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. 2023 Feb 22;18(2):e0281316. doi: 10.1371/journal.pone.0281316

Osteo-pathological analysis provides evidence for a survived historical ship strike in a Southern Hemisphere fin whale (Balaenoptera physalus)

Hannah Viola Daume 1,2,*, Helena Herr 2, Heinrich Mallison 3, Matthias Glaubrecht 1,4, Thomas M Kaiser 1
Editor: Olga Spekker5
PMCID: PMC9946264  PMID: 36812193

Abstract

The life history of a fin whale (Balaenoptera physalus) caught during whaling operations in the 1950s was partly reconstructed. 3D surface models of the bones of the skeleton curated at the Zoological Museum of Hamburg were used for an osteopathological analysis. The skeleton revealed multiple healed fractures of ribs and a scapula. Moreover, the processus spinosi of several vertebrae were deformed and arthrosis was found. Together, the pathological findings provide evidence for large blunt trauma and secondary effects arising from it. Reconstruction of the likely cause of events suggests collision with a ship inflicting the fractures and leading to post traumatic posture damage as indicated by skeletal deformations. The injured bones had fully healed before the fin whale was killed by a whaler in the South Atlantic in 1952. This study is the first in-detail reconstruction of a historical whale—ship collision in the Southern Hemisphere, dating back to the 1940s, and the first documentation of a healed scapula fracture in a fin whale. The skeleton provides evidence for survival of a ship strike by a fin whale with severe injuries causing long-term impairment.

Introduction

The Zoological Museum of Hamburg houses a large collection of marine mammal skeletons, of which only a few are currently on display. The fin whale (Balaenoptera physalus) skeleton (ZMH S 10261/10616) was brought back to Hamburg by Dr. Kurt Schubert [1] who accompanied the second whaling campaign financed by Aristoteles Onassis in 1952. Past World War II Hamburg-based whaling campaigns were financed by Greek shipowner Aristoteles Onassis using Panama and Honduran whalers to evade the ban on German whaling imposed as part of the Potsdam Conference [2,3]. Due to space limitation, the skeleton could not be mounted and displayed until in 2015 new exhibition space became available and efforts were undertaken to partly mount the skeleton for display.

On this occasion, we investigated the skeleton and documented several deformations and abnormalities, such as healed bone fractures, deformations of the neural spine and arthrosis. These findings called for a more thorough investigation of the skeleton in order to answer anatomical as well as osteopathological questions, ultimately in an attempt to reconstruct the life history of this particular specimen of fin whale.

Only a few attempts have been made to reconstruct the cause of traumata in whale skeletons from museum collections. Hellier, Hufthammer [4] investigated a fin whale and a humpback whale (Megaptera novaeanglia) skeleton collected in the 19th century, which both exhibited notable osteological pathologies. Based on visual inspection, they concluded, that the fin whale had likely suffered trauma from a ship collision. However, their investigations were restricted to visual inspection of the individual damaged bones.

Today, 3D modelling is a useful tool for surface digitising, allowing visual as well as quantitative analysis of bone specimens. The digital representation then serves as a reference of the surface morphology, while the original specimen can remain untouched. Besides of the ease in handling bulky or delicate specimens just in the virtual world, 3D models also help perfect visualisation and may serve educational purposes among many others. 3D models can then be used for e.g., for the digital assembly of a skeleton and for an animated visualisation and size-reduced 3D printouts. In this study, we used different 3D surface modelling techniques to build a virtual model of the entire fin whale skeleton and to employ those models to document its osteopathology. We then combined this virtual skeleton with a digital ship model to virtually reconstruct a plausible scenario of a ship—whale collision explaining the observed injuries and consequences.

Materials and methods

The bones of the fin whale skeleton (ZMH S 10261/10616) were first visually inspected and arranged. We then measured and described all bones according to Buchholtz and Schur [5] and Carrillo, Alcántara [6]. Subsequently, digital surface models of each individual bone or bone fragment were created applying two different methods of 3D surface data acquisition: 1) Structured light scanning using Artec 3D scanning systems (https://www.artec3D.com) was performed by 3D Manufactur Seevetal, and 2) photogrammetry was conducted using Agisoft Photoscan (www.agisoft.com) and Meshlab [7]. The latter technique was used for the cranium and other bones that proved difficult to handle or even not suitable for Artec 3D scanning gear. All surface models were converted to stereolithography format (*.STL) and digitally mounted in Rhinoceros 5.0. [8]. For the mounting we followed the protocol of Mallison [9] in order to avoid the influence of preconceived notions on the mount. We then adjusted the skeleton based on our experience from whale dissections and aerial and subaqueous photographs of fin whales. Subsequently, the skeleton was visualised, measured in full length and the pathologies described. Based on the virtually mounted model of the skeleton a virtual scenario of a ship—whale collision was generated as the most plausible cause of the observed trauma. Given the size of fin whales, only a virtual skeleton allowed a holistic assessment of the spatial distribution of the pathologies in relation to a ship hull.

Results

Based on the 3D model of the skeleton (Fig 1) we reconstructed a total body length of 19.54 m, indicating an adult individual [10]. 170 bones were preserved, including 62 vertebrae (C7 T14 L16 Ca25) and 14 pairs of ribs. The hyoid bone, the pelvis, the bullae and some of the digital bones were found missing.

Fig 1. 3D model of fin whale (Balaenoptera physalus) skeleton ZMH S 10261/10616).

Fig 1

Bones showing healed fractures are marked in red.

All epiphyses were fused with their respective vertebral bodies indicating the specimen was physically mature [11–13]. Abnormalities of different types were detected in 19 out of the 170 bones available, including healed fractures, arthrosis and deformations. The anomalies are described in detail in the following:

Scapula

The right scapula body (Figs 2–4) exhibits a simple extraarticular fracture that descends sagittally through the entire bone. The resulting two fragments moved one upon the other and healed ad longitudinem cum contractione [14] resulting in an offset of ca. 10 cm (see Fig 5 for comparison with unfractured left scapula). On the distal end of the fracture line a large mass of callus tissue formed.

Fig 2. The right scapula of fin whale ZMH S 10261/10616 from the external side.

Fig 2

Note the healed fracture.

Fig 4. The right scapula of fin whale ZMH S 10261/10616 in dorsal view.

Fig 4

Note the healed fracture.

Fig 5. Comparison of the unfractured left scapula and fractured right scapula of fin whale ZMH S 10261/10616 in external view.

Fig 5

From the measurements a 10 cm offset is evident.

Fig 3. The right scapula of fin whale ZMH S 10261/10616 from the internal side.

Fig 3

Note the healed fracture.

Ribs

A simple extraarticular fracture is evident in the proximal sections of both the first (Fig 6) and second (Fig 7) right rib, respectively. The fractures descend transversally and separate both ribs into a short proximal and a longer distal fragment each. Significant calli formed during the healing process, encapsulating the fractures. In both ribs the fragments healed in slightly displaced positions ad latus [14].

Fig 6. The first right rib of the fin whale skeleton ZMH S 10261/10616 in caudal view.

Fig 6

Fig 7. The second right rib of the fin whale skeleton ZMH S 10261/10616 in caudal view.

Fig 7

Vertebrae

Lumbar vertebrae L8, L9 and L10 (Fig 8) show fractures on their distal portion of their processus spinosi. These fractures descend sagitally through the individual processus forming a caudally descending line across the three processes (Fig 9). Lumbar vertebra L8 healed ad latus, lumbar vertebrae L9 and L10 healed displaced ad longitudinem cum contractione and thereby had shortened the processus.

Fig 8. Lumbar vertebrae L8, L9 and L10 of the fin whale ZMH S 10261/10616 in left lateral view.

Fig 8

The processus spinosi show healed fractures in a line descending from front to back.

Fig 9. Lateral view of lumbar vertrebrae L8, L9 and L10 of the fin whale ZMH S 10261/10616 in right lateral view.

Fig 9

Red line indicating the line of excessive bending moment imposed on the whale’s spine likely causing the fractures on the spinal processes.

Several lumbar vertebrae anterior (L5-L7) and posterior (L11-L15) of the fractured vertebrae (L8-10) show deformations in their processus spinosi (Figs 10 and 11). L5-L7 and L11-L13 are bended towards the left, L14-L15 are bended to the right (Fig 12).

Fig 10. Skeleton of the fin whale ZMH S 10261/10616.

Fig 10

Lumbar vertebrae anterior and posterior of the fractured vertebrae with deformations of their processus spinosi (marked in blue), likely the result of a post traumatic posture damage. Vertebrae with arthrosis are marked in yellow. Bones showing healed fractures are marked in red.

Fig 11. Lumbar vertebra 14 of fin whale ZMH S 10261/10616 in posterior view.

Fig 11

Note strong left lateral bend in processus spinosus, likely the result of a post traumatic posture damage.

Fig 12. Deformed vertebrae of fin whale ZMH S 10261/10616.

Fig 12

L5-L7 and L11-L13 surrounding the broken vertebrae L8-L10 are bended towards the left, L14-L15 are bended to the right.

Some thoracic vertebrae showed proof of arthrosis in their zygapophysial joints (Fig 13). The postzygapophysial joints were affected by zygarthrosis [15] in the thoracic vertebrae T2 and T3, both post- and praezygapophysial joints were affected in the thoracic vertebrae T4 and T5. The thoracic vertebrae T6 showed evidence of zygarthrosis in its praezygapophysial joints.

Fig 13. The contact between thoracic vertebrae T3 and T4 of fin whale ZMH S 10261/10616 in dorsal view, showing evidence of arthrosis in the zygapophysial joints (red arrow).

Fig 13

Discussion

The skeleton shows two major areas of severe trauma that cannot be easily explained without assuming a major external force acting on the bones. A fractured scapula is a rare observation in necropsied cetaceans and cetacean skeletons, and a comparatively rare injury in vertebrates because the scapula bone is particularly sturdy [16]. In small cetaceans fractured scapulae occur mainly as a result of intra-interspecific interaction [17]. In large baleen whales, to our knowledge only one case of a fractured scapula has been described from a necropsy of a North Atlantic right whale (Eubalaena glacialis, [18]). Killer whales (Orcinus orca) have been shown to cause rib fractures also in large baleen whales [19,20], but a fracture to the scapula, which is a much stronger and more robust bone than the ribs, is unlikely to have been caused by a killer whale attack. Therefore, we can exclude other marine organisms as the culprit.

The most plausible agent for this kind of blunt force trauma causing such fractures is a ship strike [21]. Trauma resulting from a vessel collision is characteristically marked by a premortem fracture or displacement of skeletal elements [21]. In fact, Campbell-Malone, Barco [21] describe comparable injuries, a broken mandible and cranium of a North Atlantic right whale.

We therefore reconstructed a possible ship strike scenario explaining the observed pattern of injuries and deformations in the skeleton. In this scenario, the fin whale would have been hit laterally by the bow of a ship, which caused fractures of the scapula and the ribs (Figs 14 and 15). The localities of the fractures and the straight break of the scapula, suggest collision with a vessel featuring a vertical steven or straight edged bow as commonly seen in military vessels built in the early decades of the 20th century, before the bulbous bow became more common in the early 1950s [22]. Whether this bow was a straight and narrow construction, sharp enough to cause penetrating trauma to soft tissues and bone, or a broader construction that resulted in blunt but localized trauma that snapped the upper part of the scapula blade via dislocation remains unclear. The breaks in a cranium and mandible of a Northern right whale reported by Campbell-Malone, Barco [21] resulted from blunt trauma, as evidenced by the bruised but not lacerated soft tissues on the carcass. Still, the breaks are straight and look as if they had been caused by a sharp blade impacting the bone directly. In the absence of histological data on either soft tissue we cannot determine the exact type of impact that caused the broken scapula in ZMH S 10261/10616.

Fig 14. Reconstruction of a ship strike as the most plausible cause for the blunt force trauma causing a fracture of the scapula and the ribs. The fin whale is hit laterally by a ship with a straight bow.

Fig 14

Generated in Rhinoceros 5.0. Ship model represents HMS Ajax (http://www.naval-history.net/xGM-Chrono-06CL-Ajax.htm), a ship of the time period estimated for the collision. Model taken from: http://tf3dm.com/3d-model/british-light-cruiser-hms-ajax1939-39659.html.

Fig 15. Front view of the reconstruction of the ship strike as the most plausible cause for the blunt force trauma causing fractures to the scapula and ribs.

Fig 15

Generated in Rhinoceros 5.0. Ship model represents HMS Ajax, a ship of the time period estimated for the collision. Model taken from: http://tf3dm.com/3d-model/british-light-cruiser-hms-ajax1939-39659.html.

The two broken ribs (Figs 6 and 7) were probably fractured by this same impact, when the upper part of the scapula was pushed deep into the body of the whale. The rib fractures are located directly medially to the scapula fracture line (Figs 1, 14 and 16).

Fig 16. Fractured scapula and ribs of the fin whale ZMH S 10261/10616.

Fig 16

Note: The rib fractures are located directly medially to the scapula fracture line.

Arthrosis observed in the thoracic vertebrae T2-T6, which are located directly above the scapula, may be a long-term consequence of a joint contusion caused by the same impact. Joint trauma can induce a contusive chondropathy responsible for post traumatic degenerative joint disease [23].

This scenario however does not yet explain the fractures in the processus spinosi of lumbar vertebrae L8-10. These could have either resulted from another traumatic incident, making the fin whale a survivor of two ship strikes. However, the similar healing stage of the injuries points to infliction at the same point in time rather than two independent collision events. The injuries to the lumbar vertebrae likely stem from a second collision with the same ship within the same incident. In this scenario, the whale would have been forced down by the initial impact or may have attempted to dive away from the ship. This motion and the continued forward motion of the ship may have brought the lumbar region of the whale into contact with the keel (Figs 17 and 18) or, less likely, the propeller of the ship resulting in fractures in the processus spinosi of lumbar vertebrae L8-10. However, fractures of the processus of the vertebrae cannot be easily attributed to external trauma, since these bones are protected by a thick layer of blubber and muscle [24–26]. Stress fractures of the processus may however be caused by excessive tension on the osteo-articular tendon structures, because the most potent locomotor muscles of cetaceans have their origins in these processus, especially in the thoracic region and lumbar region [25]. Therefore, an alternative hypothesis may be that, at the moment of the collision, the whale was forcibly flexed by (or around) the ship immediately after the impact. This excessive bending moment imposed on the whale’s spine may have caused the fractures on the spinal processus of the lumbar vertebrae L8-10 [27].

Fig 17. Front view of the reconstruction of a possible second collision as a cause for the blunt force trauma causing fractures to the lumbar vertebrae L8-L10.

Fig 17

Generated in Rhinoceros 5.0. Ship model represents HMS Ajax, a ship of the time period estimated for the collision. Model taken from: http://tf3dm.com/3d-model/british-light-cruiser-hms-ajax1939-39659.html.

Fig 18. Side view of the reconstruction of a possible second collision as a cause for the blunt force trauma causing fractures to the lumbar vertebrae L8-L10.

Fig 18

Generated in Rhinoceros 5.0. Ship model represents HMS Ajax, a ship of the time period estimated for the collision. Model taken from: http://tf3dm.com/3d-model/british-light-cruiser-hms-ajax1939-39659.html.

As a consequence of these severe injuries, it is likely that the fin whale had to maintain a relieving posture for a extended period of time. Deformations in the vertebrae posterior and anterior of the fractured vertebrae may be the result of a prolonged posture damage. Likely, the fractured scapula impaired the movement particularly of the right flipper. In addition, broken ribs and haemorrhaging probably caused pain and may have caused the whale to attain a relieving posture, sparing the right side and dominantly using the left. This may have led to an uneven muscular strain onto spinal processus of the lumbar vertebrae in which the locomotor muscles have their origin [25] particularly of the vertebrae around the broken vertebrae in order to relieve strain on these, causing a bending over the prolonged time of the relieving posture.

Assuming, that the fin whale was hit by a ship, this would be the first detailed documentation of a historical ship strike in a Southern Hemisphere fin whale. Ship strikes evolved as a major threat for whales over the last century [28] as the size, speed and quantity of ships increased [29,30]. Fin whales are one of the species most frequently affected by ship strikes [30], with ship strike rates giving reason for concern in areas such as the Mediterranean [31,32]. In our case, the fin whale survived the ship strike despite the severity of the injuries, as indicated by the fact that the fractures had completely healed by the time it was killed by the whaler. The relatively low rate of remodeling in cetacean bone [33] and the necessity for continued use of the extremity suggests, that the time of the collision must have been several years before the fin whale was shot in 1952, placing the ship collision in the 1940s.

The consequences of the injury for the individual can only be hypothesized. It is likely that the whale’s movements were considerably impaired, as suggested by the evidence for a relieving posture, potentially affecting feeding efficiency, energy budgets and swim speed. It remains unknown, if the pre-condition made the individual an easier target for the whalers than other individuals.

While sharp trauma caused by propellers and rudders during ship strikes usually provides obvious external evidence of injury, blunt trauma like in the case of our fin whale typically leaves little external evidence of the injury, even in the most severe cases, because of the thickness of soft tissue and the dark pigment of much of the epidermis, which tends to obscure evidence of swelling and bruising [21]. Even severe injuries like the one reported here may go unnoticed in visual observations of live animals as well as stranded carcasses that are not necropsied. Therefore, cases like the fin whale in this study may contribute to the overall number of ship strikes, but remain undetected.

This study highlights the value of museum collections for research on threatened species and insights into the life history of individuals.

Acknowledgments

We would like to thank Matthias Preuß and Steffen Lässle, taxidermists at the Zoological Museum of Hamburg, for providing their expertise during the digital assembly of the fin whale skeleton.

Data Availability

All relevant data are within the paper.

Funding Statement

The author(s) received no specific funding for this work.

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Decision Letter 0

Olga Spekker

16 Nov 2022

PONE-D-22-27969Osteo-pathological analysis provides evidence for a survived historical ship strike in a Southern Hemisphere fin whale (Balaenoptera physalus)PLOS ONE

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The manuscript must describe a technically sound piece of scientific research with data that supports the conclusions. Experiments must have been conducted rigorously, with appropriate controls, replication, and sample sizes. The conclusions must be drawn appropriately based on the data presented.

Reviewer #1: Yes

Reviewer #2: Partly

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2. Has the statistical analysis been performed appropriately and rigorously?

Reviewer #1: N/A

Reviewer #2: N/A

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3. Have the authors made all data underlying the findings in their manuscript fully available?

The PLOS Data policy requires authors to make all data underlying the findings described in their manuscript fully available without restriction, with rare exception (please refer to the Data Availability Statement in the manuscript PDF file). The data should be provided as part of the manuscript or its supporting information, or deposited to a public repository. For example, in addition to summary statistics, the data points behind means, medians and variance measures should be available. If there are restrictions on publicly sharing data—e.g. participant privacy or use of data from a third party—those must be specified.

Reviewer #1: Yes

Reviewer #2: Yes

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4. Is the manuscript presented in an intelligible fashion and written in standard English?

PLOS ONE does not copyedit accepted manuscripts, so the language in submitted articles must be clear, correct, and unambiguous. Any typographical or grammatical errors should be corrected at revision, so please note any specific errors here.

Reviewer #1: Yes

Reviewer #2: Yes

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5. Review Comments to the Author

Please use the space provided to explain your answers to the questions above. You may also include additional comments for the author, including concerns about dual publication, research ethics, or publication ethics. (Please upload your review as an attachment if it exceeds 20,000 characters)

Reviewer #1: This is a compact and clear case study, providing results for basic research into cetacean pathology and its applied aspect in the welfare of whales. It also shows the increasing importance of "old fashoned" museum collections in research concerning threatened species which may be worth emphasizing in the text.

The article is clearly written, supported by high quality illustrations obtained through sophisticated imaging methods.

Perhaps the description of directions could be clarified (actually simplified) in the foollowing figures:

Line 109: Figure 2: ... in lateral view from the external side

Line 113: Figure 3: ... in lateral view from the internal side.

I find the use of "lateral" confusing here as I suspect that the "external" side is lateral and the "internal" side is medial. I presume the original meaning was a reference to the lateral view of the skeleton in general. If I misunderstood something, a brief explanation would help other readers.

Line 140-143: Figures 8-9: ... left dorsolateral view

I think this is simply the lateral view as I do not see the vertebra being shown from "slightly above".

There seems to be superfluous wording here:

Line 147: Several lumbar vertebrae anterior (L5-L7) and posterior (L11-L15), [posterior and anterior of the...]

The text in square brackets could be deleted.

A reference could be added to a concrete parameters of HMS Ajax here:

Line 203: represents HMS Ajax, a ship of the time period estimated for the collision.

Although in the discussion proper references are made to a variety of accidents, it would be worth devoting a sentence in the text on how many similar traumatic cases are known. My impression is that this observation is quite unique, but it would be worth placing it in a slightly broader, concrete perspective.

Reviewer #2: The submitted manuscript [PONE-D-22-27969] concerns the recognition of traumatic pathologies on the skeleton of a whale (Balaenoptera physalus) and the reconstruction of the traumatic event. The fractures were probably caused, according to the authors, by a collision with a ship in the 1940s. This whale was killed by a whaler in the South Atlantic in 1952 and its skeleton (ZMH S 10261/10616) belongs to the Hamburg Zoological Museum, which has been exhibiting it since 2015.

The authors used 2 types of 3D surface imaging: structured light scanning and photogrammetry. The interest of the 3D surface imaging used in this article lies mainly in the creation of a virtual model of the whale and the reconstruction of the traumatic event. Indeed, it does not help in retrospective diagnosis, as it does not offer more significant information than a set of good photographic images. A X-ray study of the damaged bones, for example, would have been interesting to visualize the internal bone structures, bringing additional information to the diagnosis of these consolidated fractures.

The adjustment of the different elements of the skeleton to propose a general 3D reconstruction of the whale, carried out by the authors according to their experience is interesting and useful to better understand both the location of the lesions and the history of the trauma. It seems to me to be well done, even if I am not a specialist in the anatomy of the whale skeleton.

My main comments are about some of the skeletal lesions and the interpretations:

- although there are no supporting radiographs, I agree with the authors on the diagnosis of healed fractures of the right scapula, the right ribs of ranks 1 and 2, and the lumbar vertebrae of ranks 8 to 10. The signs visible on the external surface of the 3D reconstructions of these bones are quite characteristic of traumatic conditions and do not require a differential diagnosis.

- the authors observed deformations of the spinous processes of the lumbar vertebrae located on either side of the fractured vertebrae, i.e. lumbar vertebrae of ranks 5 to 7 and 8 to 10. These deformations are interpreted by the authors as a condition due to posture. I would like the authors to better argue this interpretation : would they not be consolidated fractures of the spinous processes ? Indeed in figure 11, the orientation of the spinous process is very clear but could as well result from direct impact with the bow of the ship and not secondary to the animal’s posture due to pain.

- similarly, the authors interpret the origin of the degenerative joint disease observed in the upper thoracic vertebrae (T2 to T6) as postural in origin. It would be interesting to illustrate these osteoarthritic vertebrae on the figures of the reconstructions of the animal's bone lesions (figures 1, 10, 13 and 14). This would make it possible to discuss an alternative hypothesis: the degenerative phenomenon observed on the posterior vertebral joints at the thoracic level could be due, not to a postural cause, but to a joint contusion linked to the same trauma responsible for the fractures of the scapula and the ribs. Indeed, any joint trauma can induce a « contusive chondropathy » responsible for post-traumatic degenerative joint disease.

- the authors mention in lines 194-195, that in the absence of bone histology data, it is difficult to determine the type of impact that caused the scapula fracture. If the authors believe that bone histology would differentiate between a blunt and a penetrating agent, I would like them to support their assertion here.

- the authors propose an alternative hypothesis to the presence of two fracture sites on the whale at the scapula and the lumbar vertebrae (lines 224 to 227): indirect trauma by excessive flexion of the spine. I am not convinced by this hypothesis. The interpretation of a first impact on the whale's scapula with the ship and then a second impact when the whale swung its tail to dive seems more plausible.

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6. PLOS authors have the option to publish the peer review history of their article (what does this mean?). If published, this will include your full peer review and any attached files.

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Reviewer #1: Yes: László Bartosiewicz

Reviewer #2: No

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Decision Letter 1

Olga Spekker

8 Jan 2023

PONE-D-22-27969R1Osteo-pathological analysis provides evidence for a survived historical ship strike in a Southern Hemisphere fin whale (Balaenoptera physalus)PLOS ONE

Dear Dr. Daume,

Thank you for submitting your manuscript to PLOS ONE. After careful consideration, we feel that it has merit but does not fully meet PLOS ONE’s publication criteria as it currently stands. Therefore, we invite you to submit a revised version of the manuscript that addresses the points raised during the review process.

Comments from the Academic Editor:

Dear Dr. Daume,

We appreciate you submitting your manuscript to PLOS ONE and thank you for giving us the opportunity to consider your work.

I have completed my evaluation of your revised manuscript, which has been reviewed by two highly qualified reviewers all of whom agree it is worth to be published in PLOS ONE. Nevertheless, I kindly ask you to review your in-text citations and reference list, as they do not always follow the guidelines of PLOS ONE (https://journals.plos.org/plosone/s/submission-guidelines#loc-references). The references should be numbered in the order that they appear in the text. In the text, the reference number should be cited in square brackets. As for the formatting of the references, PLOS ONE uses the so-called „Vancouver style”. (I copied below the relevant parts of the guide for authors.)

„References are listed at the end of the manuscript and numbered in the order that they appear in the text. In the text, cite the reference number in square brackets (e.g., “We used the techniques developed by our colleagues [19] to analyze the data”). PLOS uses the numbered citation (citation-sequence) method and first six authors, et al.”

„PLOS uses the reference style outlined by the International Committee of Medical Journal Editors (ICMJE), also referred to as the “Vancouver” style. Example formats are listed below. Additional examples are in the ICMJE sample references…”

In addition to the above, I kindly ask you to follow the guidelines of PLOS ONE regarding the figure captions (e.g., Fig 1. This is the Fig 1 Title. This is the Fig 1 legend) and in-text citation of figures (e.g., Fig 1, Figs 1 and 2, Figs 1, 2, and 3) (https://storage.googleapis.com/plos-published-prod/9cba/PLOS%20Manuscript%20Body%20Formatting%20Guidelines.pdf?X-Goog-Algorithm=GOOG4-RSA-SHA256&X-Goog-Credential=wombat-sa%40plos-prod.iam.gserviceaccount.com%2F20230107%2Fauto%2Fstorage%2Fgoog4_request&X-Goog-Date=20230107T075032Z&X-Goog-Expires=86400&X-Goog-SignedHeaders=host&X-Goog-Signature=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).

Furthermore, figure files should be uploaded separately, instead of being included in the main manuscript file. (I copied below the relevant part of the guide for authors.)

Do not include figures in the main manuscript file. Each figure must be prepared and submitted as an individual file.

Cite figures in ascending numeric order at first appearance in the manuscript file.

Figure captions must be inserted in the text of the manuscript, immediately following the paragraph in which the figure is first cited (read order). Do not include captions as part of the figure files themselves or submit them in a separate document.

At a minimum, include the following in your figure captions:

  • A figure label with Arabic numerals, and “Figure” abbreviated to “Fig” (e.g. Fig 1, Fig 2, Fig 3, etc). Match the label of your figure with the name of the file uploaded at submission (e.g. a figure citation of “Fig 1” must refer to a figure file named “Fig1.tif”).

  • A concise, descriptive title

The caption may also include a legend as needed.”

Finally, please find below some minor comments:

1) There are inconsistencies in the writing of some terms: e.g., life history or life-history; 1950ies/1940ies or 1950s/1940s; post traumatic or post-traumatic; ship-strike or ship strike; 3D models, 3D-surface or 3-D model - please, revise the text and eliminate these inconsistencies;

2) There are some typos or grammatical issues: e.g., line 76 – „or even not not suitable”; line 92 – „digital digiti”; line 166 – instead of its, their or the should be used; line 242 – „locomotors muscles” should be changed to locomotor muscles; lines 246–247 – „on the vertebra’s spinal processus” should be rephrased; lines 269–270 – „as the ships size, speed and quantity” should be rephrased (e.g., as the size, speed and quantity of ships) - please, eliminate the aforementioned issues; and

3) Figures and their captions: in the caption of figure 2, it seems that a space is missing between the specimen number and from; the caption of figures 6 and 7 should be rephrased to unify and simplify the captions (“The first right rib of fin whale ZMH S 10261/ 10616 in caudal view”; and “The second right rib of fin whale ZMH S 10261/ 10616 in caudal view.”); in the caption of figures 15, 17, and 18, “view” should not be capitalised; in the caption of figure 2, “of” is in a different font type than the rest of the caption; in the caption of figure 5, maybe external should be used instead of lateral, as in figure 2; in the caption of figure 9, one of the reviewers already mentioned that “dorso” should be deleted; and in figure 10, the “green” seems to me more like blue, and maybe the scapula should not be highlighted in red, as in the legend, only the vertebrae are mentioned.

Based on the above, I invite you to resubmit your manuscript after amending it following the aforementioned comments and suggestions. Please, note that your revised submission may need to be re-reviewed.

PLOS ONE values your contribution and I look forward to receiving your revised manuscript.

Yours sincerely,

Dr. Olga Spekker

Please submit your revised manuscript by Feb 22 2023 11:59PM. If you will need more time than this to complete your revisions, please reply to this message or contact the journal office at plosone@plos.org. When you're ready to submit your revision, log on to https://www.editorialmanager.com/pone/ and select the 'Submissions Needing Revision' folder to locate your manuscript file.

Please include the following items when submitting your revised manuscript:

  • A rebuttal letter that responds to each point raised by the academic editor and reviewer(s). You should upload this letter as a separate file labeled 'Response to Reviewers'.

  • A marked-up copy of your manuscript that highlights changes made to the original version. You should upload this as a separate file labeled 'Revised Manuscript with Track Changes'.

  • An unmarked version of your revised paper without tracked changes. You should upload this as a separate file labeled 'Manuscript'.

If you would like to make changes to your financial disclosure, please include your updated statement in your cover letter. Guidelines for resubmitting your figure files are available below the reviewer comments at the end of this letter.

If applicable, we recommend that you deposit your laboratory protocols in protocols.io to enhance the reproducibility of your results. Protocols.io assigns your protocol its own identifier (DOI) so that it can be cited independently in the future. For instructions see: https://journals.plos.org/plosone/s/submission-guidelines#loc-laboratory-protocols. Additionally, PLOS ONE offers an option for publishing peer-reviewed Lab Protocol articles, which describe protocols hosted on protocols.io. Read more information on sharing protocols at https://plos.org/protocols?utm_medium=editorial-email&utm_source=authorletters&utm_campaign=protocols.

We look forward to receiving your revised manuscript.

Kind regards,

Olga Spekker, Ph.D.

Academic Editor

PLOS ONE

Journal Requirements:

Please review your reference list to ensure that it is complete and correct. If you have cited papers that have been retracted, please include the rationale for doing so in the manuscript text, or remove these references and replace them with relevant current references. Any changes to the reference list should be mentioned in the rebuttal letter that accompanies your revised manuscript. If you need to cite a retracted article, indicate the article’s retracted status in the References list and also include a citation and full reference for the retraction notice.

[Note: HTML markup is below. Please do not edit.]

Reviewers' comments:

Reviewer's Responses to Questions

Comments to the Author

1. If the authors have adequately addressed your comments raised in a previous round of review and you feel that this manuscript is now acceptable for publication, you may indicate that here to bypass the “Comments to the Author” section, enter your conflict of interest statement in the “Confidential to Editor” section, and submit your "Accept" recommendation.

Reviewer #1: (No Response)

Reviewer #2: All comments have been addressed

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2. Is the manuscript technically sound, and do the data support the conclusions?

The manuscript must describe a technically sound piece of scientific research with data that supports the conclusions. Experiments must have been conducted rigorously, with appropriate controls, replication, and sample sizes. The conclusions must be drawn appropriately based on the data presented.

Reviewer #1: Partly

Reviewer #2: Yes

**********

3. Has the statistical analysis been performed appropriately and rigorously?

Reviewer #1: N/A

Reviewer #2: N/A

**********

4. Have the authors made all data underlying the findings in their manuscript fully available?

The PLOS Data policy requires authors to make all data underlying the findings described in their manuscript fully available without restriction, with rare exception (please refer to the Data Availability Statement in the manuscript PDF file). The data should be provided as part of the manuscript or its supporting information, or deposited to a public repository. For example, in addition to summary statistics, the data points behind means, medians and variance measures should be available. If there are restrictions on publicly sharing data—e.g. participant privacy or use of data from a third party—those must be specified.

Reviewer #1: Yes

Reviewer #2: Yes

**********

5. Is the manuscript presented in an intelligible fashion and written in standard English?

PLOS ONE does not copyedit accepted manuscripts, so the language in submitted articles must be clear, correct, and unambiguous. Any typographical or grammatical errors should be corrected at revision, so please note any specific errors here.

Reviewer #1: Yes

Reviewer #2: Yes

**********

6. Review Comments to the Author

Please use the space provided to explain your answers to the questions above. You may also include additional comments for the author, including concerns about dual publication, research ethics, or publication ethics. (Please upload your review as an attachment if it exceeds 20,000 characters)

Reviewer #1: (No Response)

Reviewer #2: The authors of the manuscript PONE-D-22-27969R1 have answered all the questions asked in my first review. They have taken into account all my remarks and suggestions, including the addition of a new figure (12) representing in 3D the concerned vertebrae, which clearly supports their interpretations.

This revised version seems to me now acceptable for publication in the journal Plos One.

**********

7. PLOS authors have the option to publish the peer review history of their article (what does this mean?). If published, this will include your full peer review and any attached files.

If you choose “no”, your identity will remain anonymous but your review may still be made public.

Do you want your identity to be public for this peer review? For information about this choice, including consent withdrawal, please see our Privacy Policy.

Reviewer #1: No

Reviewer #2: Yes: Helene COQUEUGNIOT

**********

[NOTE: If reviewer comments were submitted as an attachment file, they will be attached to this email and accessible via the submission site. Please log into your account, locate the manuscript record, and check for the action link "View Attachments". If this link does not appear, there are no attachment files.]

While revising your submission, please upload your figure files to the Preflight Analysis and Conversion Engine (PACE) digital diagnostic tool, https://pacev2.apexcovantage.com/. PACE helps ensure that figures meet PLOS requirements. To use PACE, you must first register as a user. Registration is free. Then, login and navigate to the UPLOAD tab, where you will find detailed instructions on how to use the tool. If you encounter any issues or have any questions when using PACE, please email PLOS at figures@plos.org. Please note that Supporting Information files do not need this step.

Decision Letter 2

Olga Spekker

20 Jan 2023

Osteo-pathological analysis provides evidence for a survived historical ship strike in a Southern Hemisphere fin whale (Balaenoptera physalus)

PONE-D-22-27969R2

Dear Dr. Daume,

We’re pleased to inform you that your manuscript has been judged scientifically suitable for publication and will be formally accepted for publication once it meets all outstanding technical requirements.

Within one week, you’ll receive an e-mail detailing the required amendments. When these have been addressed, you’ll receive a formal acceptance letter and your manuscript will be scheduled for publication.

An invoice for payment will follow shortly after the formal acceptance. To ensure an efficient process, please log into Editorial Manager at http://www.editorialmanager.com/pone/, click the 'Update My Information' link at the top of the page, and double check that your user information is up-to-date. If you have any billing related questions, please contact our Author Billing department directly at authorbilling@plos.org.

If your institution or institutions have a press office, please notify them about your upcoming paper to help maximize its impact. If they’ll be preparing press materials, please inform our press team as soon as possible -- no later than 48 hours after receiving the formal acceptance. Your manuscript will remain under strict press embargo until 2 pm Eastern Time on the date of publication. For more information, please contact onepress@plos.org.

Kind regards,

Olga Spekker, Ph.D.

Academic Editor

PLOS ONE

Additional Editor Comments (optional):

Reviewers' comments:

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