The Auk 124(4):1294–1304, 2007 © The American Ornithologists’ Union, 2007. Printed in USA.

A TODY (ALCEDINIFORMES: TODIDAE) FROM THE EARLY OLIGOCENE OF GERMANY Gea Ma1,3 an Chae W. Knf2 1Forschungsinstitut Senckenberg, Sektion Ornithologie, Senckenberganlage 25, D-60325 Frankfurt a.M., Germany; and 2Deutsches Krebsforschungszentrum, F030, Im Neuenheimer Feld 242, D-69120 Heidelberg, Germany

Atat.—We describe a postcranial skeleton of a stem-group tody (Aves: Alcediniformes: Todidae) from the early Oligocene of Germany. The fossil is tenta- tively assigned to Palaeotodus itardiensis Mourer-Chauviré, 1985, which previously was known only from a few incomplete bones from the early Oligocene of France. It is the most substantial fossil record of a Paleogene stem-group tody described so far and shows many previously unknown skeletal details. The specimen confi rms the occurrence of Todidae, which are today restricted to the Caribbean Greater Antilles, in the Paleogene of Europe. The biogeographical signifi cance of these fi nds depends on the phylogenetic relationships between Todidae and other alcediniform birds. If Todidae are not the sister taxon of Momotidae (motmots) as suggested by a recent phylogenetic analysis of molecular data, occurrence of stem-group Todidae in the Old World fossil record actually follows from the currently assumed Old World ori- gin of alcediniform birds. Received 29 August 2006, accepted 19 November 2006.

Key words: biogeography, evolution, fossil birds, Palaeotodus.

Ein Todi (Alcediniformes: Todidae) aus dem unteren Oligozän Deutschlands

Zaenfan.—Wir beschreiben ein postcraniales Skele eines Stamm gruppenvertreters der Todies (Aves, Alcediniformes, Todidae) aus dem frühen Oligozän von Deutschland. Das Fossil wird unter Vorbehalt Palaeotodus itardiensis Mourer-Chauviré, 1985 zugeordnet, einer Art, die bisher nur von wenigen unvollständigen Knochen aus dem frühen Oligozän von Frankreich bekannt war. Es ist der bisher vollständigste Fossilnachweis eines paläogenen Stammgruppenvertreters der Todies, und zeigt viele bisher unbekannte osteologische Einzelheiten. Das Exemplar bestätigt das Vorkommen von Todidae, die heute auf die karibischen Großen Antillen beschränkt sind, im Paläogen Europas. Die biogeographische Bedeutung dieser Funde hängt von den Verwandt- scha sbeziehungen zwischen Todies und anderen alcediniformen Vögeln ab. Falls Todies nicht das Schwestertaxon der Momotidae (Motmots) sind, wie eine kürzlich veröff entlichte phylogenetische Analyse molekularer Daten nahelegt, folgt das Vorkommen von Stammgruppenvertretern der Todidae in der Alten Welt vom gegenwärtig angenommenen altweltlichen Ursprung der alcediniformen Vögel.

The five extant species of the alcediniform the evolutionary history of these birds, but it has todies (Todidae), which occur only on the Greater been recognized that their extant distribution Antilles, are very small, brightly colored, and is diff erent from that of the “total group” (pan- predominantly insectivorous birds that inhabit monophylum, i.e., the clade including stem- and various kinds of woodlands, from rainforests to crown-group representatives), because all fossil arid brush (Kepler 2001). Li le is known about stem-group representatives were found far out- side the Caribbean region. The fi rst fossil record of the Todidae was 3E-mail: [email protected] identifi ed by Olson (1976), who described the

1294 October 2007] Oligocene Tody from Germany 1295 stem-group tody Palaeotodus emryi from the previously unknown osteological details of this early Oligocene Brule Formation in Wyoming taxon. (~30 mya; Olson 1976). The original description of P. emryi was based on a skull and the proxi- Mateia an Meth mal portion of a humerus only, and although postcranial remains of another individual of Osteological terminology follows Baumel and this species, from the lowest part of the Brule Witmer (1993). Measurements are in millimeters Formation, exist in the collection of the National and, unless indicated otherwise, represent the Museum of Natural History in Washington, maximum length of the bones along their lon- D.C., these have not yet been described. The gitudinal axis; the ungual phalanges were mea- humerus of the holotype of P. emryi is pro- sured from the tip to the processus extensorius. portionally larger than that of extant Todidae, Institutional abbreviations are as follows: SMF = and Olson (1976:114) hypothesized that “in Forschungsinstitut Senckenberg, Frankfurt am the Oligocene, the Todidae were possibly more Main, Germany; USNM = National Museum of diverse than at present and probably included Natural History, Washington, D.C. larger, more actively fl ying forms with be er developed wings than the strictly sedentary Systematic Paleontology modern todies.” Tody fossils are also known from the Old Alcediniformes (sensu Mayr 1998) World: two species of Palaeotodus, P. escamp- Todidae Vigors, 1825 siensis and P. itardiensis, were reported by Palaeotodus Olson, 1976 Mourer-Chauviré (1985) from the Late Eocene Palaeotodus cf. itardiensis Mourer-Chauviré, 1985 (P. escampsiensis: Mammalian Paleogene [MP] stratigraphic level 19, i.e., 35 mya) and early Referred specimen.—SMF Av 505: disarticu- Oligocene (P. itardiensis: MP 23, i.e., 31 mya) of lated skeleton on a slab, lacking skull and le France. Palaeotodus itardiensis is similar in size to wing (Fig. 1); the counter-slab is transferred on P. emryi and distinctly larger than extant todies, the same resin slab but contains very few bone but the crown-group tody-sized P. escampsien- fragments. sis shows that some Paleogene stem-group Locality and horizon.—Frauenweiler south Todidae were as small as their extant relatives. of Wiesloch (Baden-Wür emberg, Germany), Unfortunately, these Old World species of former clay pit of the Bo -Eder GmbH (“Grube Palaeotodus, which apparently were overlooked Unterfeld”), Rupelian, Lower Oligocene by Overton and Rhoads (2004), are known from (Micklich and Hildebrandt 2005). only a few bones: the description of P. escamp- Measurements.—Numbers in parentheses are siensis is based on an incomplete humerus, and measurements of an undescribed specimen that of P. itardiensis on a proximal ulna, distal (USNM 334940) of P. emryi. Coracoid, 14.8 (le ). tibiotarsus, and proximal tarsometatarsus. Sternum (spina externa to trabecula intermedia), Thus, the morphology of Paleogene stem-group ~15.8. Humerus, 19.2 (right) (20.9). Ulna, 24.7 Todidae is still very incompletely known, which (right) (26.7). Carpometacarpus, 10.0 (right) severely limits comparisons with crown-group (10.1). Femur, 15.3 (right) (16.6). Tibiotarsus, Todidae on the one hand and between the 23.4 (le ), 23.4 (right) (24.9). Tarsometatarsus, North American and European fossil species 16.5 (le ), 16.6 (right) (>14.7). Pedal phalanges on the other. (le /right): I1, 4.5/4.3; I2, –/3.3; II1, 2.7/–; II2, Here, we describe a postcranial skeleton of 3.2/–; II3, 2.5/2.5; III1, 3.9/–; III2, 3.8/–; III3, 3.6/ Palaeotodus from the Lower Oligocene (MP 22, 3.7; III4, –/3.5; IV1, 2.7/–; IV4, –/2.2; IV5, –/2.4. i.e., 32 mya; Micklich and Hildebrandt 2005) of Remarks on phylogenetic assignment.— Frauenweiler in Southern Germany, a former Classifi cation of the Frauenweiler fossil clay pit whose marine, near-shore deposits have into Alcediniformes (i.e., a clade including yielded several other avian taxa (Mayr 2000, todies, motmots [Momotidae], kingfi shers 2004a, b, 2005a, 2006b; Mayr et al. 2002; Mayr [Alcedinidae], and bee-eaters [Meropidae]) is and Manegold 2004, 2006). This specimen is supported by the following derived features: the most completely preserved fossil record of (1) distal end of ulna with very large condylus Palaeotodus and allows the recognition of many dorsalis (except Meropidae); (2) os metacarpale 1296 Mayr and Knopf [Auk, Vol. 124

Fig. 1. Palaeotodus cf. itardiensis Mourer-Chauviré, 1985, from the Lower Oligocene of Frauenweiler, Germany (SMF Av 505). (A) Sternum, shoulder girdle, and right wing with interpre- tive drawing. (B) Pelvis and feet with interpretive drawing. Abbreviations: fur = furcula, lco = left coracoid, lfe = left femur, lsc = left scapula, ltb = left tibiotarsus, ltm = left tarsometatarsus, ocu = os carpi ulnare, pdm = phalanx distalis digiti majoris, pel = pelvis, ppm = phalanx proximalis digiti majoris, r = rib, rcm = right carpometacarpus, rfe = right femur, rhu = right humerus, rra = right radius, rtb = right tibiotarsus, rtm = right tarsometatarsus, rul = right ulna, st = sternum, v = vertebra. Coated with ammonium chloride to enhance contrast. Scale bars equal 5 mm. minus of carpometacarpus protruding distally derived characters with crown-group Todidae: far beyond os metacarpale majus (Mayr 1998); (1) scapi clavicularum of furcula very thin; (2) (3) trochleae metatarsorum cylindrical and on the proximal end of humerus reaching far ventrally same dorsoplantar level (Mayr 1998); and (4) and infl ected so that almost entire caput humeri proximal phalanx of hallux with (albeit small) situated farther ventrally than ventral margin of lateral projection (Mayr 1998). sha ; (3) carpometacarpus with large processus The specimen agrees with extant Momotidae intermetacarpalis; (4) tarsometatarsus greatly and Todidae but diff ers from Alcedinidae and elongated and slender, measuring almost the Meropidae in its long hindlimbs and the com- length of the humerus; and (5) plantar surface plete reduction of the processus procoracoideus of trochlea metatarsi III bearing a marked sulcus. of the coracoid; it further shares the following It is identifi ed as a stem lineage representative October 2007] Oligocene Tody from Germany 1297 of Todidae by, for example, its greater size and referred specimen of P. emryi (USNM 334940) the plesiomorphic morphology of the distal tar- diff ers from that of the Frauenweiler fossil in sometatarsus (see description). that the processus acrocoracoideus has a slightly The Frauenweiler fossil is distinctly larger hooked, rather than rounded, outline (Fig. 2). than P. escampsiensis (width of distal end of Only the cranial portion of the scapula is humerus 3.8 mm vs. 2.9 mm in P. escampsiensis preserved (Fig. 3); as in all alcediniform birds, [a er Mourer-Chauviré 1985]) and diff ers from it has a bifurcated acromion with an additional P. emryi in the more rounded processus acro- medial process. coracoideus of the coracoid (hooked in P. emryi The U-shaped furcula (Fig. 3) resembles that [USNM 334940]; see Fig. 2). Also, it is slightly of crown-group Todidae and diff ers from other smaller than P. emryi (width of proximal end of alcediniform birds in that the scapi clavicularum humerus 6.1 mm vs. 6.7 mm in the holotype of are very thin, not only at the extremitas sterna- P. emryi [a er Olson 1976]; see above concern- lis, but over more than half of their length. An ing the measurements of the referred specimen apophysis furculae is absent, and details of the USNM 334940) and, thus, seems to be of similar extremitas omalis cannot be discerned. size to P. itardiensis, which was also consid- The sternum (Fig. 3) is about as long as it is ered slightly smaller than P. emryi by Mourer- wide and is similar to that of extant Todidae in Chauviré (1985). However, direct comparisons its proportions. The preserved (le ) processus of measurements and morphological features craniolateralis apparently lacks the tip, and three with P. itardiensis are diffi cult, because the over- processus costales can be counted. The carina lapping skeletal elements are poorly preserved sterni is preserved in its original position, pro- in the Frauenweiler fossil. Although there may jecting perpendicular to the corpus sterni. It is of have been a diff erence in the shape of the distal similar height to that of crown-group Todidae tibiotarsus (see description), we prefer assign- and has a convex cranial margin (straight in ment to the contemporaneous P. itardiensis to Momotidae and Alcedinidae). As in other naming a new species. We note, however, that alcediniform birds, there is a long and unbi- this classifi cation is tentative, pending the dis- furcated spina externa, which is visible through covery of additional specimens. the transparent reverse of the slab. The caudal Description and comparison.—Four thoracic margin of the corpus sterni bears four incisions, and two caudal vertebrae (situated next to the of which the incisura lateralis is much deeper distal end of the right tibiotarsus), as well as than the small incisura medialis, reaching more several ribs, are preserved but do not allow than half the length of the corpus sterni. In all for description of phylogenetically informative crown-group Momotidae, the medial incisions details. in the caudal margin of the sternum are closed The coracoid (Fig. 3) is a slender bone with a to fenestrae (the lateral incisions also are closed wide extremitas sternalis. The extremitas omalis of in Momotus and Eumomota but caudally open in the le coracoid has been completely released Aspatha). from the matrix so that both the dorsal and the As in crown-group Todidae, the proximal ventral surfaces are visible; most of the incom- end of the humerus (Fig. 4) reaches far ventrally plete right coracoid is hidden in the matrix but and is ventrally infl ected so that almost the can be seen through the reverse of the transpar- entire caput humeri is situated farther ventrally ent slab. The bone is similar to that of extant than the ventral margin of the sha . The crista Todidae and Momotidae (Fig. 2) in that the pro- bicipitalis is hardly developed, and the ventro- cessus procoracoideus is completely reduced. The distal margin of the extremitas proximalis meets elongated processus acrocoracoideus is dorsoven- the sha at an almost right angle. The crista trally compressed and has a rounded tip, which deltopectoralis is short and hardly protrudes. is not bent ventrally as in crown-group Todidae; The original degree of curvature of the sha the impressio ligamenti acrocoracohumeralis is cannot be ascertained; in Todus, P. emryi (USNM marked. The facies articularis scapularis is shallow. 334940), and P. escampsiensis, it is markedly The medial margin of the extremitas sternalis is curved; as in these taxa, the distal end is crani- convex and the processus lateralis well developed ally infl ected. The distal humerus of SMF Av 505 as in crown-group Todidae, whereas this process closely resembles that of P. escampsiensis and, as is reduced in Momotidae. The coracoid of the in that species, the condylus ventralis is elongate, 1298 Mayr and Knopf [Auk, Vol. 124

Fig. 2. Palaeotodus cf. itardiensis Mourer-Chauviré, 1985. Selected skeletal elements in comparison: (A–D) left coracoid, (E–G) distal end of right ulna, (H–J) sternum, (K–M) right carpometacarpus, (N–P) right tarsometatarsus. (A, E, H, K, N) Palaeotodus cf. itardiensis, (B) P. emryi (USNM 334940, reversed to facilitate comparisons), (C, F, I, L, O) Todus multicolor (Todidae), (G) Coracias garrulus (Coraciidae), (D, J, M, P) Momotus momota (Momotidae). Abbreviations: cnd = condylus dorsalis, dep = depression, imc = processus intermetacarpalis, pla = processus lateralis, ppc = processus procoracoi- deus, spe = spina externa. Tarsometatarsal trochleae are numbered. Scale bars equal 5 mm. with a depression on its cranial surface, and the been of a similar shape. The proximal ulna of tuberculum supracondylare ventrale is small. SMF Av 505 further agrees with extant Todidae The ulna is the longest limb element and dis- in the strongly protruding tuberculum ligamenti tinctly exceeds the humerus in length. Because collateralis ventralis, which is less prominent in the bone is severely crushed, comparisons with other alcediniform birds. The well-preserved the proximal ulna of P. itardiensis are not pos- distal end of the bone exhibits a morphology sible, though the olecranon appears to have characteristic of Todidae, Momotidae, and October 2007] Oligocene Tody from Germany 1299

Fig. 3. Palaeotodus cf. itardiensis Mourer-Chauviré, 1985. (A) Sternum in ventral view. (B) Left coracoid in ventral view, furcula, and extremitas cranialis of scapula. (C) Extremitas omalis of left coracoid in dorsal view. Abbreviations: cor = coracoid; fur = furcula; sca = scapula. Coated with ammonium chloride to enhance contrast. Scale bars equal 5 mm.

Fig. 4. Palaeotodus cf. itardiensis Mourer-Chauviré, 1985. (A) Right humerus in cranial view. (B) Distal end of right ulna and radius in ventral view and right carpometacarpus in dorsal view. Abbreviations: cmc = carpometacarpus, ocu = os carpi ulnare, ppm = phalanx proximalis digiti majoris, rad = radius, uln = ulna. Coated with ammonium chloride to enhance contrast. Scale bars equal 5 mm. 1300 Mayr and Knopf [Auk, Vol. 124

Alcedinidae, in that the condylus dorsalis is very The pelvis (Fig. 5) is of similar proportions to large, twice as long as the condylus ventralis, and that of extant Todidae; its mediolateral width strongly protruding (Figs. 2, 4). across the antitrochanters is slightly less than As in crown-group Todidae, Momotidae, and the length of the synsacrum. The alae praeac- Alcedinidae, the radius bears a marked tubercle etabulares ilii are short, and the well-developed on the ventral surface of its distal end (facies cristae iliacae dorsales and crista spinosa synsacri articularis radiocarpalis; Fig. 4). The sha of the delimit short sulci iliosynsacrales, which measure bone is straighter than the strongly curved less than one-third of the length of the pelvis. radius of extant Todidae (T. multicolor). As in extant Todidae, there are no foramina inter- The carpometacarpus (Fig. 4) most closely transversaria. resembles that of extant Todidae. In contrast Except for the trochlea fi bularis, the femur is to other alcediniform birds, there is a well- poorly preserved and does not allow for mean- developed processus intermetacarpalis, which ingful description. The bone appears to have reaches the os metacarpale minus. In other alce- been slightly more robust than the femur of diniform birds, this process is smaller and situ- crown-group Todidae, but because the sha is ated farther proximally. A processus dentiformis severely crushed, its original proportions are is situated at the proximal end of the sulcus diffi cult to estimate. tendinosus. The os metacarpale minus markedly The tibiotarsi expose their lateral (right) and exceeds the os metacarpale majus in length, but its caudolateral (le ) surfaces and almost reach distal section is not craniocaudally constricted the length of the humerus (in Alcedinidae and as in crown-group Todidae; the dorsal margin Meropidae, the tibiotarsus is very short). As in exhibits three small, equally spaced nodular extant Momotidae and Todidae, but unlike in swellings. There is a marked depression on Meropidae and Alcedinidae, the crista cnemia- the dorsal surface of the caudodistal portion lis lateralis is well developed. The distal end of of the os metacarpale majus, which also occurs the bone (Fig. 5), however, diff ers from that of in crown-group Todidae but is less marked in crown-group Todidae and is more similar to Momotidae (Momotus momota; Fig. 2). Momotidae in being mediolaterally narrower The phalanx proximalis digiti majoris is some- and with a less cranially protruding condylus what narrower craniocaudally than in crown- lateralis. With regard to the shape of the distal group Todidae, with a less convexly curved tibiotarsus, SMF Av 505 agrees with the North caudal margin. American P. emryi (USNM 334940) but diff ers

Fig. 5. Palaeotodus cf. itardiensis Mourer-Chauviré, 1985. (A) Pelvis in dorsal view. (B) Left tarso- metatarsus in medial view. (C) Right tarsometatarsus in plantar view. Abbreviation: tbt = tibiotar- sus. Coated with ammonium chloride to enhance contrast. Scale bars equal 5 mm. October 2007] Oligocene Tody from Germany 1301 from the very tody-like distal tibiotarsus of taxa, it measures less than half the humerus the French specimen of P. itardiensis. The distal length). The proximal end of the bone is poorly tibiotarsi of SMF Av 505 are, however, not very preserved but appears to have been wide, as well preserved and are somewhat crushed, so in the French specimen of P. itardiensis. The that we currently refrain from pu ing too much hypotarsus is strongly plantarly protruding taxonomic signifi cance on these diff erences and, judging from its shape, probably enclosed from the French specimen of P. itardiensis. The a bony canal as in extant alcediniform birds; short fi bula reaches about one-third the length there is also a sulcus along its plantar surface. of the tibiotarsus. As in other alcediniform birds, the fossa parahy- The tarsometatarsus (Figs. 2, 5, 6) is an elon- potarsalis medialis is marked and the proximal gate and slender bone that, as in extant Todidae, part of the margo medialis forms a sharp ridge. measures almost the length of the humerus. In The crista medianoplantaris extends over the crown-group Todidae, this bone also is slightly proximal fourth of the tarsometatarsus. The shorter than the humerus (T. multicolor) or even cross-section of the central segment of the tar- exceeds it in length (T. angustirostris). In the sometatarsal sha is subtriangular. The distal momotid taxon Aspatha, the tarsometatarsus end of the tarsometatarsus of Palaeotodus has reaches ~80% of the humerus length, whereas not been described and is preserved for the it is distinctly shorter in Momotus, Electron, fi rst time in the Frauenweiler specimen (Figs. and Baryphthengus (Momotidae) as well as in 2, 6). As in crown-group Todidae, but unlike Alcedinidae and Meropidae (in the la er two in Momotidae, the fossa metatarsi I is very large. The same is true of the foramen vasculare distale, which is small in Momotidae. The trochleae metatarsorum II and IV are only slightly shorter than the trochlea metatarsi III and hardly plan- tarly defl ected, whereas the trochlea metatarsi II is distinctly shorter than the other trochleae in crown-group Momotidae (Fig. 2). The plan- tar surface of the trochlea metatarsi II lacks a furrow, as it does in crown-group Todidae; it is, however, proportionally larger and more medially directed than in the la er. The trochlea metatarsi III is as long as it is wide and bears a marked furrow on its plantar surface, though this furrow is less marked than in crown-group Todidae. The incisura lateralis between the trochleae metatarsorum III and IV is very narrow. The trochlea metatarsi IV itself is cylindrical and asymmetric in plantar view, with the lateral rim being larger than the medial one; it is not as narrow mediolaterally as the corresponding trochlea of crown-group Todidae and exhibits a morphology that is probably plesiomorphic for alcediniform birds. Most pedal phalanges, especially of the right foot, are dissociated (Fig. 6). As in extant Todidae, the hindtoe is very long and its fi rst phalanx exceeds the other phalanges in length; as in other Alcediniformes (Mayr 1998), the Fig. 6. Palaeotodus cf. itardiensis Mourer- proximal end of this phalanx bears a lateral pro- Chauviré, 1985: distal end of right tarsometa- jection. The phalanges bear a well-developed tarsus and pedal phalanges. Coated with tuberculum extensorium. The ungual phalanges ammonium chloride to enhance contrast. Scale are weakly curved; that of the third toe is the bar equals 5 mm. longest and straightest. 1302 Mayr and Knopf [Auk, Vol. 124

Discussion It is generally assumed that Alcediniformes originated in the Old World (e.g., Olson 1976, The fossil specimen described here supports Kepler 2001), to which Meropidae and almost all Mourer-Chauviré’s (1985) identifi cation of Alcedinidae are today restricted. In addition to Todidae in the Old World and further confi rms a few other potential synapomorphies, Todidae that stem-group Todidae had an intercontinen- and Momotidae share a derived morphology of tal distribution in the early Oligocene, occurring the caudal end of the mandible (Olson 1976). If in both North America and Europe. Although these taxa indeed are sister groups (see above), the North Atlantic had already opened in the the occurrence of Old World stem-lineage rep- early Oligocene (Smith et al. 1994), small volant resentatives of both would be remarkable and birds apparently could still disperse between place a caveat on biogeographical scenarios that Europe and North America by that time (see are deduced from the distribution of the extant Mayr 2006a). taxa (e.g., Cracra 2001). Most authors consider todies the sister taxon However, a recent molecular analysis of fi ve of motmots, which today occur in Central nuclear genes did not support a sister-group America and northern South America and are relationship between todies and motmots but the only other alcediniform taxon whose crown- provided strong support for a clade (Todidae + group representatives are restricted to the New [Momotidae + Alcedinidae]; Ericson et al. 2006). World (e.g., Cracra 1981, Maurer and Raikow This clade was also retained in separate analy- 1981, Mayr 1998, Snow 2001). ses of three of the fi ve investigated genes (myo- Occurrence of stem-group Todidae in Europe globin intron 2, ODC, β-fi brinogen intron 7), is even more remarkable, given that Protornis and only analysis of one gene (RAG-1) showed glarniensis von Meyer 1844, from the early Momotidae and Todidae to be sister taxa, albeit Oligocene of Switzerland, was identifi ed as a with weak support. The results of this analysis motmot by Olson (1976) and motmots thus have put the occurrence of stem-group Todidae and an Old World fossil record as well (the only other Momotidae in the Old World fossil record in pre-Quaternary fossil motmot was described by a new light, which then (i.e., if Todidae and Becker [1986] from the Late Miocene of Florida). Momotidae are not sister taxa) actually follows Identifi cation of P. glarniensis as a motmot from the currently assumed Old World origin of was challenged by Cracra (1980:13), who alcediniform birds. noted that an The crown-group representatives of sev- eral avian taxa that occur in the Paleogene of alternative hypothesis to be considered is a Europe are now restricted to the New World sister-group relationship between Protornis on (Mourer-Chauviré 1999, Mayr 2005b). The sur- the one hand and Todidae + Momotidae on the vival of crown-group representatives of both other. This hypothesis certainly would make more sense zoogeographically by restricting Momotidae and Todidae in the New World is, the todid–momotid lineage to the New World. however, remarkable, independent of their true interrelationships. In Europe, climatic deterio- However, although P. glarniensis diff ers from ration during the Miocene certainly led to the crown-group Momotidae in several features extinction of many insectivorous and frugivo- (proportionally shorter processus orbitalis of rous avian taxa with poor dispersal capabilities. quadratum and pars symphysialis of mandible, There must, however, have been additional be er-developed processus lateralis of coracoid), factors that prevented survival of stem-group an alternative classifi cation of this species has Todidae in the tropical parts of the Old World, not been established. The Frauenweiler fossil such as feeding competition with oscine song- is clearly distinct from P. glarniensis; apart from birds (Mayr 2005a, b) or predation by carnivo- being smaller, it has a proportionally longer tar- rous mammals. sometatarsus (length of ulna and tarsometatar- Assignment of the Frauenweiler fossil to sus 24.7 mm and 16.5 mm, respectively, versus the stem lineage of Todidae is unambiguous, 34 mm and 15 mm in P. glarniensis [a er Peyer because we could not identify derived char- 1957]), and the carpometacarpus bears a much acters supporting an alternative phylogenetic be er-developed processus intermetacarpalis than placement. However, there have been some that of P. glarniensis. major morphological transformations on the October 2007] Oligocene Tody from Germany 1303 lineage leading to crown-group Todidae, and Maurer, D., and R. J. Raikow. 1981. the diff erent shape of the distal end of the tar- Appendicular myology, phylogeny, and clas- sometatarsus in particular (see above and Fig. sifi cation of the avian order Coraciiformes 2) indicates diff erent locomotory adaptations in (including Trogoniformes). Annals of the Paleogene and extant Todidae. Carnegie Museum 50:417–434. Mayr, G. 1998. “Coraciiforme” und “piciforme” Acknowledgments Kleinvögel aus dem Mi el-Eozän der Grube Messel (Hessen, Deutschland). Courier We thank O. Vogel for the painstaking prepa- Forschungsinstitut Senckenberg 205:1–101. ration of the tiny specimen and S. Tränkner for Mayr, G. 2000. A new mousebird (Coliiformes: taking the photographs. G.M. thanks S. Olson Coliidae) from the Oligocene of Germany. for enabling study of the referred specimen of P. Journal für Ornithologie 141:85–92. emryi during a visit in 1996. C. Mourer-Chauviré Mayr, G. 2004a. A partial skeleton of a new is acknowledged for unpublished information fossil loon (Aves, Gaviiformes) from the on P. itardiensis. Finally, we thank C. Mourer- early Oligocene of Germany with preserved Chauviré, H. F. James, and an anonymous stomach content. Journal of Ornithology reviewer for comments on the manuscript. 145:281–286. Mayr, G. 2004b. Old World fossil record of Literature Cited modern-type hummingbirds. Science 304: 861–864. Baumel, J. J., and L. M. Witmer. 1993. Osteologia. Mayr, G. 2005a. A tiny barbet-like bird from the Pages 45–132 in Handbook of Avian Lower Oligocene of Germany: The smallest Anatomy: Nomina Anatomica Avium (J. J. species and earliest substantial fossil record Baumel, A. S. King, J. E. Breazile, H. E. Evans, of the Pici (woodpeckers and allies). Auk and J. C. Vanden Berge, Eds.). Publications of 122:1055–1063. the Nu all Ornithological Club, no. 23. Mayr, G. 2005b. The Paleogene fossil record Becker, J. J. 1986. 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