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A standardized method of describing fossils,

using Echinoidea as an example

D.N. Lewis & S.K. Donovan

Lewis, D.N. & Donovan, S.K. A standardized method o describing ossils, using Echinoidea as an ex-ample. Scripta Geologica, 134: 109-118, 1 table, Leiden, March 2007.David N. Lewis, Department o Palaeontology, The Natural History Museum, Cromwell Road, London,SW7 5BD, England ([email protected]); Stephen K. Donovan, Department o Geology, Nationaal Natuur-historisch Museum, Postbus 9517, NL-2300 RA Leiden, The Netherlands ([email protected]).

Key words – systematics, description, Echinoidea.The main purpose o a research paper in palaeontology, or any scientifc discipline, is to spread inorma-

tion within the feld. In this paper, suggestions are made concerning the structure and organisation osystematic palaeontological papers, using the Echinoidea as an example. A comprehensive table o echi-noid morphological eatures is presented. It is intended that authors ollowing these guidelines will beacilitated in writing research papers that enable expeditious retrieval o inormation.

Contents

Introduction ............................................................................................................................................................ 109Methodology (1): Supporting inormation .......................................................................................... 110Methodology (2): The description proper ............................................................................................ 115

Acknowledgements ........................................................................................................................................... 117Reerences ................................................................................................................................................................ 117

Introduction

The purpose o this communication is to demystiy the description o ossils, usingthe echinoids as an example, and to explain some o the steps involved on the path topublication. Although other papers with broadly similar aims have been publishedwidely (e.g., Riedel, 1978; van der Hammen, 1986), our experiences show that some,perhaps many, authors would still beneft rom more instruction in this, one o the basic

methodologies o our science. In part, it should be regarded as a supplement to the ‘In-structions to Authors’ o Scripta Geologica, although we trust that it will be o wider useor the authors o systematic papers in neontology and palaeontology.

Geology is a descriptive science. Description - and curation o the material - startsin the feld with the ubiquitous feld notebook (Tucker, 1982, pp. 12-13). This is used torecord diverse details o, or example, the surace geomorphology o an area - a hillside,

beach, quarry or whatever - as well as the solid geology. Measured sections throughsedimentary sequences, orientation data pertaining to structure or palaeocurrent direc-tions would be included, together with frst impressions and interpretations o speci-mens ound, collected and saely wrapped up or transport. All these data help with the

analysis o the specimens and interpretation o what they mean.Description continues in the laboratory, whether it be a research acility in a museumor university, or the corner o a room at home where you keep your rock collection.



110 Lewis & Donovan. A standardized method o describing ossils. Scripta Geol., 134 (2007)

More detailed observations can be made using a stronger lens than you had in the feldor, i available, a microscope. Those specimens that are signifcant, unusual and uniquedemand our special attention, and are worthy o detailed description(s). The process odescription itsel gives a rigour to our methodology o observation, and can lead, ol-lowing urther research and perhaps the hunt or additional specimens, to publicationin the scientifc literature.

The main purpose o a scientifc paper is to spread inormation to others workingin that feld, in this case palaeontology. In order to achieve this, the inormation has to

be easily accessible and in an intelligible orm, or we write both or those who speakour own language, and or those who do not and have to rely on translations using adictionary. I the task can be made simpler by a standard presentation, inormationdissemination and retrieval will be much easier, and rapid comparisons can be made

between taxa.

It is also important or curators in museums to be able to extract inormation romgeological and palaeontological publications, in order to bring their databases (bothelectronic and paper) o specimens up-to-date. Having to search undierentiated, some-times turgid, descriptions looking or such elementary data as a locality or a geologicalhorizon is irritating and time-consuming. It is, thereore, desirable to establish a stand-ard layout or this kind o inormation. There is also a need to develop a standard lay-out ‘ pro orma’ checklist or each group o taxa, in which all possible characters are listedin a regular order and each character is dealt with in turn (Table 1).

Methodology (1): Supporting information

Although this paper is written as a general introduction, it is nevertheless helpul toillustrate some o the discussion below by reerence to particular examples. Althoughwe ocus on the echinoids (Table 1), they are only illustrative; the recommended meth-odology is applicable to other palaeontological taxa. First and oremost, taxonomicpublications concerning animals or plants must conorm to the laws laid down in theInternational Code o Zoological Nomenclature (1999) or Greuter et al. (2000), respec-tively. A new taxon has to have a detailed, accurate description and (at least) adequatefgures. A proper sequence in the description o new taxa would include the ollowingsections, with related sub-headings.

Taxonomy – Each frst description o a taxon in a published work has the taxonomygiven in ull, with subsequent related taxa requiring only the parts that dier. Thissaves space in the publication, something which journal publishers, editors and readersappreciate. For example (ollowing the classifcation o Smith & Wright, 1989):-

Class Echinoidea Leske, 1778

Order Cidaroida Claus, 1880

Family Cidaridae Gray, 1825

Subfamily Psychocidarinae Ikeda, 1935

Genus Tylocidaris Pomel, 1883Subgenus Tylocidaris Pomel, 1883

[Species] Tylocidaris (Tylocidaris) clavigera (Mantell, 1822)



Lewis & Donovan. A standardized method o describing ossils. Scripta Geol., 134 (2007) 111

The author is included with the date o the frst description, ensuring that readersare aware o the taxonomic priorities. This also helps with the identifcation o reer-ences in the bibliography. The frst citation o any species in the text should be associ-ated with the name o the author(s) and date o publication, which are treated as reer-ences and will appear in the reerence list (Kelt & Palma, 1992). Within the body o thetext the genus (and, i so subdivided, subgenus) can be reduced to the capital letter T. only - T. (T.) clavigera - except at the start o a sentence. The author can at this stage beomitted. The next taxon within the cidarids might then be:-

Subfamily Cidarinae Gray, 1825

Genus Prionocidaris A. Agassiz, 1863

etc.

This avoids repeating the higher taxonomy (class, order, amily), which is the sameor both genera. Bengtson (1988) provided much sensible advice regarding the use andormat o open nomenclature.

Synonymy list – This is a list that includes all previously published identifcationsand reerences to a given taxon. For example, the Chalk echinoid Gauthieria radiata (So-rignet, 1850) has been previously called:-

Cyphosoma radiatum Sorignet, 1850 [this was the original designation]Cyphosoma subradiatum Sorignet, 1850 [originally perceived as a separate species,

but subsequently synonymised]Phymosoma heberti Desor, 1858 [Desor thought dierently]

and several other designations until 1888 when Lambert erected the genus Gauthie-ria, with Gauthieria radiata (Sorignet, 1850) designated as the type species.

Synonymy lists can be exhaustive or selective; in the latter case, only the most no-table reerences are listed. There are taxa whose complete synonymy lists would re-quire a whole paper in themselves, such as the ossil bird Archaeopteryx. The preerredormat o a synonymy list or Scripta Geologica should ollow the style used by Taylor

& McKinney (2006). For detailed comments on the construction o synonymy lists, seeMatthews (1973).

Diagnosis – The diagnosis o a taxon is expressed in concise sentences which defnethe essential eatures o that taxon alone, using characters that are appropriate to theparticular taxonomic level. No two species can have identical diagnoses. The diagnosisis a kind o shorthand whose purpose is to present all the salient characters or the tax-on under consideration, whether species, genus or higher group. The amount o detailin the diagnosis will depend on the relevant characters present.

Etymology (derivatio nominis) – This gives the background to the name being given tothe new taxon. This is useul to know, apart rom the obvious interest, because it willhelp with the pronunciation o the taxonomic name, especially i given in honour or




Table 1. An example o a checklist o morphological eatures that should be included in a description oa member o a particular group o ossils, in this case an echinoid. Although not all eatures may bepresent or preserved, noting the absence o a eature may be as important as describing those structuresthat are apparent. For explanations o morphological terms, see Melville & Durham (1966) and Smith

(1984, 2003). Note that, under sections (1, 2, 15, 16, 17, 19), contrasting characters such as ‘tall/low’,‘large/medium/small’, ‘wide/narrow’, ‘large/small’, ‘long/short’ and ‘slender/at’ appear; in all othese examples, some sort o quantifcation is required o the author o a taxon.

1. Test regular

Periproct within apicalsystem

centraloset rom centre

2. Test irregular

Periproct not within apicalsystem

apicalmarginalsupramarginalinramarginaloralposterior end

3. Shape

oblate spherical (sub-spherical)globularplano-convexconical

attened oro-apicallyoral surace depressedoral surace inatedoral surace planetalllowelongated (axis III-5)quadrateoutline

circularangular

heart-shapedrostratetruncate

4. Size

largemediumsmallsexual dimorphism (yes/no)

5. Apical Disc

circularelongated

disjunctposition

central

ad-anteriorad-posterior

monocyclicdicyclichemiolicyclic

6. Ocular plates (I-V)insertexsert

7. Genital plates (1-5)fve presentone plate/gonoporepresenttetrabasal (1-4)monobasal (1-5)madreporite (2)

largesmallethmolyticethmophract

5 genital pores

4 genital pores (1-4)8. Suranal plate

present and tesselated intodiscnot tesselated into disc

9. Peristome

shapecircularovalD-shapedasymmetric

positioncentralanterior

perignathic girdleauricles/apophyses/mixedbuccal clets (“gill slits” or buccal notches)no buccal clets

10. Lantern

absent in adultspresent in adultsdemipyramids

tops pittedtops unpitted

epiphyses

rotulaeteeth

groovedkeeled

compassescidaroidaulodontstirodont

camarodont11. Coronal sculpturing

sculpturedunsculptured

12. Coronal plate imbrication

imbricatedtesselatedsimple butt-jointsutural peggingexiblerigid

13. Ambulacra

non-petaloidsub-petaloidpetaloid

openclosed

depressed/sunkenraised/convexwide c. interambulcranarrow c. interambulcratwo columnsmore than two columns

(pluriserial)14. Ambulacral plates

simplepseudo-compoundcompound

diadematoidarbacioidacrosalenioidechinoidechinothurioid

reduced platesdemi-plates

occluded platesincluded plates

pore pairs




commemoration o a person, place or thing. Long species names can be immenselydaunting to newcomers, who can better understand how to pronounce the words i

they understand the origins and how the name is constructed. An invaluable reerencewhen choosing a trivial name or a new species is Brown (1956).

Material – All o the specimens used in writing the description are listed, includingthe holotype or other primary types (see below), paratypes and other material. Undereach category the ollowing should be included:-

• details o the museum(s) or other repository where the specimens are kept;• registration and/or accession number(s) o the specimen(s); and• a brie description o the preservation o the specimen(s).

The name o the collection, collector and any other such details can be included, iappropriate. Type specimens are needed to defne the species and may belong to a type

straightobliqueconjugatenon-conjugate

pore columnssimple (uniserial)bigeminate (biserial)trigeminate (triserial)polyporous

phyllodes(phyllodes + bourrelets= oscelles)

spheridial pitsoralaboralperradial

adradialone pair2-4 pairsmultiple

15. Interambulacra

widenarrow2 columnsmore than 2 columns(= pluriserial)labral plate

sternumprotosternousmeridosternousamphisternous

bourrelets(bourrelets + phyllodes= oscelles)

16. Primary Tubercles

largesmalltubercles sunken

tubercles ushregular seriesalternating seriesperoratenon-perorateparapet crenulateparapet non-crenulateneck straightneck undercutplatorm

ushimpressed (with

parapet)scrobiculebasal terrace

17. Secondary tubercles

peroratenon-peroratelargesmallscrobicular ring

contiguousconuent

completenon-conuenttangentialnon-conuentseparated

18. Other tubercles, etc.

aboral naked zone

miliariesascioles

internalperipetalous

laterallatero-analsubanalanal

granules19. Appendages

Spines (=radioles)

primary or secondary?longshortslenderat

straightcurvedclavate (clubbed)spinulessmoothornamentedsolidhollowverticillatenon-verticillate

cortex

no cortex Pedicellariae

globieroustridentateophicephaloustriphyllous

Clavulae

Spheridia




series, a group o specimens used in the description o the species. This series can beconveniently divided into the primary type(s) (see below) and the paratypes. Techni-cally, the primary type is the only specimen which can be called by the species name; allother such named specimens are merely compared with them. Assuming there has beenno misidentifcation o the type specimen(s) (taxonomic and other complications arisewhen this happens), its species name never changes. Conversely, paratypes and non-type specimens which have been identifed as one taxon can be re-named i they arere-identifed as something else. Indeed, they may even become the basis or new spe-cies. The various orms o primary types are explained below (Davies, 1961; Interna-tional Code o Zoological Nomenclature, 1999; Greuter et al., 2000).

• The holotype is the single reerence specimen or a named species. When severalspecimens are used or a description, those apart rom the holotype may be named

paratypes. These are named as such by the author, who may also reer to any ur-ther specimens used as ‘other material,’ i appropriate. In earlier literature, para-types may not have been designated as such and hence all other specimens used insuch descriptions are regarded as paratypes.

• A syntype has the same status as a holotype. A syntype(s) is one or one o severalspecimens o a type series or which no holotype was designated. All the specimensused in that original description are syntypes.

• A lectotype is a specimen which is selected rom syntypes, or rom a paratype seriesin which the holotype was not defned or has been lost. It is the specimen which bestrepresents the original concept o the species and has the same status as a holotype.

The remaining syntypes (i any) are then reerred to as paralectotypes.• A neotype (‘new type’) is a specimen which is selected when the holotype is lost orunidentifed within a collection and there are no recognised syntypes or paratypes.The neotype would be chosen rom specimens which best represent the originalconcept o the species and has the same status as a holotype. Ideally it is chosenrom specimens which came rom the same locality and horizon as the original type.This may not always be possible, however, especially i the original locality was notknown or is no longer accessible.

The term ‘type species’ (not to be conused with type specimen) is applied to the

species which characterises a genus. For example, the type species o Hyposalenia isSalenia stellulata L. Agassiz, 1838, with several other species assigned to the genus Hy- posalenia. The type species o Hyposalenia will never change, but the other species as-signed to that genus can be placed in other genera i revisions occur.

Localities – The locality details o the type and other specimens are given so thatthe reader can, ideally, exactly identiy the source site o the material. Do not assumethat the reader will know where the locality is - some may even be uncertain o thecountry - so state the obvious, or example, “Barton-on-Sea, Hampshire, England” (asdistinct rom, or example, “Barton, Devon, England”). Locality maps show rom

where the material comes. It is a good idea to include not only a detailed map o alocality, but also a much larger scale map to show where it is in relation to elsewhere,such as the country o origin.




A locality is made up o several dierent components which can be broken downinto smaller units, especially useul i the inormation is to be added to a database. Thereason or doing this is or retrieval in a variety o orms (or example, a card index),and is particularly important or more advanced nested and other relational electronicdatabases. Conventionally, the inormation is given in order o increasing rank, closingwith the country o origin. The latitude and longitude can be given when known, to-gether with, or as an alternative to, the local national grid reerence. This might be ap-propriate i copyright inormation or the grid reerence is likely to be inringed orpermission is difcult to obtain. The use o GPS inormation has greatly simplifed de-termination o latitude and longitude.

Stratigraphy – As with locality details, in order that readers unamiliar with the geo-logical data being presented can fnd their way around the stratigraphy, do not assumethat they are amiliar with the stratigraphy; once again, state what to you may appearto be obvious. The stratigraphy is given in descending rank order, and breaks down theessential inormation into three main areas and progressively smaller units:-

• Chronostratigraphy deals with time alone - period, age, stage, etc.; or example, Cre-taceous; Coniacian. It simply tells the reader how old the material is. There is noimplication o what ossils are present, nor the rock types in which they were ound.This is important inormation or readers who do not know the minutiae o thestratigraphy involved.

• Biostratigraphy deals with the details o where exactly in the geological column the

ossils occur. Here are included the ormal biozones and subzones which locate theossils within their time bands, or example, Micraster cortestudinarium Biozone (or just Zone) (note that ‘Biozone’ and ‘Zone’ take a capital letter). Again, there is noimplication o any rock types.

• Lithostratigraphy deals with the rocks themselves. Rock masses are divided intomore manageable units in descending order - group, ormation, member, bed, etc.The main groups have ormal names, but the smallest units may only be known bytheir local or inormal names.

Rawson et al. (2002) provided comprehensive advice on stratigraphic usage. The

latest in the Cambridge series o geologic time scales was recently published (Gradsteinet al., 2004). Shaw (1971) provided a succinct critique o the poor use o stratigraphicdata in palaeontology.

Methodology (2): The description proper

Gross morphology: size and shape – This is described beore the ormal description toacquaint the reader with an idea o how big the organism might be. Overall measure-ments can be given here although they are not obligatory, but details o componentparts are preerably let to the relevant section. Indeed, illustration(s) at some easily

recognisable scale (or example, × 1, × 5, × 100), in conjunction with these details inthe text, will do more to convey the gross morphology at this stage than too manymeasurements.




Morphological description o the taxon – The component parts o a taxon are describedunder easily located sub-headings, ollowing a regular and consistent order, dependenton the group being described. A checklist is developed or each main group o structuresthat may be met in that group o organisms (Table 1). This not only ensures that thewriter can describe the taxon quickly and know that all the relevant inormation has

been given, but also that the reader can compare individual taxa quickly and easily. Sucha checklist will aid observers in realising what is absent as much as what is present.

The essence o a good description is to keep it clear and simple. Remember thatsome readers have to translate using a dictionary. Technical terms may be unavoidableor preerable because they act as scientifc ‘shorthand;’ or example, “adambitally”meaning “towards the ambitus” or the widest part o an echinoid test. When simpleralternatives are available they should be used; or example, rather than write ‘cothurni-orm,’ i a ossil is ‘boot-shaped,’ then say so.

Plates, fgures and tables – An old adage says that a good photograph is worth a thou-sand words. The purpose o good illustrations in a systematic description is to acquaintthe reader with what the taxon actually looks like, and helps to bring to the attention othe reader all the relevant characters o the species. This can be achieved with photo-graphs and artwork. Specimens are photographed with the lighting conventionally bi-assed rom the top let hand side (colloquially know as ‘top let lighting’). Hand-drawnartwork is useul to emphasise a eature not necessarily obvious in a photograph. Stand-ard views o the taxon are needed, or example, top, bottom, side, ront, back and vari-ous close-ups; all should be included whenever possible.

Whether the illustrations are published as plates or text fgures, or both, depends onthe journal style. Whichever style is used, adequate explanations o the fgures areneeded, and should at least include the name o the image, its view, its museum or in-stitution reerence number, i appropriate, and a scale. More detailed inormation canalso be included (or example, the locality and geological inormation), but that mayalso depend on journal policy. It is a general rule that a good fgure can only be urtherenhanced by an inormative caption.

The scale can be expressed either as a magnifcation o the original specimen (orexample, x 5), or as a scale-bar o given length. Magnifcation can be an unreliable ex-pression, because a journal article may be photocopied to a dierent size or the publi-

cation’s page may appear on a computer screen at another size to the original; in bothcases a magnifcation given as, say, ‘× 5’ may be very inaccurate. Scale bars are arguablypreerable because the reader can determine the size o a eature by direct observationand comparison with the scale bar. Thus, i a bar represents 10 mm, it is simple enoughto subdivide the bar into smaller millimetre units and use it to judge the size o a ea-ture. This is not as easy to do ‘by eye’ with a magnifcation.

Tables present data about specimens. They can be graphs, lists o measurements orother data that can be expressed in tabular orm (e.g., Table 1).

Comparisons and discussion/remarks – Comparisons between similar taxa are made

to justiy the decisions made by the author. For example, in describing a new species,an author should explain the unique eatures that dierentiate it rom closely relatedand other taxa within the same major group that occur in the same bed or suite o beds.




Species may be compared on the basis o a number o other criteria, as considered ap-propriate by the author.

The discussion deals with other items relevant to the taxonomic descriptions andcomparisons. Here there may be inormation, or example, about the possible phylo-genetic relationships or why a character should be seen as important. Stratigraphicposition and/or geographic distribution may be discussed, with a range extended orreduced. Similarities and dierences between species rom dierent geographic loca-tions may throw light on the evolution o the taxon. There are many points which can

be talked about here, depending on the bias o the paper, whether it be primarilytaxonomic or stratigraphic.

Conclusions – This is the fnal section o the science o the paper and summarises thecontent as well as drawing together conclusions, i any, rom the discussion. Thus, the

main conclusion o this paper is that, i authors ollow the suggestions given herein, thestructure and layout o their papers will be clear, and organised so that inormation can be retrieved quickly and easily.

Acknowledgements – This pays tribute to any help the author has received rom any-one during the progress o the work, including sources o fnancial support.

Bibliography/Reerences – This gives a complete listing o all the reerences used bythe author in the paper. All reerences that appear in the text must also be included inthe reerence list and vice versa. The journal style dictates how these are expressed. For

example, some journals require the title o a reerenced publication to be given in ull,whereas others ask or an abbreviation to be used, or example, Proceedings o the RoyalSociety o London is abbreviated Proc. R. Soc.

And fnally... – Journals produce guidelines as to how an author needs to set out thepaper. This is usually known as ‘Instructions to Authors’ and can be ound printed onparts o the cover pages o the journal, as a separate guide or on a website. Authors arewell-advised to ollow these, because re-ormatting manuscripts is time consuming,and a poor initial ormat could result in rejection o the paper.

Acknowledgements

We thank the very many colleagues who have, over a combined period approaching70 years, discussed the many concepts that we have tried to elucidate in this contribu-tion. Many thanks to our reviewers, Drs F.H.C. Hotchkiss and D.L. Pawson, or theirconstructive comments.

References

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Paleontology, Part U, Echinodermata 3(1): U220-U251. Geological Society o America & University oKansas Press, New York & Lawrence.

Pomel, A. 1883. Classifcation Méthodique et Genera des Échinides Vivants et Fossiles. Adolphe Jourdan, Alger:131 pp. [Not seen.]

Rawson, P.F. & sixteen others. 2002. Stratigraphic Procedure. Geological Society, London: vi+57 pp.Riedel, W.R. 1978. Systems o morphologic descriptors in paleontology. Journal o Paleontology, 52: 1-7.Shaw, A.B. 1971. The butterfngered handmaiden. Journal o Palaeontology, 45: 1-5.Smith, A.B. 1984. Echinoid Palaeobiology. George, Allen & Unwin, London: 190 pp.

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