645.fm Palaeontologia Electronica palaeo-electronica.org Picking up the pieces: the digital reconstruction of a destroyed holotype from its serial section drawings Julien Benoit and Sandra C. Jasinoski ABSTRACT Serial grinding, a popular but destructive technique in the early twentieth century, allows for the detailed tomographic study of vertebrate fossils. Specimen BP/1/1821 (formerly BPI/1/346) is a procynosuchid cynodont (Therapsida) that underwent serial grinding by A.S. Brink in 1961, resulting in a detailed and insightful study of its skull anatomy. However, BP/1/1821 was also designated by Brink as the holotype, and only specimen, of a new (now a junior synonym of Procynosuchus delaharpeae) species of cynodont: ‘Scalopocynodon gracilis’. This species has subsequently been recognised as a junior synonym of Procynosuchus delaharpeae, but the destruction of a holotype remains an irreversible loss. Brink built an enlarged wax model from the serial sec- tions, but it is degrading rapidly. In this article, we explain how we retrieved Brink’s orig- inal drawings of the sections, and how we were able to build a new digital model of this specimen using a scanner, virtual stack alignment, and 3D imaging. Comparison with previously published drawings demonstrates the accuracy of this digital model. With a 3D printer, we then re-created a more accurate replication of BP/1/1821 with resin. This life-sized replica now helps to complete the collections of the Evolutionary Studies Institute (University of the Witwatersrand, Johannesburg, South Africa) and replaces the long lost original specimen. The possibility to re-use these old data for palaeonto- logical research is also addressed. Julien Benoit. Evolutionary Studies Institute (ESI); School of Geosciences, University of the Witwatersrand, PO Wits, 2050, Johannesburg, South Africa; School of Anatomical Sciences, University of the Witwatersrand, 7 York Road, Parktown, 2193, Johannesburg, South Africa. julien.benoit@wits.ac.za Sandra C. Jasinoski. Evolutionary Studies Institute (ESI); School of Geosciences, University of the Witwatersrand, PO Wits, 2050, Johannesburg, South Africa. sandra_jas@hotmail.com Keywords: Cynodont; Serial Grinding Tomography; Therapsida; Digitization; Curation; Skull Submission: 12 February 2016 Acceptance: 26 July 2016 Benoit, Julien, and Jasinoski, Sandra C. 2016. Picking up the pieces: the digital reconstruction of a destroyed holotype from its serial section drawings. Palaeontologia Electronica 19.3.3T: 1-16 palaeo-electronica.org/content/2016/1478-reconstructing-scalopocynodon Copyright: © September 2016 Society of Vertebrate Paleontology. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. creativecommons.org/licenses/by/4.0/ BENOIT & JASINOSKI: RECONSTRUCTING SCALOPOCYNODON INTRODUCTION For each palaeontological animal species named, the International Code of Zoological Nomenclature (ICZN) stipulates that a holotype must be defined in order to serve as a reference for future works (ICZN, 1999). As a consequence, the designation of a type specimen is, by far, the most critical step when creating a new taxon, and the destruction of a type specimen constitutes an irre- placeable loss to science. Under South Africa’s heritage laws, this also represents a failure to pro- tect part of the National Estate. Unfortunately, this is what happened to the holotype of the South Afri- can procynosuchid cynodont ‘Scalopocynodon gracilis’ (this taxon name will thereafter be enclosed in quotation marks to reflect the fact that it is now considered a junior synonym of Procyno- suchus delaharpeae [Hopson and Kitching, 1972]), which was destroyed during a serial grinding tomo- graphic (SGT) study by Brink (1961), the author of the taxon. Although the SGT study allowed him to exquisitely describe the skull in great detail, this unfortunately led to the definitive loss of this unique piece of South African fossil heritage. Given the extraordinary wealth of fossils coming from the time-expansive Karoo sedimentary succession of South Africa (Rubidge and Sidor, 2001), South Afri- can therapsids were among the first and the most extensively studied species using this technique, even until recent time (e.g., Sollas and Sollas, 1914; Olson, 1937, 1944; Kermack, 1970; Fourie, 1993; Maier and van den Heever, 2002; Sigurdsen, 2006), because some of them are known from doz- ens or even hundreds of specimens. However, before it was synonymized with P. delaharpeae, ‘Scalopocynodon gracilis’ was represented by only one specimen (previously numbered BP/1/346, now BP/1/1821). The technique SGT was invented by Sollas (1904) and was intended to revolutionize the way in which palaeontologists study their fossils (Simp- son, 1933; Sutton, 2008). During SGT, a special device is used to grind the specimen at thin and regular intervals. At every step, a tomogram (i.e., drawing made either from a photographic plate or a projected image) or a photograph of the section is made so that the corpus of tomograms obtained illustrates the internal anatomy of the specimen in sequential slices (Sutton, 2008; see Appendix 1). Using this technique, it became possible to describe in great detail the internal anatomy of fos- sils. At that time, the only alternative was to wait for discoveries of naturally preserved internal struc- tures, such as an endocast of braincase and bony labyrinth (e.g., Case, 1914; Dart, 1925; Cox, 1962; Jerison, 1973; Quiroga, 1984; Court, 1992; Kielan- Jaworowska et al., 2004), isolated bony elements of the chondrocranium (e.g., Benoit et al., 2013a, 2013b), or to undergo a “dissection” of the struc- ture of interest (Dechaseaux, 1974; Gould, 1989). Serial grinding tomography was used in the twentieth century and led to a substantial improve- ment of knowledge about the deep internal mor- phological structure of extinct vertebrates (e.g., Sollas and Sollas, 1914; Stensiö, 1927; Jarvik, 1942, 1954). The technique also allows for the 3D reconstruction of the fossil using wax, cardboard, or polystyrene sheets that are cut into the shape of each fossil slice and stacked sequentially (Sutton, 2008; Cunningham et al., 2014). This kind of tomography is the methodological predecessor of the modern-day computer-assisted tomography (CT scan). Curiously, although alternative methods were invented to preserve the slices of the speci- mens on plates, and though non-destructive X-ray radiography was performed on fossils as early as the end of the 19th century (Branco, 1906), and CT scans as early as the end of the 1970s (Conroy and Vannier, 1984), SGT enjoyed great popularity and was applied in vertebrate palaeontology for a very long time with noticeable success (e.g., Sollas and Sollas, 1914; Stensiö, 1927; Jarvik, 1942, 1954; Olson, 1937, 1944; Fourie, 1993; Maier and van den Heever, 2002; Sigurdsen, 2006; and reviewed in Sutton, 2008; Cunningham et al., 2014; Laaß and Schillinger, 2015). Nevertheless the destructive aspect of this approach has gener- ally been implemented in well- sampled vertebrate taxa, where large numbers of specimens limited the impact of the destruction (e.g., the South Afri- can dicynodonts [Sollas and Sollas, 1914; Fourie, 1993], some Devonian fishes [Stensiö, 1927; Jar- vik, 1942, 1954], the well-sampled artiodactyls Merycoidodon, Poebrotherium, and Leptomeryx [Whitmore, 1953] and squalodontid whales from North America [Luo and Eastman, 1995], and mul- tituberculate mammals from Mongolia [Kielan- Jaworowska et al., 1986]). Researchers and curators remained reluctant to carry out SGT most of the time because the irre- versible damage to fossils is contrary to the princi- ples of heritage conservation. Moreover, the specimens selected for the technique tend to be well preserved since researchers want the best for their study in order to extract maximum information by destroying a minimum of fossils. The technique can be very time-consuming, sometimes taking years to carry out (Jarvik, 1942, 1954; Sutton, 2 PALAEO-ELECTRONICA.ORG 2008; Cunningham et al., 2014). Other methods were invented to inspect the interior of fossils with- out completely destroying them, such as midline sawing in order to prepare the interior of a skull, or tungsten microtomy which enable preservation of at least part of the original material, but they were expensive, time-consuming, and their use was not as widespread (Sutton, 2008; Cunningham et al., 2014). Only non-destructive CT imaging has replaced SGT in vertebrate palaeontology, but not prior to the 1990s. One of the most dramatic losses as a result of an SGT study was the destruction of the holotype specimen of ‘Scalopocynodon gracilis’. Brink intended to use serial grinding tomography (SGT) in order to study BP/1/346, the only known skull of a new genus and species he named in the same article (Brink, 1961). However, it appears that the destruction of a holotype was not intentional: Brink at first believed the specimen represented a com- mon therocephalian (Scaloposaurus) and did not realize that it was a new species of cynodont until the secondary palate became evident in the sec- tions (Brink, 1961, p. 119). At that point of realiza- tion, Brink (1961, p. 119) decided “to reconsider the implications of this misinterpretation ... It was also considered that the specimen might prove to be the type of new species of Procynosuchid, and careful thought was given to the implications of destroying a type. At this stage it was decided to reconstruct in wax the anterior half of the skull, as far as it was then sectioned. The resultant model, although exhibiting some peculiar features, suggested that the specimen very likely represented a juvenile stage of an existing species of Leavachia, Procy- nosuchus, or Galecranium, and it was decided to proceed with the sectioning.” The specimen BP/1/346 is thus to be consid- ered a lost holotype and ‘Scalopocynodon gracilis’ could be considered a nomen dubium. However, because Brink thoroughly described the morphol- ogy of the specimen and reconstructed a wax model (still available at the Evolutionary Studies Institute [ESI, University of the Witwatersrand, Johannesburg, South Africa]), Hopson and Kitch- ing (1972) were able to synonymize ‘Scalopocyno- don gracilis’ with Procynosuchus delaharpeae. The recognition of ‘Scalopocynodon gracilis’ as a junior synonym of Procynosuchus delaharpeae, a well- documented species, minimized the consequences of the loss of BP/1/346. However, as BP/1/346 will remain a holotype in perpetuity, its loss remains unfortunate, especially since a delicate wax model will not last as long as the original fossil. Here we present a new, digital reconstruction of BP/1/346 based on the original hand-drawn tomograms by Brink (1961). This 3D virtual model is compared with the wax model and the descrip- tion of the original specimen published by Brink (1961). This comparison revealed that the wax model is deformed, inaccurate in places, and has already been extensively restored. The purpose here is neither to resurrect the taxon ‘Scalopocyno- don gracilis’, nor it is to redescribe BP/1/346 since complete and comprehensive descriptions of this specimen as well as Procynosuchus delaharpeae have already been published (Brink, 1961; Kemp, 1979). Instead, the goal here is to compare the original description by Brink (1961), the wax model, and the digital model of BP/1/346 in order to dis- cuss the accuracy and reliability of this new model. Finally, we introduce the new 3D printed model of BP/1/346 based on the digital model, which now enriches the Karoo holotype collection of the ESI with a restored representation of the long lost holo- type. MATERIAL AND METHODS Brink (1961) clearly detailed how he recon- structed the wax copy of specimen BP/1/346. Drawings of the lateral, dorsal, and ventral views accompanied the description, as well as a selec- tion of 14 coronal sections and the reconstruction of a sagittal section (Brink, 1961, figures 33-35). Since 1961, the wax model was kept in the Karoo holotype room at the ESI under the catalogue num- ber BP/1/1821. The drawings received the same number, but they were labelled “Therocephalian” (Figure 1) and were not associated with the wax model. The drawings of the serial sections and the ventral and dorsal views of BP/1/346 were recov- ered in a plastic bag catalogued BP/1/1821. Draw- ings of the lateral view and sagittal section were not recovered. The cover of the serial sections stated that the drawings were those of a thero- cephalian skull possibly belonging to Aneu- gomphius (now synonymized with Theriognathus [Sigurdsen et al., 2012], but note that during the early stages of serial grinding Brink believed that it was a Scaloposaurus [Brink, 1961, p. 119]) (Figure 1), which could explain why those drawings were not stored along with the wax model. Despite this, we are confident that the drawings are those made by Brink in 1961 of BP/1/346 because: i) they are identical to Brink’s (1961) published figures and to the wax model (Figure 2), ii) the cover is labelled with the specimen number 346 (Figure 1), and iii) it is signed with Brink’s name and initials (Figure 1). 3 BENOIT & JASINOSKI: RECONSTRUCTING SCALOPOCYNODON Today, BP/1/346 now refers to a dicynodont speci- men, the holotype of Brachyuraniscus merwevillen- sis (transferred to Pristerodon by King and Rubidge, 1993), which was formerly numbered BP/ 1/218 and the skull of which is now ironically lost (King and Rubidge, 1993). Consequently, in order to avoid confusion, and since the original BP/1/346 cynodont specimen no longer exists, all the mate- rial formerly referred to ‘Scalopocynodon gracilis’ is hereby designated the number BP/1/1821. All 116 serial tomograms drawn by Brink (1961) are present. They were drawn with the help of an episcope (which projected an image of the specimen onto a sheet of mounted paper) with a four times magnification (Brink, 1961; Appendix 1). Though the paper is becoming yellowish, the pencil tracings are still clearly visible (Appendix 1). As stated in his article, Brink (1961) found a way to grind the entire specimen (usually the last centime- ters are shredded during the grinding process); hence the 116 slices represent the entire speci- men. The drawings of the serial sections were scanned with a standard office scanner. The pic- tures were then converted into 8 bit grayscale images and their contrast was improved using Fiji 64bit (Schindelin et al., 2012). The slices were aligned and stacked into a single RAW file using the “Align slices in stack” function of the Template Matching plugin under Fiji 64bit. Brink (1961) drew the serial sections with a four times magnification in order to reconstruct the wax specimen four times larger than the original. Here, the images were rescaled to natural size using the measurements given by Brink (1961). The digital reconstruction of BP/1/1821 was obtained using threshold and man- ual segmentation in Avizo 8 software (VSG). The digital model (Figure 3, Appendix 2) is a direct reconstruction of the slices drawings. No further efforts to cosmetically improve the digital model, such as resampling the slices or surface smooth- ing, were performed. Features that were crossed out on the drawings were not considered during segmentation (e.g., section 14, see Appendix 1). The specimen was printed in 3D using Zprinter 450 (Figure 4). This 3D printed model represents the first accurate, life-sized reconstruction of BP/1/ 1821 in 55 years. DESCRIPTION The digitization of the serial section drawings of BP/1/1821 illustrates how much information was lost during the SGT process. The slice edges are prominent (Figure 3), and it is clearly apparent that the slice intervals were too thick. Brink (1961) admitted that slicing the specimen every 0.5 mm was not enough for such a small skull (skull length: 60 mm) and a lot of information was lost. The jag- ged aspect of the digital specimen may also partly reflect the subjectivity of Brink's original hand- drawn tomograms. In effect, the apparent lack of alignment of some of the slices on what should be smooth surfaces (see Appendix 1) implies some degree of inaccuracy in the original drawings. Nonetheless, Brink clearly drew the sutures of most identifiable cranial bones, which can thus be digitally extracted and isolated (Figure 5). Having access to the complete series of sec- tions of the skull is quite useful. One can determine if some of the sutures remained patent or became obliterated, which might be an important indicator of ontogenetic age in basal cynodonts (see Jasi- noski et al., 2015). In addition, access to the com- plete set of sections (Appendix 1) allows for re- interpretation of the identification of bones and sutures and can highlight discrepancies with the published sections (Brink, 1961, figure 35) and/or the reconstructed skull (Brink, 1961, figures 33-34). This comparison showed that a few of the sutures in Brink’s published sections are not illustrated in his original drawings. For example, the parietal- parietal suture anterior and posterior to the pineal foramen is absent from his original drawings (Appendix 1), but it is present on his published sec- tions (Brink, 1961, figure 35, sections 84, 100). Therefore, it remains equivocal whether this mid- FIGURE 1. The label on the first page of the serial sec- tion drawings. It is written “Specimen N°1821 (FN). MN° 346. Complete Aneumogomphius? skull thero- cephalian. Sections ½ mm. Magnification x4.” The doc- ument is signed “A.S. Brink”. Abbreviations: FN, Field Number; MN°, Museum Number. 4 PALAEO-ELECTRONICA.ORG 5 FIGURE 2. Comparison of the different reconstructions of BP/1/1821 in dorsal (left) and ventral (right) views. 1, the wax model; and 2, the original illustrations by Brink, found with the serial section drawings. BENOIT & JASINOSKI: RECONSTRUCTING SCALOPOCYNODON line suture on the skull roof was partially obliter- ated. On the contrary, the nasal-frontal suture is absent (see Figure 5.1) from both the published and original drawings of the sections, but this might be due to the longitudinal orientation of the inter- digitations that is difficult to interpret in coronal sec- tion, or the thicker sections did not intersect the region of the suture, or weathering prevented detection of the suture (see Brink, 1961, p. 123). Brink estimated the position of this suture on his reconstruction of the skull (Brink, 1961, figure 33A). As for bone identifications, Brink’s (1961, fig- ure 35) section 102 through the labelled supraoc- cipital bone (‘so’) might actually be through the parietal or the interparietal. Additionally, the articu- lar (‘art’) on section 106 (Brink, 1961, figure 35) appears to be the quadratojugal (Appendix 1; Fig- ure 5.2). Apart from the size difference, the digital model and the wax model (Figure 6) look quite dif- FIGURE 3. The digital model in (1) right lateral, (2) left lateral and (3) ventral view in stereopairs. Scale bar equals 10 mm. See also the supplementary video in Appendix 2. FIGURE 4. The slices of specimen BP/1/1821 finally put back together again, as seen in the hands of the CT scan facility technician K. Jakata (ESI). This 3D printed model is the first time in 55 years that BP/1/1821 has been accurately reconstructed at life-size. 6 PALAEO-ELECTRONICA.ORG ferent from one another even though the latter model was apparently reconstructed using the same serial section drawings (Appendix 1). The digital reconstruction illustrates how the wax model of BP/1/1821 would have originally looked if it was based solely on Brink’s interpretations of the serial sections; however, it is apparent that some sort of smoothing was applied during or soon after the reconstruction of the wax model. The slices of wax used by Brink (1961) were 1 mm thick instead of the 2 mm that was required to reconstruct the specimen with a four times magnification. To com- pensate, every fifth slice was duplicated (see Brink, 1961, p. 122), which artificially increased the num- ber of sections, giving some room for this smooth- ing. Other inconsistencies, due to cosmetic improvements, are visible between both models. The morphology of the teeth is barely visible on the digital model (Figures 3, 6), whereas they are well reconstructed on the wax specimen (Figure 6). For instance, the rostral-most incisors were destroyed during the grinding of the first slice, and only their roots are visible on the serial sections (Appendix 1). This is not reflected by the wax model (Figure 6). The wax model also bears two canines on each side while only one canine was actually preserved on the left side (Figure 6; Brink, 1961). Brink (1961) also reconstructed two canines on both sides of the skull in his figures 33B and 34. Perhaps this dis- crepancy exists because the caudal-most of the two canines actually has an eroded root and was in the process of being replaced by the anterior one (Appendix 1, sections 20-28; Brink, 1961, p. 125). Post-canine teeth were also sculpted on the wax specimen. They are all conical, whereas the last four should be tricuspid (Brink, 1961). Hence, the close examination of the wax model of BP/1/1821 and the re-appraisal of its anatomy in comparison with the digital model and the original description show that some of the information that was lost during the serial grinding process was recon- structed on the wax model, especially the tooth morphology. However, the main differences between the models are due to deterioration of the wax model (Figures 6, 7). The pterygoid wings should be projecting ven- trally, as depicted in the digital reconstruction (Fig- ure 6) and the original description (Brink, 1961), but on the wax model they are nearly horizontal and oriented posteriorly (Figure 6). The slices of wax are also nearly horizontal on the pterygoid pro- cesses (Figure 7.3), which shows that this feature is due to deformation of the wax after the skull was FIGURE 5. Segmentation of the skull bones of BP/1/1821, (1) in the cranium and (2) in the lower jaw. Scale bar equals 10 mm. 7 BENOIT & JASINOSKI: RECONSTRUCTING SCALOPOCYNODON lying on its ventral side for a long period of time. The canines are also deformed in the same way (Figure 7.1). They are deflected laterally and ros- trally (Figures 6, 7.1), which also indicates that the specimen was stored on a flat surface for a pro- longed period of time. Incidentally, the rostrum is bent dorsally as it was pushed upward by the canines (Figure 6). The occipital plate and the area of the frontals, at the level of the orbits, are also collapsing (Figure 6). In addition, the whole wax model is laterally deformed when compared to the digital model (Figure 6). Finally, some parts of the wax model are bro- ken: the right zygoma (Figure 7.3-4), the right pter- ygoid process (Figure 7.3), part of the vomer (Figure 7.2), and the left mandible (Figure 7.5). Also, the reflected laminae of the angular are more pronounced on the digital model (Figures 3.1-2, 5.2) than on the wax model (Figure 7.7), presum- ably because they were broken on the latter. How- ever, no pieces of these broken parts were found in the specimen’s box. Eight nails and metallic sticks had been added to scaffold the wax, one in each zygoma (Figure 7.3), two inside each orbit (Figure 7.4), and one to hold each of the delicate quadrate process of the pterygoid (Figure 7.3). They were inserted inside the wax perhaps to prevent the model from collapsing in on itself. Six additional nails and sticks are also present in the jaws, two between each mandible (Figure 7.5) and two inside each ascending ramus (Figure 7.6). We are confi- dent that these nails and sticks were inserted inside the wax after the model had been recon- structed because: i) distinct traces of melting that have erased the limit between wax slices proves that the insertion of the nails was not synchronous with the construction of the model (Figure 7.6), and ii) because Brink (1961, p. 121) stated that such supports were absent from the original reconstruc- tion: “Although it was necessary to use supports in the process of reconstruction, the final product is entirely free from such supports. Such supports that were used were lengths or plates cut from the wax. On completion of the model all supports could be removed and the product is so stable that is it virtually like handling a modern mammalian skull.” The presence of this nail scaffolding suggests that the wax was restored at some point; however, it was impossible for us to determine exactly when. DISCUSSION Significance for Research The number of X-ray scanning and other non- destructive computer assisted tomographic studies (CT scan) of fossil vertebrates has greatly increased in the past 20 years, and the study of internal structures (e.g., palaeoneurology, bone FIGURE 6. Comparison between the digital model (top) and the wax model (bottom), in rostral (left) and lateral views (right). Arrows indicate the direction of deformation of the wax model. Scale bar equals 10 mm. 8 PALAEO-ELECTRONICA.ORG 9 FIGURE 7. The state of preservation of the wax model of BP/1/1821. 1, the right side of the snout showing the defor- mation of the canine; 2, the palate showing the broken vomer; 3, the basicranium showing multiple cracks, nails and sticks around the pterygoid; 4, view of the right orbit showing multiple cracks, nails and sticks; 5, dorsal view of the of the mandible showing multiple cracks, nails and sticks; 6, detail of the inner side of the left ascending ramus of the mandible showing a stick embedded in the wax; and 7, lateral view of the mandible showing the incomplete reflected lamina of the angular. The arrows point to the nails, dotted lines demarcate the cracks. Not to scale. BENOIT & JASINOSKI: RECONSTRUCTING SCALOPOCYNODON histology) in vertebrate palaeontology has entered a new Golden Age (see Sutton, 2008; Witmer et al., 2008; and Cunningham et al., 2014 for reviews). In this context, data such as those avail- able from old SGT sections are still valued as they contain important information about the internal structures of fossils that can supplement observa- tions based on CT scans. Figure 5 illustrates how much valuable data can still be extracted from this kind of serial section, which is particularly signifi- cant given the number of published SGT sections that are available in the literature. For example, the following cranial studies of South African therap- sids have published SGT sections and drawings or photographs of the sections that might still be avail- able in collections: dicynodonts (Sollas and Sollas, 1914; Keyser, 1975; Fourie, 1993), dinocephalians (Boonstra, 1968), gorgonopsians (Olson, 1938b); therocephalians (Olson, 1938a; Maier and van den Heever, 2002; Sigurdsen, 2006), and cynodonts (Broom, 1938; Rigney, 1938; Fourie, 1974). As we found discrepancies between some of Brink’s labels in his published sections (Brink, 1961, figure 35) and our interpretation of his original drawings, it is important to have access to the full series of hand-drawn tomograms so that reinterpretation can be implemented. In addition, the physical slices made using a microtome or grinding and peel techniques of some therapsid skulls are also housed in the fossil collections in South Africa, e.g., the Lystrosaurus skulls sectioned by Cluver (1971) at the Iziko South African Museum, the snout of a Diademodon sectioned by Grine (1978) at the ESI, or the skull of a Glanosuchus sectioned by Maier and van den Heever (2002) at the Depart- ment of Zoology of the University of Stellenbosch. Other fossil therapsids housed elsewhere in the world include, for example, the ‘Anomodont A’ sec- tioned by Olson (Olson, 1937, 1944; Angielczyk et al., 2016) and a young Galesaurus sectioned by Rigney (1938) at the Field Museum in Chicago. These fossil sections can be photographed and digitized to produce new 3D models of the once- complete specimen and can be used or re-ana- lyzed for research with no additional cost, time, or space requirements. Interestingly enough, using a similar technique described here, an intriguing ano- modont skull serially sectioned by Olson in 1937 was partially reconstructed in order to identify it and perform a revision of Brachyprosopus broomi (Angielczyk et al., 2016). The data produced from SGT studies should be valued as much as possible since specimens were sacrificed for them. As exemplified by Jasi- noski et al. (2015) and Benoit et al. (2015), data from SGT studies like those of Fourie (1974) are still relevant and often cited. There is also a flour- ishing literature in invertebrate palaeontology illus- trating how SGT data is useful for various fields such as the study of soft tissue structures (e.g., Sutton et al., 2005, 2006; Siveter et al., 2003, 2013), systematic and phylogenetic analyses (e.g., Sutton et al., 2002, 2005; Siveter et al., 2004, 2007a; Briggs et al., 2004, 2012), palaeoenviron- mental reconstruction (e.g., Lukeneder and Weber, 2014) or to infer behaviour in extinct species (e.g., Siveter et al., 2007b). The serial grinding, digital photography, and computerized restoration of the beautifully preserved Silurian fauna of the Here- fordshire deposit have allowed the virtual dissec- tion of several key invertebrate specimens in exquisite detail, which brought a new insight into the early radiation and palaeobiology of arthro- pods, brachiopods, and molluscs (Sutton et al., 2001, 2005, 2006; Briggs et al., 2004, 2012; Siveter et al., 2003, 2007a, b, 2013). Computer assisted tomography is still a rela- tively expensive technique in terms of specialized software and hardware, and the time and technical skill required to process the data (Sutton, 2008; Cunningham et al., 2014). Moreover, there are fea- tures that some CT scanning techniques cannot detect, such as phase contrast (i.e., contrast between the matrix infilling of two different internal structures) that is invisible to conventional X-ray sources and can only be detected by synchrotron beam or serial grinding tomography (Tafforeau et al., 2006; Sutton, 2008; Cunningham et al., 2014). Large and dense specimens often lack the appro- priate resolution and appear opaque on scans, except when using neutron-rays (which can make the specimen radioactive) or using SGT (Maier and van den Heever, 2002; Schwarz et al., 2005; Taf- foreau et al., 2006; Sutton, 2008; Cunningham et al., 2014; Laaß and Schillinger, 2015). Lastly, sometimes the specimens simply have insufficient contrast on CT scans while SGT offers direct access to the internal structure (Maier and van den Heever, 2002; see Cunningham et al., 2014). This is what happened with the invertebrate fossils from the Herefordshire deposit, which are preserved in calcite that is very similar to the micritic host rock (Sutton et al., 2001; Sutton, 2008; Cunningham et al., 2014). The fossils are difficult to prepare by the usual mechanical and chemical means, and X-ray techniques cannot discriminate between the matrix and the fossils (Sutton et al., 2001; Sutton, 2008; Cunningham et al., 2014). In this particular case, 10 PALAEO-ELECTRONICA.ORG SGT accompanied with digital photography of the sections and digital restoration proved to be the most effective way to access the morphology of the fossils (Sutton et al., 2001, 2005, 2006; Briggs et al., 2004, 2012; Siveter et al., 2003, 2007a, 2007b, 2013; Sutton, 2008). Significance for Curation and Archiving Fifty-five years ago, Brink (1961) created a wax model of BP/1/1821 without the need for sup- port structures, but the presence of metal sticks and nails indicates that over time it has collapsed on itself. Further evidence of this process of degra- dation is visible today through the dramatic defor- mation of the wax model in comparison to its digital counterpart that is based on Brink’s tomograms (Figure 6). Wax is very sensitive to temperature variations and the wax model has no internal sedi- ment filling, which makes it very susceptible to damage during physical handling. The digital reconstruction based on Brink’s drawings of the raw serial sections show that the wax model deteri- orated relatively quickly and, apart from the description (Brink, 1961), it is all that testifies to the morphology of the original specimen. What remains of the wax model of BP/1/1821 could be preserved through digitization using surface scan- ning, but the internal anatomy of the specimen would not be recorded. The use of X-ray scanning is excluded because of the low absorbance of the wax compared to the metallic sticks, which would not give enough contrast and would create artifacts on the radiographs. Moreover, the heat produced by the engine could further damage the wax. As such, digitization of Brink’s drawings in order to reconstruct the specimen in silico seems to be the most effective and easiest way to represent his interpretations of the original fossil. This is certainly the most important step for curation of the draw- ings, as they are also deteriorating (the paper is yellowish and the pencil marks are fading away). With the re-discovery of these drawings, it became possible to reconstruct a digital model that revealed how the wax model was affected by the vicissitudes of time (Figure 6). The recreation of a model of BP/1/1821 in sil- ico highlights three new opportunities for future conservation efforts: first, as a substitute to the wax model, a digital model offers fewer problems in terms of preservation since the quality of the digi- tized drawing and reconstruction will not decline with time or manipulations (however questions of data standardization, computer power, ethics, costs, and the implementation of sustainable, long- term digital archiving still remain important issues for digital palaeontological data; see Gilbert and Carlson, 2011; Mallison, 2011; and Cunningham et al., 2014 for a complete discussion). Second, a new model was 3D-printed in resin, and could be printed again at any time using any 3D printer. In contrast to the wax model, this model is the same size as the original (although this can easily be adjusted during printing), and its morphology accu- rately reflects that illustrated in Brink’s drawings of the serial sections (Figure 4). Lastly, a 3D-printed model can serve as a guide for the future resto- ration of the wax model. Nevertheless, one should keep in mind that Brink’s drawing are subjective and thus our new digital reconstruction, like the original wax model, is based on an interpretative medium and will never replace the lost specimen. CONCLUDING REMARKS Here we provide an easy and affordable tech- nique that assisted us to reconstruct BP/1/1821 from serial section drawings of the 55 year old holotype of the formerly recognised species ‘Scalo- pocynodon gracilis’. This study illustrates the value of old original drawings/ photographs of SGT sec- tions and peels which should not be neglected. Conservation of these data through digitization (e.g., scanning, digital photography) should be a priority, as they can provide important new per- spectives to long destroyed specimens. In the case of BP/1/1821, this technique has provided a tool to assess the state of preservation of the old wax model. To archive a digital copy of these drawings is therefore as important as curating original fossil specimens. If photographic plates (e.g., Sollas and Sollas, 1914), physical sections (e.g., Cluver, 1971), peels (e.g., Angielczyk et al., 2016), or drawings (e.g., Brink, 1961) are housed in collec- tions, then these items should be appropriately accessioned and linked to the record of the source fossil. Attempts to digitize them should be under- taken, and original notes from the studied speci- men should also be located and subsequently preserved. In addition, SGT data in vertebrate palaeontology has recently been overlooked because of the advent of non-destructive digital tomography (Cunningham et al., 2014). However, data already acquired could still potentially prove useful for modern comparative works and enable internal anatomical detail of a given fossil species at no additional cost. What happened to specimen BP/1/1821 is not an isolated case. According to Thackeray et al. (1998), the holotype skull of Lystrosaurus murrayi 11 BENOIT & JASINOSKI: RECONSTRUCTING SCALOPOCYNODON was sectioned by Huxley in 1859. At that time, scanning techniques did not exist, but more recently, the holotype of Stygimoloch spinifer was sectioned in order to study the cranial ontogeny in pachycephalosaurids (Horner and Goodwin, 2009). Among invertebrates, many holotypes from the Herefordshire locality underwent serial grinding because anatomical details were only accessible through this method (e.g., Sutton et al., 2002, 2005; Briggs et al., 2004, 2012; Siveter et al., 2007a); however, even in this particular case, opti- cal non-destructive tomographic techniques (e.g., synchrotron and CT scanning) might produce valu- able results (Sutton, 2008; Siveter et al., 2014). Serial grinding and sectioning can still be useful techniques to study fossil anatomy and evolution- ary processes (e.g., Maier and van den Heever, 2002; Sigurdsen, 2006), but we recommend per- forming X-ray, neutron, or surface scanning of the specimens before sectioning in order to retain a digital copy of the original (just like Maier and van den Heever [2002] did for their specimen of Glano- suchus). Destruction of a specimen, particularly a holotype, must remain a last resort, as this threat- ens the very premise of the taxonomic endeavour and, given the rapidly changing technological land- scape, prevents future analysis. ACKNOWLEDGMENTS The authors would like to thank Kudawashe Jakata for assisting with printing of the specimen. We also thank B. Zipfel, curator of the palaeonto- logical collections of the Evolutionary Studies Insti- tute (University of the Witwatersrand, Johannesburg), the assistant curator S. Jirah, and B.S. Rubidge for their help and access to the col- lections of the ESI. The authors also thank the two anonymous reviewers for their help to improve the manuscript. This research was conducted with financial support from PAST and its scatterlings projects; the NRF; and DST/NRF Centre of Excel- lence in Palaeosciences. REFERENCES Angielczyk, K.D., Rubidge, B.S., Day, M.O., and Lin, F. 2016. A Reevaluation of Brachyprosopus broomi and Chelydontops altidentalis, Dicynodonts (Therapsida, Anomodontia) from the Middle Permian Tapinoceph- alus Assemblage Zone of the Karoo Basin, South Africa. Journal of Vertebrate Paleontology, 36(2):e1078342. Benoit, J., Abdala, F., Van den Brandt, M.J., Manger, P.R., and Rubidge, B.S. 2015. Physiological implica- tions of the abnormal absence of the parietal foramen in a Late Permian cynodont (Therapsida). The Sci- ence of Nature (Naturwissenschaften), 102:69. 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In Endo, H. and Frey, R. (eds.), Anatomical Imaging: Towards a New Morphology. Springer Verlag, Tokyo. 14 PALAEO-ELECTRONICA.ORG APPENDIX 1. Movie of the aligned serial section drawings of BP/1/1821 made by A.S. Brink. The 116 sections represent the entire specimen. Note that Brink numbered the sections from 1 to 118 because he missed number 83 and duplicated 110; however Brink also labelled every section with the corre- sponding 0.5 mm interval so we are confident that no section is missing. Because of this misla- belling, the section numbers in the movie are shifted between 83 and 110 with respect to Brink’s numbered sections (see Brink, 1961, figure 35). See: palaeo-electronica.org/content/2016/1478-reconstructing-scalopocynodon 15 BENOIT & JASINOSKI: RECONSTRUCTING SCALOPOCYNODON APPENDIX 2. Movie of the virtual 3D model of BP/1/1821 that was reconstructed from the drawings made by A.S. Brink (1961). See: palaeo-electronica.org/content/2016/1478-reconstructing-scalopocynodon 16 Picking up the pieces: the digital reconstruction of a destroyed holotype from its serial section drawings Julien Benoit and Sandra C. Jasinoski Introduction Material and methods Description Discussion Significance for Research Significance for Curation and Archiving Concluding remarks Acknowledgments References << /ASCII85EncodePages false /AllowTransparency false /AutoPositionEPSFiles true /AutoRotatePages /None /Binding /Left /CalGrayProfile (Dot Gain 20%) /CalRGBProfile (sRGB IEC61966-2.1) /CalCMYKProfile (U.S. Web Coated \050SWOP\051 v2) /sRGBProfile (sRGB IEC61966-2.1) /CannotEmbedFontPolicy /Error /CompatibilityLevel 1.4 /CompressObjects /Tags /CompressPages true /ConvertImagesToIndexed true /PassThroughJPEGImages true /CreateJobTicket false /DefaultRenderingIntent /Default /DetectBlends true /DetectCurves 0.0000 /ColorConversionStrategy /CMYK /DoThumbnails false /EmbedAllFonts true /EmbedOpenType false /ParseICCProfilesInComments true /EmbedJobOptions true /DSCReportingLevel 0 /EmitDSCWarnings false /EndPage -1 /ImageMemory 1048576 /LockDistillerParams false /MaxSubsetPct 100 /Optimize true /OPM 1 /ParseDSCComments true /ParseDSCCommentsForDocInfo true /PreserveCopyPage true /PreserveDICMYKValues true /PreserveEPSInfo true /PreserveFlatness true /PreserveHalftoneInfo false /PreserveOPIComments true /PreserveOverprintSettings true /StartPage 1 /SubsetFonts true /TransferFunctionInfo /Apply /UCRandBGInfo /Preserve /UsePrologue false /ColorSettingsFile () /AlwaysEmbed [ true ] /NeverEmbed [ true ] /AntiAliasColorImages false /CropColorImages true /ColorImageMinResolution 300 /ColorImageMinResolutionPolicy /OK /DownsampleColorImages true /ColorImageDownsampleType /Bicubic /ColorImageResolution 300 /ColorImageDepth -1 /ColorImageMinDownsampleDepth 1 /ColorImageDownsampleThreshold 1.50000 /EncodeColorImages true /ColorImageFilter /DCTEncode /AutoFilterColorImages true /ColorImageAutoFilterStrategy /JPEG /ColorACSImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /ColorImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /JPEG2000ColorACSImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /JPEG2000ColorImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /AntiAliasGrayImages false /CropGrayImages true /GrayImageMinResolution 300 /GrayImageMinResolutionPolicy /OK /DownsampleGrayImages true /GrayImageDownsampleType /Bicubic /GrayImageResolution 300 /GrayImageDepth -1 /GrayImageMinDownsampleDepth 2 /GrayImageDownsampleThreshold 1.50000 /EncodeGrayImages true /GrayImageFilter /DCTEncode /AutoFilterGrayImages true /GrayImageAutoFilterStrategy /JPEG /GrayACSImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /GrayImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /JPEG2000GrayACSImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /JPEG2000GrayImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /AntiAliasMonoImages false /CropMonoImages true /MonoImageMinResolution 1200 /MonoImageMinResolutionPolicy /OK /DownsampleMonoImages true /MonoImageDownsampleType /Bicubic /MonoImageResolution 1200 /MonoImageDepth -1 /MonoImageDownsampleThreshold 1.50000 /EncodeMonoImages true /MonoImageFilter /CCITTFaxEncode /MonoImageDict << /K -1 >> /AllowPSXObjects false /CheckCompliance [ /None ] /PDFX1aCheck false /PDFX3Check false /PDFXCompliantPDFOnly false /PDFXNoTrimBoxError true /PDFXTrimBoxToMediaBoxOffset [ 0.00000 0.00000 0.00000 0.00000 ] /PDFXSetBleedBoxToMediaBox true /PDFXBleedBoxToTrimBoxOffset [ 0.00000 0.00000 0.00000 0.00000 ] /PDFXOutputIntentProfile () /PDFXOutputConditionIdentifier () /PDFXOutputCondition () /PDFXRegistryName () /PDFXTrapped /False /CreateJDFFile false /Description << /ARA /BGR /CHS /CHT /CZE /DAN /DEU /ESP /ETI /FRA /GRE /HEB /HRV (Za stvaranje Adobe PDF dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. 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