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Je (ed. O. Kudrna),liption price (when

~ can agree with some(ample, that "None thes of rare or region~lIylccessful, entirely mission"! Who has missedla.';t realized that agri-;lhle for the. production)1' nature, ~~d they areI requests to introduce; or that animal group".bioindicators, "whose

hose of the "rest" of the~an be agreed that bUI-should we forget othercrved together with the:Juld we not usc sevemllindicatorsofa valuablenolnC{;essarily such a'alion (rcd data books,mmnnity conservation

lic part of the book wi 11cs to cladistics secm to•many trivial detaiis onsubspecies should be

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GE/.R E. E. 8("/ 1

Anna/es Entom%gici Fennici 54:107-113.1988

Rocetelio'n, a new Holarctic genus of the Keroplatidae (Diptera,Mycetophiloidea): Description, phylogenetic and biogeographicnotes

Loi'c Matile

Matile, L. 1988: Rocetelion, a new Holarctic genus of the Keroplatidae (Diptera,Mycetophiloidea): Description, phylogenetic and biogeographic notes. - Ann. Entomol.Fennici 54: i07-113.

The genus Rocetelion is proposed for the three nearctic species Cerotelion fascia/urnGarrett, Keroplatusfasciolus Coquilletl and K.fenestralis Fisher, and for the palaearctic K.hllmeralis Zetterstedt. The new genus is mainly distinguished by its male gonostyle, whichsuggests asister-group relationship with the afrotropical genusParacerotelion MatHe, ratherthan with Cerute/ion Rondani or Eliceroplatus Edwards. The main characters of the genusand a key to the species are given. The phylogenetic relationships established, and thebiogeographic conclusions which may be drawn from them, a~sign to the species of the genLL~a minimum age of 40 million years.

L. Matile,Laboratoired' Enlomologie, Museum nationald' Histoire naturel!e, 45, rUl!. deBuffon, F-75005 Paris, France

Index words: Taxonomy, Keroplatidae, Holarctic, Rocetelion gen. n., new combina·tions, key to species, phylogeny, biogeography

'ors or lapses. It is se-:1110t normally inhahitnage. could extend il~ufly species is known1'0 pages laler Polyom-;l-:

108 Matile: Rocetelion, a new Holarctic genus of the Keroplatidae ...

"Mapewate", Arkansas. Laffoon (1965) did not retainthis locality (see below) and added California to thedistribution of the species. No other records are given,but the distribution ofthe known specimens is typicallywestern, a fact which makes me very doubtful abouteither the identification or the locality of the specimenofR .jasciolum cited by Fisher from Arkansas. She wasunable to find Mapewate on any map and as the labelis badly written, she suggests that it could be Maple-vale, Pulaski Co. There are several Pulaskis in the U.S.,but I suggest that the real locality ofthe specimen mightbe Maple Valley, in the State of Washington.

Cerotelion 'Jasciatus" was described by Garrettfrom British Columbia. Fisher, in her thesis (1937), leftthe species in Cerotelion, but later (1941) placed it inKeroplatus (Euceroplatus), adding to this distributiontwo localities from the States of Washington andCalifornia.

Fisher (1937) had been unable to place Cerotelionjasciatum in any of the subgenera ofher"Ceroplatus",a group which she recognized as polyphyletic. In 1938,she descri~~ Keroplatus fenestralis from Michiganand placed it in the subgenus Euceroplatus. In 1941,having been able to study a male of C.jasciatum, sheconsidered herjenestralis to be a "variety" ofGarrett'sspecies (but nomenclatorially treated it as a sub-species). Moreover, she put forward the hypothesisthat both jasciatwn and fenestralis might be onlyvarieties of K. fasciolus, the male of which was then,and still is, unkown. She placed these taxa inEuceroplatus on the basis of the arrangement of thetibial setulae in regular rows.

The genitalia of the two North American speciesfor which males are known, and of the Europeanspecies K. humeralis, are very different from those ofEuceroplatus, Cerotelion and a new genus erected forthe species-group "Euceroplatus" bellulus Williston,as defined in MatHe, 1986. I propose to erect the newgenus Rocetelion for these species. The new taxon ischaracterized by the dorsoventral flattening of the malegonostyles, which also bear an internal sclerotisedplate, by the small cerci, and the presence of a largemembranous area ventrally dividing the gonocoxalsynsclerite. Moreover, it can be distinguished fromEuceroplatus and Cerotelion by the scutellum bearingseveral rows of short apical setae, and by the elongatefore tarsus.

Like Fisher, I was able to study the type of R.jasciolum, some of the metatypes ofR.fasciatum, anda specimen of R. jenestrale (from Alaska), whichagrees perfectly with the original description and thedrawing of the hypopygium given by Fisher (1938).

The male genitalia ofR.fpsciatum andR.fenestrale aredistinctive (cf. Figs. 11-12 and 13-14). As regards R.fasciolum, it differs from the other species of Roce-telion in the presence ofa distinct antennal apicule, andseveral other characters, and cannot therefore beconspecific with any of them.

Rocetelion gen. n.

Type-species: Cerotelion fasciatus Garrett.

Derivatio nominis: anagram of Cerotelion. Gen-der: neutral.

Species: R. fasciatum, comb. n. (Cerotelion fas-ciatus Garrett, 1925:12); R.fasciolum, conlb. n. (Cero-platus fasciolus Coquilleu, 1894:126); R. fenestrale,comb. n. (Ceroplatusfenes~ralis Fisher, 1938:197); R.humerale, comb. n. (Ceroplatus humeralis Zetterstedt,1850:3445).

While not fonnally named, the genus was de-scribed and illustrated in detail in Matile, 1986. Ashorter diagnosis is given here.

Head (Figs. 1-2) wider than high. Three ocelli, themedian one smaller, the lateral large, their distancefrom the eye margin 1.5 times their own diameter (R.jasciatum, R .jasciolum) or almost twice their diameter(R .fenestra Ie,R. humerale). Eyes deeply emarginated,pilosity short. Antennae: scape and pedicel short andcylindrical. Fourteen flagellomeres, widened andflattened (Fig. 3). Last flagellomere much longer thanwide, without tenninal apicule, except in R.fasciolum,in which it is small and rounded. Face wide, bare andweakly sclerotised, except in R.fasciolum, in which itis more strongly sclerotised, and bears a few ventralsetae. Labella short, membranous on the inside.Palpifer small and well sclerotised. First palpomerevery small, second large and porrect.

Thorax (Fig. 5): scutum weakly arched, evenlycovered with short setae, the lateral and prescutellarsetae longer. Scutellum bare on disc, except in R.jenestrale and R. humerale, where it bears one or twopairs ofdiscal setae, marginal setae numerous, short, inseveral rows. Mediotergite strongly jutting out fromscutellum, rounded at apex, bare. Pleurae bare exceptproepisternum and antepronotum, and mesan-episternum, which bears a group of short dorsal setae.

Coxae with posterior apical setae"except II-III inR.fasciatum and R.fasciolum. Tibial setulae regularlyand irregularly arranged. Tibia I with regular rowsexept at base in R. humerale and R.fenestrale, on theapical half of the anterior face and almost all of the

Figs. 1-5. Roc~(Garrett). 1-2) hea~view; 3) antenna)view; 4) wing. dorcoxae and frrst allateral view.

outer face inR.jand outer faceswith regular rcappearing as (magnification,regular rows, althe anterior fa'fasciolum (tibiavailable specirrows, some clo~species. Spurs ]length of the ouII with a reduclbetween spurs, lcomb, a smalllarge, but madeprotarsus I 2.3from 1.6 to 1:serrulated, witl

Wing (Fig.lobereduced;nR.fasciolum ananterior margin(R.fasciatum) 4weak, situatedoblique (a litt]reaching wingR4+5, R5 and .Iside.

Male abdonfollowing prog

Ann. Entomol. Fennici 54:3. 1988 109

and R fenes tra/e are1-14). As regards R.er species of Roce-llltennal apicule, andannot therefore be

IS Garrett.

)f Cerotelion. Gen-

n. (Cerotelion fas-rum, comb. n. (Cero-:126); R.fenestra/e,;isher, 1938: 197); R.uneralis Zetterstedt,

Figs. 1-5. Rocetelion fasciaturn(Garrell). 1-2) head,frontaland lateralvicw; 3) antennal flagellum, lateralview; 4) wing, dorsal view; 5) thorax,coxae and first abdominal segment,lateral view.

Figs. 9-10. Rocetelionfasciatum (Garrett), structure of femalegenitalia (setae not figured). 9) ventral view; 10) lateral view. -Scale line 0.3 mm.

109

apical wider than the middle ones. Female: abdomenwidened from the second segment on, then stronglynarrowed at the seventh.

Figs. 6-8. Rocetelion fasciatum (Garrett), structure of malegenitalia (ordinary setae not figured). 6) dorsal view; 7) ventralview; 8) phallosome. - Scale line 0.3 mm.

outer face inR fascio/um, and on almost all the anteriorand outer faces in R. fasciatum. T II almost entirelywith regular rows, some of them more closely set,appearing as conspicuous black lines under weakmagnification, in R. fasciatum and R. humera/e;regular rows, also with black lines, on the apical halfofthe anterior face and almost all the outer face in R.fascio/um (tibiae and tarsi II broken on the onlyavailable specimen ofR fenestrale). T III with regularrows, some closely set, on apical third ofall faces, in allspecies. Spurs 1 : 2 : 2, the inner II-III about twice thelength of the outer. T I with a well-developed comb; TII with a reduced inner comb and a very small combbetween spurs, outercomb lost. T III also without outercomb, a small comb between spurs, and inner comblarge, but made of isolated setae. Tarsi long and thin,protarsus I 2.3 the length of tibia in the type-species,from 1.6 to 1.7 in the other. Male claws thick andserrulated, with basal spines, female claws less thick.

Wing (Fig. 4) narrow, shorter than abdomen, anallobe reduced; more orless smoky at apex (especially inR.fascio/um and R. humera/e), sometimes a spot at theanterior margin (R .fenestra/e). Sc short, ending before(R.fasciatum) or at the level of apex of basal cell. Sc2weak, situated before middle of Sc. R4 short andoblique (a little longer in R. fascio/um). All veinsreaching wing margin. Dorsal side of veins C, Rl,R4+5, R5 and An with setae; all veins bare on ventralside.

Male abdomen: segments I-II long, cylindrical, thefollowing progressively flattened and shortened, the

igh. Three ocelli, thelarge, their distance:ir own diameter (R.twice their diameterdeeplyemarginated,ld pedicel short anderes, widened andre much longer thancept in R fascio/um,Face wide, bare andIScio/um, in which itbears a few ventralIllS on the inside.:d. First palpomere:ct.kly arched, evenlyr-al and prescutellardisc, except in R.

:it bears one or twonumerous, short, in

;ly jutting out fromPleurae bare excepturn, and mesan-f short dorsal setae.tae, except II-III inial setulae regularlywith regular rows

1.fenestra/e, on thed almost all of the

the genus was de-in Matile, 1986. A

110 Matile: Rocetelion, a new Holarctic genus of the Keroplatidae ...

Figs. 11-16. Male genitalia (ventral view) and tergite IX and proctiger (dorsal view, setae not figured) ofRocetelion. 11-12)R.fenestrale(Fisher); 13-14) R.fasciatum (Garrett); 15-16) R. humerale (Zetterst.cdt). - Scale line 0.3 mm.

Antennal f1~without apicwing faintlytinctly before13-14 .

Discussion

Distance ofown diamet,bristles; melplcurites; hi!dian fusion s

2. Scutum bro-posterior codiomedian flfork. Male gScutum yellcthe median s,infuscated 0much shortetalia: Figs.

3. AntennalOa!a distinct, rolwing strongl:level of apex

The phylo!!and related ge(1986) monogrthe characters,

Rocetelionloss of the innelthe posterior c(unique among

At the specthat the male geis easy to distinlgenus. The filhumerale, whotemite strongl)coxal area widmerous and clo~The second cowith fiv~ synap'farther from miloss of discal ~setae, and lengt;of the male ofhypothesis.

At the gene]more closely altropical genus,on the basis of

15

sclerotised plate in the type-species, the plate smallerin Rfenestra/e andR. humerale. Ejaculatory apodemeshort and well sclerotised.

Female genitalia (Figs. 9-10, Rfasciatum) almostcompletely invaginate in seventh segment, from whichproject, laterally and ventrally, only the apex of tergiteX, two-thirds of sternite X and the cerci. Tergite VIIIand IX entirely invaginate and membranous. SternitcVIII entirely divided into two halves, all the apicalmargin ciliate. Sternite IX entirely internal, sclero-tised, with hollow laterodorsal expansions. Cerci verylong and narrow. Sternite X fonning a well-developedplate.

Larva and biology unknown.

1. Distance of ocelli from eye margin at most 1.5 theirown diameter; basistemite and scutellum without dis·cal bristles; metepistemite strongly darkened; hindcoxae without posterior bristles; radiomedian fusion aslong as stem of anterior fork 2

Key to the Holarctic species of Rocetelion

12

Male genitalia (Figs. 6-8,11-16): tergite IX large,more or less rounded at apex. Cerci small, rounded.Gonocoxopodites almost entirely divided ventrally bya longitudinal membranous area, extended on eachside along apical margin in R. fenestrale and R.humerale (Figs. II, 15). Ventral face of the synscleriteattached to the dorsal face by a narrow bridge,sclerotised and bearing setae, the gonocoxal tube beingtherefore very short, its internal side almost transverse;at this level, some rows of dark, conspicuous spinules.

Gonostyles dorsoventrally flattened, simple, lat-erally inserted, without apical or preapical teeth, butwith a thicker internal plate. Pointed at apex in R.fasciatum, rounded in R.fenestrale and R. humerale.No modified setae; in R. fenestrale, the ciliation issparser and shorter (Fig. 11).

Phallosome ofmedium size. Gonocoxal apodemeslong, narrow and well sclerotised. Phallosome almostentirely membranous dorsally and ventrally: only anarrow apical plate connecting the parameres (Fig. 8).Dorsal and ventral parameres connected by a large

Ann. Entomo!. Fennici 54:3. 1988 111

·on.11-12)R·fenestrale

s, the plate smallerJaculatory apodeme

l.fasciatum) almost~gment, from whichy the apex of tergitecerci. Tergite VIII

mbranous. Stemite[Yes, all the apical.ly internal, sclero-ansions. Cerci veryg a well-developed

~ete/ion

at most 1.5 their~llum without dis-y darkened; hindomedian fusion as........................... 2

Distance of ocelli from eye margin almost twice theirown diameter; basistemite and scutellum with discalbristles; metepistemite of the same colour as rest ofpleurites; hind coxae with posterior bristles; radiome-dian fusion shorter than stem of anterior fork............. 3

2. Scutum brown, without distinct longitudinal stripes;posterior coxae faintly infuscated on basal third; ra-diomedian fusion slightly shorter than stem of anteriorfork. Male genitalia: Figs. 11-12 ..... fenestrale (Fish.)Scutum yellow with three distinct longitudinal stripes,the median sometimes fainter; posterior coxae stronglyinfuscated on almost basal half; radiomedian fusionmuch shorter than stem of median fork. Male geni-talia: Figs. 15-16 humerale (Zett)

3. Antennal flagellum light brown, last flagellomere witha distinct, rounded apicule; mid coxae entirely yellow;wing strongly darkened at apex; Sc long, reaching thelevel of apex of basal cell. Male unknown ................................................... /asciolum (Coq.)Antennal flagellum dark brown, last flagellomerewithout apicule; mid coxae with a dark basal spot;wing faintly darkened at apex; Sc short, ending dis-tinctly before apex of basal cell. Male genitalia: Figs.13-14 fasciatum (Garr.)

Discussion

The phylogeny and biogeography of Rocetelionand related genera have been studied in MatHe's(1986) monograph. This includes a detailed analysis ofthe characters, to which the reader is referred.

Rocetelion shows only two autapomorphies, theloss of the inner tibial combs II-III and the reduction ofthe posterior combs of the tibiae; these characters arunique anlong the Keroplatini.

At the species level, and notwithstanding the factthat the male genitalia ofR.fasciolum are unknown, itis easy to distinguish two couples ofsister species in thegenus. The first is fonned by R. fenestrale andhumerale, who share four synapomorphies; metepis-ternite strongly coloured, male membranous gono-coxal area widened at apex, gonocoxal spinules nu-merous and closely set and gonostyles rounded at apex.The second consists of R. fasciatum and fasciolum,\vith fiv~ synapomorphies: situation of the outer ocelli,farther from middle of head, basisternite bare on disc,loss of discal scutellar setae and of posterior coxalsetae, and length ofradiomedian fusion. The discoveryof the male of R. fasciolwn will of course test thishypothesis.

At the generic level, Rocetelion was inferred to bemore closely allied to Paracerotelion MatHe, an afro-tropical genus, than to any other genus of Keroplatini,on the basis of three synap~morphies which appear

nowhere else in the tribe, and relate to the malegonostyles: dorsoventrally flattened, with the innerside modified into a thin sclerotised blade, and withoutapical teeth (this last character has been hypothesizedto be a loss, the teeth being present in the three othergenera of the group, Tolletia, Mallochinus andCerotelion).

The sister-group relationship inferred between theeastern nearctic R. fenestrale and the western palae-arctic R. humerale implies that the separation of thetwo species has a terminus post quem non situated atthe defmitive break of terrestrial connexions betweenEurope and North America. This event has been datedto the Upper Cretaceous (Dietz & Holden, 1970), to thePalaeocene (Smith & Briden, 1977), to the LowerEocene (McKenna, 1975) or even to the Upper Eocene(Talwani & HeIdorn, 1977; Hallam, 1981). This givesthe couple and its sister-group an age ranging fromabout 40 to about 60 million years.

Other Keroplatidae following the North Atlantictrack are the Macrocera of the nobilis group (Vocke-roth, 1976), the two species (one fossil) ofHesperodes(MatHe, 1980), the three Cerotelion species of thejohannseni group, the couple fonned by Keroplatusclausus Coquillett and K. reawnurii Dufour, theOrjelia of the discoloria group, the lsoneuromyia ofthe semirufa group and the genera Platyzua,Macrorrhyncha, Asindulum and Urytalpa (Matile,1986).

Many other Mycetophiloidea follow the sametrack: all the Bolitophilidae, the Diadocidiidae (butwith one or two neotropical species and a probablyintroduced Tasmanian species), the Ditomyiidae of thesubgenus Symmerus s. str., the Mycetophilidae of thegenera Paratinia, Baeopterogyna, Syntemna,Anaclileia, Ectrepesthoneura, Speolepta, Gnoriste,Novakia and Tarnania (MatHe, 1986). The same hasbeen noted for species of Mycetophila by Laffoon(1957) and Lastovka (1972), Phronia and Trichonta byGagne (1975, 1981), Pseudexechia and Epicypta byChandler (1978, 1981), Acnemia, Monoclona andAllodia of the subgenus Brachycampta by Zaitsev(1982a, 1982b; 1983; 1984), Sciophila by Zaitsev(1982c) and MatHe (1983), and Mycomya by Vaisanen(1984).

The North Atlantic track is therefore one of themost common among the Mycetophiloidea; its antiq-uity has been confmned by Baltic amber fossils (UpperEocene to Lower Oligocene) of the genera Hespe-rodes, Syntemna and Symmerus.

On the suprageneric level, the sister-group rela-tionship existing between the holarctic Rocetelion and

112 Matile: Rocetelion, a new Holarctic genus of the Keroplatidae ...

the afrotropical Paracerotelion implies that the com-mon ancestor of the couple cannot be posterior to theLate Cretaceous.

According to current plate tectonics, the separationof North America and Africa goes back to the UpperTriassic, an age which seems far too great for a coupleofpresent-day genera. If the geographic distribution ofthese taxa is to be interpreted in the light of currenttectonics*, one must then suppose that the ancestralstock was located in Africa and western Laurasia. Theancestral population was split when Europe wasseparated from Africa in the Palaeocene (and NorthAmerica from South America), the African stockgiving Paracerotelion, the Laurasian stock being at theorigin of the cool-adapted Rocetelion.

This scenario is consistent with the estimated age,Palaeocene or Eocene, of the species-groups.

Acknowledgements. I thank Dr. R. Gagne, of the U.S. Na-tional Museum, Washington, for the loan of most of the nearcticspecimer s studied by Garrett and Fisher, and of the type of R.fasciolum, and Mr. A. M. Hutson, of the British Museum (Nat.Hist), London, for the loan of a specimen of R. hwnerale. Mrs.Anna DamstrOm has revised the English text, for which I expressmy sincere thanks.

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Received 19.XI.1987Printed 15.VIII.1988