Paleontology. - A femur of palaeojavanica Dubois from Western Java. By D. A. HOOIJER. (Communicated by Prof. H. BOSCHMA.)

(Communicated at the meeting of March 29, 1947.)

STEHN and UMBGROVE (1929) have recorded fossil remains of Bos, Sus and Cervus from the Tji Taroem valley W. of Batoedjadjar, Preanger, W. Java. The collection was presented to the Geological Museum at Leiden by Prof. Dr. J. H. F. UMBGROVE. The exact geological age of the fossiIs could not be established; they might have originated from the neighbouring Neogene limestone. VAN Es (1931, p. 15) states that the absence of proboscideans points to a younger age than that of the otheroccurrences of vertebrates. VON KOENIGSWALD (1933, pp. 44, 47) refers the Sus speci~ men, a right ramus with P3-M2' to his species Sus terhaari. I cannot see the reason, however; in a later paper (VON KOENIGSWALD, 1935, p. 87) the fossil is left specifically undetermined. The cervine remains, of which two teeth have been figured by STEHN and UMBGROVE (l.c., pI. IV, figs. 1-3), were referred to Cervus unicolor Kerr (c. hippelaphus Smith) by VON KOENIGSWALD (1933, p. 74). Afterwards, however, they are identified as "Cervus (ex aH. zwaani)" (the upper molar) and "cf. Antilope modjoker~ tensis v. K." (Iower molar). The bovine fossiIs should belong to Bubalus palaeokerabau Dubois (VON KOENIGSWALD, 1935, p. 87/88). To this fauna only Rhinoceros sondaïcus Desmarest has been added since by VON KOENIGSWALD (1935, p. 87). who regards the age of the fossils as very probably lower Pleistocene. Rhinoceros sondaïcus Desma~ rest, however, ranges from the lower Pleistocene (Djetis fauna) to the present day (HOOIJER, 1946, pp. 3, 34-81 ) ; Bubalus palaeokerabau Dubois was not known previously from the lower Pleistocene (cf. VON KOENIGS~ WALD 1934, p. 191). but indeed the deer remains may belong to species which are said to be typically of lower Pleistocene age. To my great surprise I found a right femur in the STEHN and UMBGROVE collection (Geol. Mus. Leiden, no. 28069) which does not belong to either of the genera mentioned above. It was catalogued as "Felis sp.". The peculiar flatness of the shaft and the comparatively slight prominence of the condyles, however, show this identification to be incorrect. The bone belongs to:

Manis palaeojavanica Dubois.

Manis palaeojavanica Dubois, Tijdschr. Kon. Ned. Aardr. Gen., ser. 2, vol. 24, 1907, p. 455; STREMME, N. Jahrb. f. Min., 1911. p. 55; STREMME4 in: SELENKA-BLANCKEiNHORN, Die Pithecanthropus-Schichten auf Java, Leipzig, 1911. p. 141; MARTIN, Unsere pa!aeo­ zoologische Kenntnis von Java, Leiden, 1919, p. 107; VAN Es, ll1e Age of Pithecanthropus, 'The Hague, 1931, p. 31; VON KOENIOS'WALD, De Ing. in Ned. Ind., vol. I, part. 11. 414 sect. IV, 1934, pp. 190, 191, 196; KORMOS. Folia Zool. et Hydrob., vol. 6, 1934, p. 88; VON KOENIGSWALD, Quartär, vol. 2, 1939, p. 35; BOK, Bijdrage tot de kennis van de raseigenschappen van het Javaansche volk, thesis Utrecht, 1940, p. 4; VON KOENIGSWALD, 'Wet. Med. Dienst Mijnbouw Ned. Indië, no. 28, 1940, p. 56. Manis palaejavanica Dubais, Tijdschr. Kon. Ned. Aardr. Gen., ser. 2, vol. 25, 1908, p. 1267, Versl. Verg. Afd. Natuurk. Kon. Akad. v. Wetenseh., Amsterdam, Vol. 35, 1926, p. 949, Proc. Kon. Akad. v. Wetenseh., Amsterdam, vol. 291, 1926, p. 1233; VAN DER MAAREL, Leidsche Geol. Med., vol. 5, 1931, p. 472; HOOI}ER, Zool. Med. Museum Leiden, voJ. 26, 1946, p. 107. M[anis] palaejavanico Weber, Die Säugetiere, vol. 2, Jena, 1928, p. 185. [Manis] paleojavanica Raven, Bull. Am. Mus. Nat. Hist., vol. 68, 1935, p. 238.

The present giant was reported upon first by DUBOlS in 1892 (ANONYMUS, 1892, p. 14) on the evidence of remains of a skeleton from Kedoeng Broeboes, a fossil locality in the Kendeng Beds in Central Java which two years earlier had provided DUBOlS with a mandible fragment which he finally attributed to Pithecanthl'opus erectus (Dubois) but the systematic position of which is still uncertain (WEIDENREICH, 1945, p. 103/104). The total length of the pangolin is statecl to have been 2t m, and it also must have been more heavily built than the recent Manis jal'a~ nica Desmarest. The same statements we find in another paper (DUBOIS, 1907, p. 455) in which the fossil form is designated as Manis palaeojava~ nica. A more eIaborate description and comparison with other forms is given in 1908 under the name "Manis palaejavanica n. sp." (DUBOIS, 1908, p. 1267). In his final description of the remains DUBOlS (1926) only rders to the latter paper and name. Manis palaeojavanica Dubois is the best known of the fossil . We know parts of its skull, atlas, caudal vertebrae, radius, ulna, femur anèl tibia, as well as metacarpals, phalanges, astragalus, calcaneum, navicular, ectocuneiform and metatarsals. Fram DUBOIS' paper of 1926 it is clear that the species displays the greatest resemblance to Manis javanica Desmarest, though in size it comes nearest to Manis gigantea IIIiger which is about one~third smaller. Fram the characters of the endocranial cast and of the foot bones DUBOlS inferred that in all probability the fossil form was covered with scales, that it had large claws on its pes as weil as on its manus, and that it walked permanentlyon the outside of its feet. Some other fossil pangolins have been based only on isolated phalanges, e.g., Manis lydekkeri Trouessart (olim Manis gigantea Lydekker) from the Pleistocene Karnul caves near Madras in India and Manis sindiensis Lydekker from the Siwaliks of Sind (LYDEKKER, 1886, p. 50). as well as Manis hungarica Kormos (1934) from the Villafranchian of VilIány in Hungary. HELBING (1938) assembles previous records of Tertiary pango~ !ins in Europe (Quercy, Solnhofen and Wintershof West) and describes remains of Manis of Oligocene age from St. André near Marseilles, Mon~ taigu~le~Blin (Allier) and Weisenau near Wiesbaden. None of the species, named or unnamed, is as large as Manis palaeojavanica Dubois. 415

The newly~discovered femur is of the right side. The proximal portion has broken oH; the medial surface of the bone is slightly damaged below. The corpus is flattened from before backwards. At the narrowest part, about 4 cm below the broken edge, the width is one~half greater than the antero~posterior diameter. The shaft increases in width distally, its medial border is somewhat more rounded than the lateral and ends in the medial epicondyle which is somewhat less prominent and placed on a slightly higher level than the epicondylus lateralis. Just above the latter the border of the bone is very prominent, rather a crest. The bone belonged to an adult individual, but the suture of the dis tal epiphysis is still partly shown. Measured across the epicondyles the bone is twice as broad as at the narrowest part of the shaft. The trochlea is much more prominent medially than laterally; the facies pateIlaris is rounded oH above with its highest point to the medial side of the axis of the corpus. Of the two condyles the media I is by far the larger; it projects more backwards and is placed slightly higher than the lateral. lts articular surface is more strongly convex from side to side than in the lateral condyle, in which it is narrower and slopes down obliquely to the inside. The posterior surface, just above the intercondyloid fossa, presents a faint depression, but otherwise the posterior surface is slightly convex from side to side, especially towards the middle of the shaft. The only parts of the hind limb of Manis palaeojavanica Dubois that were known already from DUBOIS' descriptions consist of the medial half of the proximalextremity of a right femur with almost the entire head and the lesser trochanter, and the proximal half of a Ie ft tibia. The type speci~ mens are not fully adult; the epiphyses are not yet fused with the shaft. The Preanger femur thus fills up a hiatus in our knowledge of the osteology of this interesting species, and I was anxious to see wh ether the characters that can be drawn from this part of the skeleton would be in accord with those already found by DUBOlS or not. DUBOIS compared his fossil specimens with a subadult skeleton of Manis gigantea IIliger fr om Liberia in the Leiden Museum (cat. a) which I have also at my disposal, and with a more than middle~sized skeleton of Manis javanica Desmarest in his private collection. It is about one~third larger than the almost adult female skeleton of the same species in the Leiden Museum (reg. no. 1775, from the Rotterdam zoological garden, 13~9-1929) of which the measurements (in mm) are given in the third column of the table below. Prom this table it results that the index pilastricus of palaeojavanica is intermediate between those of gigantea and javanica; the figures for the index poplitaeus and the distal width index are greater, and that for the condyle index seems to be smaller than in the recent species. Unfortunately the lack of the proximal part of the fossil femur makes it impossible to compute more and better indices. 416

Manis Manis Manis palBeo- Femur gigBntea jBvanica javBnica IIIiger Desmarest Dubois

1. Antero-posterior diameter of caput . . 40 28 11 2. Antero-posterior diameter at trochanter minor .... . •..... 28 14 7i 3. Distance from top of trochanter minor to top of caput ...... 52 40 16 4. Width at narro\vest part of shaft 32l 22 9 5. Antero-posterior diameter at same level 21~ 13 7 meas. no. 5 X 100) 6. Index pilastricus ( meas. no. 4 66 59 78 7. Width of distal end of corpus . . . . 60 49 19 8. ft.ntero-posterior diameter of the latter . 41 29 12 meas. no. 8 X 100) 9. Index poplitaeus ( meas. no. 7 68 59 63 10. Greatest distal width ...... 66 53 20 meas. no. 4 X 100) 11. Distal width index ( 10 12 45 meas. no. 49 12. Width of condyles . • ...... 57 46 18 13. Greatest width of facies patelIaris 37 31 10 14. Distal antero-posterior diameter from medial ridge of trochlea to line connecting posterior surfaces of condyles . . . . 54 39 16 15. Height of condylus medialis . . . . . ca. 27 24 9 meas. no. 15 X 100) 16. Condyle index ( meas. no. 12 ca. 46 52 50

In search for other recent species of Manis in which the distal extremity of the shaft of the femur is Ie ss compressed antero~posterior1y , and in which the distal epiphysis is more broad than gigantea and javanica, I found Manis temminckii Smuts (Leiden Museum, cat. a) to have a distal width index of 48; unfortunately the index poplitaeus could not he computed hecause the sutures are invisihle. In th is adult specimen and in an adult of Manis longicaudata (Brisson) (Leiden Museum, cat. a) the index pilastri~ cus is 65 to 66. Both femora also possess a distinct supracondyloid crest which has not so weIl developed in the other, younger, specimens I had for comparison. In Manis aurita Hodgson (Leiden Museum, cat. a) the popli~ teal index of the femur is 56, the same figure I found in Manis tricuspis Rafinesque (Leiden Museum, cat. a). Thus a distinctive feature of the femur of Manis palaeojavanica Duhois, as compared to the other species .of its genus, seems to he the thickness of the lower end of the shaft. But certainly th ere is an amount of variation in this character; so, e.g., in two specimens of Tamandua tetradactyla (L.) in the Leiden Museum (reg. nos. 1163 and 1773) the index poplitaeus is 63 and 56 respectively. Both specimens are females. The diHerence between the distal antero~posterior diameter (meas. 417

no. 14) and the condyle width (no. 12 in the tabIe) is less in the fossil form than that in the recent. This ratio is 0.95, against 0.85 and 0.89 in gigantea and javanica. This is remarkably weIl in accord with the shape of the approximal surface of the tibia of palaeojavanica in the Dubois collection. The proximal extremity of the tibia, just like the distal extremity of the femur here described, is thicker in proportion to its width than in gigantea and javanica. The ratio's of the antero-posterior to the transverse diameter of the upper epiphysis of the tibia (see DUBOIS, 1926, p. 1238) for the three species in the above given order are 0.69, 0.60 and 0.63 respectively. The difference between the figures found for the ratio's of palaeojavanica and gigantea is more than twice as great as that between the figures in the former and javanica, i.e., exactly the same result as I arrived at with the Preanger femur. DUBOIS' type specimens are from Kedoeng Broeboes, a locality with both lower Pleistocene (Djetis-) fauna and middle Pleistocene (Trinil-) fauna (VON KOENIGSWALD, 1934, p. 188). VON KOENIGSWALD (l.c., p. 190) records a phalanx of Manis palaeojavanica Dubois from the Djetis deposits of Goenoeng Boetak, which is very near the type locality. He states that, the species not being found in any of the rich fossillocalities with Trinil or Ngandong (upper Pleistocene) fauna, it is not to be doubted at that DUBOIS' specimens originated from the Djetis deposits. If Manis palaeo­ javanica Dubois indeed is typical of the Djetis fauna, the femur in the STEHN and UMBGROVE collection supports VON KOENIGSWALD's view that the Tji Taroem valley fauna belongs to the lower Pleistocene.

LlTERATURE.

ANONYMUS, 1892. Palaeontolcgische Onderzoekingen op Java. Verslag Mijnwezen 2e kwartaal 1892, pp. 14-17. DUBOIS, E., 1907. Eenige van Nederlandsehen kant verkregen uitkomsten met betrekking tot de kennis der Kendeng-Fauna (Fauna van Trinil) .ITijdschr. Kon. Ned. Aardr. Gen., ser. 2, vol. 24, pp. 449-458. 1908. Das Geologische Alter der Kendeng- oder Trinil-Fauna. Ibid., vol. 25, pp. 1235-1270. pl. 39. ---. 1926. Manis Palaejavanica. the of the Kendeng Fauna. Proc. Kon. Akad. v. Wetens eh .• Amsterdam. vol. 29. pp. 1233-1243. 1 plate. 2 figs. Es. L. J. C. VAN, 1931. The Age of Pithecanthropus. The Hague, 142 pp., 11 maps and sections. HELBING, H., 1938. Nachweis rnanisartiger Säugetiere irn stratifizierten europäischen Oligocaen. Eclcgae Geol. Helv., vol. 31. pp. 296-303, 4 figs. HOOIJER, D. A.. 1946. Prehistorie and Fossil Rhinoceroses frorn the Malay Archipelago and India. Zool. Med. Museum Leiden. vol. 26. pp. 1-138. pIs. I-X. 1 fig., 8 tables. KOENIOS:WALD, G. H. R. VON, 1933. Beitrag zur Kenntnis der fossilen Wir'beltiere Javas I. Wet. Med. Dienst Mijnb. Ned. Ind .. no. 23, 127 pp. 28 pIs .. 9 figs. 1934. Zur Stratigraphie des javanischen Pleistocän. De Ing. in Ned. Ind., vol. 11, se ct. IV, pp. 185-201. pIs. I-IV. ---, 1935. Bernerkungen zur fossilen Saugetierfauna Javas. Il. Ibid., vol. 2, no. 10, sect. IV, pp. 85-88, 3 + 10 figs. 418

KORMOS. TH .. 1934. Manis hungarica n. sp .. das erste Schuppentier aus dem europäischen Oberpliozän. Folia Zool. et Hydrob .• vol. 6. pp. 87-94. 14 figs. LVDEKKER. R.• 1886. The Fauna of the Karnul caves. Mem. Geol. Surv. Ind., ser. 10. vol. 4. part 2. pp. 23-58. pls. VII-XI. 1 fig. STf1l:N. CH. E. and J. H. F. UMBClROVE. 1929. Bijdrage tot de geologie der vlakte van Bandoeng. Tijdschr. Kon. Ned. Aardr. Gen., ser. 2, vol. 46. pp. 301-314, 2 maps. 5 pIs .. 3 figs. WEIDENREICH, F., 1945. Giant Early Man from Java and South China. Anthr. Papers Am. Mus. Nat. Hist.. vol. 40, part 1. pp. 1-134. pIs. 1-12. 28 figs .• 19 tables.

Rijksmuseum van Natuurlijke Historie. Leiden. D. A. HOOIJER: A femur of Manis palaeojavanica Dubois from Western Java.

Manis palaeojauanica Dubais, distal portion of femur dext. , Tji Taroem valley W. of Batoedjadjar. Preanger, Western Java. STEHN and UMBüROVE coll .. Geological Museum Leiden no. 28069. Anterior, posterior, lateral and distal views, 4/ 5 natural size.