The Neotropical genus Stibadocerina Alexander and its phylogenetic relationship to other...

Systematic Entomology (2009), 34, 324333 DOI: 10.1111/j.1365-3113.2008.00454.x

The Neotropical genus Stibadocerina Alexander and itsphylogenetic relationship to other Stibadocerinaegenera: further evidence of an ancestral trans-Pacificbiota (Diptera: Cylindrotomidae)

GU I LHERME C . R I B E I RODepartamento de Biologia, Faculdade de Filosofia, Ciencias e Letras de Ribeirao Preto, Universidade de Sao Paulo,Ribeirao Preto, SP, Brazil

Abstract. Stibadocerina Alexander, a monotypic genus, includes the only knownNeotropical species of the family Cylindrotomidae, S. chilensis Alexander, 1929,from South Central Chile (ca. 368509S428179S). In this paper, Stibadocerinachilensis is redescribed and illustrated in detail. A study of wing-vein homology inthe subfamily Stibadocerinae is provided, to identify the components of thereduced radial sector in Stibadocerina and related taxa. The proposed hypothesesof wing-vein homology are tested, and the systematic position of Stibadocerina isassessed through a cladistic analysis of 13 characters of the male imago, scored forexemplar species of the four genera included in the Stibadocerinae. A single mostparsimonious tree supports the monophyly of the Stibadocerinae and thefollowing relationships among its included genera: Stibadocerodes [Stibadocera(Stibadocerella Stibadocerina)]. The subfamily includes one example of a vicar-iant distribution with a sister-group relationship between South Central Chileanand East Asian taxa, and supports a biogeographical interpretation of an ancestraltrans-Pacific biota.

Introduction

The Cylindrotomidae is the smallest of the four families ofTipulomorpha sensu stricto, with 71 extant species. Most of

the genera and species belong to the subfamily Cylindrotomi-nae, which is distributed mainly in the Nearctic and Palae-arctic regions. The Stibadocerinae includes four genera:

Stibadocera (12 species; OrientalAustralasian), Stibadocerel-la (four species; OrientalEastpalaearctic), Stibadocerodes(three species; Australasian) and Stibadocerina (one species;

Neotropical) (Oosterbroek, 2008). Only the subfamily Cylin-drotominae is known in the fossil record, mostly fromTertiarystrata from North America and Europe (Evenhuis, 1994).

Stibadocerina chilensis is the sole member of the genusStibadocerina Alexander, 1929, and the only species ofCylindrotomidae from the Neotropical region. The specieswas described mostly from specimens collected during an

expedition to South Chile led by F. W. Edwards and R. C.Shannon, between November and December of 1926. TheDiptera collected by the expedition were studied in a series

of monographs published by the British Museum (NaturalHistory), the Diptera of Patagonia and South Chile series, ofwhich the first volume, dealing with the crane flies, was

published in 1929 (Alexander, 1929).My main purpose is to revise Stibadocerina chilensis,

providing a more detailed morphological study of the taxon,and to assess its phylogenetic relationship to other genera

within the Stibadocerinae. The wing venation in S. chilensisis quite distinct and reduced compared with the patternscommonly found in the Cylindrotomidae. Therefore, to

understand the possible identities of the wing veins of theradial sector in S. chilensis and related taxa better, a study ofthe wing-vein homology in the Stibadocerinae is presented.

Correspondence: Guilherme C. Ribeiro, Departamento de Biol-

ogia, Faculdade de Filosofia, Ciencias e Letras de Ribeirao Preto,

Universidade de Sao Paulo, Avenida Bandeirantes 3900, 14040-901

Ribeirao Preto, SP, Brazil. E-mail: [email protected]

324

# 2009 The AuthorJournal compilation # 2009 The Royal Entomological Society

SystematicEntomology

The systematic position of Stibadocerina is assessed, and apreliminary phylogeny of the Stibadocerinae is provided,followed by a short discussion of biogeographical implications.

Materials and methods

Studied specimens of Stibadocerina, Stibadocera and Stiba-docerella belong to the Natural History Museum, London,

U.K. (BMNH). Other specimens used in the comparativestudy are from the collections of the Zoologisch Museum,Amsterdam, The Netherlands (ZMAN), the United StatesNationalMuseum, Smithsonian Institution, U.S.A. (USNM),

and the Departamento de Biologia, FFCLRP-Universidadede Sao Paulo, Ribeirao Preto, Brazil (DBRP). Details on theexamined specimens of Stibadocerina are provided under the

redescription of the species. Details on the other specimensused for phylogenetic analysis are given in Appendix 1.

For most characters, the descriptive terminology followsMcAlpine (1981), with terminology for male gonostylusstructures following Ribeiro (2006). The adopted terminol-

ogy for the wing veins accords with the results of thehomology study below.Male terminalia were cleared with warmed KOH and

mounted in non-permanent slides with glycerol. After study

and illustration, the dissected parts were transferred tomicrovials and pinned with their corresponding specimens.Illustrations were produced with drawing tubes attached to

stereoscopic and compound microscopes. Measurementswere taken with an ocular reticule.To test the phylogenetic information of the primary

homology hypotheses for the wing veins, and to elucidatethe phylogenetic position of Stibadocerina, 13 characters fromthe male imago were scored for species representing the fourgenera of Stibadocerinae as the in-group, plus two genera of

the subfamily Cylindrotominae and six exemplars of thefamilies Limoniidae and Tipulidae as out-groups. Characterswere scored from the direct observation of specimens for

all taxa except Stibadocerodes zherikhini and Phalacrocerareplicata, which were based on literature (Brodo, 1967;Krzeminski, 2001). All characters were considered as unor-

dered. Character polarity was determined a posteriori withrooting using the out-group method. The characters aredescribed in Table 1, and the datamatrix is shown in Table 2.

The data matrix was analysed in TNT (Goloboff et al., 2003)with tree bisectionreconnection (TBR) branch swapping,random stepwise addition and 1000 replications holding upto 10 trees. The matrix was analysed using both prior (equal)

weights and implied weights (with k varying from 2 to 6).

Results and discussion

Homology of wing veins in Stibadocerinae

In crane fly systematics, the homology and nomenclatureof the wing veins has been a contentious issue, with different

systems currently in use by different authors. The situation

Table 1. Characters scored for phylogenetic analysis. Consistency

index within square brackets.

1. Proportion between the length and width of first and second

flagellomeres: less than 3 longer than wide (0); more than3 longer than wide (1). [0,5].2. Vein Sc: reaching the wing margin (0); atrophied and not

reaching the wing margin (1). [1,0].

3. Vein R1: reaching the wing margin (0); atrophied at tip and not

reaching the wing margin (1). [0,5].

4. Vein R1: ending on C at a more distal position (0); ending on

C in a more proximal position (1). [1,0].

5. Vein r-r: transversal in position (0); oblique in position, but still

distinguishable (1); aligned with R1 and distal section of R2 (2). [1.0].

6. Vein R23: shorter than one-half the length of R3 (0); longer thanone-half the length of R3 (1); absent (2). [0,66].

7. Basal section of R2 (bR2): long (0); short (1). [1,0].

8. Basal section of R2 (bR2): sinuous (0); straight (1). [1,0].

9. Basal section of R2 (bR2): longitudinal in position (0);

sub-perpendicular in position (1). [1,0].

10. Vein R45: present (0); absent (1). [1,0].11. Vein M12: bifurcated (0); not bifurcated (1). [0,5].12. Aedeagus: simple (0); trifid (1). [1,0].

13. Gonostylus: 2-branched with well-developed clasper and lobe

(0); 2-branched, but with a reduced lobe (1); with a single

branch (2). [0,66].

Table 2. Data matrix for phylogenetic analysis. Inapplicable characters coded as .

1 2 3 4 5 6 7 8 9 10 11 12 13

Edwardsomyia chiloensis 1 0 0 0 0 0 0 0 0 0 0 0 0

Tinemyia margaritifera 0 0 0 0 0 0 0 0 0 0 0 0 0

Ptilogyna sp. 0 0 0 1 1 1 0 0 0 0

Leptotarsus (Longurio) gymnocerus 0 0 0 0 1 1 0 0 0 0

Cylindrotoma distinctissima 0 1 0 1 1 1 0 0 1 2

Phalacrocera replicata 0 1 0 1 1 1 0 1 1 2

Stibadocerodes zherikhini 1 0 1 1 2 0 0 0 1 1 1 0

Stibadocera sp. 1 0 0 1 1 0 1 0 1 1 1 1 1

Stibadocerella sp. 1 0 1 2 1 1 1 1 1 1 1 0

Stibadocerina chilensis 1 0 1 2 1 1 1 1 1 1 1 2

The Neotropical genus Stibadocerina 325

# 2009 The AuthorJournal compilation # 2009 The Royal Entomological Society, Systematic Entomology, 34, 324333

is especially problematical regarding the identity of the veins

of the radial sector, which shows much plasticity and high

levels of homoplasy.Theproposal of homology relations forwing veins, and the

understanding of the possible modifications leading from

more generalized to highlymodified patterns are biased, after

all, by the proponents conceptions of what constitutes the

plesiomorphic condition for the various wing-vein charac-

ters, and what would constitute plausible changes. When

phylogentic studies are lacking, such ground-plan formula-

tions can be quite subjective and intuitive. In the absence of

information on the developmental paths and the underlying

mechanisms behind changes in the venation, judgments on

the plausibility of certain transformations are also very

subjective and difficult to test more directly.A recent study of the early phylogenetic patterns of crane

flies (Ribeiro, 2008) allowed a better understanding of the

possible ground-plan condition of the wing venation in theTipulomorpha sensu stricto. Regarding the radial sector, the

following conditions seem likely to be present:

1. The vein R1 reaches the wing margin.2. The cross-vein r-r is present, perpendicular in position,

and is closer to the mid-point of vein R2 than to either itsorigin or its apex.

3. Veins R2 and R3 both reach the wing margin and runmore or less in parallel or only gradually diverging fromeach other.

4. The petiole of cell r2 (vein R23) is short, or moreprecisely, shorter than one-half the length of vein R3.

5. Veins R4 and R5 are fused in a single element R45reaching the wing margin. There is some doubt whetherthis condition for R45 is the ground-plan condition ofthe entire Tipulomorpha sensu stricto, because at least

one species in the family Pediciidae (Tricyphona protea,

Figs 19. Proposed homology for the

wing veins. 1, Edwardsomyia chiloensis

(Limoniidae: Limnophilinae); 2, Stibadocer-

odes australensis (Cylindrotomidae: Stibado-

cerinae); 3, Stibadocerodes tasmanensis

(Cylindrotomidae: Stibadocerinae); 4, Stiba-

docerodes zherikhini (Cylindrotomidae:

Stibadocerinae); 5, Stibadocera sp. (Cylin-

drotomidae: Stibadocerinae); 6, Stibadocer-

ella sp. (Cylindrotomidae: Stibadocerinae);

7, Stibadocerina chilensis (Cylindrotomidae:

Stibadocerinae); 8, Phalacrocera replicata

(Cylindrotomidae: Cylindrotominae); 9,

Cylindrotoma distinctissima (Cylindrotomi-

dae: Cylindrotominae).

326 G. C. Ribeiro


figured in Alexander & Byers, 1981, figure 38) seems tohave retained a free vein R5. However, with the likely

position of Pediciidae as the sister group of all otherTipulomorpha sensu stricto (Ribeiro, 2008), it seemsquite safe to assume that a single element R45 is presentin the ground-plan of the other Tipulomorpha exceptPediciidae. The conditions described above are preservedin many Limoniidae genera, of which Edwardsomyia isa good example (Fig. 1).

Instead of considering previous hypotheses of homology

and the nomenclature applied to the Cylindrotominae (e.g.Brodo, 1967; Alexander & Byers, 1981), the comparisonsmade here start from the acceptance of the ground-plan

conditions as described above.Among all Stibadocerinae genera, Stibadocerodes

(Figs 24) is the one in which the characters of the radial

sector match most closely that of the ground-plan. Withoutassuming much transformation from the basic pattern, themost obvious difference is the lack of the element R45 in allspecies, and the loss of the petiole of cell r2 (vein R23) inS. tasmanensis (Fig. 3) and S. zherikhini (Fig. 4). Further-more, the tip of R1 obviously is lacking in all species. InS. australensis (Fig. 1), a fragment of the tip of R1 remains,

but with vein r-r positioned as in the original configuration.However, in S. tasmanensis and S. zherikhini, (Figs 3, 4) thecross-vein r-r seems captured by the basal remnants of R1,

producing a single element (R1 r-r).In Stibadocera (Fig. 5), the situation is more distant

from the original conditions. Although still present, the tipof R1 is positioned in a much more proximal location, and

the vein r-r assumes a more oblique position. The petiole ofcell r2 (vein R23) is still present. The anterior displace-ment of R1 seems to have caused a distortion in the basal

section of vein R2 (bR2), which is more inclined and shorterthan in Stibadocerodes. Nonetheless, there seems to belittle doubt, within the framework of the comparisons

being made here, that the sinuous, sub-perpendicularelement in the radial sector of Stibadocera is the basalsection of vein R2 (bR2).

We now approach the more apomorphic conditions of theradial sector as found in Stibadocerella (Fig. 6) and Stiba-docerina (Fig. 7). Understanding the identities of these veinsin these genera would be very difficult if they were taken in

isolation; however, the study of the identity of the veins inrelated genera furnishes evidence of what could havehappened in these derived groups.

Starting from a condition similar to that of Stibadocera,a simple change, the loss of the tip of R1, could result ina dramatic change in the overall appearance of the radial

sector. The vein r-r would, as in the case of Stibadocerodestasmanensis and S. zherikhini, be captured by the basalremnants of R1. In this case, as a result of the more inclined

position of r-r, its capture would result in a single elementR1 r-r almost continuous with, or aligned with, the apicalsection of vein R2. These transformational events, togetherwith the elongation of the vein R23, are all that needs to beassumed to conclude that the first longitudinal vein of the

radial sector in Stibadocerella and Stibadocerina formsa composite vein comprising an element (R1 r-r) alignedor almost aligned with the distal section of R2. The sub-perpendicular or almost transversal element of the sector isthe basal section of vein R2 (bR2). The second longitudinal

vein is the primitive R3.The wing venation of other Cylindrotominae, Phalacro-

cera replicata (Fig. 8) andCylindrotoma distinctissima (Fig. 9)are shown for comparison.

Taxonomy

Genus Stibadocerina Alexander, 1929

Stibadocerina Alexander, 1929: 66. Type species: Stiba-docerina chilensis Alexander, 1929.

Diagnosis. Stibadocerina differs from Stibadocerodes andStibadocera mainly by having the vein r-r indistinguishable,

aligned with both R1 and the distal section of R2. It differsfrom Stibadocerella by keeping the second anal vein (lost inStibadocerella) and having a single, unbranched gonostylus.

Stibadocerina chilensis Alexander, 1929

(Figs 7, 1016)

Stibadocerina chilensis Alexander, 1929: 66 (originaldescription); Plate II, figure 37 (wing venation). Alexander& Alexander, 1970: 4.44 (catalogue citation); Oosterbroek,

2008 (catalogue citation).

Colour (male and female). Head dark brown; antenna,rostrum and palpus brownish; pronotum light brown; legs

mostly light brownyellowish, with last 43 tarsal segmentswhite. Scutum brown; lateral thoracic sclerites mostlybrown, light brownyellowish near attachment of wing

and halter; wing with a brownish tinge; mesothoracic andmetathoracic coxae and base of first abdominal segmentlight brownyellowish; abdomen from second segment to tip

uniformly brown.

Dimensions (male; maximum lengths and widths in

mm). Head length, 0.53; head width, 0.530.60; winglength, 5.625.93; wing width, 1.31; gonocoxite length,0.22; gonocoxite width, 0.15; gonostylus length, 0.16.

Morphology. Head and appendages (Fig. 10): antennalonger in male than in female; flagellum 13-segmented,covered with verticils shorter than individual flagellomeres;

flagellomeres cylindrical, decreasing in length towards tip ofantenna; first flagellomere ca. 10 longer than wide in male,ca. 5 longer than wide in female; pedicel ca. 1.7 longerthan scape; palpus 4-segmented; palpomeres more or lesscylindrical; last palpomere almost as long as precedingsegments together; rostrum (including labella) ca. 0.30the total length of head; compound eyes widely separated



dorsally and ventrally. Thorax and appendages: thoraxalmost as long as high; pleural sclerites as figured (Fig. 10);tibial spurs absent; tarsal claw simple. Wing (Figs 7, 11, 12):

vein h situated at mid-length between the origin and the forkof M Cu; Sc running very close to R, ending on C just

distally of the origin of Rs; sc-r faint or absent, when presentplaced at tip of Sc; Rs almost straight, except for slightcurvature at origin; two elements of Rs (R2 and R3) reaching

wing margin; vein R23 almost as long as R3; basal sectionof R2 (bR2) straight, sub-perpendicular; vein M with tree

Fig. 10. Stibadocerina chilensis. Head

(anterolateral view) and thorax (lateral

view). Abbreviations: anatg, anatergite;

anepm, anepimeron; anepst, anepister-

num; anepst cleft, anepisternal cleft; aprn,

antepronotum; comp eye, compound eye;

cx, coxa; kepm, katepimeron; kepst, kate-

pisternum; ktg, katatergite; lbl, labella; ltg,

laterotergite; mr, meron; mtanepst, meta-

nepisternum; mtepm, metaepimeron; mtg,

mediotergite; mtkepst, metakatepister-

num; ped, pedicel; plp, maxillary palpus;

pprn, postpronotum; scp, scape; sct, scu-

tum; sctl, scutellum.

Fig. 11. Stibadocerina chilensis. Wing venation.

328 G. C. Ribeiro


branches, M12, M3 and M4; m-cu placed slightly proximalto mid-point of discal cell; Cu strongly curved; A1 running

very close to Cu at its basal section, gently curved andreaching the wing margin at the level of the origin of R23;A2 slightly sinuous, reaching wing margin well before the

origin of Rs. Male terminalia (Figs 1315): ninth tergumand sternum separated, not forming a contiguous ring; ninthtergum produced into a single small median lobe; gonocox-ite globular, almost as long as high, with a large ventro-

medial hairy projection; gonostylus simple, unbranched,gradually narrowed towards tip, slightly curved at apex;

aedeagus trifid, with medial branch slightly longer thanlateral branches; lateral process of aedeagal sheath welldeveloped and stout, branching off from the sheath

obliquely, not in parallel with the aedeagus; interbaseslender, almost straight from base to apex, bearing a rela-tively stout and well-developed lateral extension articulatingwith gonocoxite apodeme, and a similar posterior extension

Fig. 12. Stibadocerina chilensis. Detail of

wing venation, showing the position of the

tip of Sc vein in relation to other veins, not

visible in Fig. 11.

Fig. 13. Stibadocerina chilensis. Male terminalia, dorsal view.

Abbreviations: aed, aedeagus; goncx, gonocoxite; gonst, gonostyle;

t9, ninth tergite.

Fig. 14. Stibadocerina chilensis. Male terminalia, dorsolateral view.

Abbreviations: aed, aedeagus; goncx, gonocoxite; gonst, gonosty-

lus; s8, eighth sternite; t8, eighth tergite; t9, ninth tergite.



articulating with paramere; interbases connected to each other

medially. Female terminalia (Fig. 16): tenth tergite approxi-mately as long as cercus,more or less triangular in lateral view;cercus likea stout curvedblade; hypogynial valve reachingmid-

length of cercus, with its tip aligned with apex of tenth tergite.

Distribution. Stibadocerina chilensis is known to occuronly in South Central Chile, with its northernmost limit at

Concepcion (ca. 368509S, 738009W) and its southernmostlimit at Mechuque Island (ca. 428189S 738159W).Examined material (label information in italics; informa-

tion of different labels separated by a vertical line; geo-graphical coordinates within brackets): HOLOTYPE. #. S.Chile: Llanquihue prov. F.&M.Edwards. B.M. 1927-63. |Peulla. 12-13.xii.1926. | BMNH(E)#246146. (418039S718019W); ALLOTYPE. $. S. Chile: Llanquihue prov.,F.&M. Edwards., B.M.1927-63. | Peulla., 12-13.xii.1926.(418039S 718019W); PARATYPES. 3 #. S. Chile: Llanquihueprov. F.&M.Edwards. B.M. 1927-63. | Peulla. 12-13.xii.1926.(418039S 718019W); PARATYPE. #. S. Chile: Llanquihueprov. F.&M.Edwards. B.M. 1927-63. | Casa Pangue, 4-

10.xii.1926. (418029S 718519W) PARATYPE. $. S. Chile:Chiloe I., F. & M. Edwards., B.M.1927-63 | Mechuque I.,23.xii.1926(428189S 738159W).

Phylogenetic position of Stibadocerina

The parsimony analysis of the data matrix in TNT (Goloboffet al., 2003) using both prior (equal) weights and implied

weights (with any value of k) results in the same single mostparsimonious tree (21 steps; CI 0.761; RI 0.833), of whichthe relationships for the in-group taxa are shown in Fig. 17.The monophyly of the Cylindrotomidae (Clade A) is

corroborated by the trifid aedeagus, a synapomorphyunique to this family within the Tipulomorpha sensu stricto.The monophyly of the Cylindrotominae (Clade B) is

supported also by at least two synapomorphies: the loss of

Fig. 15. Stibadocerina chilensis. Aedeagus andassociated structures.

Abbreviations: aed, aedeagus; aed apod, aedeagus apodeme; interb,

interbase; lp, lateral process of aedeagal sheath; pm, paramere.

Fig. 16. Stibadocerina chilensis. Female

ovipositor. Abbreviations: cerc, cercus;

hyp vlv, hypogynial valve; s8, eighth ster-

nite; t10, tenth tergite.

330 G. C. Ribeiro


one of the gonostylar branches (character 2, state 1) and theatrophy of the tip of vein Sc (character 13, state 2).

The monophyly of the Stibadocerinae (clade C) is cor-roborated by the long flagellomeres (character 1, state 1)and the loss of vein R45, a character state unknown to haveoccurred in any other lineage within the Tipulomorpha.Stibadocerodes constitutes the sister-group to all other

Stibadocerinae. The clade Stibadocera (Stibadocerela Stibadocerina) (Clade D) is grouped on the basis of the

reduction and inclination of the basal section of vein R2(character 7, state 1, and character 9, state 1).The monotypic Stibadocerina is placed as the sister-group

of Stibadocerella (Clade E). The synapomorphies unitingthese taxa are the alignment of the vein r-r with R1 and thedistal section of R2 (character 5, state 2), and the straight

basal section of vein R2 (bR2) (character 8, state 1). Thispicture is consistent with the opinion of Alexander (1929)who, in describing Stibadocerina, pointed to Stibadocerellaas its most closely related taxon.

The sister-group relationship between Stibadocerina andStibadocerella seems well supported. The monophyly ofStibadocerella also is well supported by the loss of the

second anal vein. However, to test the monophyly ofStibadocerodes and Stibadocera a taxonomic revision andphylogenetic study including more taxa is necessary.

Trans-Pacific disjunction

The subfamily Cylindrotominae is widespread in the Holarc-tic Region, but the Stibadocerinae has a muchmore restricted

distribution: Stibadocerina is endemic to South Central Chilebetween ca. 368509S and 428189S; Stibadocerella is knownfrom the eastern part of the Oriental region; Stibadocera hasan East OrientalAustralasian range; and Stibadocerodes is

restricted to New South Wales and Tasmania (Fig. 18).Recent studies have reinforced the biogeographical affin-

ities between components of the faunas and floras on both

sides of the Pacific Ocean (for a recent review and examples,

Fig. 17. Cladogram depicting relationships among exemplar spe-

cies of Cylindrotominae and Stibadocerinae genera. Unique and

homoplastic characters represented by closed and open circles,

respectively.

Fig. 18. Distributions of the genera Sti-

badocerodes (1), Stibadocera (2), Stibado-

cerella (3) and Stibadocerina (4), with

cladogram superimposed on it. The map

is schematic and not to scale.



see Grehan, 2007). Among these groups there are examplesof taxa with limited dispersal capabilities, indicating vicar-

iance as a major cause for multiple trans-Pacific disjunctions(McCarthy et al., 2007).A list of putative trans-Pacific sister areas is given by

McCarthy (2003), based on evidence from several studies,including examples from the Diptera (revised in Cranston,2005). According to McCarthy (2003), the area labelled asSouth Central Chile (358S428S) is probably the sister-areaof Tasmania (408S438S), whereas Northern Australia andIndochina (238N238S) (i.e. the area including the distribu-tional ranges of both Stibadocera and Stibadocerella) is the

sister group of northern South America and southern NorthAmerica (238N238S).As pointed out by Cranston (2005), there are few examples

of trans-Pacific disjunctions within the Diptera, and this studyprovides one particular case in the Tipulomorpha. The trans-Pacific clade Stibadocerella Stibadocerina adds furtherevidence in favour of an ancestral biota centred around amore

spatially restricted Pacific basin, as opposed to the conven-tional interpretation with a Panthalassan Ocean occupying thenon-continental half of the globe (and thus implying trans-

Pacific disjunctions as derived from long-distance dispersals orrelicts of prior vicariance events). However, the closestrelationship as indicated by this study, namely that between

the northern part of the Australasian and the eastern part ofthe Oriental regions (ca. 238N88S) to South Central Chile(368S468S), does not fit exactly in the framework of trans-Pacific sister areas given by McCarthy (2003). This may beevidence of a more complex picture, and indicate that addi-tional evidence is necessary for a better understanding of thecomplex relationships of the areas around the Pacific basin.

Acknowledgements

I am very indebted to Dr Erica McAlister (Natural HistoryMuseum, London, U.K.), Dr Pjotr Oosterbroek (Zoolog-isch Museum, Amsterdam, the Netherlands) and Dr WayneMathis (Smithsonian Institution, Washington DC, U.S.A.)

for the loan of the specimens used for this study, and to DrJaroslav Stary for the exchange of literature. Special thanksto Dr John R. Grehan for advice concerning other cases of

trans-Pacific disjunctions. At the time this research wasconducted, I was supported financially by a post-doctoratefellowship from FAPESP.

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Accepted 30 October 2008

First published online 22 January 2009

Appendix 1.

Information on the specimens used for comparative study.Each entry is a different specimen. The specimens ofStidabocerina chilensis are listed elsewhere in the text.

Limoniidae

1. Edwardsomyia chiloensis Alexander, 1929. Paratype, #,CHILE: Ancud, 18.xii.1926 (Shannon) (USNM).2. Edwardsomyia chiloensis Alexander, 1929. 1#, CHILE:

Chiloe I., Aucar, 6-15.i. 1952 (Pena) (USNM).

332 G. C. Ribeiro


Tipulidae

3. Leptotarsus (Longurio) gymnocerus (Alexander, 1938).Paratype, #, BRAZIL: Marambaia, 1100 m, 2.xii.1935(Zikan) (USNM).4. Leptotarsus (Longurio) gymnocerus (Alexander, 1938).

Paratype, #, BRAZIL: Marambaia, 1100 m, 2.xii.1935(Zikan) (USNM).

5. Ptilogyna sp. 1#, BRAZIL: Sao Paulo, Salesopolis, E.B. Boraceia, Ponte Rio Claro, 14.xi.2003 (G. C. Ribeiro)(DBRP).

6. Ptilogyna sp. 1#, BRAZIL: Sao Paulo, Salesopolis, E.B. Boraceia, Ponte Rio Claro, 14-16.xii.2003 (G. C. Ribeiro)(DBRP).

Cylindrotomidae (Cylindrotominae)

7. Cylindrotoma distinctissima (Meigen, 1818). 1#,FRANCE: 23km N Sospel, pine forest, 1500 m, 14.vi.1997(P. Oosterbroek and C. Hatveld) (ZMAN).

8. Cylindrotoma distinctissima (Meigen, 1818). 1#,FRANCE: 23km N Sospel, pine forest, 1500 m, 14.vi.1997

(P. Oosterbroek and C. Hatveld) (ZMAN).

Cylindrotomidae (Stibadocerinae)

9. Stibadocera sp. 1#, MALAYSIA: Sarawak, Mt. Dulit,R. Koyan, Primary Forest, 2500 ft, 21.xi.1932 (B. M. Hobbyand A. W. Moore) (BMNH).10. Stibadocera sp. 1#, MALAYSIA: Sarawak, R. Kapah

trib., of R. Tinjar., 25.x.1932 (B. M. Hobby and A. W.Moore) (BMNH).11. Stibadocera sp. 1#, MALAYSIA: Penang, Penang

Hills, Ayer Itam 1000, 18.xii.1963 (H. T. Pagden) (BMNH).12. Stibadocerella sp. 1#, MALAYSIA: Sarawak, Mt.

Dulit, Moss forest, 4000 ft, 25.x.1932 (B. M. Hobby & A. W.Moore) (BMNH).

13. Stibadocerella sp. 1#, JAVA: Tjibooas, 4000 ft. i.1936. (L. E. Cheesman) (BMNH).



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