Created February 1998; revised February 1998 with summary posted in Feb. 1998 on ELASMO-L. Please send comments, corrections, and updates to:

Henry Mollet or Jim Bourdon


Ichthyologists and Paleontologists

  • Are we continuing to throw darts at dentitions and pick terminology accordingly? There are basically 3 directions (mesial-distal, labio-lingual, and for pavement-like dentitions an in-between direction looks well defined) and 3 terms (row, file and series). Based on prior usage just about any combination of these terms can be used. We've done a fairly thorough but not complete literature search, included dictionary definitions of the terms, had feedback from John Bruner, Leonard Compagno, Malcolm Francis, Bill Heim, John McEachran, Sandy Moss, Gavin Nayler, Bob Purdy, Matthias Stehmann, Michael Siverson, Adam Summers, David Ward, and Sabine Winter but no agreement. We are proposing the following terminology for ichthyologists and paleontologists (with full or partial endorsements by M. Francis, J. McEachran, G. Naylor, S. Moss, R. Purdy, and M. Siverson):

  • Row for teeth in the mesial-distal direction (mesio-distal, mesodistal). They are at the same developmental stage. In a dentition with imbricated/pavement teeth, adjacent teeth in a row may have slightly different ages.

  • File (tooth family, transverse row) for teeth in the labio-lingual direction. They are at different developmental stages and derived from the same tooth bud position.

  • Diagonal file for teeth in in-between direction. They are at different developmental stages and derived from different tooth bud positions.

  • In our opinion, there can be little doubt that for the mesial-distal direction the term row is best. In the often quoted Applegate (1965), row is used for both directions and today S. Applegate is using it for the mesial-distal direction. (R. Purdy pers. comm.)

    What term to use for the labio-lingual direction is more difficult to assess. After careful evaluation of prior usage, temporal and positional meaning of the terms file and series we propose to retain file. Any comments appreciated.

    Summary of Literature Search

  • Andre, W. 1784. A description of the teeth of Anarrhinchas lupus...to which is added an attempt to prove that the teeth of cartilagnious fishes are perpetually renewed. Phil. Trans. Roy. Soc. 74 (The sting of a sting ray embedded in the jaw of a Galeocerda, pp. 279-281; Pl. 13).

    Row is used for labio-lingual direction according to orginal text quoted in Gudger 1937.

  • Leidy, J. 1877. Description of Vertebrate Remains, chiefly from the Phosphate Beds of South Carolina. Journal of the Academy of Natural Sciences of Philadelphia 8: 209-261.

    The only specimens which consisted of more than isolated teeth were those of myliobatids. No reference was made to a mesio-distal set of teeth other than the tooth count of median teeth. Row was the most often used term to describe a labio-lingual line of teeth. On page 235, "The dental pavement consists of about a dozen median teeth together with part of the contiguous two rows of lateral teeth at the sides." Occasionally, series was employed for a labio-lingual line of teeth (page 235), "... seven median occupy an extent equal to the same number in the preceding specimen. The first and last of the series measure ..."

  • Lohberger, J. 1910. Ueber zwei riesige Embryonen von Lamna. Inaugural-Dissertation zur Erlangung der Doktorwuerde der Hohen Philosophischen Fakultaet der UniversitŠt Leipzig. Sonderabdruck aus den Abhandlungen der math.-phys. Klasse der K. Bayer. Akademie der Wissenschaften, IV. Suppl. Bd, 2. Abhandlg., 1-45 (and 5 plates with 34 figures).

    Lohberger introduced row for mesial-distal direction. Later, the possibility of ambiguous use arises unless we assume two typographical errors i.e. 4 (instead of 14) and 3 (instead of 13) replacement rows in upper and lower jaw, respectively (suggested by Sabine Winter).

  • Garman, S. 1913. The Plagiostomia (Sharks, Skates and Rays). Memoirs of the Museum of Comparative Zoology at Harvard College, 36:528 pages. Reprinted by Benthic Press (1997), Summers A. P. ed.

    Garman consistently used row to refer to a labio-lingual line of teeth and series was used for both the mesial distal and the labio-lingual direction . Garman reported tooth counts on p. 20 as row of teeth 40-46/36-40. Isistius (page 238) was described as "31 rows, a single series in function, ...". "Series" however, although employed for a mesio-distal line of teeth in the preceding example, was also used for a labio-lingual description in the myliobatids (page 427): "... median series commonly broader than the laterals."

    Accoording to Adam Summers, the terminology in Garman 1913 is consistent with the terminology used in earlier papers (likely a 1888? publication on myliobatids).

  • Bolk 1913. Odontologische Studien. I. Die Ontogenie der Primatenzaehne. Versuch einer Loesung der Gebissproblem.

    Not seen by us. See Vorstman 1922 and Edmund 1960. Likely used row for mesial-distal direction according to Vorstman 1922.

  • Woerdeman, M. 1919. Zur Entwicklungsgeschichte von Zaehnen und Gebiss der Reptilien. Beitrag I. Die Anlage und Entwicklung des embryonalen Gebisses als Ganzes und seine Beziehung zur Zahnleiste. Beitrag II. Ueber die Anlage des Ersatzbegisses und den Zahnwechsel. Archif. fuer Mikroskop. Anat. Bd. 2 Abt.1.

    Not seen by us, see Vorstman 1922 and Edmund 1960. Used row for labio-lingual direction and series for the mesial-distal direction according to Vorstman 1922 and Edmund 1960.

  • Woerdeman, M. 1921. Zur Entwicklungsgeschichte von Zaehnen und Gebiss der Reptilien. Beitrag IV. Ueber die Anlage und Entwicklung von Zaehnen. Beitrag V. Ueber die Beziehung der MundhoehlendrŸsen zum Zahnsystem. Archif. fuer Mikroskop. Anat. Bd. 95 Abt.1

    Not seen by us, see Vorstman 1922.

  • Vorstman, A. G. 1922. Ueber die Anordnung und die Entwicklung der Zaehne bei Teleostiern. Verhandlungen der Koninkliijke Akademie van Wetenschappen te Amsterdam (Tweede Sectie) Deel 22: 3-41.

    Adriana Vorstman is clear about the use of Zahnreihe (tooth row) for the mesial-distal (mesio-distal) direction. She is also discussing the different definitions used for the term "row" by Woerdeman and Bolk and summarized it as follows: Woerdeman used an entirely different definition for Stichos (pg. 206). Wordeman probably used row for labio-lingual direction) compared to the original definition used by Bolk 1913 (Bolk likely used row for mesio-distal direction).
    Original text from summary (item 7): Woerdeman hat mit seiner Definition eines Stichos (Pag. 206) ein grundsaetzlich ander Reihe ahgenommen, als diejenige, welcher Bolk urspruenglich den Name Stichos gegeben hat.

  • Leriche, M. 1927. Les poissons de la molasse suisse. Memoires de la Société Paléontologique Suisse 46: 1-119, avec 7 planches, 1 carte, 7 figures dans le texte.

    Leriche used file for a labio-lingual group of teeth, but no reference for a mesio-distal group could be found. Probably defined in different 1927 publication.

  • Gudger, E.W. 1937. Abnormal dentition in sharks, Selachii. Bulletin American Museum of Natural History 73: 249-280.

    Author used row for labio-lingual direction and transverse row for mesial-distal direction.

  • Bigelow, H. B. and W. C. Schroeder 1948. Part 1. Sharks In Fishes of the western North Atlantic. Sears Foundation for Marine Research, Yale University. pp. 59-546.

    Acccording to Compagno 1988, authors used row and series interchangeably for the mesio-distal direction.

  • Bigelow, H. B. and W. C. Schroeder 1953. Part 2. Sawfishes, Guitarfishes, Skates and Rays; Chimaeroids In Fishes of the western North Atlantic. Sears Foundation for Marine Research, Yale University, New Haven. pp. 588.

    Authors consistently used row for mesio-distal and series for labio-lingual with one deviation. On p. 442 "median row" (transverse) for labio-lingual direction instead of "median series" was used. A transverse row is indeed the same as a series but we don't understand why the authors did not use series in this instance.

  • Edmund, G. 1960. Tooth replacement in the lower vertebrates. Contribution No. 52 Life Sciences Division, Royal Ontario Museum Toronto. pp. 190.

    In the introduction the author uses the term "row" for the mesial-distal direction (teeth in approx. the same stage of development).

    In Chapter 19 embryological studies and Woerdeman's work are discussed. The term series (probably following Woerdeman is used for the mesial-distal direction) and the term "Zahnreihe" (German term is used) certainly following Woerdeman is introduced. We suggest that this is a diagonal file as per Compagno 1988 Fig. 3.2C. This is NOT the Zahnreihe as per Vorstman 1922, which was used for the mesial-distal direction. If one compares Fig. 54 or Fig. 55 with Fig. 56 (after Vorstman) one sees that Edmund used "tooth family" (file) for the labio-lingual direction, Bolk's Odontosticky for the medial-distal direction and Zahnreihe for what Compagno (1988) called a diagonal file. In Fig. 56 (after Vorstman), Edmund is now using Vorstman's Zahnreihe (medial-distal direction) in the figure caption. In summary, we suggest that Edmund's use of the term Zahnreihe for a diagonal file should be avoided. That leaves the discussion of Woerdeman 1921, which was not available. However, since Woerdeman 1921 introduced terminology in order to refute Bolk's disticky theory (presence of alternate discrete dental series produced in different locations), we suggest that he might have used terminology best suited to get the point across and which now can be abandoned.

  • Applegate, S. P. 1965. Tooth terminology and variation in sharks with special reference to the sand shark. Carcharias taurus  RAFINESQUE. Los Angeles County Museum Contributions to Science, 86:1-18.

    Author used the term "row" ambiguously, i.e. it was used to indicate a number of teeth in both mesio-distal and labio-lingual direction. Shelly Applegate is now using the term for the mesial-distal direction (R. Purdy, pers. comm.)

    We are quoting each statement in reverse order of appearance in the text to follow what we consider the weight of each.

    a. On p. 13 bottom/p. 14 top of page: "In reconstructing the dentition of fossil sharks several terms were needed. First "tooth set"; that is a single complete ROW of all types of mature teeth from both sides of the upper and lower jaw." (ROW is used to indicate teeth in mesial-distal direction in agreement with usage by and Zangerl (1981) and Cappetta (1987).

    b. On p. 13, second to last paragraph: "In some of the specimens of C. taurus the addition of posterior teeth during the life of the shark may be demonstrated, several single teeth were followed in the replacement series by a double row of immature teeth." (series is used to indicate mesial-distal direction; row is apparently now used to indicate position in mesial-distal direction, i.e. labio-lingual direction. Author here discusses what we would call FILE splitting following Zangerl (1981) and Cappetta (1987), in his terminology it would be ROW splitting.

    c. On p. 12, Replacement and Number of Teeth, end of first paragraph: "At any given time there are usually two functional ROWS of anterior, intermediates and laterals followed by from 4 to 6 rows of small posteriors." (Here, authore is apparently using the term row to indicate a partial tooth set. It might be considered in conflict with the term row used in his definition of a tooth set. How can we talk of rows of anterior if referring to the mesial-distal direction? That would be only 2 teeth. Perhaps author here also thinks in terms of the labio-lingual direction?

  • Applegate S. P. 1967. A survey of shark hard parts. In Sharks, skates and rays. P. W. Gilbert, R. F. Mathewson, and D. T. Rall, eds. John Hopkins Press, Baltimore, Maryland. pp. 37-67.

    Just as ambiguous as Applegate 1965 w.r.t to the use of the term row. On p.47 the following sentence appears: "Recent evidence from the hornshark kept at Marineland of the Pacific suggests that an adult advances its tooth row one position every 3 to 4 weeks.. There are twelve teeth that are undergoing calcification in a single row or file; once calcifications in a tooth is visible in a section, the tooth lasts about a year and is then lost. Since the teeth generally form an interlocking series, the whole row must move at approximately the same rate.

  • Peyer, B. 1968. Comparative odontology. Translated and edited by R. Zangerl. University of Chicago Press. Chicago, Illinois. 468 pp.

    Peyer clearly differentiates between cross-rows (tooth family as per Bolk 1913-1919) and longitudinal rows (occupying the edge of the jaw).

  • Moss, S. A. 1972. Tooth replacement and body growth rates in the smooth dogfish, Mustelus canis (Mitchill). Copeia 1972, 808-811.

    Intermixes the usage of row to mean labio-lingual and mesio-distal

  • Zangerl, R. 1981. Handbook of Paleoichthyology. Chondrichthyes I: Paleozoic Elasmobranchii. Gustav Fischer Verlag, Stuttgart and New York, 115 pages.

    Pg 15 "In addition to being arranged in rows parallel to the jaw, each tooth in concert with its replacements, forms a transversal row, across the jaw, sometimes called a tooth battery, tooth file, or tooth family."

  • Romer, A. S and T. S. Parsons 1986. The Vertebrate Body: Sixth Edition. Saunders College Publishing, 679 pages.

    Page 331, "... functioning teeth limited to a single marginal series along each upper and lower jaw." On page 336, a labio-lingual tooth set is referred to, "... a whole series of successional teeth can ..."

  • Cappetta, H. 1987. Handbook of Paleoichthyology. Chondrichthyes II: Mesozoic and Cenozoic Elasmobranchii. Gustav Fischer Verlag, Stuttgart and New York, 193 pages.

    author uses file for labio-lingual direction and row for mesio-distal direction on pages 13-19.

  • Herman, J., M. Hovestadt-Euler, and D. C. Hovestadt 1988. Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites in living supraspecific taxa of chondrichthyan fishes. Part A: Selachii. No 2a: Order: Carcharhiniformes - Family: Triakidae. Bulletin de L'Institut Royal des Sciences Naturelles de Belgique, Biologie, 58:99-126.

    Terms generally not employed within text, but included in glossary. File: Tooth row from symphysis toward end of jaw (longitudinally). Row: Tooth row from inner face to outer face of jaw (transversally). This series of publications started in 1987 and has maintained these definitions through the 1996 installment.

  • Compagno L. J. V. 1988. Sharks of the Order Carcharhiniformes. Princeton University Press, 486 pp.

    Author relied on Applegate 1965 and 1967 and Springer 1966 and used series for mesio-distal direction and row for labio-lingual direction. Author's "row" is a transverse row and "series" is a row according to our proposed terminology.

  • Welton, B. J. and R. F. Farish 1993. The Collector's Guide to Fossil Sharks and Rays from the Cretaceous of Texas. Before Time, Texas. 204 pp.

    Followed Applegate 1965 and Compagno 1988. Authors' "row" is a transverse row and "series" is a row according to our proposed terminology.

  • Kent, B. W. 1994. Fossil Sharks of the Chesapeake Region. Egan Rees & Boyer, Maryland. 146 pp
    Glossary defines: Row - mesio-distal line of teeth, consisting of one tooth from each tooth file or position. File - labio-lingual line of teeth, consisting of functional tooth and all replacement teeth for a single tooth position.

  • Hubbell, G. S. 1996. Using tooth structure to determine the evolutionary history of the white shark. In The Biology of the White Shark, Carcharodon carcharias . A. P. Klimley & D. G. Ainley eds. Academic Press, San Diego CA, USA. pp. 9-18.

    Used row for a mesio-distal set of teeth in illustrations.

  • Long, D. G. and B. M. Waggoner 1996. Evolutionary Relationships of the White Shark: A Phylogeny of Lamniform Sharks Based on Dental Morphology. In The Biology of the White Shark, Carcharodon carcharias. A. P. Klimley & D. G. Ainley eds. Academic Press, San Diego CA, USA. pp. 37-47.

    Used series for a mesio-distal set of teeth in illustrations.

  • Gottfried, M. D., L. J. V. Compagno and S. C. Bowman 1996. Size and Skeletal Anatomy of the Giant "Megatooth" Shark Carcharodon megalodon. In The Biology of the White Shark, Carcharodon carcharias . A. P. Klimley & D. G. Ainley eds. Academic Press, San Diego CA, USA. pp. 55-66.

    Page 58 reference "... in the lower anterior and lateral rows .." (labio-lingual)

  • Shimada, K. 1997. Dentition of the Late Cretaceous Lamniform Shark, Cretoxyrhina mantelli, from the Niobrara Chalk of Kansas. Journal of Vertebrate Paleontology, 17(2): 269-279.

    Shimada acknowledges that he followed the terminology of Applegate (1965). Row was generally used for a labio-lingual line of teeth, but occasionally, these were referred to as tooth family or file. Teeth running mesio-distally were referred to as a dental series

  • Purdy, R. W. 1998. (personal communication to HFM)

    "I think I must have mis-communicated about our usage of row and file. As you pointed out row is mesial-distal and file is labio-lingual. This is Shelly's current usage also. Thanks for your efforts to straighten this out."

  • Webster Dictionary. Definitions according to Merriam Webster's Collegiate Dictionary, tenth edition, 1994:

    Row n 1 a : a number of objects arranged in a straight line 4 a : a continuous strip usually running horizontally or parallel to a base line b : horizontal arrangement of items -- in a row: one after another : successively

    File n 1:a row of persons, animals, or things arranged one behind the other.

    Series n 1 a : a number of things or events of the same class coming one after another in spatial or temporal succession.

  • Intuitive understanding.

    Row. Two familiar usages of "row" are for theater seating and spreadsheets (or ledger pads). In each case, the row is a left to right assemblage of people/data in a two dimensional environment. [A row in front to back direction is a file (think file cabinet) or up and down direction is a column].

    File. Other than the hand tool, file is usually associated with some item being filed (placed in order) or items/people standing/moving single-file -- a front to back image.

    Series. A group of items or events (World Series), directional connotation associated with time -- before and after.

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