Pal~iontologische Zeitschrift 74 (1/2) 187-194 7 Abb., 2 Tab. Stuttgart, Mai 2000

A remarkable new 'gruiform' from the Middle 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 '') 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 und einigen non-'gruiform' neognathous . 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 is assumed to have had a predomi- the 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 , , 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- (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 '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, , gallinules), Gruidae lated to the Eurypygidae). The fossil birds presented (cranes), Aramidae (), (finfoots, herein add a very distinctive new taxon to this list. They sungrebes), Rhynochetidae (), 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 (), Otididae (), 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 , 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 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; 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 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 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 (). 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, ) 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; = 10mm. A remarkable new 'gruiform' bird from the Middle Eocene of Messel (Hessen, Germany) 189 190 GERALD MAYR

Fig. 2. Salmila robusta n. gen., n. sp. - referred specimen SMF- Fig. 3. Salmila robusta n. gen., n. sp. (SMF-ME 3014). - De- ME 617. - Scale bar -- 10 mm. tail of Fig. 1 showing the pectoral region. - Scale bar = 10 mm.

Description and comparison: Coracoid (Fig. 3): The coracoid is fairly long and slen- Skull: The skull is rather small in comparison to the re- der. The tuberculum brachiale is well developed and bent mainder of the body. Due to preservation of both known far medially which gives the extremitas omalis a hooked specimens, no details of the cranium can be discerned and appearance, like in the Cariamidae, Bathornithidae and only the bill, which in comparison with recent birds Idiornis (not Elaphrocnemus). Contrary to the Cariami- seems to have been most similar to the of the dae and Idiornis, the processus acrocoracoideus does not Otididae, allows a more detailed description. It is long seem to have been connected by an osseous bridge with and slender, measuring approximately half of the overall the processus procoracoideus (which itself is not visible). length of the skull. The culmen is fairly straight in the The processus lateralis of the extremities sternalis is proximal two thirds of the beak and curved on the level short, the angulus medialis pointed. of the rostrum maxillae. The narial openings (visible through the reverse of the transparent slab) reach ap- Furcula (Fig. 3): The furcula is U-shaped like that of the proximately two third of the entire length of the bill, but Rallidae, but the scapus claviculae and especially the it is not clearly visible if the beak of Salmila robusta has extremitas omalis are more robust and cranio-caudally been holorhinal (like in the Rallidae, Otididae, Caria- much wider than in most other 'Gruiformes'. An apophy- midae, and Psophiidae) or schizorhinal (like in all other sis furculae seems to be absent. 'gruiform' birds). The processus praemaxillaris of the os Sternum (Fig. 4A): The sternum is large and has a very nasale is slender. The rami mandibulae are straight and characteristic shape. Like in the Turnicidae, Rallidae, and of average height, a fenestra mandibulae is not visible. Mesitornithidae (Figs. 4B-D) the margo caudalis is in- Vertebrae: The cervical vertebrae are short (SMF-ME cised by two deep notches which reach more than half 617), but details of their morphology cannot be dis- the length of the sternum. Yet, in contrast to the sternum cerned. of the Mesitornithidae the trabecula mediana is narrow A remarkable new 'gruiform' bird from the Middle Eocene of Messel (Hessen, Germany) 191

Fig. 4. Sterna in comparison. - A: Salmila robusta n. gen., n. sp.; B: Turn& tanki (Turnicidae); C: Gallinula chloropus (Rallidae); D: Mesitornis unicolor (Mesitornithidae, after LowE 1924); E: Cariama cristata (Cariamidae); F: Psophia viridis (Psophiidae). - Scale bar = 10 mm.

and has a triangular outline, and unlike the Rallidae it reaches farther caudally than the lateral trabeculae (but not as far as in the Turnicidae). The lateral trabeculae are slender, their caudal end is transversely widened, like in Turnix (Turnicidae). Details of the cranial end of the ster- num are not visible. The carina sterni is low. Humerus: The humerus is stouter and more robust than that of most other 'gruiform' birds. In its proportions, especially concerning the proximal end, it is closest to the humerus of the Otididae. The crista deltopectoralis which measures less than one third of the entire length of the humerus has a convex dorsal margin. The caput hu- meri is large and only slightly inflected caudally. The crus Fig. 5. Salmila robusta n. gen., n. sp. (SMF-ME 3014). - De- ventrale fossae is proximo-distally wide when seen from tail of Fig. 1 showing the distal end of the right wing. Covered ventrally, like in Psophia or Opisthocomus (SMF-ME with ammonium chloride. - Scale bar = 10 mm. 3014, right side). Details of the distal end of the humerus are hardly visible, but the processus flexorius is short and a processus supracondylaris dorsalis absent. grallator (see WETMORE 1944), and bears a short claw Ulna: Like in many other 'gruiform' birds (except (Fig. 5). The phalanx digiti minoris is short. Gruidae, Otididae), the ulna of S. robusta does not ex- ceed the humerus in length. In its proportions it is similar Pelvis: The pelvis is wider than that of most recent to the corresponding bone of Psophia. The olecranon is 'gruiform' birds except the Mesitornithidae and Otididae. short, the tuberculum ligamenti collateralis ventralis ap- As far as comparable it is most similar to the pelvis of pears to have been low. galliform birds in the general proportions, the shape of the broad alae ischii, and the length of the os pubis. A Carpometacarpus (Fig. 5): The short carpometacarpus short processus terminalis ischii is present, the fenestra most closely resembles that ofPsophia (Psophiidae) and ischiopubica is narrow and slit-like. A spina dorsolate- Idiornis (Idiornithidae). The os metacarpale minus is ralis ilii seems to have been absent. bowed and its proximal end dorso-ventrally wide; like in Psophia, Cariama, Ameghinornis, and Idiornis it bears Femur: The femur is only poorly preserved. It is long and a small tubercle on the proximal end of its ventral side. becomes markedly wider towards its distal end. The extremitas proximalis of the carpometacarpus is Tibiotarsus: The tibiotarsus is the longest limb element. large. The processus pisiformis is stubby and shifted Both cristae cnemiales are enlarged (the crista cnemialis craniad; its tip also projects cranially. The portion of the cranialis is visible at the right tibiotarsus of the holotype). trochlea carpalis between the processus pisiformis and As far as comparable, the distal end of the bone is similar the os metacarpale minus is distinctly raised. The sym- to that of the Middle Eocene putative rail Fulicaletornis physis metacarpalis distalis is wide. A processus inter- venustus (MARSH 1872), compared to recent birds it most metacarpalis is absent. closely resembles that of Psophia sp. The condylus me- Other elements of the wing: The phalanx digiti alulae is dialis is proximo-distally slightly shorter than the con- very long, like is that of (= 'Neocathartes') dylus lateralis, the latter is also medio-laterally much 192 OERALDMAYR

wider than the condylus medialis. The epicondylus me- dialis is raised. The incisura intercondylaris is fairly wide, a pons suprantendineus cannot be recognised but might have been lost during preparation.

Tarsometatarsus (Fig. 6): The tarsometatarsus is shorter than the humerus and measures two thirds of the length of the tibiotarsus (in contrast thereto the long tarso- metatarsus of the Cariamidae, Idiornithidae, Bathorni- thidae, and Psophiidae exceeds the humerus in length). As far as it can be compared it closely resembles the cor- responding bone ofAenigmavis sapea PETERS 1987 (Fig. 7), but apart from being shorter and stouter it is also simi- lar to the tarsometatarsus of Idiornis. The shaft is most narrow on the level of the fossa metatarsi I and becomes gradually wider towards the proximal end of the bone. Like in Aenigmavis (and Idiornis) the cotyla medialis is situated farther proximally than the cotyla lateralis. The eminentia intercotylaris is prominent. The fossa infra- cotylaris dorsalis is well developed. Some features of the plantar surface of the tarsometatarsus can be seen through the reverse of the transparent slab: A well developed crista medialis hypotarsi is visible, and apart from a crista medianoplantaris both, a crista plantaris medialis and a crista plantaris lateralis are present which converge to- wards the distal end of the tarsometatarsus. The distal tarsometatarsus itself resembles that of Idiornis and Psophia. The foramen vasculare distale is small. The tro- chlea metatarsi III reaches farther distally than the other two trochleae metatarsorum and is longer than wide; its Fig. 6. Salmila robusta n. gen., n. sp. (SMF-ME 3014). - De- tail of Fig. 1 showing the left foot. Covered with ammonium articular surface extends farther proximally than in chloride. - Scale bar = 10 mm. Aenigrnavis. The trochleae metatarsorum II and IV are on the same level and both trochleae bear a plantarly di- recting process (visible through the reverse of the slab). The dorsal surface of the trochlea metatarsi II is rounded, like in Aenigmavis and Psophia.

Toes: The toes are robust and resemble those of galliform birds in their relative length. The third toe is the longest, the second and the fourth toe are distinctly shorter; the fourth toe is slightly shorter than the second. The hallux corresponds well with that ofldiornis tuberculata PETERS 1995; it is short like in most recent 'gruiform' birds, and bears a stubby claw. The claws of the other three toes are only slightly curved and resemble those of Rhynochetos jubatus ('Gruiformes', Rhynochetidae). The tuberculum flexorium is situated as far proximally as the tuberculum extensorium. The claw of the fourth toe is smaller than those of the second and third toe. The processus articularis tarsometatarsalis of the os metatarsale I is short, like in Cariama cristata. : If the interpretation of the dark shadow on the slab of the type specimen as remains of the tail feathers is correct, the long tail (140 ram) distinguishes S. robusta from all other 'gruiform' birds except the Mesitornithi- dae. The maximum length of the wing feathers measures as preserved N110 mm. Fig. 7. Aenigmavis sapea PETERS 1987 (SMF-ME 1819), left tarsometatarsus. - Scale bar = 10 mm. Dimensions: see Tabs. 1 and 2. A remarkable new 'gruiform' bird from the Middle Eocene of Messel (Hessen, Germany) 193

Tab. 1. Salmila robusta n. gen., n. sp. - length of the limb bones in comparison (left/right, in ram).

humerus ulna carpo metacarpus femur tibiotarsus tarsometatarsus SMF-ME 3014 53.4 / 56.0 -53 / -52 / 27.0 / N45 69.7 / 64.4 42.9 / 44.7 SMF-ME 617 -47 / -46 -46 / -48 -23 / -23 -37/ -41.5 /

Tab. 2. Salmila robusta n. gen., n. sp. - length of the pedal phalanges (in mm).

I 1 I2 I11 II2 II3 III 1 1II2 III3 III4 IV 1 IV2 IV3 IV4 IV5 SMF-ME3014 7.6 5.0 11.8 8.2 6.7 11.2 9.3 8.4 8.3 7.6 4.2 4.2 5.1 5.6

Discussion are unknown. Having been greatly reduced like in all other phorusrhacid birds, the wing skeleton of A. sapea Resembling that of galliform birds, the foot of Salmila cannot be compared with that of S. robusta. robusta n. gen., n. sp. suggests that this species had a pre- All Phorusrhacidae have at best been weak flyers and dominantly terrestrial habit. The small hallux with its in most cases were even entirely flightless. The recogni- stubby claw probably largely prevented it from perching. tion of this family, which formerly mainly has been According to the shape of its bill, S. robusta presumably known from South America (a single genus was found in had a similar diet like recent bustards, consisting of vari- the Pleistocene of North America), in the Early Tertiary ous seeds and berries as well as of small vertebrates and of France and Germany thus was a very unexpected dis- invertebrates (in the type specimen a thick carbonised covery (MOURER-CHAUVIRI~1981, 1983; PETERS 1987). layer of the putative former stomach content is preserved To explain the paleobiogeography of these birds it has which has not yet been closer investigated). been suggested that the Phorusrhacidae are either a very The new species from Messel has been classified into old group which originated prior to the break-up of the 'Gruiformes' in this study and within this taxon it is Gondwanaland and dispersed to Europe via Africa, or tentatively referred to the Cariamae. An exhaustive phy- that members of this family migrated to Europe via North logenetic analysis, however, is aggravated by the poorly America (MOURER-CHAUVIRI~1982; PETERS & STORCH understood relationships of recent 'gruiform' birds and 1993). by the lack of an updated revision of the Phorusrhacidae. Yet, concerning the skeletal remains of the 'Quercy- The recent members of the Cariamae as well as the phorusrhacid' Ameghinornis MOURER-CHAUVIRt; 1981, Idiornithidae and Bathornithidae are long-legged birds, OLSON (1985: 146) already pointed out, that there "is a but the comparatively short tarsometatarsus of S. robusta possibility that these elements of the wing and shoulder is not in general conflict with an assignment to these taxa, girdle of Ameghinornis evolved their similarities to South since at least the Phomsrhacidae comprise short-legged American phorusrhacids in parallel, and that their simi- species, too. larities are due in part to degenerative reduction". Fur- In some features, however, Salmila robusta distinctly ther better preserved specimens ofAenigmavis sapea and differs from all other Cariamae. Especially the extremitas a revision of the Phorusrhacidae will likewise have to omalis of the furcula and the proximal end of the humerus evaluate the possibility, that A. sapea is a flightless (or are much more robust than the corresponding elements nearly so) relative of S. robusta which convergently of other 'gruiform' birds. The very wide pelvis is unu- evolved some phorusrhacid-like features (e. g. the raptor- sual for 'gruiform' birds, too, and the shape of the caudal like claws). margin of the sternum is unique among all avian taxa known so far and differs greatly from the sternum of the Cariamidae, and Bathornithidae (see Fig. 4E and Acknowledgements CRACRAFT 1968: fig. 8). In the latter two features and in I would like to thank D. S. PETERS (Forschungsinstitut the long tail S. robusta more closely resembles the enig- Senckenberg, Frankfurt am Main) for comments on the manu- script, and S. TRA.NKNER (Forschungsinstitut Senckenberg, matic Madagascan Mesitornithidae which on the other Frankfurt am Main) for taking the photographs. hand in many characters clearly are distinguished (e. g. in the well-developed hallux and in the morphology of the pectoral and wing skeleton). References The similarities between the tarsometatarsus of S. BAUMEL, J.J. & WITMER, L.M. 1993. Osteologia. - [In:] BAU- robusta and that of the 'Messel-phorusrhacid' Aenig- MEL, J.J.; KING, A.S.; BREAZILE, J.E.; EVANS, H.E. & mavis sapea PETERS 1987 are especially noteworthy (see VANDEN BERGE, J.C. [eds.] Handbook of avian : Nomina Anatomica Avium. - Publications of the Nuttall description and Figs. 6 and 7), but the only known sub- Ornithological Club 23: 45-132, Cambridge/Mass. stantial skeleton of A. sapea is rather poorly preserved BRODKORB, R 1967. Catalogue of fossil birds. Part 3 and particularly the shape of the beak and the sternum (Ralliformes, Ichthyornithiformes, ). - 194 GERALD MAYR

Bulletin of the Florida State Museum (Biological Sciences) MOURER-CHAUVlRI~, C. 1981. Premibre indication de la pr6- 11 (3): 99-220, Gainesville. sence de Phorusrhacid6s, famille d'oiseaux g6ants d'Am6- CRACRAFT, J. 1968. A Review of the Bathornithidae (Aves, rique du Sud, dans le Tertiaire europ6en: Ameghinornis Gruiformes), with Remarks on the Relationships of the nov. gen. (Aves, Ralliformes) des Phosphorites du Quercy, Suborder Cariamae. - American Museum Novitates 2326: France. - G6obios 14 (5): 637-647, Lyon. 1-46, New York. -- 1982. Les oiseaux fossiles des Phosphorites du Quercy -- 1973. Systematics and of the Gruiformes (Class (Eoc6ne sup6rieur ?t Oligoc6ne sup6rieur): Implications Aves). 3. Phylogeny of the suborder Grues. - Bulletin of pal6obiog6ographiques. - [In:] BUFFETAUT,E.; JANWER, P.; the American Museum of Natural History 151: 1-127, New RAGE, J.-C. & TASSY, P. [eds.] Phylog6nie et Pal6o- York. biog6ographie. Livre jubilaire en l'honneur de Robert -- 1982. 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181-199, Frankfurt am Main. Annahme durch die Schriftleitung am 5. August 1999.