J. Ornithol. 142, 2±6 (2001) ã Deutsche Ornithologen-Gesellschaft/Blackwell Wissenschafts-Verlag, Berlin ISSN 0021-8375

The earliest fossil record of a modern-type piciform bird from the late Oligocene of Germany Gerald Mayr

Forschungsinstitut Senckenberg, Sektion fuÈr Ornithologie, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany, Email: [email protected]

Summary Presented here is a record of a piciform bird from a late Oligocene fissure filling from Ger- many. This is the earliest known fossil record of a modern-type piciform bird and the only one from deposits earlier than Miocene. The specimen, an isolated tarsometatarsus, is simi- lar in size and proportions to the tarsometatarsus of the recent Cardinal-Woodpecker (Dendropicos fuscescens, Picidae). However, owing to its fragmented state a reliable as- signment to anyof the recent piciform families is not possible.

Keywords: Piciformes, Pici, late Oligocene, fossil birds, earliest record.

Zusammenfassung Der aÈlteste Nachweis eines modernen Spechtvogels aus dem spaÈten OligozaÈn Deutschlands Ein Nachweis eines Spechtvogels wird aus einer spaÈtoligozaÈnen SpaltenfuÈllung Deutsch- lands beschrieben. Es ist der bisher aÈlteste Nachweis eines modernen Spechtvogels und der einzige aus vor-miozaÈnen Ablagerungen. Das Exemplar, ein isolierter Tarsometatar- sus, aÈhnelt in GroÈûe und Proportionen dem Tarsometatarsus des rezenten Kardinalspechts (Dendropicos fuscescens, Picidae). Aufgrund seiner fragmentarischen Erhaltung ist je- doch keine zuverlaÈssige Zuordnung zu einer der rezenten Familien spechtartiger VoÈgel moÈglich.

Introduction (Simpson & Cracraft 1981, Mayr 1998a). Hardlycorroborated with derived characters, Recent Piciformes (woodpeckers and allies) however, is the monophyly of the taxon (Gal- are generallyclassified within two ªsubor- bulae + Pici), i. e. the traditional Piciformes, dersº, the Galbulae (including the neotropic and the Galbulae possiblyare more closelyre- Galbulidae, jacamars, and Bucconidae, puff- lated to some coraciiform taxa (Olson 1983, birds) and the Pici (Capitonidae, barbets [in- Sibley& Ahlquist 1990). cluding ªRamphastidaeº, toucans, see Prum 1988], Indicatoridae, honeyguides, and Pici- Feduccia & Martin (1976) classified several dae, woodpeckers, wrynecks, piculets). A fossil birds from the Eocene of North America monophyly of both, Galbulae and Pici, is well within the Piciformes, but with the possible ex- supported with derived osteological characters ception of ªNeanisº kistneri Feduccia 1973

U.S. Copyright Clearance Center Code Statement: 0021-8375/2001/14201±02 $ 15.00/0 G. Mayr ´ A modern Piciform from the Oligocene 3

(which might be a representative of the Galbu- Material and methods lae) all of these were misidentified in order (see Houde & Olson 1989, 1992, Mayr 1998a, Comparisons have been made with skeletons of the 1998b, Mayr & Peters 1998). The earliest rep- following recent piciform taxa (all in the ornitholog- resentatives of the Piciformes belong to the ical collection of Forschungsinstitut Senckenberg): extinct familyPrimoscenidae Harrison & Capitonidae: Aulacorhynchus derbianus, Capito squamatus, Lybius minor, L. vieilloti, Megalaima Walker 1977 and are among the predominant armillaris, M. asiatica, M. javensis, M. mystaco- small perching birds in the earlyEocene (about phanos, M. virens, Pogoniolus bilineatus, P. sco- 50 million years ago) of Europe. These birds lopaceus, Psilopogon pyrolophus, Pteroglossus show a mosaic of piciform and passeriform castanotis, Ramphastos ambiguus, R. toco, R. vitel- characters (Mayr 1998a) and with regard to the linus, Selenidera maculirostris, Trachyphonus mar- morphologyof their distal tarsometatarsus garitatus; Indicatoridae: Indicator variegatus; Pici- most closelyresemble the earlyMiocene pici- dae: Campethera abingoni, C. cailliautii, C. nivosa, form family Zygodactylidae Brodkorb 1971. Chrysocolaptes lucidus, Colaptes auratus, Dendro- Mayr (1998a) described another extinct pici- copos major, D. medius, Dendropicos fuscescens, form family, the Miopiconidae, which is the Dryocopus martius, Jynx ruficollis, J. torquilla, Melanerpes carolinus, Picus canus, P. viridis, sister taxon of the Pici and which is known Sphyrapicus varius. In addition, representatives of from the Middle Miocene of Morocco. all other higher avian taxa have been investigated. The earlyTertiaryfossil record of the mod- The osteological terminologyused in this studyfol- lows Baumel & Witmer (1993), the dimensions are ern-type Piciformes of the suborder Pici is in millimeters. The fossil specimen is deposited in verysparse. Specimens from the earlyto Mid- Forschungsinstitut Senckenberg, Frankfurt/M., Ger- dle Miocene of Europe have been classified many(SMF). within the Capitonidae, in the genus Capito- nides Ballmann 1969b (Ballmann 1969a, 1969b, 1983). Laybourne et al. (1994) as- Systematic paleontology signed a feather preserved in amber from the Piciformes (Meyer & Wolf 1810) Dominican Republic to the Picidae; the exact Pici (sensu Simpson & Cracraft 1981) age of this specimen is unknown, according to the authors its minimum age is Lower Mio- The following combination of characters pre- cene. Olson (1985) mentioned Middle Mio- sented bythe specimen described below is di- cene fossils of a ªmedium-sized woodpeckerº agnostic for the Pici: retinaculum extensorium from New Mexico (USA) which have not been tarsometatarsi ossified (except some Capitoni- figured or described so far. Ballmann (1976) dae); foramen vasculare distale verysmall; described a record of the Picidae from the dorsal side of distal tarsometatarsus flat, troch- Upper Miocene of the Gargano Peninsula leae metatarsorum arranged in the same plane; (Italy). All other fossil Piciformes are from trochlea metatarsi IV turned plantarly(in the younger, Pliocene or Pleistocene, deposits (see fossil specimen the trochlea metatarsi IV is Olson 1985, Feduccia 1996). broken, but the remaining parts clearlyindi- cate that it was retroverted). Outgroup compar- Presented here is a record of a piciform bird isons with putativelybasal neognathous birds from the late Oligocene fissure filling Herrlin- (e. g. Anseriformes, Galliformes) suggest that gen 9 near Ulm (Baden-WuÈrttemberg, Ger- all of these characters are derived. many) which is the earliest known fossil re- The specimen differs from the Miopiconidae cord of a modern-type piciform bird. The few in that the part of the shaft which abuts on the other avian remains which have been found at plantar surface of the trochlea metatarsi III is this localitybelong mainlyto the Passeri- less vaulted, the trochlea metatarsi III is more formes. deeplygrooved. 4 Journal fuÈr Ornithologie 142, 2001

Fig. 1. Specimen SMF Av 429 (A, C) in comparison with the right tarsometatarsus of the recent Cardinal- Woodpecker Dendropicos fuscescens (Picidae) (B, D). A, B, plantar view; C, D, dorsal view. Abb. 1. Exemplar SMF Av 429 (A, C) im Vergleich mit dem rechten Tarsometatarsus des rezenten Kardi- nalspechts Dendropicos fuscescens (Picidae) (B, D). A, B, Plantaransicht; C, D, Dorsalansicht. familyincertae sedis Description and comparison: The specimen gen. et. sp. indet. is similar in size and proportions to the tarso- metatarsus of the recent Dendropicos fusces- Referred specimen: SMF Av 429 (incomplete cens (Cardinal-Woodpecker, Picidae). right tarsometatarsus, lacking the lateral part of the proximal end and the trochleae metatar- The tuberositas musculi tibialis cranialis is sorum II and IV) (Fig. 1). situated at the medial side of the sulcus exten- sorius. Most of the hypotarsus is broken, but Locality: Fissure filling Herrlingen 9 near two adjacent sulci hypotarsi can be recog- Ulm (Baden-WuÈrttemberg, Germany). nized. The fossa parahypotarsalis lateralis ap- Horizon: Late Oligocene, MP 29 (Engesser pears to have been verydistinct, although only 1999: tab. 30.2), absolute age about 25 million its distal part is visible. The foramina vascu- years (Legendre & LeÂveÃque 1997). laria proximalia are verysmall, centrallyposi- tioned, and situated close together (in some re- Dimensions: Maximum length, 14.7; medio- cent Capitonidae, e. g. Lybius and Psilopogon, lateral width of trochlea metatarsi III, 0.8; me- the medial foramen vasculare proximale is dio-lateral width of shaft at narrowest section, much larger than the lateral one). The retinacu- 1.0. lum extensorium tarsometatarsi is ossified and G. Mayr ´ A modern Piciform from the Oligocene 5 forms an arcus extensorius. The margo medi- fossil species. Recent trunk-climbing wood- alis bears a mediallyprotruding bulge on the peckers are characterized bya verydeeply level of the foramina vascularia proximalia. grooved trochlea metatarsi III, this feature is This bulge is present in all the recent Picidae I also present in Jynx but absent in the Capitoni- investigated, but is absent in some Capitonidae dae. As stated above, in the fossil specimen the (including Capitonides), and in Indicator and sulcus on the trochlea metatarsi III appears to Jynx. If viewed medially, the proximal end of be intermediate in its depth between that of re- the margo medialis is verynarrow in dorso- cent Picidae and that of recent Capitonidae. ventral direction. The shaft of the bone slightly This might indicate that the fossil species was widens towards its proximal end. As in all re- not as specialized for trunk climbing as recent cent Pici, the foramen vasculare distale is very Picidae. small. Some recent Picidae (e. g. Picus viridis, Sphyrapicus varius, Campethera spp.) also ex- The fossil record of piciform birds suggests hibit a second, medial foramen on the distal that at least in Europe the radiation of modern- tarsometatarsus which is absent in the fossil type Piciformes took place contemporaneously specimen. The fossa metatarsi I is shallow. with the radiation of passeriform birds, at the The trochleae metatarsorum II and IV are bro- end of the Oligocene. Obviously, at this time ken, but what is remaining from the trochlea manyecological niches for small perching metatarsi IV clearlyindicates that it was retro- birds were vacant in Europe. The fossil evi- verted. The trochlea metatarsi III is small and dence further supports an Old World origin of directed somewhat medially. Unfortunately, the Pici, which is in concordance with the fact this trochlea is slightlydamaged in the fossil that their sister taxon, the Miopiconidae, is specimen, as far as can be discerned it is not as known from North African deposits only. deeplygrooved as the trochlea metatarsi III of recent trunk-climbing woodpeckers, but deep- er than that of recent Capitonidae. Acknowledgements I thank S. TraÈnkner (Forschungsinstitut Senckenberg, Frankfurt a. M.) for taking the photographs. Discussion

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