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'Gruiform' Bird from the Middle Eocene of Messel (Hessen, Germany)

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'Gruiform' Bird from the Middle Eocene of Messel (Hessen, Germany) Pal~iontologische Zeitschrift 74 (1/2) 187-194 7 Abb., 2 Tab. Stuttgart, Mai 2000 A remarkable new 'gruiform' bird from the Middle Eocene of Messel (Hessen, Germany) GERALD MAYR, Frankfurt am Main With 7 figures and 2 tables Kurzfassung: Salmila robusta n. gen., n. sp. (Aves: phylogenetic analysis performed by ERICSON (1997), for 'Gruiformes') wird aus dem Mittel-Eoz~in von Messel (Hes- example, resulted in a sister-group relationship between sen, Deutschland) beschrieben. Die neue Art war vermutlich vorwiegend bodenbewohnend und teilt abgeleitete Merkmale the Mesitornithidae and a number of other 'gruiform' and des Carpometacarpus mit den rezenten Psophiidae und einigen non-'gruiform' neognathous birds. After studying the Taxa der Cariamae. Sie unterscheidet sich jedoch in einigen phylogeny of 'gruiform' birds with molecular techniques, Merkmalen deutlich von anderen 'gruiformen' VGgeln (z.B. im HOUDE et al. (1997) also concluded that this taxon prob- grGfSeren proximalen Ende des Humerus und in der Form des ably is not monophyletic. Nevertheless, cited in quota- caudalen Sternumrandes). Salmila robusta wurde unter Vorbe- halt zu den Cariamae gestellt, aber ihre genaue systematische tion marks, the term 'Gruiformes' is used in a traditional Stellung mul3 noch weiter untersucht werden. Der recht kurze sense (e. g. WETMORE 1960) herein to facilitate com- Tarsometatarsus der neuen Art ~ihnelt dem des 'Messel- parisons. Phorusrhaciden' Aenigmavis sapea PETERS 1987. A phylogenetic analysis becomes much more difficult, if the numerous extinct 'gruiform' families which have Abstract: Salmila robusta n. gen., n. sp. (Aves: 'Gruiformes') is described from the Middle Eocene of Messel (Hessen, Ger- been described so far are also included (for a survey over many). The new species is assumed to have had a predomi- the fossil taxa see CRACRAET 1973; OLSON 1985; nantly terrestrial habit. It shares derived features of the carpo- FEDUCCIA 1996). Yet, most authors agreed on a mono- metacarpus with the recent Psophiidae and some taxa of the phyly of the 'suborder' Cariamae including the Caria- Cariamae. In several features, however, it distinctly differs midae and the extinct Idiornithidae, Bathornithidae, and from other 'gruiform' birds (e. g. in the larger proximal end of the humerus and in the shape of the margo caudalis of the ster- Phorusrhacidae (e.g. BRODKORB 1967; CRACRAFT 1968; num). Although Salmila robusta has been tentatively assigned MOURER-CHAUVIRE 1981, 1983; OLSON 1985). Compar- to the Cariamae in this study, its exact systematic position de- ing the extant birds, STRESEMANN (1927-34), STEGMANN serves further investigations. The rather short tarsometatarsus (1978), and CRACRAFT (1982) considered the Psophiidae of the new species resembles that of the 'Messel-phorusrhacid' to be the closest relatives of the Cariamidae, but this has Aenigmavis sapea PETERS 1987. not been confirmed by SmLEY & AHLQUIST (1990) and HOUDE et al. (1997). Three 'gruiform' families have been identified from the Introduction Middle Eocene deposits of Messel (Hessen, Germany) so The avian order 'Gruiformes' is a quite heterogeneous far: the Idiornithidae, Phorusrhacidae, and the Messel- assemblage which generally comprises the following re- ornithidae (which HESSE 1990, believed to be closely re- cent families: Rallidae (rails, coots, gallinules), Gruidae lated to the Eurypygidae). The fossil birds presented (cranes), Aramidae (limpkins), Heliornithidae (finfoots, herein add a very distinctive new taxon to this list. They sungrebes), Rhynochetidae (kagu), Eurypygidae (sun- share several features with some of the birds included in bittern), Mesitornithidae (roatelos), Psophiidae (trumpet- the Cariamae and Psophiidae, but in others distinctly dif- ers), Cariamidae (seriemas), Otididae (bustards), and fer from all 'Gruiformes' known so far. (For additional Turnicidae (hemipodes). A review of the history of clas- information on the avifauna of Messel see HESSE 1990; sification is given by SIBLEY 8z AHLQUIST (1990). MAYR 1998a, 1998b; MAYR & DANIELS 1998; MAYR & The monophyly of the 'Gruiformes' has not been suffi- PETERS 1998; PETERS 1987, 1988, 1995; general informa- ciently supported with derived characters so far and a tion on the site can be found in SCHAAL & ZIEGLER 1988). Address of the author: Dr. GERALD MAYR, Forschungsinstitut Senckenberg, Division of Ornithology, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany. E-mail: [email protected] 0031-0220/00/0074-0187 $ 4.00 2000 E. Schweizerbart'sche Verlagsbuchhandlung, D-70176 Stuttgart 188 GERALD MAYR Material and methods - furcula U-shaped with a cranio-caudally wide extre- mitas omalis; The fossil specimens are deposited in the Forschungsinstitut -sternum large with only two deep incisions in the Senckenberg (SMF), Frankfurt am Main (Germany). The ana- margo caudalis of the corpus sterni; trabecula mediana tomical terminology follows BAUMEL & WITMER (1993). The narrow, of triangular shape and reaching farther dimensions represent the maximum length of the bone along its longitudinal axis; concerning the length of the claws the caudally than the lateral trabeculae; carina sterni low; distance between the tuberculum extensorium and the apex - humerus stout and with large proximal end; phalangis has been measured. - carpometacarpus short with bowed os metacarpale mi- nus (other distinctive features of this bone have been listed above); Systematics - pelvis large and medio-laterally wide; -tibiotarsus with condylus lateralis medio-laterally Class Aves LINNAEUS 1758 much wider than condylus medialis, raised epicondylus Order 'Gruiformes' (BONAPAkTE 1854) medialis, and fairly wide incisura intercondylaris; ? Suborder Cariamae F~RBRINGER 1888 -tarsometatarsus shorter than humerus and robust; Family incertae sedis cotyla medialis reaching farther proximally than cotyla R e m a r k s : Apart from being most similar to these taxa lateralis; foramen vasculare distale small; trochleae in their 'overall morphology', too, the fossil birds from metatarsorum II and IV shorter than trochlea metatarsi Messel described in this study share features of the car- III, reaching equally far distally and bearing a process pometacarpus with the Psophiidae, Cariamidae, Bathor- directing to palmar side of bone; dorsal surface of tr. nithidae, and most Idiornithidae which most likely are mt. II rounded; derived (due to their absence in most other neognathous - hallux short and with stubby claw. birds): R e m a r k s : Based on fragmentary isolated bones, sev- - the processus pisiformis is stubby and shifted craniad eral 'gruiform' birds have been described from the Lower with its tip also projecting cranially; and Middle Eocene of North America and Europe (see -the portion of the trochlea carpalis between the CRACRAFT 1973; HARRrSON & WALKER 1977, 1979; processus pisiformis and the os metacarpale minus is HARRISON 1984). As far as comparable, the distal end of distinctly raised; the tibiotarsus of Salmila n. gen. especially resembles - the os metacarpale minus is bowed and has a dorso- that of the putative rail Fulicaletornis venustus (MARSH ventrally wide proximal end (see ERICSON 1997); 1872) from the Middle Eocene of Wyoming (CRACRAFT -- the proximal end of the os metacarpale minus bears a 1973: fig. 5). Yet, since F. venustus is known from the small tubercle on its ventral side (I could not check the distal tibiotarsus only, which in the new genus from presence of this character in the Bathornithidae; it is Messel is only visible from its cranial side and partially absent in the idiornithid genus Elaphrocnemus, but covered by the tarsometatarsus, a detailed comparison is present in the phorusrhacid Ameghinornis; see not possible. F. venustus is smaller than the new species MOURER-CHAUVIRI~ 1981, 1983). from Messel described below. As mentioned in the introduction, modern phylogenetic studies could not confirm a monophyly of the Psophiidae and Cariamae, and therefore the features of the carpo- Salmila robusta n. sp. metacarpus listed above obviously evolved two times Figs. 1-6 independently within the 'Gruiformes'. A tentative as- E t y m o I o g y : The specific name has been derived from signment of the Messel-birds to the Cariamae might be 'robustus' (Lat.) = 'robust' and refers to the robust physique of supported with the well developed tuberculum brachiale the species. of the coracoid which in the new taxon is bent far medi- Holotype: SMF-ME 3014 (almost complete articulated ally like in the Cariamidae, Bathornithidae and Idiornis skeleton; Fig. 1). Type locality: Grube Messel (Hessen, Germany). (but not Elaphrocnemus). Ty p e h o r i z o n : Geiseltalium, lower Middle Eocene. Referred specimen: SMF-ME 617 (incomplete and poorly preserved articulated skeleton lacking the right leg; Salmila n. gen. Fig. 2). Type species: Salmila robusta n. sp. E t y m o 1o g y : The generic name is an anagram of 'masilla', D i a g n o sis: Only species of the genus, therefore same the old Latin name of Messel, and is feminine in gender. diagnosis as for genus. Salmila robusta n. sp. has ap- proximately the size ofLagopus rupestris (Galliformes). D i ag n o s i s : Salmila n. gen. comprises medium-sized birds with limb proportions similar to Galliformes (e. g. Crax alector, Cracidae) and is characterised by the fol- lowing characters: Fig. 1. Salmila robusta n. gen., n. sp. - type specimen (SMF- - bill long and slender with an, except for the curved tip, ME 3014). Covered with ammonium chloride. - Scale bar straight culmen;
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