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Home , Digitalis minor, Priotelus, Trogon, Trogon (genus)

The Auk 116(2):427-434, 1999

A NEW FROM THE MIDDLE OLIGOCENE OF Ct2RESTE,FRANCE

GERALD MAYR • ForschungsinstitutSenckenberg, Sektion fiir Ornithologie,Senckenberganlage 25, D-60325 Frankfurta.M., Germany

ABSTRACT.--Anew and of trogon( Trogoniformes, Trogonidae) is describedfrom the Middle Oligocene(approximately 33 million yearsago) of C•reste, France.The new , Primotrogon wintersteini, is the oldest trogon that has been named so far andclosely resembles living species; differences mainly concern proportions of theskull. The speciesmost likely is the sistertaxon of the extanttrogons. In contrastto mostof its recentrelatives, Primotrogon wintersteini lived in an arid environment.Received 17 December 1997, accepted21 September1998.

THE ORDERTROGONIFORMES comprises a sin- tons are extremelyrare (Stemvers-vanBemmel gle family,the Trogonidae,with 37 to 39 living 1984, Mourer-Chauvir• 1995b). species(Morony et al. 1975,Sibley and Monroe 1990).Extant trogonsare found in the tropics SYSTEMATICS and subtropicsof Asia, , and America. They feed on insectsand fruits, and apart from The measurementsreported below represent size and coloration,all membersof the overalllength along the longitudinalaxis the orderare quitesimilar morphologically. exceptfor the length of the carpometacarpus, Until recently,several species of the Archaeo- which representsthe distancebetween the trogonidaefrom the Upper Eoceneto Upper proximal end of the trochleacarpalis and the Oligocene(40 to 20 million ago) Phos- distal end of the os metacarpalemajus. Ana- phoritesdu Quercy,France, were consideredto tomical terminologyfollows Baumeland Wit- be the earliesttrogons (e.g. Mourer-Chauvir• mer (1993). 1980). Membersof this family lackedthe het- Membersof the Trogonidaeare character- erodactylfoot of the Trogoniformes,however, ized by their uniqueheterodactyl foot in which and Mourer-Chauvir•(1995a) proposed a clos- the second toe and the trochlea metatarsi I! of er relationshipto the .The the tarsometatarsusare turned plantad.Within fossilrecord of trogonsis very scanty,with this family, the monophylyof the eight living only two known early Tertiary taxa. Paratrogon genera(Trogon, , Euptilotis, Priote- gallicus(Milne-Edwards, 1867-1871) is known lus, Temnotrogon,, Heterotrogon, and from the Lower (22 million yearsago) )is supportedby the following two of Franceand closelyresembles living species. featuresof the coracoid:(1) a small processon Olson (1976) identified an incompleteskeleton faciesarticularis clavicularis overhanging the of a trogonfrom the early Oligocene"Glarner sulcusmusculi supracoracoidei; and (2) a notch Fischschiefer"of Matt, Switzerland, which was on the medial side of extremitas sternalis. originally assignedto the Alcedinidae(Peyer These two characterscertainly are derived 1957).This undoubtedlyis a trogonbut it within neognathousbirds, because character 1 is poorly preservedand lacks important ele- is not foundin anyother recent bird, andchar- ments(e.g. the skull). acter2 only occurswithin the Momotidae(Cor- The new specimenpresented here is the most aciiformes)and a few other . completefossil trogon known and also is the first bird describedfrom the Middle Oligocene Primotrogon gen. nov. (ca. 33 million years ago) depositsof C•reste, France.This localityis well knownfor impres- Typespecies.---Prirnotrogon wintersteini Mayr. sionsof isolatedfeathers, but completeskele- Diagnosis.--Prirnotrogonexhibits the hetero- dactylfoot of all trogoniforms,but the two de- E-mail:[email protected] t.de rivedcharacters of the livinggenera mentioned

427 428 GERALDMAYR [Auk, Vol. 116

FIG. 1. Primotrogonwintersteini, holotype (BSP 1997 1 38). Scale(lower left) in mm. aboveare absent.This suggeststhat the new Remarks.--Theunnamed trogon from the genusis the sistertaxon of extanttrogons. Pri- "GlarnerFischschiefer" (see above) might also motrogonfurther differs from all recenttrogons belongto the genusPrimotrogon, but the only in the smaller orbitae, the narrower base of the known specimenis too poorly preservedfor bill, the lessprotruding facies articularis digi- detailed comparisons.It is larger than P. win- talis minor, and the shorterprocessus exten- tersteini;according to Peyer (1957), the mea- soriusof the carpometacarpus.The statusof surements(in mm) are ulna, about 35; tibiotar- these characters(i.e. if they are derived or sus, about 36; and tarsometatarsus, about 19. primitive within trogoniforms)is unknown and can only be solvedby an exhaustivephy- Primotrogon wintersteini sp. nov. logeneticanalysis including most of the"high- er" land birds;such an analysisis beyondthe Holotype.--SeeFigures 1 and 2. Slightlydis- scopeof this paper. integratedincomplete skeleton, which is de- Etymology.The genericname is Latin for positedin the BayerischeStaatssammlung fiir "first trogon." Paliontologieund HistorischeGeologie (BSP), April 1999] OligoceneTrogon from Cdreste 429

4

FIG.2. X-ray photographof Prirnotrogonwintersteini holotype. Numbers indicate left coracoid(1); right coracoid(2); left scapula(3); unidentifiedbone (4); proximalend of mandible(5); scapusclaviculae (6); and ala ischii (7). Scale = 10 mm.

Mtinchen,Germany (collection number 1997 I Typehorizon. The fossilsfrom C•reste are 38). found in the "Calcaires de Montfuron" or in the Diagnosis.--Sameas for genus. Primotrogon "Calcairesde Vach•res"(C. Mourer-Chauvir• wintersteiniis slightlysmaller than the extant pers. comm.).These formations are of Middle Trogonviolaceus. Oligocene(MP 24) age (Ducreuxet al. 1985). Measurements.--Skull(total length), about 34 Referredspecimens.--None. mm; left coracoid, 19.8 mm; humerus, 27.2 mm Etymology.The speciesis namedafter Wil- (left), about27.2 mm (right); ulna, 31.0 mm helm Winterstein,whose generousfinancial (left), 31.0 mm (right); carpometacarpus,14.7 supportmade it possibleto purchasethe spec- mm (left), 14.9mm (right);left tibiotarsus,24.9 imen. mm; left tarsometatarsus,12.7 mm; left pedal phalanges,I1, 3.5 mm; II1, 5.5 mm; II2, about REMARKS 4.3 mm; III1, 4.0 ram; III2, 4.1 mm; III3, 4.7 mm; IV1, 2.6 mm; IV2, about 2.6 mm; IV3, about 2.6 The taphonomyof the skeletonis very odd. mm; longestfeather of wing, about90 mm. Both coracoidsand the left scapulaare sepa- Typelocality.--C•reste, D•partement Alpes- ratedfrom the pectoral girdle and displaced far de-Haute-Provence, France. caudally(Fig. 2, nos. 1 to 3). This would be 430 GERALDMAYR [Auk, Vol. 116 PrimotrogonPharomachrus

F[C. 3. Skull of Primotrogonwintersteini (on left; generalizedreconstruction that is not to be reliedon forScaledetails) = 10 mm.and Pharomachrus pavoninus(onright). quite unusualif it was due to decomposition, Pharomachrus Prirnotrogon becausein birds the pectoralgirdle, the ster- num, and the wingsare joinedtightly to each F•C. 4. Left coracoid of (A) Pharomachruspavoni- other It is alsostriking that both wings and the nusand (B) Primotrogonwintersteini. Numbers indi- left leg are preservednear to their originalpo- cateprocess overhanging the sulcusmusculi supra- sition,although the elementsconnecting them coracoidei(1); processusprocoracoideus (2); notch to the restof the body are eitherabsent (left fe- on medial side of extremitas sternalis (3); and an- mur) or displaced(coracoids). Some of theskel- gulusmedialis (4). Scale= 5 min. etal elements(e.g. vertebralcolumn, left scap- ula, right coracoid,and furcula) are incom- species,too. The bill is moreelongated, and al- plete,and the right leg and the tail are entirely thoughits tip is not clearlyvisible, it seemsto absent.It is difficult to interpretthis tapho- be lesspointed than that of the recenttrogons. nomicsituation. Possibly, the skeletonwas dis- Thenarial openings are large and oval-shaped. turbedby an animalfeeding on the carcass. The mandible does not articulate as far caudal- A large bonelying next to the left coracoid ly asit doesin living (its proximalend (Fig. 2, no. 4) certainlydoes not belongto P. is visibleon the x-ray photograph; Fig. 2, no.5). wintersteini,yet I havebeen unable to identify Most of the cervical and some thoracic ver- it. It looks somewhat like the caudal end of an tebraeare preserved,but no detailsare visible. avian ramus mandibulae, but besides the fact The caudalvertebrae including the pygostyle that it would be too large for P. wintersteini, havebeen lost entirely. Six left (?)vertebral ribs both rami mandibulae are connected to the can be counted. skull as can be seenon the x-ray photograph The coracoid is similar to that of extant tro- (Fig. 2, no. 5). gons in its proportions,yet it differs in some Descriptionand comparison.--Probably P.win- details(Figs. 4A, B). The faciesarticularis clav- tersteinidiffers most stronglyfrom living tro- icularisof the processusacrocoracoideus does gonsin the proportionsof the skull (Fig. 3), not exhibitthe small projectionoverhanging which resemblesome passeriforms (e.g. Bom- the sulcusmusculi supracoracoidei(Fig. 4A, bycillagarrulus). In the extanttrogons, the or- no. 1), and the processusprocoracoideus (Fig. bitaetake up mostof the cranium,whereas the 4A, no. 2) is broader.The extremitassternalis cranial cavity is distinctly larger and more differsfrom that of the living generain the ab- roundedin P.wintersteini. The distalpart of the senceof a notchon the medialmargin (Fig. 4A, cranium (os frontale and os lacrimale)and the no. 3) and in the form of the angulusmedialis proximalpart of thebeak are narrowerin this (Fig. 4A, no.4),which doesnot taper off to a April 1999] OligoceneTrogon from Cdreste 431

FxG.5. Right manusof (A) Pharomachruspavoninus and (B) Primotrogonwintersteini. The digitus alulaeis not shown.Numbers indicate processus extensorius (1); faciesarticularis minor (2); andtubercle on phalanxproximalis digiti majoris(3). Scale= 5 mm. point;the anguluslateralis of the processusla- is dorsalisis small, and the condylusdorsalis teralis is lesspronounced. and the condylusventralis resemble those of Muchof the corpusof the left scapula(Fig. 2, extant genera (see Fig. 2). no. 3) is preservedbut doesnot show any pe- The ulna is longerthan the humerus.Papil- culiarities.A singlescapus claviculae is visible lae remigalesare absent,and the olecranonis near the cranialend of the sternum(Fig. 2, no. large.Other details of thisbone are not visible. 6). It becomesbroader toward its extremitas The carpometacarpusis comparable to that of omalis. living trogons(Figs. 5A, B). The spatiuminter- The sternumis shortand very wide (its cau- metacarpaleis wide, and the symphysismeta~ dal width evenexceeds the length of the hu- carpalisdistalis is short.The distalpart of the merus).The processuscraniolaterales and the os metacarpaleminus is slightlytilted. The margo cranialisare not visible.Three proces- processusextensorius (Fig. 5B, no. 1) is shorter sus costalescan be countedon the x-ray pho- than in extant genera.The faciesarticularis tograph.The margo caudalisshows four inci- digitalisminor is in line with the os metacar- surae,of whichthe incisuraelaterales are deep- pale majus,whereas it protrudesfarther dis- er than the incisurae mediales. The trabeculae tally in extanttrogons (Fig. 5A, no.2). Thesul- lack transverseprocesses at their caudalend. cus tendinosusopens in a shallow funnel- Like in the recentgenera, the trabeculamedi- shapeddepression. ana is triangular The digitusminor is large.In recenttrogons, The humerus differs in some details from the phalanxproximalis digiti majorisexhibits thatof recenttrogons. An enlargedcrista bicip- a tubercleon its caudalside opposite to thedig- italis, which is characteristicof the humerus of itus minor(Fig. 5A, no.3) that apparentlyis ab- living genera,apparently is absent.The fossa sentin P.wintersteini (Fig. 5B). The oscarpi ul- pneumotricipitalisis single,whereas in recent nare resemblesthat of the extanttrogons and trogonsa shallowsecond fossa ends beneath manyother birds; its crusbreve is short. the caputhumeri. The tuberculumdorsale is The alaeischii of the pelvisseem to be very weak.There is a slightnotch between the tu- slenderlike in the extantgenera (Fig. 2, no. 7); berculum ventrale and the crus ventrale fossae. the synsacrumshows no peculiarities. Likein living trogons,the cristadeltopectoralis The tibiotarsus is stout and similar to the cor- seems to be bent toward the cranial side of the respondingbone of Pharomachrussp., whereas humerus.It is not visiblein the specimenor in it is muchmore slenderin other trogons,e.g. the x-rayphotograph (both humeri are slightly Trogonviridis and temnurus. The con- turned in dorsocranialdirection). The proces- dylus lateralisprotrudes only slightlytoward susflexorius is short;the sulcusscapulotricip- the cranial surface of the bone. italis is shallow.The processussupracondylar- The tarsometatarsus is short and robust. It 432 GERALDMAYR [Auk, Vol. 116

doesnot prove an Old Worldorigin for theTro- goniformes,this notionis supportedby a re- cent cladisticanalysis of living species(Espi- nosade los Monteros1998). Primotrogon winter- steiniis the earliestknown trogonid, but a tar- sometatarsusof an unquestionablyheterodactyl bird doselyresembling that of a trogoniform hasbeen collected in the Lower (53 mil- lion yearsago) LondonClay of Essex,England (private collection M. Daniels). Surprisingly, trogonshave not beenfound in the extensive and well-studiedmaterial of the Phosphorites du Quercy.This is odd becausethe localityis geographicallyand stratigraphicallyclose to FIG. 6. Left foot of Primotrogonwintersteini. The C•reste. digits are numbered from dl to d4; 1 is trochlea Palaeobiology.--Mostrecent authors postulate metatarsi II. an arid climatefor C•resteduring the Oligo- cene.Lutz (1984)assumed that the vegetation was openwith only few bushesand trees,and dosely resemblesthat of most extant species, Schmidt-Kittlerand Storch(1985) consideredit althoughin Priotelustemnurus it is distinctly longerand more narrow.In lateralview the to be a "semi-arid open habitat on a large central sectionof the shaft is broad. The hy- scale."According to Mai (1995:423),there were semi-evergreenopen forests interspersed with potarsusis very prominent and seemsto ex- hibit two sulci on its lateral side. The trochlea sclerophyllousbushes and sparsepines. Be- metatarsiII (Fig. 6) is turned as far plantad as cause trogon feet are especially adapted to in recenttrogons. The trochleametatarsi IV is perching,and recenttrogons nest in treeholes, small and round when seen from its lateral the presenceof at least isolatedwoodlands in side. and near C•reste is a reasonableconjecture. The arrangementof the toesclearly is het- The presenceof a trogon in an arid environ- erodactyl.The phalangeformula and the pro- mentmight be surprising,because most living portionsof the phalangesare normal.The hal- trogonsoccur in humidsubtropical or tropical lux is weak, and no clawshave been preserved forests.However, some recent species (e.g. Tro- on any of the toes.The processusarticularis gonelegans, T. violaceus,and T. citreolus) also oc- tarsometatarsalis of the os metatarsale I is cur in open woodlandsand arid scrub(Sibley short. and Monroe 1990). The impressionsof the remigescorrespond Phylogeny.--Aspointed out in the diagnosis, in relativelength with thoseof recenttrogons. Primotrogonwintersteini most likely is the sister However,it is not dear whetherthe distalpri- taxonof the recenttrogons. The speciesis re- mariesare shortenedlike in recentspecies (a markablysimilar to living trogons,and only a shortfeather on the outerside of the left wing few other avian taxa are known to have under- is not the mostdistal primary because it does goneso little morphologicalalteration during not inserton thephalanx distalis digiti major). the past33 millionyears. Differences concern- Trogonspossess very long tail ;unfor- ing therelative size of theorbitae and the more tunatelythose of P.wintersteini have been lost narrowbeak might indicate that the Oligocene togetherwith thepygostyle. specieswas lessspecialized at catchingflying .In its proportions,P wintersteinifalls DISCUSSION within the rangeof recenttrogons and is clos- estto the Asianspecies Harpactes ardens (Table Contraryto their extantrelatives, early Ter- 1). All New World trogonsseem to have a tiary trogonsare known only from Europe. The slightlylonger ulna than P wintersteiniand the oldest New World are from African and Asian trogons,but this must be depositsin the DominicanRepublic and Brazil confirmedwith a larger numberof skeletons (Brodkorb1971). Although the fossilrecord than was available to me. April 1999] OligoceneTrogon from C&este 433

BecausePrimotrogon wintersteini is very close to extanttrogons, it doesnot give us additional informationon the higher systematicposition of the order Trogoniformes.Feduccia (1975) discoveredthat trogoniformsand somecora- ciiforms (Meropidae,Alcedinidae, Momotidae) sharea derivedmorphology of the columella. A very similar of columella,however, also occursin the suboscinepasserines (Feduccia 1975), and no other charactersare known to support a relationshipbetween trogons and coraciiforms.

ACKNOWLEDGMENTS

I am indebtedto E Wellnhofer(Bayerische Staats- sammlungfiir Pal•ontologieund HistorischeGeo- Iogie)for makingthe C6restespecimen available for study. B. Stephankindly loanedbones of somere- cent trogonsfor comparison.M. Daniels informed me aboutthe trogoniformspecimen in his collection. I alsothank G. Arratia, J. Gregor,H. Lutz, C. Mayr, C. Mourer-Chauvir6, H.-E Schultze, and G. Storch for providing information on C•reste; and D. S. Pe- ters, R. M. Zink, and an anonymousreviewer for commentson the manuscript.S. Tr•nkner took the photographs,and H. Zetzscheprepared the x-ray picture.

LITERATURE CITED

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