF] Forstmeier | Does song reflect age and viability? A comparison between two populations of the great reed warbler Acrocephalus arundinaceus[END_REF]). In nightingales Luscinia megarhynchos two-year or older males did not increase their repertoire size in the subsequent year [START_REF] Kipper | Long term persistence of song performance rules in nightingales (Luscina megarhynchos): a longitudinal field study on repertoire size and composition[END_REF]. By cross-sectional comparison [START_REF] Kiefer | First-year common nightingales (Luscinia megarhynchos) have smaller song-type repertoire sizes than older males[END_REF] showed that one-year old males had smaller repertoires than older males. This was confirmed by a longitudinal study [START_REF] Kiefer | Tuning towards tomorrow? Common nightingales Luscinia megarhynchos change and increase their song repertoires from the first to the second breeding season[END_REF]) that showed, that one-year old males had an increased repertoire the next year.

Song repertoires may be costly to produce and maintain [START_REF] Catchpole | Song and female choice: good genes and big brains?[END_REF][START_REF] Gil | The honesty of bird song: multiple constraints for multiple traits[END_REF], and evidence for a link between song repertoire size and brain size has been found in some species [START_REF] Szekely | Evolutionary changes in a song control area of the brain (HVC) are associated with evolutionary changes in song repertoire among European warblers (Sylviidae)[END_REF][START_REF] Airey | Song, sexual selection, and a song control nucleus (HVc) in the brains of European sedge warblers[END_REF][START_REF] Buchanan | Developmental stress selectively affects the song control nucleus HVC in the zebra finch[END_REF]), but not always [START_REF] Leitner | Syllable repertoire and the size of the song control system in captive canaries (Serinus canaria)[END_REF]. The present study clearly indicates that one-year old whitethroat males do not express a full repertoire, which may suggest that one-year old whitethroat males may have insufficient brain capacity to express large song repertoires. Song repertoire size in whitethroats could thus provide a reliable cue to male age. The relation between age and repertoire size could have implications for female mate choice, because age may indicate the indirect (genetic) quality of the male (Manning 1988[START_REF] Kokko | Good genes, old age and life history trade-offs[END_REF]. Female whitethroats indeed seem to prefer older males as all old males succeed in attracting a female, whereas half of the one-year old males never attracted a female (Balsby 2000a). Song repertoire size may, therefore, be a sexually selected trait in whitethroats.

In whitethroats differential survival of males with large song repertoire size [START_REF] Espmark | Variations in the song of the pied flycatcher within and between breeding seasons[END_REF]) cannot explain the repertoire difference between age-classes, as the total repertoire size (Fig. 2) of one-year old and two-year old whitethroat males does not overlap in this study and in a cross sectional study (Balsby 2000a). Song length and number of different elements per song did not differ between first and second year in cross-sectional (Balsby 2000a) and longitudinal studies (the present study).

Song length and number of different elements in the song thus hold no potential for indicating age in whitethroat males.

Song repertoire development

Despite a considerable interest in the function of and the evolution of large song repertoires, their development between years has not been studied. We investigated the development of song element repertoire in the whitethroat by quantifying the degree of element sharing between the two years and the use of shared elements. In whitethroats a substantial part of the element repertoire consisted of elements shared between years. Both one-year and two-year old males, except one male, used shared song elements more frequently than expected from the proportion of shared song elements in the song repertoire. Shared elements also came in sequences, and at least dyads (the sequence of two elements) seemed to be conserved between years. Long identical sequences of song elements between years suggest that longer sequences of shared elements to some degree are transferred between first and second year. Male whitethroats, therefore, partly build their song element repertoire as two-year old on the song element repertoire and sequences used as one-year old. Repertoire turnover between years has also been found in sedge warblers and pied flycatchers, which drop some and include other syllables between years (Nicholson et al. 2007, Espmark and[START_REF] Espmark | Variations in the song of the pied flycatcher within and between breeding seasons[END_REF]. None of these studies, however, proceed to look at how the repertoires develop, but they suggest that turnover of part of the repertoire between years occur in several species besides whitethroats. Our song recordings of two-year old males have not sampled all elements in the repertoire of individual males, so there might be more shared elements. However, more shared elements would only support the conclusion that males base their repertoire as two-year old on their repertoire in the first year.

The large change in repertoire size and the recombination of the sequence of song elements between one-year and two-year old males suggest that males modify and to some degree rebuild their repertoire before the second breeding season. The rebuilding of the repertoire from first to second year would be in accordance with the seasonality of the size of various brain centres. In canaries the song controlling parts of the brain grow prior to the start of the breeding season [START_REF] Nottebohm | Developmental and seasonal changes in canary song and their relation to changes in the anatomy of song-control nuclei[END_REF][START_REF] Ball | Seasonal plasticity in the song control system. Multiple brain sites if steroid hormone action and the importance of variation in song behavior[END_REF]).

The majority of songs given by whitethroats starts with a motif and ends with a variable part (Balsby 2000b). These motifs are conserved within years. Elements, shared between years, tended to occur earlier in the song than non-shared elements, suggesting that many of the shared elements were motif elements. The lack of complete motif sharing between years suggests that males modify their motif repertoire each year. Males may make this modification in order to adjust to the motifs of neighbours, thus allowing matched countersinging. Such adjustment to the song types of neighbours are known from sedge warblers [START_REF] Nicholson | Song sharing and repertoire size in the sedge warbler, Acrocephalus schoenobaenus: changes within and between years[END_REF].

Furthermore, in great tits, wrens (Troglodytes troglodytes), dusky warbler (Phylloscopus fuscatus) and nightingales males share a substantial part of their repertoire with their neighbours [START_REF] Mcgregor | Song learning in adult great tits (Parus major) the effect of neighbours[END_REF][START_REF] Catchpole | Song sharing and local dialects in a local population of the European wren Troglodytes troglodytes[END_REF][START_REF] Forstmeier | Why mated dusky warblers sing so much: territory guarding and male quality announcement[END_REF][START_REF] Griessmann | Song sharing in neighbouring and non-neighbouring thrush nightingales and its implication for communication[END_REF]. Whitethroats are generally philopatric [START_REF] Hansen | Tornsangeren på Hjelm[END_REF]Cramp 1992). One of the males in this study used shared elements less often than the other four males. In the second year this male (#5) settled in a territory about 1 km from his old territory. By moving, male #5 might have had to change a much larger part of his repertoire than males that returned to the same territory as the year before. The other four males returned to the same area in both years.

Whitethroats might have learned all the song elements in an early sensitive phase, but without the capability to express them all as one-year old. Such a scenario would be similar to that found in nightingales, where males learn more song types than they express [START_REF] Todt | Age dependent effects of song exposure: song crystallization sets a boundary between fast and delayed vocal imitation[END_REF]. However, whitethroat males stop singing once mated and only males with small repertoires, which did not attract a female, would continue singing (Balsby 2000b). Opportunities for learning song from large repertoire males would, therefore, be quite limited. A sensitive song learning period shorter than one year thus seems unlikely. Male whitethroats could acquire parts of their repertoire in the winter quarters or during migration similar to the marsh warbler (Acrocephalus palustris) [START_REF] Dowsett-Lemaire | The imitative range of the song of the marsh warbker Acrocephalus palustris, with special reference to imitations of African birds[END_REF]. All these possibilities require whitethroat males to be open-ended learners or to have a sensitive period for song learning that lasts at least two years. The current data set does not allow discrimination between these two possibilities. However, it is clear that the song element repertoire size is a delayed matured trait [START_REF] Dale | Population divergence in sexual ornaments: the white forehead patch of Norwegian pied flycatchers is small and unsexy[END_REF], as whitethroats only can express a large repertoire when two-years old or older, and as they partly build their second year song element repertoire on the first year repertoire.

Zusammenfassung

Altersabhängige

  Entwicklung und Veränderung von Gesangselementen und -repertoiren bei Dorngrasmücken Sylvia communis Gesangsrepertoire sind häufig von Bedeutung für sexuelle Selektion. In einigen Vogelarten haben ältere Männchen größere Repertoire als einjährige Männchen. Die Entwicklung eines großen Repertoires innerhalb Individuen ist jedoch weniger gut untersucht. Wir haben Veränderungen von Elementen im Gesangsrepertoire von individuellen, ein-und zwei Jahre alten Dorngrasmücken untersucht. Männchen vergrößerten ihr Elementrepertoire zwischen dem ersten und dem zweiten Jahr. Gesanglänge und Anzahl unterschiedlicher Elemente pro Gesang veränderten sich nicht zwischen den Jahren. Deshalb könnte die Größe des Gesangsrepertoirs in Männchen zuverlässig Alter anzeigen. Im Schnitt wurden 44.3% der Gesangselemente der Repertoire von einjährigen Vögeln auch im jeweils zweiten Jahr gefunden. Elemente die zwischen den Jahren gleich blieben traten früher in den Gesängen gefunden und traten auch in Sequenzen auf. Solche Sequenzen von geteilten Elementen scheinen zwischen den Jahren erhalten zu bleiben. Unsere Arbeit lässt vermuten, dass das Gesangselementrepertoire im zweiten Jahr teilweise auf das Repertoire im ersten Jahr basiert. Dies kann möglicherweise erklären warum große Gesangsrepertoire vorwiegend in Männchen die zumindest zwei Jahre alt sind, gefunden werden. Wir schließen daraus dass das Gesangsrepertoire ein zuverlässiger Indikator für das Alter eines Männchens sein kann.

Figure legends Figure 1 .

 legends1 Figure legends

Figure 2 .

 2 Figure 2. Cumulative graph of the number of new song elements as a function of the

Figure 3 .

 3 Figure 3. Change between one-year old and two-year old males in rep1000 (a),

Figure 4 .

 4 Figure 4. Number of elements found only in the first year (white), in both years (dots),

Figure 5 .

 5 Figure 5. Observed (grey) and expected (white) number of shared elements used as

Acknowledgement.

The bird banding was done under permission from the ringing centre at the Zoological Museum, Copenhagen, Denmark. The ringing centre also provided the metal rings for ringing the whitethroats. We thank the landowners and the hunters in Venning for allowing us to study the whitethroats on their properties. TJSB is supported by a Steno stipend from the Danish Natural Science Research Council. This project complies with the laws of Denmark. The authors wish to thank the reviewers for their comments.


Giuseppe Pagliarulo
email: giuseppe.pagliarulo@unito.it

Keywords: Gospel, Lindisfarne Glosses, Anglo-Saxon, Northumbrian, word-order, auxiliaries

The author examines all instances of independent use of passive and progressive periphrases in the Lindisfarne Glosses. After having classified the relative positioning of their elements according to frequency and having considered it in its co-occurrence with various contextual variables, especially the presence of pronominal subjects, he suggests that the word order of the text's language can be regarded as being predominantly of the OV type, while at the same time showing clear signs of evolution toward the more modern VO pattern.

The issue of word order in the tenth-century Northumbrian text of the Lindisfarne Glosses (LG) has never attracted much attention from scholars, for reasons that may seem obvious. Being a literal, word-for-word translation, it is commonly assumed not to offer useful clues for research in the field.

Callaway made this clear as early as 1918: "As to the normal order of words, this Northumbrian gloss, like most interlinear translations, gives next to no evidence of value, since, as a rule, the glossator adheres strictly to the order of his Latin original" (Callaway 1918: iv). More recent studies, while acknowledging the LG's occasional departures from the original's word order, fail to analyze these in a systematic fashion. [START_REF] Ross | The Errors in the Old English Gloss to the Lindisfarne Gospels[END_REF] explains a number of such departures as the result of the glossator's use of more than one manuscript as a source text, or as plain errors. A more focused approach is that of [START_REF] Nagucka | Glossal Translation in the Lindisfarne Gospel according to Saint Matthew[END_REF], who, in a study on the gloss to Matthew's Gospel, examines certain cases in which the glossator seems to deliberately violate his otherwise strict adherence to the Latin original in order to meet his native language's constraints and peculiarities: thus, he sometimes reorders the genitive or the adjective, as in Mt 1:1 (dauides sunu for filii dauid) or in Mt 1:18 (of halig gaast for de spiritu sancto), or he uses double negation, something that Latin does not tolerate, as in Mt 9:30 (geseað þaette nan nyte for uidete ne quis sciat). Nagucka's analysis is, however, mostly focused on stylistic, not syntactic, aspects of the problem. Other scholars have considered the problem only partially and in passing, in studies devoted to other subjects (see, for instance, [START_REF] De Haas | The Origin of the Northern Subject Rule[END_REF][START_REF] Kroch | Verb Movement in Old and Middle English: Dialect Variation and Language Contact[END_REF].

The text, however, deserves the effort, being a major primary source on an otherwise scarcely attested Old English dialect with noteworthy original traits. It is true that an interlinear translation such as the LG is bound to suffer the syntactic influence of the source text in a heavy fashion, but it should also be noted that there are various Latin grammatical and lexical forms for which Old English lacks exact equivalents. In these cases the glossator is effectively forced to take some liberty in interpreting and paraphrasing the original. It seems reasonable that a close look at all those cases in which the glossator uses periphrases to render single Latin words will yield precious insights as to the "natural" ordering of elements in the Northumbrian phrase. The present study seeks to fill in some of our gaps in the knowledge of Northumbrian word-order by focusing on all instances in which a single Latin verb is rendered by a participle-copula complex or such a complex is used independently of the source text. This kind of periphrasis is most frequently used with a preterite participle for rendering the Latin passive, in its periphrastic as well as synthetic forms. A "progressive" periphrasis consisting of a copula and a present participle is also attested [START_REF] Scheffer | The progressive in English[END_REF]). In the LG, when used independently, it usually translates the Latin active imperfect tense. Both periphrases may appear in independent subordinate clauses rendering Latin passive infinitives, absolute ablatives, elliptical clauses and other expressions of difficult word-for-word translation. For clarity's sake, they shall be dealt with separately in what follows.

Periphrases consisting of a copula (C) and a preterite participle (P) are abundantly attested in the LG. 802 independent instances can be found in the whole of the text, and they are distributed These very raw figures may suggest that the P-C ordera typical feature of OV languagesis predominant in the LG. This would be consistent with the view, now held by many scholars, that Old English is a predominantly or underlyingly OV language. A closer look at the text, however, reveals that there are contexts in which different patterns are numerically stronger.

53 of the 62 occurrences grouped together as "other cases" (our fourth category) are renderings of single Latin verbs or verbal adjectives by periphrases accompanied by a pronominal subject or object (an example of the latter is dimittimini = iuh bið forleten, J 3:18). Of these, 46

show the addition of a pronominal subject 1 . In 35 cases out of 46 the presence of the pronominal subject is accompanied by fronting of the finite verb and thus a C-P order in the periphrasis (the type turbaris = ðu bist astyred, Lk 10:41), consistently with the tendency, nearly always observed in Old English declarative independent clauses, for the finite verb to occupy the position immediately to the right of the pronominal subject (Fischer et al., 2000: 49). It should be noted, however, that such a pattern is also detectable in the translation of Latin subordinate clauses (e. g. ut baptizarentur

= þaette hia wero gefuluad, Lk 3:7). This may be a result of the word-for-word technique adopted by the glossator or it may be the signal of a levelling of the pattern throughout clause types, not an uncommon phenomenon in the earliest Middle English documents (Francovich Onesti, 1993: 91).

We will return to this issue later.

Only three cases with an added pronominal subject show a P-C order in the periphrasis. One of these is clearly recognizable as a final clause: ut uideantur = þaet hia gesene sie, Mt 6:5. In Mt 23:12 we have exaltabitur = he ahaefen bið, and in Mt 23:28 paretis = ge gesene bið.

Three of the cases with an added pronominal subject show a C-subject-P order. One of these is immediately preceded by a negative particle: ne turbemini = ne se gie gestyred, Mt 24:6; one is immediately preceded by a WH-word: quid turbamini = forhuon arogie gestyred, Mk 5:34; the third has baptizabimini = se gie gefulwad, Mk 10:39. The most parsimonious explanation for the ordering of elements in Mt 24:6 and Mk 5:34 is that it is due to the influence of the preceding word, as it is an unexceptionable rule of Old English syntax for sentences to show subject-verb inversion in presence of an immediately preceding negative particle or in independent questions (Fischer et al., 2000: 49-50;van Kemenade 1997a: 93). This, however, would imply that the glossator was rather less "mechanical" in his work than one would expect the author of a mere word-for-word gloss to 1 Mt 3:13, 4:1, 5: 19,5:25,6:1,6:2,6:5,6:16,6:18,6:31,7:2,8:25,10:18,12:37,12:37,22:3,23:5,23:10,23:12,23:12,23:28,23:34,24:6,26:2,27:26;Mk 5:32,10:39,13:9;Lk LXXIIII,XCIIII,1:74,3:7,6:21,10:41,12:50,13:32,15:29,21:12,21:16,23:32,24:49;J 1:50,14:28,16:32,18:36,21:18. be. It is therefore pertinent to consider Mt 24:6 in the wider context of all renderings in which a Latin finite verb is translated by a Northumbrian finite verb accompanied by a pronominal subject and it is immediately preceded by a negative particle (e. g. non ueni = ne cuom ic, Mt 5:17). There are 194 such cases 2 . Of these, 182 (94.3% of the total) show subject-verb inversion. Of the 12 divergent cases, 8 appear to be mechanical translations (the type non sitiam = ne ic ðyrsto, J 4:15)

while the others place ne after the subject (the type non habeo = ic ne hafu, J 4:17). In some cases the glossator gives two alternative renderings, a literal one and a "free" one, of the same expression:

in J 4:38 the Latin non ad te pertinet is translated first by ne to ðe byreð and then by ne reces ðu. In the only case of conjunctive negation there is no inversion: neque nubent neque nubentur = ne hia maensumiað ne hia biðon gemaensumad (Mk 12:25), the common Old English word order in such cases being SOV. The periphrasis, however, has a C-P order, contrary to the general Old English tendency (see Mitchell 1985: § 3911 ff.). The presence of the pronominal subject hia probably accounts for this discrepancy, as will be suggested in greater detail in what follows.

Let us now consider all cases in which a Latin finite verb in a "yes/no" question or immediately following an interrogative pronoun or adverb is translated by a Northumbrian finite verb accompanied by a pronominal subject (e. g. quem quaeritis = huoelcne soecas gie, J 18:4).

2 Mt 3:11,5:17,5:21,5:26,5:36,6:1,6:24,7:3,8:8,8:10,10:23,10:28;10:34,11:17,11:17,12:7,13:13,13:13,14:17,15:17,14:24,15:32,15:32,16:3,16:7,16:23,17:27,18:3,20:13,20:22,21:32,22:12,23:10,23:30,23:37,24:44,25:12,25:13,25:26,25:42,25:42,25:43,25:43,25:43,25:44,25:45,26:29,26:40,26:70;Mk 3:9,4:13,4:38,7:4,7:18,8:2,8:16,8:17,8:18,8:18,8:18,8:33,10:38,11:31,12:14,12:14,12:24,12:25,12:26,12:34,13:33,14:25,14:37,14:68,14:68;Lk 1:34,3:16,5:32,6:41,6:42,6:46,7:7,7:9,7:32,7:32,7:44,7:45,7:46,11:7,12:17,12:40,12:59,13:7,13:25,13:27,13:34,14:20,15:19,15:21,16:2,16:3,17:9,18:4,18:11,19:44,21:14,22:16,22:18,22:53,22:57,22:58,22:67,22:68,22:68,24:18;J 1:21,3:8,3:11,4:15,4:15,4:17,4:17,4:48,5:7,5:30,5:38,5:38,5:40,5:41,5:42,5:43,5:44,6:36,6:39,6:53,7:8,7:28,7:34,7:35,7:36,8:14,8:16,8:19,8:41,8:43,8:45,8:49,8:50,9:12,9:21,9:25,9:29,9:41,10:26,10:33,11:15,11:50,12:49,13:8,13:8,13:11,13:33,13:36,14:9,14:10,14:11,14:18,15:5,15:15,15:19,16:1,16:4,16:10,16:12,16:16,16:18,16:23,16:24,16:32,17:11,17:15,18:9,18:17,18:20,18:25,18:36,19:6,19:10,19:11,19:15,19:36,20:2,20:13,20:25,21:25. Negative interrogatives are excluded.

There are 110 such renderings 3 . Of these, 92 (83.6% of the total) show subject-verb inversion (note that the percentage drops to 40.6% when other elements intervene between the WH-word and the subject). The case numquid ego = hueðer ic see, Mk 14:19, has been counted as a divergent instance but it should be remembered that questions introduced by hueðer do not usually show subject-verb inversion (Fischer et al., 2000: 54;Mitchell 1985: §3895) 6:24,7:18,8:17,10:17,10:36,10:38,10:51,11:3,11:5,12:10,14:12,14:19,14:68,15:9,15:34;Lk 1:65 ,2:48,3:12,3:14,4:30,4:34,5:22,6:2,6:42,7:19,7:24,7:25,7:26,7:44,8:25,9:54,10:26,12:17,12:41,12:42,12:49,12:51,13:2,13:4,16:3,16:5,16:7,18:8,18:19,18:19,19:31,19:33,22:9,22:46,22:49,24:5,24:41;J 1:38,1:38,4:12,4:27,4:27,5:6,5:44,6:5,6:28,6:30,6:68,8:53,11:26,11:47,11:34,14:5,18:4,18:22,18:25,18:35,18:39,21:5,21 ,10:34,18:25,23:8,27:64;Mk 10:4,10:38,13:4,13:10,14:49;Lk II,III,XI,LXV,LXVI,1:54,5:15,6:37,6:37,9:22,12:4,12:50,12:56,12:58,15:19,15:19,15:23,16:19,17:2,17:25,21:8,21:20,21:31,22:7,22:37,23:2,23:8,24:7,24:7,24:44,24:47;J I,II,VI,3:4,12:34,15:11. 5 Mt 3:16,14:19,21:35,26:56;Mk 1:31,3:23,5:26,9:20,9:32,10:22,11:18,14:3,15:44;Lk IIII,XVII,XXXVIII,2:43,3:21,4:2,8:29,9:1,16:5,20:45;J Arg.,IIII,7:14. one from Mk 14:32 (ðe is genemned predium, Mk 14:32; ðe is sua genemned, Mk 12:41; ðio forleteno bið = dimissam, Lk 16:18).

In the 47 final or complement clauses introduced by þaet(-te), word order is predominantly C-P, whether a (pronominal) subject is present (e. g. þaette ic se gefuluwad = baptizari, Lk 12:50, and 16 other cases) or not. There are only two divergent cases: þaet gesendet were = missum, J II;

þaette ofersuiðet waes = superatus, Lk XI. Subordinate clauses introduced by miððy present a much more mixed picture: 17 out of 26 cases (roughly two-thirds of the total) have a P-C order in the periphrasis. None of these has an independent expressed subject, however, and in the only case in which an independent complement is present (Mk 11:18, given above), although the complement is on the left of the periphrasis, the periphrasis has a C-P order: the same pattern that has been observed in Mt 27:26.

Let us now consider the cases falling into our first and second categories in the light of these data.

Periphrases immediately preceded by a negative particle (e. g. ne bið forgefen = non remittetur, Mt 12:31) show a strong prevalence of the C-P order, so that the finite verb immediately follows ne, consistently with what has been observed in cases like Mt 24:6. Of the 45 instances 6 of this kind, only four have a P-C order. Expressions of wish seem to make the C-P pattern preferable, too: the elliptical Latin benedictus is always rendered by the Northumbrian se gebledsod (Mt 21:9, 23:39; Mk 11:9, 11:10; Lk 1:68, 13:35, 19:38; J 12:13) even though it is always and very predictably followed by the noun or pronoun it refers to. Compare these cases to the elliptical beati in the Sermon of the Mount (Mt 5:3-12), which is always rendered by eadge biðon.

6 Mt 12:2,12:4,12:31,12:32,12:39,14:4,16:4,17:21,24:2,24:2,24:34,Mk 2:24,4:22,6:9,6:18,7:4,8:12,9:44,9:46,9:48,13:2,13:2,13:31;Lk 2:26,6:4,7:30,8:17,8:18,8:27,10:42,11:29,12:2,21:6,21:18,21:34,24:23;J 3:20,5:10,7:23,12:42,14:1,14:27,18:31. The same may be said of periphrases immediately preceded by a pronominal subject (e. g. ic beom gefulwuad = ego baptizor, Mk 10:39). One can count 13 such cases 7 . Only two of these have a P-C order in the periphrasis: the divergent instances are ðés gefroefred bið = hic consolatur, Lk 16:25; ða gefroefred biðon = ipsi consolabuntur, Mt 5:5, the latter introduced by forðon = quoniam and probably meant as a subordinate. Renderings like the one from Mk 10:39 may be compared to ðu arð ðaes ðegn = tu discipulus illius es, J 9:28; ic am tó ðon accenned = ego in hoc natus sum, J 18:37: here the glossator is brought to violate his source text's word order to introduce verbseconding in presence of a pronominal subject. In Lk 20:16 we find a rendering that seems particularly interesting. Here the ablative absolute quo audito is translated, as usual, by a finite subordinate clause: miððy þaet waes gehered. The periphrasis shows a C-P order, which, as we have seen, is not the majority pattern for subordinates introduced by miððy. This particular rendering finds an exact parallel in miððy ðas woeron gecuedno = his dictis, Lk 19:28. This, along with what we have observed in independently formed subordinates like the example from Lk 12:50, above, makes it tempting to say that the presence of a (pronominal) subject plays some role in the fronting of the finite verb even in subordinates like these, which would be another early sign of the levelling of the word order pattern throughout clause types observed in later English.

33 periphrases of interest are immediately preceded by þaet(-te) = ut 8 (e. g. þaette sie gefylled = ut adimpleretur). The C-P order has a slight majority in these cases, with 18 instances against 15 instances of P-C. But again, note that when a pronominal subject is independently added in such a rendering (e. g. þaette hia wero gefuluad = ut baptizarentur, Lk 3:7, and even þaet ðu ne se gesene = ne uidearis, Mt 6:18) the C-P order prevails greatly: of 13 such renderings 9 , only one shows a P-C 7 Mt 5:5,26:9;Mk,10:38,10:39;Lk 5:38,16:25,16:25,18:14,19:28,20:16;J 1:42,13:14,21:25. 8 Mt 1:22,2:22,2:23,4:14,5:13,8:17,12:17,13:35,20:33,21:4,26:56,27:35;Mk 10:38,12:33,15:15;Lk 2:21,3:12,9:44,11:50,14:23,21:22,23:23;J 1:31,3:17,3:21,9:2,11:4,12:23,13:18,14:13,15:25,18:9,19:16,19 Again, the raw numbers are in favour of the P-C pattern. However, the contextual variables we have contemplated seem to exert their influence here too.

Nine of the "other cases" show the independent addition of a pronominal subject (e. g. he waes spraecend = loquebatur, Mk 8:32). In seven of these (Mt 13:1,17:5,21:9;Mk 3:11,8:32,11:25;J 11:19) the periphrasis has a C-P order, even when a subordinating conjunction precedes dum fabularentur, Lk 24:15. A relative pronoun precedes the periphrasis in þaette cuoeðend waes = quod dixerat, Mk 14:72; ðaðe bidendo woeron = qui expectabant, Lk 2:38.

(
It seems that three points can be made in the light of what we have observed. The first is that the glossator didn't carry out his work on a purely word-for-word basis but he took context into account to an appreciable degree. The second is that he seems to show a preference for the P-C pattern except when certain contextual elementsmost interestingly, the presence of a pronominal subjectdrive him to adopt different solutions, and that these elements seem to exert their influence regardless of the clause type in which they are found. The third is a consequence of the second: the

LG seem to show the signs of a levelling out of word order distinctions between subordinate and main clauses, thus anticipating a development that would become evident in the Middle English period.

This analysis, as I stated, is partial, and I do not doubt that I have missed many contextual variables that can be usefully considered in relation to periphrastic patterns in the text. Further research is clearly required, and I hope that this paper may serve as motivation for new studies.

EDITION

Skeat, Walter W. (ed.),

The Gospel according to Saint Matthew and according to Saint Mark, Cambridge, 1871-1887.

The Gospel according to Saint Luke and according to Saint John, Cambridge, 1874-1878.

  .The C-P pattern appears again with a pronominal subject in the only independently formed subordinate introduced by ða: ða hia weron faerend = egressis, Mt 9:32. An ambiguous case is ðende waes he spreccende = athuc illo loquente, Mt 26:47. Here ðende probably functions as an adverb, as the subject-copula inversion shows.The only case with an exhortative function shows a C-P pattern: waes milsende = misertus, Mk 9:22. Two periphrases are preceded by a negative particle and, as we would expect, they show a C-P pattern: ne is sellennde = non dandum, Mt XXII; ne waes spreccend = non loquebatur, J 4:34.The periphrasis is immediately preceded by þaet = ut in þaet fore-ondetande weron = ut profiterentur, Lk 2:3, and by miððy = cum/dum in miððy gehatend waes = cum iussisset, Mt 14:19; miððy stigende weron = cum ascendissent, Mt 14:32; miððy gie biðon stondende = cum stabitis, Mk 11:25 (note the pronominal subject); miððy waes faerende = cum processisset, Mk 14:35; miððy woeron spellendo = 11 After writing this article I became aware that a similar case is found in the fifth Blickling Homily (Dominica V in Quadragesima), 40.72-75 in Richard J. Kelly's edition: forþon hi ne besceawiaþ [...] on hwylcum sare hi acennede fram medder waerun. I am grateful to dr. Giuseppe Donato De Bonis for pointing this out to me.

  . Inversion is regularly present in negative questions: quare non abduxistis = forhuon ne gebrohtongie, J 7:45, and the glossator may even find it necessary to independently add a post-verbal pronominal subject in a negative question in which the Latin text puts the subject in preverbal position, as in J 8:46: quare vos non creditis = forhuon gie ne gelefeð gie. In Mk 11:3 and Lk 7:19 tu es (interrogative) is given the alternative renderings ðu arð and arð ðu, the latter obviously against the original's word order. It thus seems quite clear that the glossator did take context into account, though not to a "literary" extent, of course, and that he tried to mediate between his adherence to the source text and the constraints of his own language,

	3	Mt 6:31, 6:31, 6:31, 7:4, 8:29, 9:4, 9:28, 11:3, 11:7, 11:8, 11:9, 12:27, 12:34, 12:48, 13:51, 15:16, 16:8, 18:1,
	20:21, 20:22, 20:32, 21:16, 23:33, 24:45, 26:15, 26:17, 26:50, 26:53, 27:17, 27:46; Mk 1:24, 2:24, 3:33, 4:41, 5:39,

sometimes with clumsy results. The ordering of elements in Mt 24:6 and Mk 5:34 can be thus accounted for by normal Old English syntactic rules. In Mk 10:39 the inversion is probably due to the verb being understood as an exhortative form: it is not unusual for the Latin future tense to be rendered by the Northumbrian imperative mood (e. g. geceig ðu = uocabis, Mt 1:21; worða ðu = adorabis, Lk 4:8) and the Northumbrian gloss, like the West Saxon Gospel version, regularly has inversion in renderings of Latin imperatives and in other expressions of command and wish, consistently with general Old English rules

(Mitchell 1985: § 3897)

.

  the four cases showing P in initial, apparently topicalizing position (gesald he woere = tradendum, Lk LXXIIII; gefreouad we se = liberati, Lk 1:74; aoehtad gé biðon = persequentur, Lk 16:18; gesáld gie biðon = trademini, Lk 21:16) the finite verb retains its post-subject position, as is common in Old English when the subject is a pronoun, although it should be remembered that Northumbrian seems less coherent than West Saxon in this respect, and the LG do show a few divergent cases such as fulle biðon/aro gie = pleni estis, Mt 23:28. It has been estimated that in 10-20% of the cases where the Latin text can be interpreted as having a preposed topic, the pronoun subject inverts with the verb in the LG(Kroch-Taylor 1997: 321-322).The only case showing the independent addition of a pronominal subject and a noun complement is he on rode were genaegled = crucifigeretur, Mt 27:26. The clause is introduced by þaette = ut and is clearly a subordinate. Although the periphrasis is found on the right of the complement, as is common in Old English subordinate clauses and would be expected in an OV Mk 14:32). One can count 47 cases like the example from Lk 9:22 4 , 26 cases like the example from Mk 11:18 5 and three passive relative clauses like the

	4	Mt Arg.

:21. Negative interrogatives are excluded.

In Seven of our "other cases" are impersonal passives showing the addition of an indirect pronominal object. The pronoun's position varies. In four cases it comes immediately before the finite verb: him bið onfoen = adsumetur, Mt 24:40; iuh bið forleten = dimittimini, Lk 6:37; iuh bið sald = dabitur (uobis), Lk 6:38; ðe sint uncuðo = ignoras, J 3:10. The remaining instances have it immediately before the participle: bið him gesald = fiet (ei), Mk 11:23; bið him forgefen = remittetur, Lk 12:10; bið ðe gedoemed = iudicatur, J 3:18. All of these can reasonably be taken as main clauses. The cases from Lk 12:10 and J 3:18 are immediately preceded by ne, whose influence on the copula's position cannot be ruled out. It should be noted, apart from that, that in all of these cases the copula appears somewhere left of the participle.

Cases involving stranded prepositions or particles are in is asaegd = infertur, Lk XXXVII; inn biðon gelaeded = inducantur, Mt 24:24; to sie gefylled = adimpleatur, Mt 13:14. The preposition or particle always precedes the whole periphrasis. It is common for the LG glossator correctly or incorrectly to "dismember", as in the quoted cases, prefixed and compound Latin verbs in translating them, giving rise to what might be taken as syntactic calques (compare eft gie toslitas = rescindentes, Mk 7:13; to gebrohte = attulit, Mk 9:17; wel gie doeð = bene-feceritis, Lk 6:33; in ic foerde = intraui, Lk 7:45). That the observed positioning of stranded prepositions is not entirely due to Latin influence, however, is shown by such instances as aed he were = esse, Mk 6:55.

The Preface to Matthew's Gospel contains two cases in which a coordinating conjunction is added to the periphrasis: & geteled is = referturque; & ec were gesaegd = enarratur. An adverb is added in eft gewoegen bið = remetietur, Mk 4:24; eft forgefen bið = remittetur, Lk 12:10; eft bið gesald = retribuetur, Lk 14:14; sie eft accenned = renatus, J VI. language, the order of its elements is C-P, contrary to the general Old English tendency

(Mitchell 1985: § 3911; Koopman 1992: 322

) and typically of VO languages.

Let us now consider those instances in which a single Latin verb is rendered by a Northumbrian subordinate clause containing a periphrasis of interest (our third category).

The glossator is often brought to render the Latin passive infinitive and ablative absolute (as well as, sporadically, other Latin forms) by means of finite subordinates introduced, respectively, by þaet(-te) and miððy (e. g. þaette sé ofslaegen = occidi, Lk 9:22; miððy ofðon waes gehered = quo audito, Mk 11:18). Relative clauses may also be independently used to clarify passages (e. g. þaet lond ðe is genemned predium = praedium,

  :28.

	Preface	1	2	1	1
	Luke
	Gospel	9	7	1	1
	Preface	0	0	0	0
	John
	Gospel	7	3	1	2
	Total	47	25	7	11

  þaette hia uoero gefroefrende = ut consolarentur, J 11:19), the exceptions being the relative clause ðaðe he wyrcende waes = quae faciebat, Mk 3:8, and biðongie hyngrendo = esurietis, Lk 6:25perhaps an instance of emphatic inversion. It is interesting to compare these renderings to hiu waes The only instance in which the periphrasis is part of a subordinate clause introduced by þaette shows a P-C order: þaette bodande wero = praedicaturus, Lk XXXIII. Four renderings in which subordinate clauses introduced by miððy are independently used can be counted: miððy wutedlice waes he útgeongende = exeunte autem illo, Mt 26:71; miððy waes faerende = egressus, Lk 22:39;miððy he waes sprecende = illo loquente, J 8:30; miððy faerende weron = exeuntes, J 5:13. The instance from Mt 26:71 is of great interest, as it shows a case of verb-seconding with an adverb in initial position, a common phenomenon in Old English main clausesthough not with pronominal subjects, as in this casebut a very unusual one in subordinates, where V2 is only found after bridge verbs (van Kemenade 1997b: 333). In J 8:30 we observe again the fronting of the finite verb in presence of a pronominal subject, as in Lk 19:28, 20:16 etc. Even more interesting is fylgende hine woere = secuturus, Lk XXXVII. This is found in the translation of the Latin docet ut eum quisque secuturus abneget semet ipsum, rendered, somewhat confusedly, thus: laereð þaette hine aelc/sua huaelc fylgende hine woere onsaeca hine seolfne. The rendering is interesting because the pronominal object is placed after the non-finite verba very infrequent and innovative pattern found only in texts later than 950(Fischer et al. 2000: 142) and because the periphrasis shows a P ... C pattern, which is extremely rare, if not unique ("the fourth possibility (V ... v) does not occur",

	Koopman 1992: 319) 11

wyrcenda = operata est, Mt 26:10; hia wero sprecendo = locuturi sint, Lk LI, where the glossator seems again driven to violate the original's word order to let the copula immediately follow the subject. In one case the glossator renders the Latin praecedebat by onfora waes geongend, Mk 10:32.

Mt 3:13, 4:1, 6:1, 6:2, 6:5, 6:16, 6:18, 23:5, 26:2; Mk 15:5; Lk 3:7, 23:32; J 16:32.

order (Mt 6:5, already mentioned). In three cases (Lk 3:21,9:51,22:45) the periphrasis is immediately preceded by miððy = cum/dum. All three have a P-C order. In 52 cases 10 the periphrasis is immediately preceded by a relative pronoun (e. g. ðaðe gihaeled biðon = qui saluantur, Lk 13:23).

The P-C order is represented in 28 of these against 23 instances with C-P, and in one instance two contrasting renderings are given: ðaðe geswoenced woere / weron geswoencde = qui uexabantur, Mk 1:34. It thus seems that the glossator hesitates a lot in such cases, so that no particular ordering of elements neatly prevails over the other: the hesitation is also evident in alternative renderings like þaet we sie dead / þaet we deado sie = quia perimus, Mk 4:38. Such a distribution of C-P and P-C clusters in these subordinates is, however, interestingly similar to that observed by Koopman in the works of AElfric and in the Alfredian translation of the Cura Pastoralis (Koopman 1992: 320).

This is what seems possible to say considering the contextual elements that have the highest probability of influencing the periphrasis's word order. Let us now turn to periphrases consisting of a copula and a present participle (e. g. foerende waes = egrediebatur, Mk 1:5). These are independently attested in 90 cases, which are distributed thus: :16,2:23,4:18,10:2,15:37,25:29,26:3,27:17,27:22,27:33;Mk 1:34,5:36,8:8,10:42,15:7;Lk 1:20,1:35,1:36,2:33,6:15,6:18,7:11,8:2,9:7,10:8,13:23,13:34,18:34,19:10,19:29,21:4,21:37,22:1,22:3,22:47,23:33,23:48;J 1:38,1:42,4:5,4:25,5:2,9:7,9:11,11:54,12:6,13:10,19:14,19:17,20:16,20:24,21:2.