THREE-DIMENSIONAL MUSCLE PRESERVATION IN JURASSIC FISHES OF CHILE

Museum 01 Natural History, and Department 01 Systematics and Ecology, HANS-PETER SCHULTZE The University 01 Kansas, Lawrence, Kansas 66045-2454, U.S.A.

ABSTRACT

Late Jurassic fishes of Northern Chile are preserved in calcareous concretions within black shales of Oxfordian age. Co-occurring invertebrates (decapod crustaceans, ostrean bivalves, and algae) indicate benthic life. Soft tissues of the fishes were impregnated by calcium phosphate during life, whereas the decay of the remaining soft tissue induced the formation of the calcareous concretions around the fishes. Occurrence of vitamin 03 or a hydroxylated form of Ihis vitamin is postulated to have occurred in Ihe Jurassic phytoplankton; vitamin 03 induced a break-down of the regulation of the calcium metabolism in the fishes as is the case in calcinosis 01 cattle. High concentration of phosphate in phytoplankton 01 the upwelling zone on the western margin of the shelf of South America could explain the additional supply of phosphate in the fossil fishes which is missing from surrounding concretions and sediment.

Key words: Preservation, 50ft tissue,lmpregnation in lite (calcinosis), Teleostean fishes, Oxtordian, Northern Chile.

RESUMEN

Los peces del Jurásico Superior del norte de Chile están preservados en lutitas negras. La presencia de invertebra­ dos (crustáceos decápodos, ostras y algas) junto con los peces es indicio de vida bentónica. Los tejidos blandos de los peces se impregnaron con fosfato de calcio en vida de los individuos, mientras que la descomposición de otros tejidos blandos indujo a la formación de concreciones calcáreas alrededor de los peces. Se postula la presencia de vitamina 03 o de un compuesto hidroxilado de ella en el fitoplancton jurásico, lo que produjo el quiebre de la regulación del metabo­ lismo del calcio en los peces; una situación similar a la presentada en la calcinosis en vacunos. El elevado contenido de fosfato, en el fitoplancton de la masa de agua vertical del borde oeste de la plataforma continental de América del Sur, explicaría el suplemento adicional de fosfato en el pez fósil, el que está faltando en las concreciones yen el sedimento.

Palabras claves: Preservación, Tejidos blandos, Impregnación en vida (calcinosis), Peces teleósteos, Oxfordiano, Norte de Chile.

INTRODUCTlON

Jurassic fishes have been known from northern and 69°12' to 69°14 W; Fig. 1) of the Quebrada del Chile since the 1950's (Biese, 1957, 1961). In a Profeta in the south and of the adjacent quebrada southern continuation of Biese's localities (Cerri­ in the north. The following is a short outline of the tos Bayos, at the northern end of Cordillera Domey­ geology, fauna and paleoenvironment of the locali­ ko), Dr. G. Chong discovered, early in the 1970's, ty. a rich new locality in the Sierra de Varas of the Cor­ The teleost fishes of the Cordillera de Domeyko dillera de Domeyko. The first fish, the type speci­ are important in a phylogenetic context for studies men of Bobbichthys opercularis, was found by R. of early teleosts (Arratia, 1984 and in press; Arra­ Arias. The diverse fish fauna was collected north tia and Schultze, 1985). In addition, they show ex­ of Quebrada del Profeta in different tributary val­ ceptional preservation. Usually, only hard parts of leys (approximate area from 24°58' to 24°59' S organisms are preserved as fossils, but the fossil

Revista Geológica de Chile, Vol. 16, No. 2, p.183-215, 21 Figs .. 6 Pis. 1989 184 THREE-DIMENSIONAL MUSCLE PRESERVATION, JURASSIC FISHES

24° 58'

24°59'

o.... __ ===J' 1km ~7 !:t==={1 5 If l ,ja A~B IY:::::] 6 l~o~-=!Jo19 2 69°14' 69° 13' 69° 12' 110 YII

3

FIGS. 1-3. Quebrada del Proleta, Sierra de Varas, Cordillera de Domeyko, northern Chile. Fig, 1. Geographic position 01 Quebrada del Proleta. Fig. 2. Outcrops 01 lish bearing layers. A·S : Section to demonstrate the reappearance 01 the same horizon by intersection between lolds and topography; C: Section (Fig. 3) south 01 Quebrada de Breas with repetition 01 horizons with concretions (1-10) ; 1. Concretions with lossil fishes; 2. Concretions with re:ltile remains . Fig. 3. Section C with vertical distribution 01 concretions: 4. Siltstone; 5. Shales; 6. Finei)rained sandstone; 7. Limestone; 8. Calcareous sandstone; 9. Concretions; 10. Black colored rocks; 11 . Gypsum; Encir­ cled numbers (1 -10): Horizons with concretions; Roman numbers (I-V): Sediment samples taken. H-P. Schultze 185

fishes of Cordillera de Domeyko, with three-dimen­ ta, Sierra de Varas, with Professor G. Arratia, and sionally preserved muscles, look as if they had Dr. A. Quinzio and Mr. L. Santander, bath of the De­ only recently died. Preserved are muscle fibers partmento de Geología, Universidad el Norte, Anto­ with mitochondria and nuclei, blood vessels, swim fagasta. Field work was organized by Dr. G. bladder, intestine and melanophores. These pre­ Chong, Departamento de Geociencias, Universi­ served soft tissues will be described, trying to ex­ dad del Norte, Antofagasta, and supported by the plain how such unusual preservation could occur. Alexander van Humboldt Foundation, Germany, In 1978, the author collected in Quebrada de and the Universidad del Norte, Antofagasta, Chile. San Pedro, Caracoles, and in Quebrada del Profe-

MATERIALS ANO METHOOS

AII the material was found in concretions and X-ray diffraction was performed on sediment collected in the Oxfordian rocks north of Quebrada samples I to V (Fig_ 3), on one concretion (LBUCH 1- del Profeta, Cordillera de Domeyko, northern Chile. 210277-13 = paratype of Domeykos profetaensis) The concretions were split in the field if they were and on muscles (LBUCH 1-210277-13 = paratype of not already opened by weathering. The material is Domeykos profetaensis and LBUCH 4277 = para­ deposited in three collections : type of Varasichthys ariasi) and banes (LBUCH 1- KUVP Division of Vertebrate Paleontology, Mu­ 210277-13 and 4277) at the Institut für Geologíe seum of Natural History, The University of und Dynamík der Uthosphare, G6ttíngen. Kansas, Lawrence, Kansas. For comparison, concretions of the Cretaceous LBUCH Laboratorio de Biología, Universidad de Santana Formation of Brazil were studied at the Chile, Santiago-Sur (most specimens). Field Museum of Natural History, Chicago, lIIinois R Departamento de Geología, Universidad (FMNH). Samples from concretions in the FMNH de Chile, Santiago, Chile. collection and from one sample from the American A few of the specimens were prepared with Museum of Natural History, New York (AMNH) were acetic acid, and scanning electron microscope pic­ processed by X-ray diffraction. tures were taken from these specimens at the Drawings were prepared by the author with Geologisch-Palaontologisches Institut, Universi­ drawing attachment to a WILD stereomicroscope .. tat G6ttingen (SEM-prepared pie ces of specimens M5A, photographs taken with a Nikkon FM 35 mm and negatives were deposited at the department in camera. G6ttingen).

GEOLOGY ANO PALEOENVIRONMENT

Concretions which contain fishes occur in and sometimes around the concretions). These different horizons (Fig. 3) of the Oxfordian (Chong, shales are interealated by fine grained calcareous 1977) sequence north of Quebrada del Profeta, sandstones which predominate in the lowar part of Sierra de Varas. The author counted 10 horizons. the section, and by thin limestone layers. The Within these horizons, the frequency of concre­ section (Fig. 3) has been measured in a tr butary of tions changes from north to south. Places with the Quebrada de Breas, north of Quebrada del Pro­ high numbers of concretions contain a low number feta, where it was not obscured by debris from high­ of fossils. (The black dots in Fig . 2 mark the occur­ er horizons. Here, the concretions (ha rizan 10) are rences of concretions with fish fossils; conere­ alllarge (over 10 em long and 4-8 em thick), numer­ tions occur along the whole outcrop of the ho­ ous and less fossiliferous than further south. In rizan .) The concretions occur in black sandy the southern part, just north of Quebrada del Profe­ shales whieh are rich in gypsum (occurs as a sec­ ta, one can distinguish a main horizon (Fig .3, No. ondary formation in thin layers within the shales 10) with large grey concretions (up to 25 cm long, 186 THREE-DIMENSIONAL MUSCLE PRESERVATION. JURASSIC FISHES

15 cm wide and 8 cm thick), from two horizons with shriveled fish seales. The abdomen, telson and small black concretions (below 10 cm long, 7 cm uropods (PI. 1, Fig. 4) are frequently found be­ wide, 4 cm thick). The horizon (Fig. 3, Nos. 7-9) tween the shriveled segment parts of crusta­ with black concretions aboye the main horizon has ceans. The ammonites (PI. 1, Fig. 5) are three­ many mere fossíl fishes than that below the main dimensionally preserved, occur alone in concre­ horizon. Reptile bones are present, but are only tions or together with other ammonites or fishes. partially enclosed in concretions. The concretio!ls They are preserved together with aptychus and were not in situ, having been collected from the siphon, sometimes the aptychi are found separa­ surface of often steep slopes and therefore of tely. Ostrean bivalves occur singly, but in most unknown relationship to specific horizons in the cases as clusters. Bivalves attached to ammo­ section. nites were not observed; in some cases large The same sequence appears on both sides of ostreans occur with small perisphinct ammonites the deeply cut valleys and in consequent tributary in the same concretion. Besides ostrean bivalves, valleys because the whole sequence is strongly 'Posídonia' (PI. 1, Fig . 3), in different sízes, cover folded (cross section A-B in Fig. 2) [and not mainly horizons which are ene/osed in the concretions. faulted as described by Chong (1977; in Arratia et Very few teuthoids (PI. 1, Fig. 2) have been al., 1975b; in Chong and Forster, 1976)]. The fold­ discovered, they seem to represent Plesiotheutís ing is caused by compression from the east where or a e/osely related form o Língula is known from a Triassic rocks overthrust the Jurassic sequence, single specimen. Common within the concretions and Paleozoic volcanic and magmatic rocks over­ and on their outer surfaces are coprolitic masses lie the Tnassic sequence (Chong, 1977). The inten­ without internal structure (PI. 1, Fig. 6). sity of folding of the Jurassic rocks increases east­ The impartant component of the fauna is fishes ward within a short distance, so that one encoun­ (Arratia, 1987) of whích the teleosts (Arratia et al., ters many repetitions of the same horizon in the 1975a, b, c; Arratia, 1981, 1982, 1984, 1986; Arra­ area where the reptile bones were found. tia and Schultze, 1985) and ene halecostome (A­ The fossiliferous beds are Oxfordian, but the rratia and Schultze, 1987) have been described. whole ammonite fauna must be described in order The fish fauna (compilations see Arratia and to identify the zone within this stage. The age of Chong, 1979; Gasparini, 1979; Arratia, 1985, the horizon was given as middle to late Oxfordian 1987) ine/udes: (Chong, 1977), but early to middle Oxfordian in a. Undescribed pycnodonts (Arratia, 1987, PI. 3, Chong and Forster (1976). Within the sequence, Fig.3) Chong (in Arratia et al., 1975b) cited different am­ b. Semionotid halecostome: Lepidotes sp. (Arra­ monites, those similar to Perísphínctes (PI. 1, Fig. tia, 1987, PI. 4, Fig. 1) 5) occur in middle Oxfordian sediments in northern c. Large undescribed pachycormid Chile (ven Híllebrandt, 1988, wrítten commun.). In d. Pachycormid halecostome (Arratia, 1987, PI. 5, Chong and Forster (1976), Chong cited Euaspído• Fig.1) ceras perarmatum for middle Oxfordian transversa­ e. Halecosteme Atacamíchthys greení Arratia and rium zone, even though it could be corralatad wíth Schultze, 1987 (Fígs. 1-15, 16B, 19B) tha early Oxfordian cordatum zone. f. Teleosts (listed alphabetically) : The fauna of the main concretion-producing ho­ - Antofagastaíchthys mandibularís Arratía, 1986 rizon (Fig. 3, No. 10) consísts of crustaceans, fish­ (Fígs. 7-12, 13A, Pis. 3B, 4; Arratía, 1987, Fíg. es, ammonites, bivalvas, teuthoids and Língula, 3D) listed in order of decreasing frequency. One speci­ - Bobbichthys opercularis (Arratía, Chang and men of a decapad crustacean, Chílenophoberus Chong, 1975) (Arratía et al., 1975a, Fígs. 2, 3, atacamensis, was described as a unique speci­ Láms. 1-2; Arratía, 1986, Figs. 1-5, 60, Pis. 1, men by Chong and F6rster (1976). In contrast, it 2, 3A; 1987, Fig. 3B, PI. 5, Fíg. 3) was found that decapad crustaceans are the most - Chongichthys dentatus Arratia, 1982 (Fígs. 2- common fossils in the concretions. They occur to­ 14, 15A; Arratia, 1987, Fígs. 4B, 50, 9, PI. 1, gether with the fishes as complete specímens (A­ Fig.6) rratia, 1987; PI. 3, Fig. 1), but in most cases, as - Domeykos profetaensís Arratia and Schultze, disintegrated parts which could be mistaken for 1985 ( Figs. 4C, 6C, 12-17, 21C, 22, 23A, Pis. 3, H-P. Schullze 187

4; Arratia, 1987, Figs. 5C, 8, PI. 1, Fig. 5, PI. 6 ties and most of them preserve calcitic hard Fig.1) parts only as impressions. Pinna (1985) pointed - ?Pholidophorus domeykanus Arratia, Chang out that the fauna of Osteno contains a large per­ and Chong, 1975 (Arratia et al., 1975b, Fig. 3, centage of benthic elements (85.5%) so that ben­ Láms. 1, 2; Arratia, 1987, PI. 6, Fig. 2) thic life must have been possible, and reducing - Protoclupea atacamensis Arratia and Schultze, conditions began within the sediment and not 1985 (Figs. 48, 58, 68, 9-11, 21 E, PI. 2) aboveit. - Protoclupea chilensis Arratia, Chang and 81ack bituminous shales are usually inte'preted Chong, 1975 (Arratia et al., 1975c, Figs. 2, 4,6, as deposits of a sea basin with stagnant, oxygen Láms. 1-4; Arratia, 1982, Fig. 158; 1986, Fig. deficient bottom waters (Seilacher and Westphal, 6F; 1987, Figs. 3C, 4A, 58, 7, PI. 2, Fig.2, PI. 5, 1971; Seilacher et al., 1985). Since bottom condi­ Fig. 4; Arratia and Schultze, 1985, Figs. 2-3, tions are hostile to most organisms, benthic forms 4A, 5A, 6A, 7, 8, 21 D, PI. 1) may have been restricted to drifting woods or oc­ - Varasichthys ariasi Arratia, 1981 (Figs. 2-19, PI. curred as epizoans on ammonites. That was also 1-7; Arratia, 1982, Fig. 15C; 1984, Figs. 1-6, 7F, the interpretation of Chong and F6rster (1976) for 1DA, 11 A; 1986, Fig. 6E; 1987, Figs. 3A, 5A, 6, the deposits at Quebrada del Profeta. However, PI. 5, Fig. 2; Carroll, 1987, Figs. 7-16; Arratia we now know that decapod crustaceans are the and Schultze, 1987, Fig. 19C). most common fossils at the loca lit Y, and their preferred mode of locomotion, that of crawling on AII teleosts, except Protoclupea, are unique to the substrate, indicates that they may have lived the localities north of Quebrada del Profeta. Proto­ in the branched algae (PI. 1, Fig. 1: cf. Cladophora, clupea is the only teleost known from further north Chlorophyceae) growing on the sea floor. A further in 8iese's localities in Cerritos 8ayos (Arratia, indication of bottom life are ostreans which are not 1987, PI. 2, Fig. 4). At the north end of Cordillera attached to ammonites; they sometimes occur de Domeyko in Quebrada de San Pedro at Cara­ with ammonites in the same concretion. When they coles (Fig. 1), the large undescribed pachycormid co-occur, the ostreans are larger than the small of Quebrada del Profeta occurs more frequently in perisphincts. Most ammonites are thought to be the Oxfordian than in Quebrada del Profeta; in addi­ bottom dwellers (Lehman, 1981, p.179) morethan tion, pycnodonts, crocodiles and ichthyosaurs free swimming forms. In summary, the presence of have been found in very large concretions at that decapods, other invertebrates and algae, favors locality. Ichthyosaur vertebrae and one large rep­ an interpretation of the paleoenvironment like that tile bone are known from Quebrada del Profeta. for the Posidonienschiefer in southern Germany The fishes in the small black concretions (Fig. 3, by Kaufmann (1981) and Lehmann (1981) with No. 7-9) are not yet studied. AII are teleosts. The bottom life. The oxic/anoxic boundary was within author found no invertebrates in these concre­ the sediment, so that benthic life was poss ble for tions. the Oxfordian fish deposits in the Sierra de Varas. The co-occurrence of crustaceans and fishes is Only a careful recording of all sedimentological not uncommon in the fossil record (Glaessner, and paleontological data throughout the sequence 1965); another recently discovered locality is the will result in a more accurate interpretation of the Jurassic site in Osteno, Italy (Pinna, 1985). These paleoenvironment. fish-crustacean shales are not concretion locali-

PRESERVATION OF SOFT ANATOMY

Fossils usually represent only the hard parts of ing stopping or retarding oxidation and bacterial an organismo Thus all additional information, such activity by freezing or drying (mummies), or embed­ as color pattern, tracing of the soft body and espe­ ding (e.g. amber). In other cases, soft tissue is cially preservation of soft tissues, gets special at­ replaced and not preserved in its original composi­ tention. The original soft tissue can be preserved tion. These replacements can occur post mortem under special conditions; the main requirement be- by mineral infiltration or bacterial activity, and dur- 188 THREE-DIMENSIONAl MUSClE PRESERVATION. JURASSIC FISHES

ing life (calcinosis). formed post mortem, the common explanation, or Preservations of original tissue is found only in in the living organism (calcinosis). very recent deposits, Holocene and Pleistocene. Famous occurrences are the frozen mammoth ca­ POST MORTEM REPLACEMENT davers with preserved muscles (Zimmermann and Tedford, 1976), but only some muscles retain Sea water has, even in upwelling zones, too low identifiable histological structures whereas most a concentration of phosphorus for precipitation of tissues are completely disintegrated or partly phosphate, therefore, the phosphorus must be en­ replaced through decomposition. Further back in riched, possibly through decomposition of organic geologic time, the Eocene deposits (Geiseltal and matter, either from carcasses or from fecal de­ Messel) of Germany present two kinds of preser­ posits (Burnett, 1977). A good example of such a vation, not including preservation of original ma­ relationship was given by Bate (1972) for the terial such as hair in Messel (Richter and Storch, replacement of original calcium carbonate and 1980) or keratin in Messel (Koenigswald et al., chitin by apatite in cstracodes from the Creta­ 1981) and Geiseltal (Voigt, 1937). Wuttke (1983) ceous Santana Formation of Brazil. Only ostra­ demonstrated that the preserved 'soft tissue' in cods feeding on a fish carcass (two concretions of the deposits of Messel is only a ghost of the origin­ thousands) show the exceptional preservation, al tissue. Mineralized bacteria have pseudomor­ whereas other ostracodes Iying outside the fish in phosed the original soft tissue. A similar process the calcareous concretions do not show phos­ of lithification has been demonstrated by Willems phatic preservation. Bate (1972) spoke of "chance and Wuttke (1987) for 'soft tissue' preservation of mineralization under unique conditions". In this Permian tetrapods of the Saar-Nahe-basin in W­ case, and other cases of exceptionally preserved Germany; there the substituted mineral is apatite. arthropods, and especially in the muscles de­ They argued that the ellipsoid shape of the spherul­ scribed in the present study, the phosphatization ite is indicative of bacteria. The vertebrates in both must have taken place befo re or immediately after localities (Messel and Saar-Nahe-basin) are flatten­ death, prior to any considerable decay (Müller, ed, and nearly two-dimensional. The same interpre­ 1979). The mineralization of muscles is not a tation was given for three-dimensional muscle pre­ simple replacement of one mineral by another as in servation in the Cretaceous concretions from the case of the ostracodes (Bate, 1972), but an Araripe, Brazil, by Martill (1988). infiltration of soft tissue by phosphate salts In contrast, a bacterial substitution is not feasi­ without distortion of the microstructure. ble for the exceptionally well preserved soft tissue from the Eocene of Geiseltal. Ce lis with nuclei, DECAY-INDUCED REPLACEMENT melanophores, muscle striation, etc., appear Usually, soft tissue preservation is considered much as they do in extant soft tissue (Voigt, 1934, the result of mineral precipitation associated with 1935, 1936, 1937, 1938a, b, 1949; Brenner, 1939). incipient decay (beginning decomposition of car­ Voigt (1937) postulated a two step process, first a casses). Calcium precipitation under such con­ kind of tanning, subsequently followed by silicifica­ ditions is known from Recent fishes (Berner, 1968; tion. Both processes are, of course, diagenetic Wilcox and Effler, 1981). Calcite precipitation is to processes found in reducing enviroments, and be expected under marine conditions because leading to nearly two-dimensionally preserved soft concentrations of bicarbonate ions exceed those tissue. of phosphate in the sea and the sediment (Gul­ Muscle fibers with striation have been de­ brandson, 1969). In contrast, phosphate precipita­ scribed not only in Pleistocene and Tertiary depos­ tion requires enrichment of the pore water in phos­ its, but also in Cretaceous (Martill, 1988), Jurassic phate. The phosphate may be derived from decom­ (Reis, 1888, 1893, 1895; Rauff, 1895; Ammon, position of marine plankton (Gulbrandson, 1969) or 1896; Schweizer, 1964; Wykoff, 1972; Pinna, other organisms (Burnett, 1977; Lucas and Pré­ 1985), Carboniferous (Traquair, 1884) and Devon­ vot, 1984), at the same time, anoxic conditions are ian (Jaekel, 1892; Dean, 1902, 1909) fishes. necessary for precipitation. This process requires These muscles are always preserved as phos­ a very fine line between decomposition of soft tis­ phate (apatite). The phosphate could have been sue to produce the needed amount of phosphate H-P. Schultze 189

and precip~ation of phosphate to preserve the soft tion, the carcass passes through the oxic/anoxic tissue befo re its complete decay. boundary where phosphate is concentrated by bacterial breakdown of the original tissue. At the INFILTRATION same time, bacteria deposit phosphate to pre­ As a first condition, Reis (1893, 1895) argued serve the fine structure of museles. that the base substances for his 'myophosphorit' Allison (1988) could not determ ine which of the (apatite) must be present shortly after the death of three post-mortem processes (1. Oecay-induced; the animal befo re decomposition starts. Oecompo­ 2. Bacterial fixation; 3. Formation of intermediate sition forms gases which distort the structures of organic/inorganic complexes) is responsible for the soft tissue. Second, Reis argued that the phos­ phosphatization of soft tissue; he argued an ex­ phate derived from the vertebrate remains in the perimental approach to develop a model. intestine of fishes with preserved muscles, and he showed a connection between phosphatized IMPREGNATION OF MUSCLES IN LlFE muscle tissue and carnivorous behavior; only carnivorous fishes or reptiles feeding on other fish The preservation of the microstructure of mus­ show muscle preservation. After discovering mus­ eles, down to the cellular level, may be more readi­ ele preservation in pycnodonts, Reis (1898) ar­ Iy explained by the phosphatic impregnation of gued then that pycnodonts with their crushing den­ muscle tissue in life. Such cases are known (calci­ tition for a bivalve diet, were partially carnivorous nosis) in extant terrestrial mammals and are caus­ or were also feeding on carcasses. ed by feeding on plants which contain vitamin 03 (Morris, 1982; Boland, 1986). V~amin 03 of plants REPLACEMENT BY BACTERIA breaks down the mechanism of phosphate regula­ Early phosphatization by bacterial activity was tion in the intestine so that the phosphate is ab­ thought to be responsible for the preservation of sorbed and deposited in various soft tissues as muscle tissue in fishes from the Cretaceous San­ apatite (Capen et aL, 1966). Vitamin 03 is present­ tana Formation of Brazil (Martill, 1988). Martill (op. Iy known to occur only in angiosperms within cit.) described a sequence from mass mortality, plants. The application of in vivo phosphatization change of bottom water to anoxic conditions by de­ as the mechanism for the preservation of phos­ composition of carcasses, to the covering of car­ phatic microstructure in marine environments pos­ casses by a cyanobacterial ? mato By sedimenta- tulates the occurrence of vitamin 03 in plankton.

JURASSIC FISHES OF NORTHERN CHILE

PRESERVED SOFT TISSUE Schultze, 1985; PI. 1, Fig. B; Arratia, 1987, PI. 2, Fig. 2; PI. 5, Fig. 4), Varasichthys ariasi (Arratia, Oifferent soft tissues are preserved in the 1981, PI. 3, Fig. B) and indet. (Arratia, 1937, PI. 2, Jurassic fishes of Quebrada del Profeta. Starting Figs. 1-3). The swim bladder can be seen n figures from outside to inside, we can observe skin, super­ of Protoclupea chilensis (Arratia et al., 1975c, ficial muscles, myomeres, museles of fins, blood Lám. 1; Arratia and Schultze, 1985, PI. 1, Fig. B), vessels, swim bladder and intestine. Not every and Varasichthys ariasi (Arratia, 1981, PI. 3, Fig. part is preserved in every fish; the most obvious 1). Intestine, blood vessels and melanophores (Ar­ structure preserved in nearly all fishes are the ratia, 1987, PI. 3, Fig. 2) are mentioned in Arratia myomere museles. Myomere muscles (Fig. 5; PI. (1987, p. 261). 2) appear in figures in the systematic descriptions of the following fishes: Atacamichthys greeni (Arra­ SKIN tia and Schultze, 1987, Fig. 1), Bobbichthys oper­ Below the scales lie a more or less structure­ cularis (Arratia, 1986, PI. 1, Fig. B), Domeykos pro­ less mass (PI. 3, Fig. 2) which melanophores in­ fetaensis (Arratia, 1987, PI. 6, Fig. 1), Protoclupea eludes (PI. 3, Fig. 1). It shows the impressions of chilensis (Arratia et al., 1975c, Lám. 4; Arratia and the scales which may represent the inner side of 190 THREE·DIMENSIONAL MUSCLE PRESERVATION, JURASSIC FISHES

FIG. 4. Teleost indet. LBUCH 15- 260972. Muscles 01 the dorsal fin. de.d: depressor dorsalis; d.ra: distal radial; e.d: erector dorsalis; f.r. 1,2: fin ray 1, 2; i.d: incli­ nator dorsalis; p. ra: pro­ 4 ximal radial; pt: pterygeio­ 1cm phore.

/1 ..... / I .- I I I I \ \ -...... \ ...... "

1cm 5

FIG. 5. Protoc/upea chilensis Arratia et al., 1975c. Arrangement 01 myomeres in R-396 (holotype) and KUVP 71205b superimposed on reconstruction by Arratia and Schultze (1985, Fig. 2). a.e: anterior cone; de.d: depressor dcrsalis; d.p.e: dorsal posterior cone; d.ra: distal radial; e.d: erector dorsalis; f.r. 1, 2: finray 1,2; h.s: hori­ zontal septum; i.d: inclinator dorsalis; mm: myomere; ms: myoseptum; sb: swim bladder; v.p.e: ventral pcsteriorcone; p.ra: proximal radial; pt: pterygiophore. the scale pocket. The author has never observed 6). The central anterior cone seems to be simple, this mass to cover the scales, so that only the low­ not double, even in the caudal region. The dorsal er part 01 the dermis with its melanophores seems posterior cone straightens out anteriorly, but is de­ to be preserved. The melanophores are always in a veloped up to the head ( Fig. 6). The myosepta are contracted phase. Below the deeper dermis, a thin not preserved, they appear as rock(lime)-lilled layer 01 subcutaneous muscle appears (PI. 3, Fig. spaces. This space between the myomeres varies 1); its libers run dorso-ventrally. between specimens. This could indicate different stages 01 contraction or different degrees 01 miner­ MUSCLES alization. The distance between myomeres within The best preserved muscles are those 01 the each specimen are the same. Equal distance la­ myomeres (Fig. 5, PI. 2). They can be observed in vors the interpretation 01 different degree 01 miner­ most lishes Irom the locality in Quebrada del alization in dillerent specimens. For each speci­ Proleta. The myomeres are typically I-shaped. men a linite amount 01 calcium-phosphate was a­ The I-shape is displayed best in the caudal vailable; however, this amount varied Irom speci­ region. The horizontal septum runs in the middle 01 men to specimen. the body in the posterior abdominal and caudal re­ Dietz (1913) introduced a ratio to express the gions; it runs anterodorsally in the anterior abdo­ lolding 01 a single myomere. He related the length minal region. Each myomere is lolded with eones 01 a myomere to the direct distance from its dorsal alternatively directed anteriorly and posteriorly to its ventral recurving. Here the shorter distance (Fig. 8). The ventral posterior eones are straighten­ Irom the apex 01 the dorsal posterior cone (P,) to ing out a:lteriorly, and in most cases they are not the apex 01 the ventral posterior cone (P2) is used, visible or preserved anterior to the pelvic lin (Fig. because the recurving ends 01 the myomeres are H-P. Schultze 191

d.p.c

h

8

o 10 mm
L-______~ ______~I

FIGS. 6-8. Protoclupea chilensis Arratia et al. , 1975. Holotype R-396. Fig. 6. Myomere 01 anterior abdominal region . Flg. 7. Myomere 01 anterior caudal region behind dorsal fin and dorsal anterior par! 01 anal lin. Fig. 8. Myomere 01 posterior caudal region. a.e: anterior cone; d.p.e: dorsal posterior cone; h.s: horizontal septum; PI: apex 01 dorsal posterior cone; P2: apex 01 ventral posterior cone; v.p.e: ventral posterior cone. To the lelt is anterior. Angles give deviation 01 direction 01 muscle libers Irom direction 01 horizontal septum . only rarely visible in fossil specimens. Consequent­ folding ratio corresponds, in part, to the body Iy, this folding ratio is lower than that of Dietz shape. In Protoclupea and Domeykos, the depth (1913), i.e. Dietz's ratio (after his figures) of Ga­ of the abdomen is reduced by about 20% from the dus morrhua is 1.15 for myotomes of the anterior region behind the head to the anterior abdominal abdominal region, 1.2 for myotomes of the poste­ region whereas in Varasichthys and Chongichthys rior abdominal region, and 1.77 for myotomes of body depth narrows by 33% between the two re­ the tail region. Comparable ratios, using the short­ gions. An even larger difference in folding ratio be­ er distance from the fossil fishes, are 1.0, 1.09 tween these two genera in the tail region is ex­ and 1.4, respectively. pected because Varasichthys has a very narrow The folding ratio in Profoclupea chilensis (Figs. peduncle. Unfortunately, no specimen of Varasich­ 6-8) increases from the apdominal region (anterior thys has a complete myomere preserved in the tail abdominal region 1.1 to posterior abdominal region region. /anterior caudal region 1.38) to the tail region The folding ratio gives only a general indication (3.29) much more than in Gadus. In comparing a of similarity or dissimilarity of myomeres between myomere of the posterior abdominal region or an­ species. The shape of the myomeres can be quite terior caudal region of four genera from Quebrada different with the same folding ratio. In Protoclu­ del Profeta (Figs. 9-12), differences in shape and pea (Figs. 7, 9) and Chongichthys (Fig. 12), myo­ degree of folding are obvious between them . Prota­ meres of the posterior abdominal region are nearly clupea and Domeykas have the lowest folding ratio symmetrically shaped dorsal and ventral to the (1 .28 and 1.23, respectively), whereas the myo­ horizontal septum, but myomeres of Chongichthys meres in this region of Chongichthys (1 .47) and are more strongly folded in the posterior abdominal Varasichthys (1.59) are folded as much as those region, comparable to the degree for folding in the in the tail region of Gadus (1.4). The difference in anterior caudal region of Protoclupea. Domeykos 192 THREE·DIMENSIONAL MUSCLE PRESERVATION. JURASSIC FISHES

h.s

o 10mm I I

L..l------'-----', I I I I

FIGS. 9-12. Myomere from !he anterior caudal region (behind dorsal fin and in front or aboye anterior par! of anal fin). Flg. 9. Protoclupea atacamensis Arratia and Schultze, 1985, KUVP 8124. Fig. 10. Domeykos profetaensis Arratia and Schultze. 1985, LBUCH 12-260912a. Fig. 11. Varasichthys ariasi Arratia, 1981, LBUCH 15- 2609729. Fig. 12. Chongichthys denta tus Arratia. 1982, KUVP 65038. h.s: horizontal septum; P,: apex of dorsal posterior cone; P2: apex of ventral posterior cone. To the left is anterior. Angles give deviation 01 direction 01 muscle libers lrom direction 01 horizontal septum .

(Fig. 10) and Varasíchthys (Fig. 11) have myo­ The direction of the muscle fibers changes a­ meres arranged asymmetrically to the horizontal long the body and within one myomere. The fibers septum. In Domeykos, the dorsal portion is more run parallel to the body axis in the abdominal re­ tolded than the ventral portion in the posterior ab­ gion, and at different angles towards the body in dominal region, the same is the case in the anterior the caudal region. In Protoc/upea, changes are abdomen. In Varasíchthys, the ventral posterior mini mal in the direction of the muscle fibers in the cone is directed caudodorsally and not just caudal­ abdominal region (Figs. 6, 7) within one myomere. Iy. It shows that the tour genera differ in the In the anterior caudal region, the muscle fibers are shape of their myomeres, which does not closely almost parallel near the horizontal septum (angle correlate with differences in body shape. Protoc/u­ between 2-5°), whereas the angle (up to 15-20°) pea shows a high increase of myomeres folding in increases towards the dorsal and ventral posterior the tail region despite an evenly decreasing body eones. The muscle fibers are directed posteriorly depth (Fig . 5), but it indicates a powerful propul­ towards the horizontal septum between horizontal sion by the tai!. Extensive folding of tail myomeres septum and posterior eones. They are directed an­ is indicated by the higher folding ratio in the pos­ teriorly towards the horizontal septum dorsal and terior abdominal region of Varasichthys and Chong­ ventral of the posterior eones at a much steeper ichthys. and what is preserved of the tail region in angle (up to 40°). The direction of muscle fibers de­ the holotype, indicates the same for Domeykos. viates even more from the body axis with stronger Thus al! four genera were active swimmers with folding of the myomeres in the tail region (Fig. 8). powerful tail propulsion. Muscle fibers in myomeres of the posterior ab- H-P. Schultze 193

dominal region of the four genera (Figs. 9-12) run nate with eaeh other. Another, not too eommon, more or les s parallel to the horizontal septum, ex­ strueture in the Jurassic fishes from northern Chile cept in the differently shaped ventral posterior is the nucleus (PI. 4, Fig. 1). The volume of the cone of Varasichthys (angle of 45°). In all four gen­ nucleus is filled with Ca-phosphate; the figured era, the direction of muscle fibers in caudal myo­ nucleus has a length of about 20 ¡..tm . meres deviates strongly from the body axis. Change in direction of muscle fibers is functionally BLOOO VESSELS necessary to permit a similar contraction of fibers Blood vessels can be observed together with in each myomere (Alexander, 1969). muscles in cross-sections (PI. 3, Fig . 5), longitu­ Fortunately, in a few cases, the whole set of dinal sections (PI. 3, Fig. 4) or below the vertebral muscles are preserved at the base of the dorsal column (dorsal aorta). Swim bladder and intestine (Fig . 4) and anal fin. From the outside to the inside, are identifiable in the abdominal eavity, but the au­ inclinator, depressor and erector are displayed. thor hesitates to interpret other partially preserved The erector is often quite obvious with its structure structu res. of cones packed into each other. These muscles attach to the proximal end of the fin rays, but the SWIM BLAOOER actual attachment point can rarely be observed. The swim bladder is preserved in three speci­ Thus far, only body (axial) muscles have been mens of Protoclupea chilensis (holotype R-396; PI. observed, muscles could not be found in the head 5, Fig . 1) and of Varasichthys ariasi (15-260972 region of any fish or around the intestine. The mi­ and LBUCH CH-3378). It ends aboye the base of crostructure of the axial muscles is preserved the pelvic fin (R-396 and LBUCH CH-3378) or aboye down to minute details. The muscle fibers are phos­ the posterior half of the fin rays of the pelvic fin. phatized but not the myosepta (PI. 3, Fig. 3). Phos­ The shape of one larger chamber is visible, but it is phate granules completely fill the muscle fibers or, not clear how far it reaches anteriorly. Impressions in other cases, copy the striation (PI. 3, Fig. 4). of blood vessels run dorso-ventrally on the lateral The granule size is variable ; within the striations, wall of the swim bladder of specimen R-396B. A two small granules (PI. 3, Fig. 6) occupy the width striated surface appears in the antero-dorsal part of the bands, whereas those which completely fill below the wall of the swim bladder in R-396B and vessels and fibers, can have twice the diameter. LBUCH CH-3378 . This may represent co ll agenous The phosphatized bands (between 0.6 and 1.15 fibers or their impressions of the tunica externa or 11m wide) may represent the overlap area between the smooth muscles of the tuniea interna. thick (myosin) and thin (actin, troponin and tropo­ myosin) filaments where cross-bridges of the thick GILLS filament connect with the thin filament. These In two specimens (LBUCH 12A-260972, Chong­ cross-bridges are Ca++ (thin filament) - Adenosine ichthys, PI. 6, Fig. 1, and in KUVP 97765, teleost phosphate (ADP, ATP) connections, here the pos­ indet. in a small black concretion), the gills are pre­ sibility for phosphate additions exists. Another ex­ served. They each surround a calcitic rod which planation is also possible. The phosphatized seems to be a diagenetic formation. They are bands may represent only the thin filaments, with closely packed on the gill arches, with the excep­ Ca++ ions as attachment points for the phosphate tion of some very long filaments under the opercu­ surplus. The double appearence of the bands lum of specimen LBUCH 12A-260972, that are could then be explained as mineralization on both spaced far from each other. sides of the Z-line. In cross-section (PI. 3, Fig. 5; PI. 4, Fig. 2), the INTESTINE muscle fibers appear blocky with cornered outline. The alimentary traet is preserved in a few spe­ The cross-section of the fibers usually appears cimens, mostly in the smaller black concretions structureless, but in a few cases a banding struc­ (Fig. 3, Nos. 7-9). One specimen (KUVP 97764, ture appears (PI. 4, Fig. 2). This may represent the Figs. 13-17; PI. 5. Fig. 2) shows nearly the com­ internal structure, or alternatively, large mitochon­ plete alimentary tract whereas only part of the dria. As in mitochondria, finger-like folds extend intestine is preserved in others (Bobbichthys from the walls towards the center where they alter- opercularis: LBUCH 2-260972, KUVP 83702; teleost 194 THREE-DIMENSIONAL MUSCLE PRESERVATION. JURASSIC FISHES

FIGS. 13-17. Alimentary tract in specimen KUVP 97764, teleost inde!. Fig. 13. Abdbminal region with alimentary trae!. Fig. 14. Anterior par! of intestine. Fig. 15. Enlargement of posterior portian of Fig. 14. Flg. 16. Posterior part of intestine. Flg. 17. Enlargement of posterior portian of Fig. 16. end: endodermal layer on folds ; es: esopha­ gus; mu: mucosa; pro: propria (mucosal center of villi); re: rectum; st: stomach.

indet.: LBUCH F5-61 , CH-35900, 6-16, 5-1212). In wards the end of the intestine, where a crystallized specimen KUVP 97764, the esophagus is filled w~h space of a gas bubble reaches into the intestine an amorphous mass. The region of the stomach is (Fig. 16). The folds do not extend to the wall of the indicated by a depression in the fossil, but definite intestine in the posterior part of the intestine boundaries cannot be determined. In contrast, the because the mucosa is also phosphatized (Fig. intestine is very well defined, it is an elongate or­ 17: mu); the mucosa reaches into the folds and gan without coiling. Internal structures (folds) and forms their center (the propria; Fig. 17: pro). The the muccsa are phosphatized whereas longitudinal layer around the center of the folds is thick relative and ring muscles are not preserved. The anterior to the whole diameter of the fold. This layer (Fig. part of the intestine (Fig. 14; PI. 6, Fig. 2) has 17: end) may represent the endodermal epithel longitudinal mucous-membrane folds which pos­ only as seen in cross-sections of the teleost teriorly beco me complicated zig-zag folds (Fig. Rutilus rutilus by AI-Hussaini (1949, Figs. 5b-g). 15). A meshwork connects the zig-zag folds. A The rectum is not preserved, ~ must have occupi­ portion with only short folds follows the zig-zag ed the short distance between the last preserved folds. These give way to circular folds that occupY part of the intestine and the body margin, just in the posterior part of the intestine (Fig. 16). Anterior front of the anal fin. to the longitudinal folds, a few circular folds also A straight intestine is primitive within actinopte­ appear (Fig. 14). Within the posterior part of the rygians (Nelson, 1972) and may be primitive within intestine, the circular folds diminish in length to- teleorcts as well (Harder, 1980). The lalter is sup- H-P. Schultze 195

ported by the specimen described aboye. It is not and food preference has not yet been found (Har­ possible to determine from which side of the sto­ der, 1964). An intestine shorter than standard mach on the intestine originates. It may pass the length is taken as an indication of carnivorous be­ stomach on the right side as in primitive actinopte­ havior. rygians and most teleosts, or on the left, as in os­ teoglossomorphs. The combination of longitudinal TVPES OF PRESERVATION with circular folds is found in some clupeomorph, whereas zig-zag folds are characteristic of Esox Soft structures have been described in such de­ and the cyprinids (Eggeling, 1908; Jacobshagen, tail to demonstrate that a diagenetic or even bac­ 1937). A satisfactory correlation between folding terial transformation is very unlikely. In both

---.---.... - ... --.... ------1ft-Ca ------18 ___~------~~~Q------~ ----Ca ------

19 - _. Ba - .1t-+-----

------~-Ba~------~-~r---~-

- 0------O 0- - 20 - - _ . .. - Q o-o A O -- -_.- ... ------.- - ._- O A A ----- .. t--- t--- O O A A III A Al j J .l.AA n '!lJB..A.t,¡j I A w AAA~ \, ~ "W'I"" ~ qr tt !JL ...- ~\Ij .""""'" -,.¡ ~~\IW' ...... "...., .... ~

.... r¡m...... _. ,_ ,__ ...... JL _. , ._ ~ ...cJ"," = m::,--,-,- .. " '-'-L.. L.. I I I I l' I I I I I I I I I I I I I I I I I I I I I I I I I I I 1111111111

FIGS. 18-21. X-ray diffractometer records. Fig. 8-20: Domeykos profetaensis Arratia and Schultze, 1985; LBUCH ,- 210277-13. Oxfordian, Upper Jurassie; Quebrada del Profeta, Sierra de Varas, Cordillera de Domeyko, northern Chile. Fig. 18. Sample from eoneretion surrounding the specimen. Flg. 19. Vertebrae (Quartz added). Flg. 20. Muscles (Quartz added). Fig. 21. Notefops brama (Agassiz, 1841); AMNH 11753) = Martill, 1988, PI. 1, Fig. 2). Santana Formation (Upper Aptian?); Chapada do Araripe, northeast Brazil. Dorsal abdominal muscles (Quartz added). A: fluorapatite; Ba: barite; Ca: calcite; M: mica; Q: quartz. 196 THREE-DIMENSIONAL MUSCLE PRESERVATION, JURASSIC FISHES

cases, one would expect indication of decay. But, (head) and posterior (tail) portions missing. Some­ in contrast, soft parts with most abundant blood times they contain the whole fish except the snout suppty are best preserved and even such delicate and the posterior, distal parts of the caudal fin. structures as gills and villi in the intestine are com­ The body is usually intact with soft and hard parts pletely preserved. These structures would decay in their anatomical position - disregarding those immediately, especially the intestine with its bac­ specimens which have been mechanically disturb­ terial flora, before transforming bacteria could ed - even though the space in between the phos­ reach the intestine. The fibers in the myomeres are phatized soft tissues is filled with lime or calcite three-dimensionally preserved throughout the crystals (PI. 6, Fig. 3). Never a case was observed depth of the myomeres. This is in contrast to the where the parts show post mortem displacement preservation of muscles in the Ceará concretions within the body. This is in contrast to that ob­ from Brazil, where muscle fibers are visible on the served by Martill (1988, Fig. 5) in the fishes from outside, whereas the mass becomes structureless Ceará, Brazil, where the vertebral column falls to with dep~h (inside). the 'bottom' of the body cavity (PI. 6, Fig. 4). The museles cover the skeletal structures of MINERALOGY the body. They have to be removed ('dissected') to The concretions occur in a calcareous sandy to see vertebrae, caudal skeleton, etc. These cut win­ shaley sequence (Fig. 3). The lower part of the se­ dows have been figured in specimens of Proto­ quence is richer in calcite. Based on X-ray diffrac­ cfupea chifensis (Arratia and Schultze, 1985, PI. 1, tion, samples V, IV and 111 (Fig. 3) can be describ­ Fig. B = Arratia, 1987, PI. 5, Fig. 4; Arratia, 1987, ed as calcareous sandstones to siltstones. The up­ PI. 3, Fig . 2), Atacamichthys greeni (Arratia, 1987, per part of the section contains more shales (sam­ PI. 4, Fig. 3; Arratia and Schultze, 1987, Fig. 1) pies 11 and 1: plagioclase, feldspar, quartz and light and Domeykos profetaensis (Arratia, 1987, PI. 6, colored micas) where the calcite is concentrated Fig.1). within the concretions. Gypsum occurs as a secon­ In cases where the fish was destroyed mechani­ dary formation resulting from the weathering of sul­ cally after deposition en the sea floor, bones and fides throughout the section; it often occurs in muscles behaved in the same way. Fragments cf veins which run with the bedding plan or across it. muscles are spread around like bones (PI. 4, Fig. Sample I contains jarosite, an iron sulfate, which 3). The fragments of muscles have sharp edges, occurs in the oxidation zone of mineral depesits. It and they do not show any sign of decay at the is also a secondary formation. margins. That indicates that the fish was exposed The concretions are calcareous (Iimestene) with to currents or wave actions before the specimen quartz and some mica (Fig. 18). They do not con­ was enelosed in the concretion, but after the mus­ tain apatite. Fluorapatite is restricted to bones eles were phosphatized. However, to induce forma­ (Fig. 19) and muscles (Fig. 20), other soft tissues tion of the calcareous concretions, some of the de­ and coprolites. Barite occurs in the vertebrae (Fig. composing products must have been present. 19) of Domeykos profetaensis (LBUCH 1-210277- Therefore, it seems unlikely that the separation of 13) and muscles of Varasichthys ariasi (LBUCH parts occurred by reworking of sediments after the 4277), whereas it has not been discovered in 10 fish was first covered by sediment followed by analyzed sediment samples nor in the concre­ phosphatization of museles. tions. The deposition of barite in bones and mus­ eles is unusual, it may indicate that barite distribu­ COMPARISONS tion in the Late Jurassic was not uniform and that, like today, its highest concentration was in the Martill (1988) described musele preservation in southern East Pacifico Cretaceous concretions from Ceará, Brazil. He supposed that these museles are preserved in PRESERVATION phosphate (p. 6: "cryptocrystalline franco lite"). The fishes in the Upper Jurassic concretions and developed a model explaining the phosphatiza­ from Quebrada del Profeta are preserved three­ tion by bacterial activity. Phosphatization has dimensionally although somewhat flattened. Most been described for ostracodes (Bate, 1972) and concretions enclose the body, with the anterior copepods (Cressey and Patterson, 1973) in the H-P. SchultZ9 197

Brazilian concretions, but these are singular 1895, 1898). Willems and Wuttke (1987) explained cases (Bate, 1972: two in thousands) where the the phosphatic preservation of a ghost of the soft phosphatized organism lies close to a phosphate tissue in Permian amphibians as resulting from bac­ source, e.g" the content of an intestine or a copro­ terial activities. The Permian amphibians do not lite. In these cases, chitin and original calcitic show three-dimensionally preserved soft tissues shells are encapsulated or replaced by phosphate. in contrast to the fishes in the Late Jurassic of In contrast, the muscles in the Brazilian concre­ southern Germany (Reis 1893, 1895, 1898) and tions are preseNed in cal cite as shown by analy­ those from northern Chile. sis of thin sections and by X-Ray diffraction of Reis (1983) explained the phosphatization of muscle tissue of Microdon penalvai (FMNH PF muscle tissue and skin in the Late Jurassic fishes 12663) and Notelops brama (Fig. 21, AMNH 11753 = and reptiles of southern Germany by diffusion of Martill, 1988, PI. 1, Fig. 2). The muscles appear Ca-phosphate from the stomach and intestine to white in contrast to the yellowish-brown of the skel­ other soft tissues at the beginning of decomposi­ etal parts and scales; muscle fibers are distinct on tion. He argued that carnivorous forms collect the outside but the white mass becomes structure­ enough bone material in their intestine to account less internally. The white mass also appears be­ for the amount of Ca-phosphate deposited within tween the scales. The calcification of superficial muscles. The postulated mechanism is diffusion tissue and the change of structure in the muscle from the intestine to the muscles. This mechanism tissue from the outside to the inside indicate a pro­ does not explain the obseNed selective phosphati­ cess of infiltration from the outside of the fish. It is zation of some but not all structures in the Chilean a presedimentary or very early syngenetic (during or German Jurassic fishes. In addition, one would sedimentation) process, because the decay of expect deposition of dissolved phosphate around muscle tissue is not complete; decomposing pro­ soft structures and not replacement of soft struc­ cesses have progressed only on the inside of the tures themselves. In contrast, Reis (1893) spoke fish so that hard structures are no longer in place of a complete anorganic replica of the microscopic (PI. 6, Fig. 4). Post mortem calcification of muscle structure of muscles. The preserved soft tissue in tissue is known from Recent fishes. Calcium is Late Jurassic fishes from southern Germany is bound by fatty acids under supersaturated cal­ thin, only 0.5 to 2.5 mm thick in fishes of up to 1 m cium conditions in an anaerobic environment (Wil­ total length. That is quite different from the Late cox and Effler, 1981) or in a high pH microenviron­ Jurassic fishes in northern Chile. In addition, the ment even without supersaturated calcium condi­ occurrence of phosphatized muscle tissue in Late tions (Berner, 1968). The occurrence of such calci­ Jurassic fishes from southern Germany should be fied fishes is known from marine environments restudied, the author has not seen muscle tissue (Wells and Erickson, 1933; Faber and Krejci-Graf, in any acid prepared specimen. In contrast to the 1936), as well as from the marine fishes such as findings of Reis (1893) for the fishes from Ger­ the alewife in freshwater lakes (Wilcox and Effler, many, muscle tissue is preserved in nearly every 1981). Preservation of muscle tissue in calcite and fish in the Upper Jurassic of northern Chile and is preservation of essentially only superficial parts of not restricted to carnivorous fishes. Only one Late muscles do not permit a comparison with the mus­ Jurassic fish (undescribed) from northern Chile cle preservation in the Late Jurassic fishes from has been obseNed with a teleost head in its sto­ northern Chile. Other localities may have phos­ macho phatized muscle preservation with microstructure, In conclusion, bacterial replacement or diffu­ as described by Martill (1988) from Ceará, but at­ sion of phosphate from intestine to muscle tissue tention is called about the use of shape and size of are doubtful mechanisms when used to explain sphaerulites as an indication of bacterial forma­ fine preservation of the soft anatomy of fishes. tion. These structures could be explained as crys­ Both mechanisms are processes active at the be­ tal formation under special environmental condi­ ginning of the decomposition process and unlikely tions. to explain the detailed preservation of the Chilean Phosphatization of soft tissue was described material. Another mechanism must have been by Willems and Wuttke (1987) and Reis (1893, active there. 198 THREE-DIMENSIONAL MUSCLE PRESERVATION, JURASSIC FISHES

INTERPRETATION

Phosphatization 01 s01t tissue has always been recently, vitamin D3 has been discovered in plants interpreted as a syngenetic (during sedimentation) (Worker and Carillo, 1967; Dobereiner et al., 1971) or diagenetic process in 10ssils. Nevertheless, and previtamin D very recently in phytoplankton published explanations (Reis, 1893, 1895; M artill , (Holick et al., 1982). Effects of vitamin D3 in phyto­ 1988; AIIison, 1988) 10r 10ssil preserved muscle plankton on fishes have not yet been described. tissue seem not to be applicable to the Late Juras­ Still, that premise in combination with high phos­ sic 1ishes from Chile. A process preceding synge­ phate production by phytoplankton in an upwelling netic phosphatization is required to explain the zone like the western coast of South America, can described circumstances of these extremely well be used to postulate that the same or a process si­ preserved 50ft tissue. milar to calcinosis in cattle, was induced in the Today, phosphatization of living muscle tissue Jurassic fishes. The proposed explanation postula­ is known as calcinosis, an endemic sickness of tes the following scenario: grazing animals in different parts 01 the world (Mor­ 1. An upwelling zone with high production of phy­ ris, 1982). Calcinosis is induced by different plants toplankton which was rich in phosphate, was which produce vitamin D3 or its compounds (Bol­ present in the Late Jurassic on the western mar­ and, 1986). In vertebrates, vitamin D3 is important gin of the shelt 01 South America, í.e., condi­ for regulating calcium metabolism. Vitamin D3 is tions similar to the present day. hydroxylated in the liver to 25-hydroxyvitamin D3 2. The phytoplankton containing vitamin D3 or a (24[OH]2D3), and then by the kidney to 1,25-dihyd­ hydroxylated 10rm of it, was consumed by the roxyvitamin D3 (1,25[OH]2D3). The lalter form is the 1ishes. Normal calcium phosphate metabolism main regulator of calcium metabolism. Its forma­ of the fishes broke down, and additional ingest­ tion corresponds to calcium and phosphorus ed phosphate-rich phytoplankton induced an needs in the body. This regulation is bypassed as even higher rate 01 precipitation 01 phosphate in soon as vitamin D3 or one of the hydroxylated the 50ft tissue of the 1ishes than occurs in cat­ forms are absorbed by the intestine. The conse­ tle. quent surplus of the metabolite leads to precipita­ 3. The fishes that died 01 calcinosis on the conti­ tion of calcium phosphate in bones and soft tis­ nental shelf, sunk anta the floor of the epiconti­ sues. Vitamin D3 plays a role in the regulation of nental sea where black shales were deposited. muscle cells Ca metabolism and P04 fluxes (Bo­ The 1ishes were, in some cases, 1irst destroyed land, 1985), so that it is understandable that a befare they were cave red by sediment devoid of change of the metabolite has effect on the 50ft tis­ phosphate. Chemical processes associated sue. An intake as low a 0.1 percent of the whole with decay of the remaining s01t tissue were suf- diet can produce calcinosis in cattle (Okada et al., 1icient to induce the 10rmation of calcareous 1977). concretions. The importance of vitamin D3 for the calcium This scenario explains co-occurrence 01 1ishes phosphate metabolism in fishes has only recently with phosphatized 50ft tissue in phosphate-1ree been shown (Henry and Norman, 1975). Also only concretions and sediment.

CONCLUSIONS

Fishes with exceptionally well preserved 50ft tis­ imbedded, but partly disarticulated be10re they sue occur in calcareous concretions in Upper Jur­ were enclosed in calcareous concretions. Muscles assic black shales 01 Northern Chile. Calcareous are spread out, as were the bones, which indicates shells of invertebrates occur together with phos­ that the muscles were phosphatized befare embed­ phatic bones and 50ft tissue 01 vertebrates. The ding. invertebrate fauna 01 the black shales indicates The three-dimensional preservation 01 50ft tis­ that bottom life existed and that the boundary be­ sue down to the cell nuclei is unparalleled in the tween oxidation and reduction was within the sedi­ fossil record. Diagenetic or syngenetic re place­ ment. The 1ishes which sunk to the floor of the epi­ ment or impregnation 01 50ft tissue by calcium continental sea were sometimes not immediately phospllate appears very unlikely because those H-P. Schultze 199 models require beginning decompasilion of the ca­ ghost of the soft tissue (Wuttke, 1983). In con­ daver lo initiate the precipitation of calcium phos­ trast, the soft tissue of the Late Jurassic fishes phate. There is no indication of incipient decay in from Northern Chile is preserved three-dimension­ the Jurassic fishes of Northern Chile. To the con­ ally. trary, even structures such as abdominal organs, Therefore, the impregnation of soft tissue by especially the intestine where the decay process calcium phosphate is explained here by a process starts and progresses very rapidly, are preserved in the living fish comparable to calcinosis in cattle down to the cellular level. The most recently pro­ (Boland, 1986). Vitamin D3 or a hydroxylated form pased model by Martill (1988) for the 'phosphatiza­ of it is proposed to have occurred in the Late Juras­ tion' of the cretaceous fishes from Brazil is un­ sic phytoplankton. Intake of that phytoplankton founded because there muscles are preserved in induced a breakdown of the calcium metabolism in calcium carbonate and not phosphate. Transforma­ the fishes, and lead to precipitation of calcium tion of freshly killed fishes into calcareous mum­ phosphate in the soft tissue. The surplus of phos­ mies is known from fishes under marine and fresh­ phate is derived from phosphate-rich phytoplank­ water conditions (Wells and Erickson, 1933; Wil­ ton in the upwelling zone at the west margin of the cox and Effler, 1981). In addition, the lack of phos­ shelf of South America in the Jurassic. The excep­ phate in the concretions and surrounding sedi­ tional preservation and the lack of phosphate in ments excludes diagenetic or syngenetic replace­ concretions and sediment indicate a predepasi­ ment. Phosphatization by bacteria can be exclud­ tional precipitation of calcium phosphate in life for ed too; such a process leads to preservation of a which calcinosis in cattle is a modern analogue.

ACKNOWLEDGMENT

The SEM photographs were taken by Mr. H. Mrs. A. Musser, Museum of Natural History, The Scholz, Geologisch-Palaontologisches Institut University of Kansas, Lawrence, Kansas. Dr. J. und Museum, Universitat Gottingen, W-Germany. Chorn, kindly revised the English. The manuscript The X-ray diffraction analyses were made by Mr. was typed by Mrs. J. Elder (Division of Biology, H. Faber, Institut für Geologie und Dynamik deí University of Kansas). Dr. L. Grande, Department Lithosphare, Universitat Gottingen, and Mr. I.J. of Geology, Field Museum of Natural History, Chi­ Rowell, Department of Geology, The University of cago, lIIinois, arranged support for a visit to the Kansas, Lawrence, Kansas, USA. Dr. H. Arendt, Field Museum of Natural History, and he and Dr. J. Institut für Geologie und Dynamik der Lithosphare, Maisey, Department of Vertebrate Paleontology, Universitat Gottingen, and Dr. A.W. Walton, De­ American Museum of Natural History, New York, partment of Geology, The University of Kansas, loaned material of concretions from the Cretace­ helped with the interpretation of the X-ray diffracto­ ous of Chapada do Araripe, Ceará, NW-Brazil. A meter records. I would like to thank Prof. Dr. O.H. seminar by Dr. R. Boland, Departamento de Biolo­ Walliser, Institut und Museum für Geologie und Pa­ gía, Universidad Nacional del Sur, Bahía Blanca, laontologie, Universitat Gottingen, for permission Argentina, held at the University of Kansas in the to use the facilities in the department. Prof. Dr. spring of 1988 on vitamin D3 metabolism in plants, A.von Hillebrandt, Institut für Geologie und Palaon­ led me to the explanation of the preservational tologie der Technischen Universitat Berlin, kindly phenomenon. In addition, Dr. R. Boland was so verified the determination of the ammonite from kind as to furnish pertinent literature on calcinosis. these localities. Prof. Gloria Arratia, Museum of Na­ I would like to thank al! these people for their tural History, The University of Kansas, Lawrence, suppart of the project. Kansas, kindly identified the fossil fishes and The manuscript was kindly reviewed by Dr. J. permitted study of al! specimens on loan to her. Maisey (Department of Vertebrate Paleontology, Photographs were taken by Dr. J. Chorn, and the American Museum of Natural History, New York). final line drafting of art figures were prepared by 200 THREE-DIMENSIONAL MUSCLE PRESERVATION, JURASSIC FISHES

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Jenaische Zeitschrift fOr Naturwissen­ Pis. New York. schaft, Vol. 43 (Neue Folge, Vol. 36), p. 417-529, Pis. Lucas, J.; Prévót, L. 1984. Synthese de I'apatite par voie 16-18. bactérienne a partir de matiere organique phosphatée Faber, w.; K rejci-Graf, K. 1936. Zur Frage des geologi­ et de divers carbonates de calcium dans des eaux schen Vorkommens organischer Kalkverbindungen. douce et marine naturelle. Chemical Geology, Vol. 42, Mineralogisch-Petrologische Mitteilungen, Vol. 48, p. p.101-118. 305-360. Martill, D.M. 1988. Preservation of fish in the Cretaceous Gasparini, Z. de. 1979. Comentarios críticos sobre los Santana Formation of Brazil. Palaeontology, Vol. 31, vertebrados mesozoicos de Chile. In Congreso Geoló• p. 1-18,4 Pis. gico Chileno, No. 2, Actas, Vol. 3, p. H15-32, 1 Lám. Morris, K.M.L. 1982. Plant induced calcinosis: A review. Glaessner, M.F. 1965. Vorkommen fossiler Dekapoden Veterinary and Human Toxicology, Vol. 24, 34-48. (Crustacea) in Fisch-Schiefern. 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Seilacher, A.; Westphal , F. 1971. Fossil Lagerstatten. In le). Nova Acta Leopoldina, Neue Folge, Vol. 6. p. 3-38, Sedimentology 01 parts 01 Central Europe. Guidebook 7Pls. 8. IntBrnatíonal Sedímentologíeal Congress, No. 8, p. Voigt, E. 1949. Mikroskopische Untersuchungen an 105- 327-335. Heidelberg. silen tierischen Weichteilen und ihre Bedeutung lür Seilacher, A. ; Reil., W.-E. ; Westphal, F. 1985. Sedimen­ Systematik und Palaobiologie. Zeitschrift der Oeut­ tological, ecologieal and temporal patterns 01 fossil schen Geologischen Gesellschaft, Vol. 101 , p. 99- Lagerstatten. Philosophíeal Transactíons of the Royal 104. Society of London, Series 8, Vol. 311, p. 5-23. Wells, R.o.; Erickson, E.T. 1933. The analysis and com­ Traquair, R.H. 1884. Description of a lossil shark (Ctena­ position 01 latly material produced by the decompo­ eanthus costellatus) from the Lower Carboniferous sition 01 herring in sea water. Journal of the American rocks of Eskdale, Dumfrieshire. Geologieal Magazine, Chemieal Society, Vol. 55, p. 338-341. New Series, Oeeade 111, Vol. 1, p. 3-8, 2 Pis. Wilcox, D.A.; Effler, S.w. 1981. Formation of alewile con­ Voigt, E. 1934. Die Fische aus der mitteleozanen Braun­ cretions in polluted Onondaga Lake. Environmental kohle des Geiseltales. Nova Acta Leopoldina, Neue Pol/ution, Ser. 8, Vol. 2, p. 203-215. Folge, Vol. 2, p. 21 -146, 14 Pis. Willems, H.; Wuttke, M. 1987. Lilhogenese lakustriner Voigt, E. 1935. Die Erhaltung von Epithelzellen mit Zell­ Dolomite und mikrobiell induzierte Weichteil-Erhartung kernen, von Chromatophoren und Corium in fossiler bei Tetrapoden des Unter-Rotliegenden (Perm, Saar­ Froschhaut aus der mittlereozanen Braunkohle des Nahe-Becken, SW-Deutschland). Neues Jahrbuch für Geiseltales. Nova Acta Leopoldina, Neue Folge, Vol. Geologie und Palaontologie, Abhandlungen, Vol. 174, 3, p. 339-360. 3 Pis. p. 213-238. Voigt, E. 1936. Weichteile an Saugetieren aus der Worker, N.A.; Carillo, B.J. 1967. 'Enteque seco', calcili­ eozanen Braunkohle des Geiseltales. Nova Acta Leo­ eation and wasting in grazing animals in the Argen­ poldina, Neue Folge, Vol. 4, p. 301-310, 2 Pis. tine. Nature, Vol. 215, p. 72-74. Voigt E. 1937. Weichteile an Fischen, Amphibien und Wuttke, M. 1983. 'Weichteil-Erhaltung' durch lithifizierte Reptilien aus der eozanen Braunkohle des Geiselta­ Mikroorganismen bei mittel-eozanen Vertebraten aus les. Nova Acta Leopoldina, Neue Folge, Vo1.5, p. 115- den Olschielern der 'Grube Messel' bei Darmstadt. 142, 5 Pis. Senckenbergiana Lethaea, Vol. 64, p. 509-527, 2 Pis. Voigt, E. 1938a. Tierische Weichteile aus der Braunkohle Wykoff, R.w.G. 1972. The biochemistry 01 animal 105- des Geiseltales bei Halle/S. Natur und Volk, Vol. 68, p. sils. Williams and Wilkins Co., 152 p. Baltimore. 11-117. Zimmermann, M.R.; Tedlord, R.H. 1976. Histologic struc­ Voigt, E. 1938b. Weichteile an fossilen Insekten aus der tures preserved lor 27,300 years . Science, Vol. 194, eozanen Braunkohle des Geiseltales bei Halle (Saa- p. 183-184. H-P. Schul1Z9 203

PLATES 1 - 6 204 THREE-DIMENSIONAL MUSCLE PRESRVATION, JURASSIC FISHES

PLATE 1

Figures Plants and invertebrates from the Oxfordian (Upper Jurassic) north of Quebrada del Profeta, Sierra de Varas, Cordillera de Domeyko, northem Chile. Scale: 1 cm.

Cladophore algae. x 1.

2 Cuttlefish Plesiotheutis sp., posterior half of the shell. x 1.

3 Bivalve "Posidonia". x 1,

4 Decapod crustacean Chilenophoberus atacamensis Chong and Forster, 1976; abdomen and uropod. x3.

5 Ammonite Perisphinctes sp, x 1.

6 Coprolite on top of small concretion. x 1. Scale: always 1 cm. H-P. Schultze PLATE 1 206 THAEE-DIMENSIONAL MUSCLE PAESAVATION, JUAASSIC FISHES

PLATE 2

Protoclupea chilensis Arratia, Chang and Chong, 1975. KUVP 71205b; postcranial portion with partly removed scale caver lo show muscle myomeres below dorsal fin, and myomeres taken off partly to show the vertebrae; Quebrada del Profeta, Sierra de Varas, Cordillera de Domeyko, northern Chile. Oxfordian, Upper Jurassic. Aboul x 2. Scale: 1 cm. H-P_Schultze PLATE 2 208 THREE-DIMENSIONAL MUSCLE PRESRVATION, JURASSIC FISHES

PLATE 3 Figures Skin and muscles in fishes from Oxfordian (Upper Jurassic) north of Quebrada del Profeta, Sierra de Varas, Cordillera de Domeyko, northern Chile.

Protoclupea chilensis Arratia, Chang and Chong, 1975. KUVP 71205b; posterior ventral abdominal region between pelvic and anal fins showing myomeres overlied by dorso-ventrally running subcutane­ ous muscles, dermis with melanophores and scale impressions. x 9. Scale : 1 mm.

2 Varasichthys ariasiArratia, 1981. LBUCH 4277a (Paratype; Arratia, 1981 , p. 11). SEM 21568, skin. x 176. Scale: 0.1 mm .

3 - 6 Domeykos profetaensis Arratia and Schullze, 1985. LBUCH 1-210277-13 (Paratype, Arratia and Schul­ Ize, 1985, p. 43), etched by acetic acid. 3. SEM 21581 ; muscle libers 01 two myomeres along a space representing !he separating myosep- tumo x 82. Scale: 0.1 mm. 4. SEM 21598; muscle fibers with and without striation. x 60. Scale: 0.1 mm. 5. SEM 21586; cross sections 01 muscle fibers and vessels (ve). x 437. Scale: 0.1 mm. 6. SEM 21588; muscle striation. x 1914. Scale: 0.01 mm. H-P. Schultze PLATE 3 210 THREE-DIMENSIONAL MUSCLE PRESRVATION, JURAS SIC FISHES

PLATE 4

Figures Muscle preservation in fishes from Oxfordian (Upper Jurassie) north of Quebrada del Profeta, Sierra de Varas, Cordillera de Domeyko, northern Chile,

1 - 2 Domeykos profetaensis Arratia and Sehultze, 1985. LBUCH 1-210277-13 (Paratype, Arratia and Sehul­ tze, 1985, p. 43); etched by aeetie aeid.

1. SEM 21584; striated muscle fiberwith nucleus. x 1030. Seale: 0.01 mm. 2. SEM 21596; mitoehondria. x 960. Seale 0.01 mm.

3 Teleost indet. LBUCH 13-260972. Museles myomeres partially intaet, partially distributed over the sur­ face lika tha bonas. x 1. Seale: 1 cm. H-P. Schultze PLATE 4 212 THREE-DIMENSIONAL MUSCLE PRESRVATION, JURASSIC FISHES

PLATE 5

Figures Organs of the abdominal cavity in fishes from Oxfordian (Upper Jurassic) north of Quebrada del Profeta, Sierra de Varas, Cordillera de Domeyko, northern Chile.

Protoclupea chilensis Arratia, Chang and Chong, 1975. Holotype, R-396B, swim bladder (sb). x 3.1. Scale: 1 cm.

2 Teleost indet. KUVP 97764; alimentaty tract (compare Fig. 13); s.int: anterior region of intestine; p.int: posterior region of intestine. x 3. Scale: 1 cm. Arrow points anteriorly. H-P. Schultze PLATE5 214 THREE-DIMENSIONAL MUSCLE PRESRVATION, JURASSIC FISHES

PLATE 6

Rgures

1 - 3 Fishes from Oxfordian (Upper Jurassic) north of Quebrada del Profeta, Sierra de Varas, Cordillera de Domeyko, northern Chile.

1. Chongichthys dentatus Arratia, 1982. LBUCH 12A-260972; opercular region with gills. x 3. Scale: 1 cm.

2. Bobbichthys opercularis Arratia, Chang and Chong, 1975. KUVP 83702 cross section through an­ terior region of intestine. x 1.6. Scale: 1 mm .

3. Teleost indet. KUVP 99300; cross section through compressed caudal region above anal fin . x 4. Scale: 1 cm. Arrow points dorsally.

4 Rhacolepis buccalis Agassiz, 1844. FMNH P12174; axial cross section (= Martill, 1988, Fig. 5) with cal­ cite filled abdomen, cavities with calcite crystals at dorsal margin, vertebrae and muscles at ventral side of body. x 9. Scale: 1 cm . Arrow points anteriorly. Santana Formation (Upper Aptian?); Chapada do Araripe, northeast Brazil. Sediment structures in oblique angles to specimen. H-P. Schultze PLATE 6