Annales Societatis Geologorum Poloniae (2015), vol. 85: 529–549. doi: http://dx.doi.org/10.14241/asgp.2015.032

CONSTRUC TION OF ICHNOGENERIC NAMES

An drew K. RINDSBERG

De part ment of Bi o log i cal & En vi ron men tal Sci ences, Sta tion 7, Uni ver sity of West Al a bama, Livingston, AL 35470; e-mail [email protected]

Rindsberg, A. K., 2015. Con struc tion of ichnogeneric names. Annales Societatis Geologorum Poloniae, 85: 529–549.

Ab stract: Ichnologists have overused the root ichn- “trace”, em ploying it in new terms and new ichnogenera alike, to the point where it can be dif fi cult to express one self clearly with out using it sev eral times in one sen tence. The root de rives from Ancient Greek ÇP<@l (ichnos), which means “foot print” or “track”, or by ex ten sion a “trace”, any sign of an an i mal’s activ ity. Perhaps it is time to explore the use of other roots to create new ichnologic terms and gen era. Al ter na tive Latin and Greek roots are given here, as well as advice on how to construct new ichnogenera in a techni cally cor rect and aes theti cally pleas ing manner.

Key words: Ichnology, trace fossils, ichnotaxonomy, ter minol ogy.

Manuscript received 16 June 2015, ac cepted, 7 Sep tem ber 2015

IN TRO DUC TION

Robert W. Frey used to remark that the phrase “ichno- ichnology (ichnologie comparée, Lessertisseur, 1952), ge nus Teichichnus” sounded awful, but there was nothing to coprolichnia (Macsotay, 1967, p. 10), cursichnion, mor- be done about it. Ichnologists have hundreds of uses for the dichnion, natichnion, volichnion (Müller, 1962), ichnidion Greek word ÇP<@l (-@LH, J`), which means “foot print” or or endichnidion (Martinsson, 1970, p. 328), ichnite (Hitch- “track”, or by exten sion a “trace”, any sign of an ani mal’ s cock, 1837, p. 175), ichnoabundance (Knaust et al., 2014, p. pass ing. Seilacher, in par tic u lar, has been re spon si ble for 2232); ichnodisparity (Buatois and Mángano, 2013), coin ing new terms in cor po rat ing ÇP<@l. Knaust (2012, p. ichnodiversity (Lockley and Gillette, 1987), ichnoguild 95) deter mined that about 44% of all valid inver te brate (Bromley, 1990, p. 211), ichnofamily (Seilacher, 1977, p. 296; ichnogenera end in ichnus. Accord ingly, this paper suggests Rindsberg, 1990, p. 61), ichnoflora (Frey, 1973, ta ble 2), al ter na tive roots that are ap pro pri ate for con struct ing new ichnolite (Hitchcock in R., 1838, p. 201), ichnolithology ichnogeneric names. (Hitchcock, 1841, p. 770), ichnospectrum (Ichnospektrum, Many of the most com monly used ichnologic terms in- Seilacher, 1964, p. 697), ichnostratigraphy (Baldwin, 1977, clude the stem ÇP<-. These include ichnocoenose or ichno- fig. 6), ichnotaxonomy (Bromley and Fürsich, 1980; Ekdale coenosis (Davitashvili, 1945; Lessertisseur, 1956, p. 10), et al., 1984, p. 17), ichnotope (Radwañski and Roniewicz, ichnofabric (Bromley and Ekdale, 1982), ichnofacies 1971, p. 53), lithichnozoa (Hitchcock, 1858, p. 1), ornithi- (Ichnofazies, Seilacher, 1963), ichnofauna (Frey, 1973, ta - chnites, ornithichnology (Hitchcock, 1836), palaeoichno- ble 2), ichnofossil (Seilacher, 1956, p. 158), ichnogenus logy (Paläoichnologie, Abel, 1935, p. 33) or palichnology (Seilacher, 1953, p. 440), ichnologist, ichnology (Buckland, (Palichnologie, Seilacher, 1953, p. 423); and the many etho- about 1830, vide Häntzschel, 1975, p. W2), ichnospecies log i cal cat e go ries re viewed by Vallon et al. (2015). I cite the (Ichnospezies, Seilacher, 1953, p. 440), ichnotaxon, neo- ear liest uses of the terms of which I am currently aware. ichnology (Neoichnologie, Seilacher, 1953, p. 473); cubich- Some of the lat ter terms are des tined to become part of the nion, domichnion, fodinichnion, pascichnion, repichnion reg u lar vo cab u lary of sci ence; some are ob so lete; some have (Seilacher, 1953, pp. 432–434); fugichnion (Frey, 1973); been used only once. agrichnion (Ekdale et al., 1984); endichnion, epichnion, The greatest use for ÇP<@l is in a large num ber of ge- exichnion, hypichnion (Martinsson, 1965). Where possi ble, neric names for fos sil traces such as Aulichnites, Cylin- I have given the cita tion for the first use of the term. All of drichnus, Diplichnites, Ichnium, Ichnocumulus, Ichnyspica, these words are dysphonious in English, in which the simi - Isopodichnus, Permichnium, Phoebichnus, and Teichichnus lar word icky means “sticky, dis taste ful”, but they are part of (Ta ble 1). More are named ev ery year. Häntzschel (1975, p. the reg u lar vo cab u lary of ichnology. W24) traced the or igin of this custom to Jardine (1853), In addi tion, there is a host of terms in less com mon use: who “proposed that the ending -ichnus be added to the ge- anichnial (Dörjes and Hertweck, 1975, p. 479), compar a tive neric names of ver tebrate trails from Scotland so that it 530 A. K. RINDSBERG

Ta ble 1 cles, but used ÇP<@l when he named Trichichnus Frey, 1971 and Schaubcylindrichnus Frey and Howard, 1991. Bromley Ichnogeneric names and other terms exhib it ing variants also con sis tently ex pressed a pref er ence for such names, de rived ul ti mately from ichnos, in clud ing its di min u tive e.g., Oichnus Bromley, 1981, Phoebichnus Bromley and ichnion (“lit tle track” or “lit tle trace”). Not all of these Asgaard, 1972. The practice has been very wide spread in vari ants are proper; some are de lib er ate misspell ings in verte brate and inver te brate ichnology; nearly half of all substi tute names for older, unavail able ichnogenera valid inver te brate ichnogenera have names end ing in -ichnus (Knaust, 2012). Variant Ichnogenus or other term Meaning Many varia tions on ÇP<@l have been employed as well of ichnos (Ta ble 1). The Ancient Greek can be transcribed in Latin Lanicoidichna Chamberlain, 1971 Lanice-like trace -ichna charac ters as ichnos or latinized as ichnus. It may be femi- Undichna Anderson, 1976 wave trace nized as ichna or neutered as ichnum, prac tices that are not ichni- Ichniotherium Pohlig, 1892 little track beast strictly cor rect grammat i cally, but which serve to vary the -ichnia Pilichnia Chamberlain, 1971 little pillar trace language. A di minu tive form may be used (ichnion or ich- -ichnion domichnion Seilacher, 1953 little dwelling trace nium, “lit tle trace”). And the root may be used to begin a Mammillichnis Chamberlain, 1971 breast trace -ichnis word instead of ending it (ichn-). Circulichnis Vyalov, 1971 circle trace The root ÇP<@l has been valu able to ichnologists in the Climactichnites Logan, 1860 ladder trace -ichnites past, but is now perhaps overused. Hitchcock him self was ichnite Hitchcock, 1837 ancient trace not so con strained as to use a single root in every new name. little trace Ichnium Pabst, 1896 The purpose of this arti cle is to suggest alter native Latin and -ichnium light (not heavy) little Kouphichnium Nopcsa, 1923 trace Greek roots for construct ing new generic names of trace fossils. But first, the Indo-Euro pean ori gin of ÇP<@l is ex - ichno- Ichnocumulus Seilacher, 1956 trace heap plored and its cognates are identi fied. Acanthichnus Hitchcock, 1858 thorn trace Chomatichnus Donaldson & mound trace Simpson, 1962 double flute trace Didymaulichnus Young, 1972 six-foot trace or hexapod ET Y MOL OGY OF }3O;?G Hexapodichnus Hitchcock, 1858 trace Imbrichnus Hallam, 1970 shingle trace Isopodichnus Bornemann, 1889 isopod trace The root most com monly used by ichnologists is the Ixalichnus Callison, 1970 leaping trace latinized Greek ichn-, deriv ing from Ancient Greek ÇP<@l Lobichnus Kemper, 1968 lobe trace (ichnos), “trace, track, footprint” . This word is of obscure -ichnus Margaritichnus Bandel, 1973 rosary trace ori gin, but may be related to @ÇPbird foot prints”); in fers chiefly to the wasp, and sev eral spe cialized words have 1837 he named sev eral new ichnogenera ending in -ichnites. derived from this usage for use in en tomol ogy. Both of In later works, Hitchcock (1858, 1865) used -ichnus as the these words entered the English language in the sixteenth to root for a few verte brate trackways (Apatichnus, Arachich- eigh teenth cen tu ries, pre dat ing the de vel op ment of ichno- nus, Toxichnus) and in ver tebrate traces (Acanthichnus, logy in the early nineteenth cen tury (Brown, 1993, 1: 1301). Cochlichnus, Hexapodichnus), but overall his names were constructed us ing many differ ent roots. Others used the form gen era Ichnium or Ichnites to in clude trace fos sils (Hänt- GOOD AND BAD WAYS TO NAME zschel, 1975, p. W24). The recom menda tion to end new ICHNOGENERA ichnogenera in -ichnus was echoed by ichnologic theo rists Seilacher (1953, p. 446) and Häntzschel (1962, p. W182; A good name is short, easy to remem ber, and easy to 1975, p. W24). Frey did not endorse it directly in his arti - pronounce. The best names are fresh, distinc tive, and mean- CON STRUC TION OF ICHNOGENERIC NAMES 531 ingful, de scrip tive of some diag nos tic as pect of the fossil. Ta ble 2 However, it is not desir able for all names to be formed in the same way; for names to be mne monic, they must be var ied. Some of the many ichnogeneric names that are based on This arti cle focuses on the mechan ics of word construc - morphol ogy of the trace fossils tion and is not intended as a complete guide to naming new ichnogenera, an outline of which was given by Rindsberg Ichnogenus Meaninh (2012). Ulti mately, new names are governed by the In ter na- Alectorurus Schimper, 1869 rooster's tail Amphorichnus Myannil, 1966 amphora trace tional Code of Zoo log i cal No men cla ture (In ter na tional Arthraria Billings, 1873 joint “group” (-aria Com mis sion on Zoo log i cal No men cla ture, 1999), which denotes groups) contains some guide lines for the construc tion of valid ge- Asterosoma Otto, 1854 star body neric names. In gen eral, the name must be in the Latin al- Baroccoichnites Vyalov, 1971 ancestral baroque trace phabet, which may include j, k, w, or y (Art. 11.2), but not Belorhaphe Fuchs, 1895 arrow seam Bifasciculus Volk, 1960 two little bundles dia critic marks or hyphens (Art. 27). It must be “used as a Bifungites Desio, 1940 two mushroom-like word” (i.e., be pronounce able; Art. 11.3). Although classi - Bifurculapes Hitchcock, 1858 two-fork foot cal Greek and Latin roots are pre ferred, and the Code rec- Calycraterion Karaszewski, 1971 calyx bowl ommends that they be latinized, words from any language Circulichnis Vyalov, 1971 circle trace Cochlichnus Hitchcock, 1858 spiral trace may serve, or indeed from no language (Art. 11.3). The name Conispiron Vyalov, 1969 cone spiral must not be a homonym, i.e, one that has been previ ously Conostichus Lesquereux, 1876 cone row used under the ICZN (Art. 52). The guidelines re garding cor- Curvolithus Fritsch, 1908 bent stone rect and incor rect spellings are complex (Arts. 19, 32, 33). Dendrina Quenstedt, 1848 tree-like Dimorphichnus Seilacher, 1955 two-shape trace Most authors continue to prefer to base new names on Granularia Pomel, 1849 granular one Greek and Latin roots. Construct ing a name correctly re- Gyrochorte Heer, 1865 turning hay quires more knowl edge than sim ply consult ing a dictio nary; Helminthopsis Heer, 1877 worm-like Bo tan i cal Latin (Stearns, 1966), and Com po si tion of Sci en - Laevicyclus Quenstedt, 1879 smooth circle Ophiomorpha Lundgren, 1891 snake shape tific Words (Brown, 1956) are es pecially help ful in this re- Palaeochorda M’Coy in Sedgwick, 1848 old string gard, along with the rec om menda tions on gram mar and Paleodictyon Meneghini in Murchison, 1850 old net translit er a tion in the third edition of the Code (ICZN, 1985). Pholeus Fiege, 1944 hole The clas si cal lan guages, es pe cially Latin, have con tin ued to Phymatoderma Brongniart, 1849 tumor skin Plagiogmus Roedel, 1929 slope furrow evolve in sci en tific us age, par tic u larly for the de scrip tion of Plangtichnus Miller, 1889 wandering trace or gan isms. Bo tan i cal Latin is a stan dard ized and flex i ble Polykampton Ooster, 1869 many bend(s) mode of commu ni ca tion that was honed over a period of Polykladichnus Fürsich, 1981 many branch trace cen tu ries. Linnaeus used it, and bot a nists still use it. Polyupsilon Howell, 1957 many upsilon(s) (U’s) Scalarituba Weller, 1899 ladder pipe Ichnologists naming new genera have found inspi ra tion Taonurus Fischer-Ooster, 1858 peacock tail in diverse ways (Häntzschel, 1975, pp. W24–W25; Pember - Taupezia Declair, 1962 tau (T) foot ton and Frey, 1982). Some have related the trace fossil to its Tomaculum Groom, 1902 sausage supposed maker, or to modern organ isms. Some have Trisulcus Hitchcock, 1865 three furrow Upsiloides Byrne and Branson, 1941 upsilon (U)-like named a trace fossil for its site of discov ery, forma tion, or Vexillum Rouault, 1850 flag age. Some have chosen to honour an ichnologist or other Xenohelix Mansfield, 1927 strange spiral scien tist. Some have even named ichnogenera af ter other ichnogenera. Humour has inspired others. Currently, the most approved method is to incor po rate some aspect of the trace’s morphol ogy in the name (Ta ble 2). Pem berton and these mistakes have been memo ri al ized in ichnologic taxon - Frey (1982, p. 847) disap proved of all of these methods ex- omy. The ending -ites was com monly used to denote ances - cept the morpho log i cal, but it is more impor tant for names tral forms of modern organ isms; it was used so often, and so to be varied than to be ideal. er ro ne ously, that the Zoo log i cal Code now dis cour ages end- A formerly popu lar method was to incor po rate the ing any new name in -ites or its vari ants, -ytes and -ithes (In - name of the de duced maker into the trace fos sil’s name (Ta- ter na tional Com mis sion for Zoo log i cal No men cla ture, ble 3). Even in cases where the tracemaker is found within 1999, Art. 20). Some ichnogenera can be traced back to the trace, it can not be de duced that this is the trace’s only names of modern sea weeds (Ta ble 4): Caulerpites to Cau- maker; still, the temp ta tion to draw anal o gies can be ir re sist - lerpa, Chondrites to Chondria. Zoo log i cal er rors in clude ible. Thus, Balanoglossites is supposed to have been made trace fos sils Arenicolites and Nereites, thought to be re- by the enteropneust Balanoglossus, Corophioides by the mains of the polychaete Arenicola and Nereis; Asteriacites, am phi pod Corophium, Pelecypodichnus by pelecypods, thought to be the starfish Asterias; and Pennatulites, mis- Lingulichnus by lingulids, Isopodichnus by isopods, and so taken for a sea pen (Table 3). Another Greek root, nL6@H, on. This method has often led to embarassment when later latinized as phycus or fñcus, means “sea weed” and was ev idence shows that sev eral di verse mak ers can ac count for commonly employed for fossil seaweeds or “fuc oids” (a one trace fossil, or when the proposed maker is misiden ti - fam ily of brown algae in cluding Fu cus). Sev eral ven er a ble fied (Ta ble 3). names include this root, includ ing Arthrophycus, Astero- Nineteenth-cen tury au thors fre quently mis took trace phycus, Dactylophycus, Fucoides, Fucusopsis, Palaeophy- fossils for body fossils of plants or ani mals, and many of cus, Phycodes, Phycosiphon, Rusophycus, Spirophycus, 532 A. K. RINDSBERG

Ta ble 3 Some ichnogeneric names that are modi fied from generic names of ani mals, either because the author thought this was a body fossil of the ani mal, or because the author concluded that it was a trace fos sil made by the ani mal. This is not a recom mended practice; aster isks (*) show exam ples of ichnogenera that are now thought to be grossly misnamed.

Ichnogenus Meaning Presumed maker Clionolithes Clarke, 1908 Cliona stone Porifera Clionoides Fenton & Fenton, 1932 Cliona-like *Alcyonidiopsis Massalongo, 1856 alcyoniid-like *Pennatulites Stefani, 1885 ancestral Pennatula Coelenterata *Protovirgularia M’Coy, 1850 first Virgularia *Rhizocorallium Zenker, 1836 root coral *Gordia Emmons, 1844 Gordius (hairworm) Nematomorpha *Nemertites MacLeay, 1839 ancestral Nemertes Nemertea Annelidichnium Kuhn, 1937 annelid trace Oligochaeta *Lumbricaria Münster in Goldfuss, 1831 earthworm (lumbricus) group Arenicolites Salter, 1857 ancestral Arenicola Diopatrichnus Kern, 1978 Diopatra trace Goniadichnites Matthew, 1891 ancestral Goniada trace Lanicoidichna Chamberlain, 1971 Lanice-like trace Maeandropolydora Voigt, 1965 meander Polydora Myzostomites Clarke, 1921 ancestral Myzostomum Polychaeta *Nereites MacLeay, 1839 ancestral Nereis Palaeosabella Clarke, 1921 ancient Sabella *Phyllodocites Geinitz, 1867 ancestral Phyllodoce Polydorites Douvillé, 1908 ancestral Polydora *Sabellarifex Richter, 1921 Sabellaria (reef) maker *Sabellarites Dawson, 1890 ancestral Sabellaria Helminthoida Schafhäutl, 1851 worm-like Helminthoidichnites Fitch, 1848 ancient wormlike trace Undifferentiated “worms” *Scolicia Quatrefages, 1849 worm (scolex) Gastrochaenolites Leymerie, 1842 Gastrochaena stone Martesites Vitális, 1961 ancestral Martesia Bivalvia Pelecypodichnus Seilacher, 1953 pelecypod trace Teredolites Leymerie, 1842 Teredo stone *Archaeonassa Fenton & Fenton, 1937 ancient Ilyanassa *Olivellites Fenton & Fenton, 1937 ancient Olivella Gastropoda *Palaeobullia Götzinger & Becker, 1932 ancient Bullia Aglaspidichnus Radwañski & Roniewicz, 1967 Aglaspida trace Anobichnium Linck, 1949 Anobium trace (beetle) Arthropodichnus Chiplonkar & Badwe, 1970 arthropod trace Asaphoidichnus Miller, 1880 Asaphus-like trace Caridolites Nicholson, 1873 (Ceratio)caris stone Corophioides Smith, 1893 Corophium-like (amphipod) Arthropoda Isopodichnus Bornemann, 1889 isopod trace Merostomichnites Packard, 1900 ancient merostome trace Myriapodites Matthew, 1903 ancient myriapod Oniscoidichnus Brady, 1949 Oniscus-like trace (isopod) Palaeobuprestis Häntzschel, 1962 ancient Buprestis (beetle) Thalassinoides Ehrenberg, 1944 Thalassinidea-like Balanoglossites Häntzschel, 1962 ancestral Balanoglossus Hemichordata Asteriacites Schlotheim, 1820 ancestral Asterias Asterozoa Chirotherium Kaup, 1835 hand beast Cincosaurus Aldrich, 1930 five (toed) lizard Erpetopus Moodie, 1929 reptile foot Ichniotherium Pohlig, 1892 little-track beast Ichthyoidichnites Ami, 1903 ancestral fishlike trace Vertebrata Ignotornis Mehl, 1931 unknown bird Megalosauropus Colbert & Merrilees, 1967 megalosaur foot Otozoum Hitchcock, 1847 ear animal Pulchravipes Demathieu et al., 1984 beautiful bird foot Tyrannosauropus Haubold, 1971 tyrannosaur foot CON STRUC TION OF ICHNOGENERIC NAMES 533

Ta ble 4 Ta ble 5 Ichnogeneric names with roots indi cat ing that they were Some ichnogeneric names that are based on geographic orig inally named as fos sil plants, es pe cially seaweeds. For names. This should be done sparingly. the most part, this practice ended about 1900. Not all of these ichnogenera are consid ered to be valid today. Ichnogenus Meaning Brookvalichnus Webby, 1970 Brookvale trace (Australia) Ichnogenus Meaning Capodistria Vyalov, 1964 Capodistria (now Koper, Slovenia) Gluckstadtella Savage, 1971 little Gluckstadt (South Africa) Archaeozostera Koriba & Miki, 1971 ancient Zostera (a seagrass) Jeholosauripes Yabe, Inai & Jehol (China) lizard foot Arthrophycus Hall, 1852 joint(ed) seaweed Shikama, 1940 Asterophycus Lesquereux, 1876 star seaweed Kiivermus Katto, 1976 Kii (Peninsula, Japan) worm Aulacophycus Heer, 1877 flute seaweed Oldhamia Forbes, 1849 Oldham (Ireland) Bostricophyton Squinabol, 1890 curl plant Polarichnus Narbonne, Gibling & (North) Polar trace Buthotrephis Hall, 1847 growing (in) the deep Jones, 1979 Cancellophycus Saporta, 1872 lattice seaweed Quebecichnus Hofmann, 1972 Québec (Canada) trace Caulerpites Sternberg, 1833 ancestral (or near) Caulerpa Steigerwaldichnium Kuhn, 1937 Steigerwald (Germany) little-trace (a seaweed) Tambia Müller, 1969 Tambach (Germany) Ceratophycus Schimper, 1879 horn seaweed Tambachichnium Müller, 1954 Tambach (Germany) little-trace Chondrites Sternberg, 1833 ancestral Chondria Tasmanadia Chapman, 1929 Tasmania (Australia) Criophycus Toula, 1906 ram seaweed Umfolozia Savage, 1971 (Swart) Umfolozi, (South Africa) Flabellophycus Squinabol, 1890 fan seaweed Fucoides Brongniart, 1822 like Fucus (a seaweed) Gyrophyllites Glocker, 1841 ancestral turning leaf Halmedides Lorenz von Liburnau, 1902 like Halimeda (a calcareous alga) Ta ble 6 Halopoa Torell, 1870 salt grass Hippodophycus Hall & Whitfield, 1872 horse-hoof seaweed Some ichnogeneric names that are based on stratig ra phy. Hormosiroidea Schaffer, 1928 like Hormosira (a seaweed) Ichnophycus Hall, 1852 track seaweed This should be done only occa sion ally. Licrophycus Billings, 1862 antler seaweed Mycelites Roux, 1887 ancient mycelium (fungus) Ichnogenus Meaning Myelophycus Ulrich, 1904 marrow or pith seaweed Archaeichnium Glaessner, 1963 old little-trace (Precambrian) Nulliporites Heer, 1865 ancestral Nullipora (a seaweed) Beaconichnus Gevers, 1973 Beacon (Group) trace (Antarctica) Palaeophycus Hall, 1852 ancient seaweed Beaconites Vyalov, 1962 ancestral Beacon (Group) Phycodes Richter, 1850 seaweed-like Graysonia Stephenson, 1952 Grayson (Marl, Texas) Phycosiphon Fischer-Ooster, 1858 seaweed tube Hartsellea Rindsberg, 1994 Hartselle (Sandstone, Alabama) Physophycus Schimper, 1869 bellows seaweed Navahopus Baird, 1980 Navaho (Sandstone, Arizona) Phytopsis Hall, 1847 plant-looking Permichnium Guthörl, 1934 Permian little-trace Rusophycus Hall, 1852 wrinkled seaweed Protichnites Owen, 1852 first trace (Cambrian) Spirophycus Häntzschel, 1962 spiral seaweed Torrowangea Webby, 1970 Torrowangee (Group, New South Wales) Spirophyton Hall, 1863 spiral plant Yakutatia Ulrich, 1904 Yakutat (Formation, Alaska) Staurophyton Meunier, 1891 cross plant Yokoia Hatai and Noda, 1975 Yoko-o (Formation, Japan) Trichophycus Miller and Dyer, 1878b hair seaweed Zoophycos Massalongo, 1855 animal plant

Some generic names honour ichnologists or other sci- and Zoophycos. Among the trace fossils mistaken for ani - entists, gener ally by adding the ending -ia to the stem of the mals, Rhizocorallium is not a coral, and Protovirgularia is per son’s name (Table 7). Haentzschelinia honours Walter not related to the octocoral Virgularia. Häntzschel, who wrote the ichnologic part of the Trea tise Sev eral ichnogenera were named for sites of discov ery, on In ver te brate Pa le on tol ogy as well as many other ar ticles strata, or geo logic age (Ta bles 5, 6). Place-derived names on traces. Topsentia (a sponge; the borings within this ge- in clude Brookvalichnus (Brookvale), Capodistria (Capo nus were later re named Topsentopsis) honours Émile d’Istria), Steigerwaldichnium (Steigerwald), and sev eral Topsent, who worked on modern sponges; Rouaultia was others (Table 5). Such geo graphic names can be tire some named for Marie Rouault, who named several ichnogenera when addi tional speci mens are discov ered far from the type in 1850. Saportia Squinabol, 1890 and Lorenzina Gabelli, area. Na tive names are of ten re freshing, e.g., Umfolozia. 1900 respec tively honour ichnologist Gaston de Saporta Stratigraphic names are less common (Table 6), but include and nat u ral ist Stefano Lorenzini. Some patronyms are less Beaconites and Beaconichnus (Bea con Group), Hartsellea ap pro pri ate. Scoyenia White, 1929 was named for E. T. (Hartselle Sand stone), and Yakutatia (Yakutat Forma tion). Scoyen, Chief Ranger of Grand Canyon National Park, Only a few ichnogenera refer to stratigraphic age, such as Lockeia for John Locke, a Cincinnati pale on tol o gist, Permichnium (Permian); one might also include Archaei- Cruziana Orbigny, 1849 for General Andrés de Santa Cruz chnium (“old little trace”, from the Precam brian). All these y Calahumana, and Berguaeria Prantl, 1946, ap parently names have a seri ous dis advan tage. At its first publi cation, play fully, for the Czech sexologist Vladimír Bergauer. Be - the full range of a genus is un likely to be known. Beaco- cause these persons did no ichnologic work, these names nichnus is now known outside the Beacon Group of have little signif i cance to ichnologists. Such names say Antarctica, and Permichnium has been reported from the nothing about the trace fossil it self. However, persons’ Tri as sic (Häntzschel, 1975, pp. W45, W91). Never the less, names are rela tively easy to re mem ber, and they pro vide re - they have the advan tages of sounding fresh and mem o rable. lief from the endless ichnogenera ending in -ichnus. 534 A. K. RINDSBERG

Ta ble 7 A selec tion of ichnogeneric names that are modi fied from the names of persons. This should be done sparingly.

Ichnogenus Meaning Person Abeliella Mägdefrau, 1937 little Abel Othenio Abel, paleobiologist Bergaueria Prantl, 1946 Bergauer Vladimír Bergauer, geologist Brachyzapfes Codez, 1957 short Zapfe Helmuth Zapfe Brooksella Walcott, 1896 little Brooks William Keith Brooks, zoologist Collettosaurus Cox, 1873 Collett lizard John Collett, geologist Condranema Bassler, 1952 Condra thread George Evert Condra, geologist Cruziana Orbigny, 1842 pertaining to Cruz Andrés de Santa Cruz y Calahumana, President of Peru and Bolivia Favreina Brönnimann, 1955 little Favre J. Favre, first describer of trace Glockerichnus Pickerill, 1982 Glocker trace F. E. Glocker; substitute name for Glockeria Haentzschelinia Vyalov, 1964 little Häntzschel Walter Häntzschel, ichnologist Harlania Goeppert, 1852 Harlan Richard Harlan, paleontologist Keckia Glocker, 1841 Keck Michael Keck von Keck, collector Kingella Savage, 1971 little King Lester King, discoverer of trace Lapparentichnus Haubold, 1971 Lapparent trace Albert-Félix de Lapparent, ichnologist Lockeia James, 1879 Locke John Locke, paleontologist Lorenzinia Gabelli, 1900 Lorenzini Stefano Lorenzini, naturalist Matthewichnus Haubold, 1970 Matthew trace George Frederick Matthew, ichnologist Mehliella Strand, 1932 little Mehl Maurice G. Mehl, vertebrate paleontologist Moodieichnus Sarjeant, 1972 Moodie trace Roy Lee Moodie, paleopathologist Muensteria Sternberg, 1833 Münster Georg von Münster, paleontologist Rauffella Ulrich, 1889 little Rauff Hermann Rauff, paleontologist Saportia Squinabol, 1891 Saporta Gaston de Saport, paleobotanist Scoyenia White, 1929 Scoyen E. T. Scoyen, park ranger Shepardia Hitchcock, 1858 Shepard Charles Upham Shepard, geologist Sillimanius Hitchcock, 1845 Silliman Benjamin Silliman, geologist Tasselia de Heinzelin, 1965 Tassel R. Van Tassel, first describer of trace Topsentia Clarke, 1921 Topsent E. Topsent, sponge biologist Volkichnium Pfeiffer, 1965 Volk trace Max Volk, ichnologist Walcottia Miller & Dyer, 1878a Walcott Charles Doolittle Walcott, paleontologist Wildeichnus Casamiquela, 1964 Wilde trace Eduardo Wilde, physician and politician Zapfella Saint-Seine, 1956 little Zapfe H. Zapfe

Some ichnogenera are named for other ichnogenera Ta ble 8 (Ta ble 8): Paratisoa for its simi lar ity to Tisoa, Megaper- michnium af ter Permichnium, Sublorenzinia for Lorenzinia, A selec tion of ichnogeneric names (not all valid) that are Parahaentzschelinia for Haentzschelinia, Neonereites af ter modi fied from other ichnogeneric names. This is not Nereites, Mixoteichnichnus for Teichichnus. The sim i lar ity a rec om mended prac tice. is often idio syn cratic and may strike other workers as super - ficial and mislead ing. Also, these names tend to be incon ve - Ichnogenus Meaning niently long. Placing a prefix like sub- (“un der, less”) be- Chondritoides Borrello, 1966 Chondrites-like fore a person’ s name, as in Sublorenzinia and Subglockeria, Eugyrichnites Ami, 1905 true Gyrichnites Mixoteichichnus Müller, 1966 mixed Teichichnus is un in ten tion ally of fen sive. In some cases, re jected hom - Neonereites Seilacher, 1960 new Nereites onyms have been re placed by sim ilar names, re tain ing no- Parahaentzschelinia Chamberlain, 1971 near Haentzschelinia men cla tor ial con ti nu ity. Glockerichnus re placed Glockeria; Paratisoa Gaillard, 1972 near Tisoa Topsentopsis re placed Topsentia. Protopaleodictyon Ksi¹¿kiewicz, 1970 first Paleodictyon Pseudobilobites Kennedy, 1967 false Bilobites Names of mis cel la neous or i gin, in clud ing hu mor ous Radionereites Gregory, 1969 radiating Nereites names, provide refresh ment and amusement (Table 9). Spirocosmorhaphe Seilacher, 1989 spiral Cosmorhaphe Ardelia (“busy body”) is a ver ti cal bur row hav ing hor i zon tal Subglockeria Ksi¹¿kiewicz, 1968 under (i.e., “not quite”) branches in all direc tions. A burrow having a sun burst Glockeria Subphyllochorda Götzinger & Becker, 1932 “not quite” Phyllochorda cross-sec tion re ceived the cog no men Phoebichnus, for Taphrhelminthopsis Sacco, 1888 ditch Helminthopsis Phoebus Apollo, the Greek sun god. Baroccoichnites refers to its bi zarrely “ba roque” curls. A fine name is Daedalus, a screw-shaped trace fos sil, hon our ing the myth i cal sci en tist Daedalus. Daimonelix is a rough transla tion into Greek of and Asteriacites gugelhupf Seilacher, 1983, a plug-shaped its com mon name, “devil’s cork screw”. Less amusing in the trace that re sembles a Ger man cake called a Gugelhupf. An long run is Hondichnus, whose au thor likened to the track occa sional joke is good, and Linnaeus himself was a great of a Honda mo tor cycle. If species-level names may be humor ist (Austin, 1993), but ichnologic taxon omy is al - introduced into this discus sion, surely the follow ing must be ready burdened by an abundance of unin tended humour. men tioned: Diplocraterion yoyo Goldring, 1962, a U-shaped A few names are of un cer tain der iva tion, e.g., Laotira burrow whose spreite shifted up and down like a toy yoyo, Walcott, 1896, Macanopsis Macsotay, 1967 (perhaps “like CON STRUC TION OF ICHNOGENERIC NAMES 535

Ta ble 9 both morphologic and etholog ic, and it is this kind of name I would like to encour age. If a new trace fossil includes more A selec tion of ichnogeneric names (not all valid) having than one kind of behav iour, and espe cially if its behav iour is mis cel la neous der i va tions in doubt, the researcher should refrain from incor po rat ing its ethology into its name. Ichnogenus Meaning Ardelia Chamberlain, 1971 busybody Cursipes Matthew, 1903 run foot Daedalus Rouault, 1850 Daedalus (mythical Greek craftsman) ETHO LOGICALLY BASED ROOTS FOR Daimonelix Barbour, 1892 demon helix (translation of common ICHNOGENERIC NAMES name “devil’s corkscrew”) Dromopus Marsh, 1894 run foot Entobia Bronn, 1838 liv(ing) within Most trace fossils are inver te brate burrows and borings, Fraena Rouault, 1850 bridle, bridge (i.e., straplike) ver tebrate trackways, and cop ro lites (for an over view see Grallator Hitchcock, 1858 stilt-walker Knaust, 2012). Inver te brate burrows can be classi fied in the Hondichnus Ausich, 1979 Honda (motorcycle) trace Incisifex Dahmer, 1937 incis(ion) maker fa mil iar etho log i cal cat e go ries (rest ing, dwellling, feed ing, Iramena Boekschoten, 1970 anagram of Annemarie crawl ing, graz ing, farming, and es cape traces), and roots Boekschoten-Van Helsdingen sug gest ing these ac tiv i ties can be in cor po rated into ge neric Phoebichnus Bromley & Asgaard, Phoebus (Apollo) trace names. Appro pri ate roots for verte brate trackways, borings, 1972 Planolites Nicholson, 1873 wandering stone and cop ro lites are also avail able in clas sical lan guages. Psammichnites Torell, 1870 ancestral sand trace The In ter na tional Code of Zoo log i cal No men cla ture Saltator Hitchcock, 1858 leaper (ICZN, 1999) en cour ages the use of clas sical (Greek and Scotolithus Linnarsson, 1871 obscure stone Latin) roots; it does not re quire them, but roots from other Sustergichnus Chamberlain, 1971 pig-back (i.e., hogback) trace (from local football team “Arkansas languages are less likely to be under stood. Although it has Razorback”) been a long time since sci en tific pa pers were rou tinely writ- Talpina Hagenow, 1840 little mole ten in Latin, and the typi cal student no longer studies classi - Tisoa Serres, 1840 Tisoa, a sea nymph in Greek cal lan guages, ev ery sci en tist is nev er the less fa mil iar with at mythology Walpia White, 1929 Hualapai (Native American tribe) least the common est classi cal roots, such as the bio- of bi ol - ogy and biogeography and bioerosion, and can consult dic- tio naries to obtain others. The root ichn- is rare out side of the studies of trace fossils and ichneu mon wasps, but its ex- a macana”, i.e., “club” in Spanish), and Palaxius Brönni- tensive use in ichnology has made it easily under stood to mann and Norton, 1980. Occa sion ally, an author does not geol o gists. The prem ise of this paper is that addi tional roots reveal the ety mol ogy of his cre ation, al though it is good can become famil iar to ichnologists with use. practice to do so. Further, authors may dub a new ichnoge- With the aid of standard sources such as Liddell et al. nus with any com bina tion of letters (e.g., Pirandikus Holub (1940), Brown (1956), and Stearn (1966), I have com piled and Kozur, 1981) as long as they follow a few sen sible lists of roots that may be used in creat ing new ichnogeneric rules. The name must con sist of at least three Latin let ters, names (Tables 11, 12). While all can be used to describe a be pronounce able, not previ ously used, and so forth (ICZN, trace fos sil mor pho log i cally, some can ad di tion ally be used 1999). to sug gest an etho log i cal in ter pre ta tion. Thus, tubus, “tube”, The most appro pri ate names for ichnogenera are pri- can be used to denote trace fossils that are tubu lar in form, marily morphologic (Ta ble 2), because the cri teria for iden- while nidus, “nest”, could be incor po rated within the name tify ing ichnotaxa are morphologic (Bromley, 1970, 1996; of an ichnogenus that is inter preted as a calichnion. Sarjeant and Kennedy, 1973; Fürsich, 1974; Häntzschel and Frey, 1979; Pem berton and Frey, 1982, p. 847). These names are very nu merous and most are still appro pri ate. A few ex am ples are Cylindrichnus (“cyl in der trace”), AES THETIC AND PRAG MATIC Asterosoma (“star body”), Ophiomorpha (“snake shape”), CONSID ER ATIONS Cochlichnus (“screw trace”), Unisulcus (“one furrow” ), Diplocraterion (“dou ble cup”), Dimorphichnus (“two shape In ex pe ri enced bi ol o gists and ge ol o gists of ten em bark trace”). Morphologic names are mnemonic in any language on nam ing a new organ ism or trace fos sil in the spirit of ad- that de rives its scien tific ter minol ogy from Latin or Greek, ven ture, vying for a name that will be dis tinc tive in its own in clud ing all ma jor Eu ro pean lan guages. sake: the longest, the shortest, the hardest to pronounce; the Only a handful of names re fer to the bi o logi cal activ ity most enter tain ing, the most demean ing, the most memo ra - rep re sented by the trace fos sil. Such names in clude Pha- ble. This is why we have para sites named for polit i cal fig- gophytichnus (“eat plant trace”) for bite marks on leaves, ures, plankton for rock stars. These kinds of jokes tend to Fascifodina (“bun dle eat”), and Limulicubichnus (“limu- pall with age, but sci en tists will be stuck with them indef i - li[de] rest ing trace”). Although the danger exists that such nitely. If a joke is called for, it should be one that can be ap - names may misrep re sent the ac tiv ity, such names are eas ily pre ci ated de cades or even cen tu ries hence. re mem bered and fall neatly into the etho log ic cate go ries of Ex pe ri enced tax on o mists tend to oc cupy the mid dle Seilacher (1953) as revised by later authors (for a review ground, striving for names that are appro pri ate, useful, and see Vallon et al., 2015). Names can be constructed that are con sid er ate of oth ers. 536 A. K. RINDSBERG

Ta ble 10 Appro pri ate Latin roots for ichnogenera and other ichnologic terms. Meanings are selected from among those given by Simpson (1968). Abbre vi a tions: m. = mascu line noun, f. = femi nine noun, n. = neuter noun, v. = verb, a. = adjec tive.

Latin word and stem Selected meanings Examples Roots for general use signum n., sign- sign, distinguishing mark tubus m., tub- tube, pipe, trumpet Micatuba Chamberlain, 1971; Scalarituba Weller, 1899; Vermitubus Hatai, Murata & Kawakami, 1972 vestigium n., vestigi- footprint, trace Punctatumvestigium Butts, 1891; Triavestigia Gilmore, 1927 Roots for resting trace celare v., cel- to hide fovea f., fove- small (dug) pit stratum n., strat- covering tectum n., tect- shelter, roof Roots for dwelling traces atrium n., atri- chief room in Roman house baculum n., bacul- staff, stick caula f., caul- hole Caulostrepsis Clarke, 1908 caverna f., cavern- hollow, cavern Cavernaecola Bentz, 1929 cavea f., cave-; den, cavity, hollow nucleocavia Richter & Richter, 1930 cavus m., cav-; cavum n., cav- culmus m., culm- stalk, stem (use for shafts) domus f., dom- home domichnion Seilacher, 1953 aedes f., aed- room, house Roots for locomotive traces canalis m., canal- channel, canal fossa f., foss- ditch Rodocanalis Schloz, 1972 iter m., itiner- way Fossil ligula f., ligul- strap lira f., lir- furrow ridge motus m., motu- motion semita f., semit- path, way sulcus m., sulc- furrow Trisulcus Hitchcock, 1865; Unisulcus Hitchcock, 1858 taenia f., taeni- ribbon, headband Taenidium Heer, 1877 via f., vi- highway, road Roots for trackways digitus m., digit- digit, finger or toe gradus m., grad- footstep manus f., manu- hand palma f., palm- palm (of the hand) Circapalmichnus Gand, 1977; Palmichnus Schmidt, 1959 pes m., ped- foot Bifurculapes Hitchcock, 1858; Hamipes Hitchcock, 1858; Pachypes Leonardi et al., 1975; Quadropedia Aldrich, 1930 planta f., plant- sole (of the foot) Roots for grazing traces pascuum n., pascu- pasture pascichnion Seilacher, 1953 pastus m., pastu- pasture Roots for feeding traces fodio v., fodi- dig (dug = foss-) Fascifodina Osgood, 1970; fodinichnion Seilacher, 1953 quareia f., quarei- quarry (Medieval Latin) spica f., spic- ear (of grain; Ichnyspica Linck, 1949 use for “ear-shaped traces” Roots for traces including a spreite folium n., foli- leaf lamina f., lamin- sheet, thin layer Laminites Ghent & Henderson, 1966 lamna f., lamn- Roots for farming traces hortus m., hort- garden Roots for nesting traces nidus m., nid- nest CON STRUC TION OF ICHNOGENERIC NAMES 537

Table 10 continued

Latin word and stem Selected meanings Examples Roots for coprolites faex f., faec- lees, impure residue (hence faeces in the modern sense) stercus n., stercor- dung fimus m., fim-; dung fimum n., fim- globus m., glob- ball glomus n., glomer- ball pila f., pil- ball pilula f., pilul- pill Roots for borings foramen n., foramin- (bored or pierced) hole foris f., for- door Conchifora Müller, 1968; Foralites Rouault, 1850; Vermiforichnus Cameron, 1969 noxa f., nox- harm, injury rodere v., rod- gnaw, erode Calciroda Mayer, 1952; Filuroda Solle, 1938; Rodocanalis Schloz, 1972 specus m., f., n., specu- cave, hole, hollow Lapispecus Voigt, 1970; Specus Stephenson, 1952 terebra f., terebr- borer terebro v., terebr- to bore Terebripora Orbigny, 1842

Con sult a dic tio nary Incis-i-fex. Greek roots should be con nected by -o-, e.g., Arthr-o-phycus, Collett-o-saurus. Several guides to the construc tion of new scien tific terms are available. The most valu able are Bo tan i cal Latin (Stearns, 1966) and Com po si tion of Sci en tific Words Keep the name short (Brown, 1956). De scrib ing Spe cies (Winston, 1999) is also Save those extra sylla bles for the diag no sis. It is incon - useful, though it is unre li able with regard to ichnotaxono- sider ate to give an organ ism a monstrously long name, how- mic prin ci ples. The clas si cal lan guages, es pe cially Latin, ever appro pri ate the name may be. There is no need to com- have con tin ued to evolve in sci en tific us age, par tic u larly for press an entire di agno sis into an ichnogenus; an ac curate the de scrip tion of organ isms. Botan i cal Latin is a standard - name is good, but a brief name is better. Greek is often more ized and flexi ble mode of commu ni ca tion that was honed suc cinct than Latin; compare Greek ichnos and Latin over a period of centu ries. Linnaeus used it, and bota nists vestigium, “trace”. A good limit for most words is five sylla - still use it. bles (Table 13); Bromley (oral comm., 1998) rec om mended Beware of “false friends”, words whose meaning has three in clud ing -ichnus. Remem ber that the ichnogeneric changed over the centu ries. Rarus is “sparse”, not “rare”; name will often be used together with specific names, and if intricatus is “en tan gled”, not “in tri cate.” In these cases, the one is very long, the other should be short. meaning has shifted in English while remain ing constant in Latin. Use a va riety of con struc tion meth ods

Consult an expert on correct language use Morphologic and etholog ic roots are best, but an occa - sional excep tion is good to make names memo ra ble. Names If you know no Latin or Greek, have someone else check of ani mals, per sons, places, and forma tions are ac cept able, your work. A bota nist is the preferred choice, because Botan - though not ideal. Names can also be constructed from ana- i cal Latin dif fers con sid er ably from Clas si cal Latin. The In - grams or random combi na tions of letters. ter na tional Code of Zoo log i cal No men cla ture (ICZN, 1999) has spe cific rec om men da tions on the trans lit er a tion and Say the name out loud latinization of Greek roots. It is all too easy to make embar - rassing but lasting mistakes, e.g., Ophiomorpha irregulaire It may be that the proposed name is a pun or joke, and Frey, Howard and Pryor, 1978 incor rectly used French not even an inten tional or clever one. Aim for euphony; if irregulaire in stead of Latin irregularis, but once published you cannot pronounce it, it is not a good name. the name could not be changed. Do not com bine unlatinized Greek roots with Latin Check to see if the name has al ready been used ones in the same word. While it is not forbid den by the Code, it is in poor taste. If a combi na tion of such roots is de- In the precybernetic era, this meant a trip to a good ref- sired, the Greek root may be latinized, a prac tice be gun by erence library for an after noon with compi la tions such as the Romans them selves. the Trea tise on In ver te brate Pa le on tol ogy, Fossilium Cata- When combin ing Latin roots, use the vowel -i- to con- logus, Nomenclator Zoologicus, and Zoo log i cal Re cord. nect them if one is not supplied by the root itself, e.g., Today, it is more likely to involve searches of online data - 538 A. K. RINDSBERG

Ta ble 11 Appro pri ate Greek roots for ichnogenera. Translit er a tion and latinization are done accord ing to recom men da tions of the In ter na tional Code of Zoo log i cal No men cla ture (In ter na tional Com mis sion for Zoo log i cal No men cla ture, 1999); ei ther version is accept able, although Greek and Latin should not be mixed. Latinized Greek roots are treated as if they were Latin; they may be combined with Latin roots. Se lected meanings are those that are most appro pri ate to ichnology. Abbre vi a tions: m. = mascu line noun, f. = femi nine noun, n. = neuter noun, v. = verb, a. = adjec tive.

Greek Transliterated Greek Latinized Greek Selected meanings Examples Roots for general use Aulacophycus Heer, 1877 Aulichnites Fenton & Fenton, 1937a "L8`H aulos m., aul- aulus m., aul- flute Cymataulus Rindsberg, 1994 Didymaulichnus Young, 1972 Tropidaulus Fenton & Fenton, 1937a {`D:@H hormos m., horm- hormus m., horm- chain, cord, necklace, collar Hormosiroidea Schaffer, 1928 ÇP<4@< ichnion n., ichni- ichnium n., ichni- little track See Table 1. ÇP<4@H ichnos n., ichn- ichnus m., n., ichn- trace, track, footprint, spoor See Table 1. Cylindrichnus Toots in Howard, 1966 6L84<*D@H kylindros m., kylindr- cylindrus m., cylindr- cylinder Cylindricum Linck, 1949 Kulindrichnus Hallam, 1960 Belorhaphe Fuchs, 1895 Chondrorhaphe Seilacher, 1977 {D"n\H rhaphis f., rhaphid- rhaphis f., rhaphid- needle Helicorhaphe Ksi¹¿kiewicz, 1970 Helminthorhaphe Seilacher, 1977 F0:" sema n., semat- semum n., semat- sign F6VBJT- skaptein v. scapt- dig, hoe Syringodendron Fuchs, 1895 FLD4(> syrinx f., syring- syrinx f., syring- pipe Syringomorpha Nathorst, 1886 Roots for resting traces {X*D" hedra f., hedr- hedra f., hedr- seat, abode {X*D4@< hedrion n., hedri- hedrium n., hedri- little seat 6"8bBJ`H kalyptos a., kalypt- calyptus a., calypt- covered, hidden 6"8bBJD" kalyptra f., kalyptr- calyptra f., calyptr- covering, veil, lid 68\<0 kline f., klin- clina f., clin- couch 6DLBJ`H kryptos a., krypt- cryptus a., crypt- hidden, secret cryptobioturbation Howard and Frey, 1975 `DL(:" orygma n., orygmat- orygma m., f., orygmat- (dug) pit, hole, trench, tunnel Diorygma Biernat, 1961 @DbFF@T orysso v., oryss- oryss- v., oryss- dig, dig up, dig through, bury @Db>4H oryxis f., oryx- oryxis f., oryx- digging F6"n0 skaphe f., skaph- scapha f., scaph- trough, skiff, dugout FJX(" stega f., steg- stega f., steg- roof, room, shelter FJ4(:",-"J@H stigma f., stigmat- stigma m., f., stigmat- tattoo, brand Roots for dwelling traces V

Greek Transliterated Greek Latinized Greek Selected meanings Examples @46@H oikos m., oik- oecus m., oec- house, room, home Ctenopholeus Seilacher & Hemleben, 1966 nT8g`H pholeos m. pholeus hole, cave, den Pholeus Fiege, 1944 nDg"D phrear n., phreat- phrear n., phreat- well Dictyoporus Mägdefrau, 1937 B`D@H poros m., por- porus m., por- opening, pore Spathipora Fischer, 1866 Terebripora Orbigny, 1842 Phycosiphon Fischer-Ooster, 1858 F\nT< siphon m., siph- siphus, sipho m., siphon- tube, pipe Siphodendron Saporta, 1884 Siphonites Saporta, 1872 F6XB"H skepas n., skepa- scepas m., scep- covering, shelter F6gBJD@< skeptron n., skeptr- sceptrum n., sceptr- staff, cane F6bn@H skyphos m., skyph- scyphus m., scyph- cup, can F6LJV"0 skytale f., skytal- scytala f., scytal- staff, club, cudgel F@8Z< solen m., solen- solen m., solen- pipe, channel, gutter FJ@:" stoma n., stomat- stoma m., f., stomat- mouth Myzostomites Clarke, 1921 2V8":@H thalamos m., thalam- thalamus m., thalam- inner room Thalamophaga Rhumbler, 1911 2060 theke f., thek- theca f., thec- box, case, chest 2@8@H tholos f., thol- tholus m., thol- round building Roots for grazing traces choria, chorea f., chori-, P@Dg\" khoreia f., khore- round dance chore- noma f., nom- pasture (food) <@:Z nome f., nom- nomas f., nomad- pasturing, hence nomad <@:`H nomos m., nom- nomus m., nom- pasture (place) Roots for feeding traces *g4B<@< deipnon n., deipn- dipnum n., dipn- dinner, meal, food @DL6J`H oryktos a., orykt- oryctus a., oryct- dug, dug up, dug out n"(g4< phagein v., phag- phageo v., phag- eat Phagophytichnus van Ameron, 1966 {D"$*@H rhabdos f., rhabd- rhabdus m., rhabd- rod, stick, wand Rhabdoglyphus Vassoevich, 1951 {D"$*\@< rhabdion n., rhabdi- rhabdium n., rhabdi- little rod {D`B"8@< rhopalon n., rhopal- rhopalum n., rhopal- club (heavy stick) Ropalonaria Ulrich, 1879 ear (of grain; to be used for FJVPLH stakhys m., stakh- stachys m., stach- "ear-shaped traces" Roots for traces having a spreite X8LJD@< elytron n., elytr- elytrum n., elytr- (rolled) sheath nb88@< phyllon n., phyll- phyllum n., phyll- leaf Gyrophyllites Glocker, 1841 any sweeping motion, e.g., a snake FLD:`H syrmos m., syrm- syrmus m., syrm- trail Roots for locomotion traces in general *D`:@H dromos m., drom- dromus m., drom- (running) course Helicodromites Berger, 1957 {@*`H hodos f., hod- hodus m., hod- road, way {@86`H holkos f., holk- holcus m., holc- furrow, hence trace 6\

Greek Transliterated Greek Latinized Greek Selected meanings Examples Roots for trackways Dactyloidiscus Œl¹czka, 1965 Dactyloidites Hall, 1886 Dactylophycus Miller & Dyer, 1878b *6JL8@H daktylos m., daktyl- dactylus m., dactyl- finger, toe Harpedactylopus Hitchcock, 1858 Peratodactylopus Sarjeant & Mossman, 1978 Rotodactylus Peabody, 1948 Pg\D kheir f., kheir- chir f., chir- hand Chirotherium Kaup, 1835 Bg*\@< pedion n., pedi- pedium n., pedi- metatarsus Quadropedia Aldrich, 1930 BX8:" pelma n., pelmat- pelma n., pelmat- sole (of the foot) Copeza Hitchcock, 1858 BX." peza f., pez- peza f., pez- foot, instep Sphaeropezium King, 1845 Xiphopezia Hitchcock, 1848 nV8"(> phalanx f., phalang- phalanx f., phalang- finger bone, toe bone Phalangichnus Schmidt, 1959 B@*4@< podion n., podi- podium n., podi- little foot Crossopodia M’Coy, 1851 Agialopous Branson & Mehl, 1933 Diplopodichnus Brady, 1947 Harpepus Hitchcock, 1865 B@bH pous m., pod- pus m., pod- foot Octopodichnus Gilmore, 1927 Podichnus Bromley & Surlyk, 1973 Sauropus Lea, 1849 BJgD<\H pternis f., pternid- pterna f., pternid- base (of a dish) FJ\$@H stibos m., stib- stibus m., stib- footprint, trackway, path flat basket or mat, hence flat of the J"DF`H tarsos m., tars- tarsus m., tars- foot (between toes and heel) 2X<"D thenar m., thenat- thenar m., thenat- palm (of the hand) Thenaropus King, 1846 Roots for farming traces (Graphoglyptids) $"88\.T ballizo v., balliz- ballizo v., balliz- to dance, jump about $4$"F4H bibasis f., bibas- bibasis f., bibas- Spartan dance P@D`H khoros m., khor- chorus m., chor- circle dance Desmograpton Fuchs, 1895 *gF:`H desmos m., desm- desmus m., desm- band (for fastening) Spirodesmos Andrée, 1920 Dictyodora Weiss, 1884 Dictyoporus Mägdefrau, 1937 Glenodictyum Marck, 1863 *\6JL@< diktyon n. dictyum n., dicty- net Irredictyon Vyalov, 1972 Paleodictyon Meneghini in Murchison, 1850 Ramidictyon Seilacher, 1977 Squamodictyon Vyalov and Golev, 1960 bioglyph Vassoevich, 1953 Fustiglyphus Vyalov, 1971 Petaloglyphus Vyalov, Gorbach & Dobrovolska, 1964 (8nZ glyphe f., glyph- glypha f., glyph- carving, a cut hole Rhabdoglyphus Vassoevich, 1951 Stelloglyphus Vyalov, 1964 vermiglyph Fuchs, 1895 cf. graphoglyptid Seilacher, 1977 graphoglyptid Seilacher, 1977 (D"nZ graphe f., graph- grapha f., graph- writing Lithographus Hitchcock, 1858 Desmograpton Fuchs, 1895 (D"BJ`H graptos a., grapt- graptus a., grapt- written, engraved Megagrapton Ksi¹¿kiewicz, 1968 P`DJ@H khortos m., khort- chortus m., chort- grass, feeding lot Gyrochorte Heer, 1865 6`D*"> kordax n., kordak- cordax m., f., cordac- lively comic dance :@((VH mongas n., mongad- mongas m., f., mongad- wild dance :`<4:@H monimos m., monim- monimus m., monim- stable (for animals) @DP0F4H orkhesis f., orkhes- orchesis f., orches- dancing, the dance @DPZFJbH orkhestes m., orkhest- orchestus m., orchest- dancing, the dance CON STRUC TION OF ICHNOGENERIC NAMES 541

Greek Transliterated Greek Latinized Greek Selected meanings Examples Roots for farming traces @DF\J0H orsites m., orsit- orsitus m., orsit- Cretan dance Acanthorhaphe Ksi¹¿kiewicz, 1970 Belorhaphe Fuchs, 1895 Cosmorhaphe Fuchs, 1895 {D"n\H rhaphe f., rhaph- rhapha f., rhaph- seam Helicorhaphe Ksi¹¿kiewicz, 1970 Spirorhaphe Fuchs, 1895 Strobilorhaphe Ksi¹¿kiewicz, 1968 F6"\DT skairo v., skair- scaero v., scaer- to dance, skip, caper Roots for coprolites 6@BD\" kopria f., kopri- copria f., copri- dunghill coprolite Buckland, 1829 Coprolithus Paréjas, 1948 coper, coprus m., coper-, Coprulus Richter and Richter, 1939 6`BD@H kopros f., kopr- dung copr- Prethocoprolithus Elliott, 1962 Scolecocoprus Brady, 1947 Syncoprulus Richter and Richter, 1939 BV88" palla f., pall- palla f., pall- ball F6TD skor n., skat- scor n., scat- dung scatology Roots for borings and other bioerosion structures $8V$0 blabe f., blab- blaba f., blab- harm, damage Ostreoblabe Voigt, 1965 4H nyxis f., nyx- nyxis f., nyx- pricking, stabbing salpinx f., salping- salpinx f., salping- trumpet 2LD\H thyris f., thyrid- thyris f., thyrid- small door, window JDg:" trema n., tremat- trema m., f., tremat- hole, orifice (rubbed) Conchotrema Teichert, 1945 JD0F4H tresis f., tres- tresis f., tres- perforation (pierced) JDf(80 trogle f., trogl- trogla f., trogl- hole (gnawed), mousehole JDbB"T trypan v., tryp- trypo v., tryp- bore, pierce Trypanites Mägdefrau, 1932 JDbB0:" trypima n., trypimat- trypima m., f., trypimat- hole (bored) bases in clud ing the Zoo log i cal Re cord, Google Scholar, and REF ER ENCES PaleoDatabase. Bear in mind that these are incom plete; if you select a name based on a single Greek or Latin word, or Abel, O., 1935. Vorzeitliche Lebensspuren. Jena: Gustav Fischer, even an obvi ous combi na tion of two classi cal roots, it is 644 pp. rather likely to be unorig i nal. Early names, names published Aldrich, T. H., Sr., 1930. De scrip tion of tracks. In: Aldrich, T. H., in obscure works, and names published in works written in Sr. & Jones, W. B., Foot prints from the coal mea sures of Ala - non-Euro pean languages are frequently overlooked in data - bama. Al a bama Mu seum of Nat u ral His tory, Mu seum Pa per, bases. 9: 64 pp. Uni ver sity, Al a bama. Ameron, H. W. J. van, 1966. Phagophytichnus ekowskii nov. And finally, ichnogen. & nov. ichnosp., eine Missbildung infolge von Insektenfrass, aus dem spanischen Stephanien (Provinz Leon). Leidse geologische Mededelingen, 38: 181–184. Use your imag i na tion! Ami, H. M., 1903. De scrip tion of tracks from the fine-grained sili - ceous mudstones of the Knoydart For mation (Eo-De vonian) of Antigonish County, Nova Sco tia. Pro ceed ings and Trans- ac tions of the Nova Sco tian In sti tute of Science, 10 [for Ac knowl edge ments 1898–1902]: 330–332. Hal i fax. I am grate ful to Rich ard G. Bromley, who hosted the First Ami, H. M., 1905. Prelim i nary list of the fossils collected by Workshop on Ichnotaxonomy at Limensg¯de Mill (kirkeby, Profesor L. W. Bailey from vari ous local i ties in the Prov ince Bornholm, Den mark), where I de liv ered the first ver sion of this of New Brunswick dur ing 1904. Summary Re port of the Can - pa per as a talk in 1998. I thank the Ed i tor, Al fred Uchman, for en - ada Geo log i cal Sur vey for the Cal en dar Year 1904, 289–292. cour ag ing me to write a piece for this issue even though I was un - Ander son, A. M., 1976. Fish trails from the Early Permian of able to at tend the One Ichnology con fer ence in 2014. Lothar H. South Africa. Palae on tol ogy, 19: 397–409. Vallon kindly ar ranged to have Häntzschel’s copy of Jar dine’s Andrée, K., 1920. Über einige fossile Problematika. I. Ein (1853) Ichnology of Annandale scanned at the Universität Ham- Problematikum aus dem Paläozoikum von Battenberg an der burg. I also thank Max Wisshak (Senckenberg Re search In sti tute, Eder und das dasselbe beherbergende Gestein. Neues Frank furt am Main) and Dirk Knaust (Statoil ASA, Stavanger) for Jahrbuch für Mineralogie, Geologie, und Paläontologie, thought ful and thor ough re views, though it should not be as sumed 1920(1): 55–88. that they share all of the views expressed here. Ausich, W. I., 1979. Hondichnus monroensis n. gen. n. sp.: a new 542 A. K. RINDSBERG

Ta ble 12 sand stones in Kan sas. Uni ver sity of Kan sas Paleontological Con tri bu tions, Pa per, 18: 1–13. The length of ichnogeneric names expressed as number of Barbour, I. H., 1892. No tice of new gi gan tic fos sils. Science , 19: sylla bles. How long is too long? 99–100. Bassler, R. S., 1952. Tax o nomic notes on gen era of fos sil and re - Number of syllables Examples of ichnogenera cent Bryozoa. Jour nal of the Wash ing ton Acad emy of Sci - Fraena Rouault, 1850 ences, 42: 381–385. Bentz, A., 1929. Fossile Röhrenbauten im Unterneokom des Specus Stephenson, 1952 2 Isterberges bei Bentheim. Jahrbuch der Preussische Stichus Etheridge, 1904 Geologische Landesanstalt, 49(2) [for 1928]: 1173–1183. Truncus Fritsch, 1908 Berger, W., 1957. Eine spiralförmige Lebensspur aus dem Rupel Chondrites Sternberg, 1833 der bayrischen Beckenmolasse. Neues Jahrbuch für Mine- Eubrontes Hitchcock, 1845 ralogie, Geologie, und Paläontologie, Monatshefte, 1957: 538–540. Gordia Emmons, 1844 3 Biernat, G., 1961. Diorygma atryophilia n. gen. n. sp. – a par a sitic Oichnus Bromley, 1981 or gan ism of Atrypa zonata Schnur. Acta Palaeontologica Skolithos Haldeman, 1840 Polonica, 6: 17–28. Tisoa Serres, 1840 Bill ings, E., 1861–1865. Palaeozoic Fossils, vol. 1: Con tain ing De scrip tions and Fig ures of New or Lit tle Known Spe cies of Arthrophycus Hall, 1852 Organic Re mains from the Si lu rian Rocks. Geo log i cal Sur vey Belorhaphe Fuchs, 1895 of Can ada, 426 pp. Chelonipus Rühle v. Lilienstern, 1939 Bill ings, E., 1873. Palaeozoic Fossils. Vol. 2: Geo log i cal Sur vey 4 Nereites MacLeay, 1839 of Can ada, 144 pp. Otozoum Hitchcock, 1847 Boekschoten, G. J., 1970. On bryo zoan bor ings from the Danian at Fakse, Denmark. In: Crimes, T. P. & Harper, J. C. (eds), Planolites Nicholson, 1873 Trace Fos sils. Geo log i cal Jour nal, Spe cial Is sue, 3: 43–48. Asterosoma Otto, 1854 Bornemann, J. G. 1889. Über den Buntsandstein in Deutsch land Buchholzbrunnichnus Germs, 1973 und seine Bedeutung für die Trias. Beiträge zur Geologie und Circapalmichnus Gand, 1977 Paläontologie, 1: 1–61. 5 Didymaulichnus Young, 1972 Borrello, A. V., 1966. Paleontografia Bonaerense. Fasc. 5, Tra- zas, restos tubiformes y cuerpos problematicos de la Forma- Ophiomorpha Lundgren, 1891 cion La Tinta, Si er ras Septentrionales – Provincia de Bue nos Taphrelminthopsis Sacco, 1888 Ai res. Provincia de Bue nos Aires, Gobernación, Comisión Arenicolites Salter, 1857 Investigaciónes Científica, Notas, La Plata, Argen tina, 42 pp. Diplocraterion Torell, 1870 Brady, L. F., 1947. In ver te brate tracks from the Coconino sand - stone of north ern Ar i zona. Jour nal of Pa le on tol ogy, 21: 466– Jeholosauripes Yabe, Inai & Shikama, 1940 6 472. Spirocosmorhaphe Seilacher, 1989 Brady, L. F., 1949. Oniscoidichnus, new name for Isopodichnus Steigerwaldichnium Kuhn, 1937 Brady 1947 not Bornemann 1889. Jour nal of Pa le on tol ogy, Tyrannosauropus Haubold, 1971 23: 573. Alcyonidiopsis Massalongo, 1856 Branson, E. B. & Mehl, M. G. 1932. Footprint re cords from the Pa leo zoic and Me so zoic of Mis souri, Kan sas, and Wy o ming. Parahaentzschelinia Chamberlain, 1971 Geo log i cal So ci ety of Amer ica Bul le tin, 43: 383–398. Peratodactylopus Sarjeant & Mossman, 1978 Brönnimann, P., 1955. Microfossils incertae sedis from the Upper 7 Protovirgularia M’Coy, 1850 Juras sic and Lower Cre ta ceous of Cuba. Micropaleontology, Pseudopolydorites G³azek, Marcinowski & 1: 28–51. Wierzbowski, 1971 Brönnimann, P. & Norton, P., 1980. On the classi fi ca tion of fos sil Punctatumvestigium Butts, 1891 fae cal pellets and de scrip tions of new forms from Cuba, Gua- temala and Libya. Eclogae Geologicae Helvetiae, 53: 832– 842. Bromley, R. G., 1970. Borings as trace fos sils and Entobia Early Mis sis sip pian trace fos sil. Jour nal of Pa le on tol ogy, 53: cretacea Portlock, as an ex ample. In: Crimes, T. P. & Harper, 1155–1159. J. C. (eds), Trace Fos sils. Geo log i cal Jour nal, Spe cial Is sue, Austin, D., 1993. The nu ance and wit of Ca ro lus Linnaeus. The 3: 49–90. Pal metto, 13(4): 8. Bromley, R. G., 1981. Concepts in ichnotaxonomy il lustrated by Baird, D., 1980. A prosauropod di nosaur trackway from the Na - small round holes in shells. Acta Geológica Hispánica, 16: vajo Sand stone (Lower Ju ras sic) of Ar i zona. In: Jacobs, L. L. 55–64. (ed.), As pects of Ver te brate His tory: Es says in Honor of Bromley, R. G., 1990. Trace Fos sils: Bi ol ogy and Taphonomy. Edwin Harris Colbert. Mu seum of North ern Ar i zona Press, Unwin Hyman, Lon don, 280 pp. pp. 219–230. Flag staff, Ar i zona. Bromley, R. G., 1996. Trace Fos sils: Bi ol ogy, Taphonomy and Baldwin, C. T., 1977. The stra tig ra phy and fa cies as soci a tions of Ap pli ca tions, 2nd Edi tion. Chap man & Hall, Lon don, 361 pp. trace fos sils in some Cambrian and Or do vi cian rocks of north Bromley, R. G. & Asgaard, U., 1972. Notes on Greenland trace west ern Spain. In: Crimes, T. P. & Harper, J. C. (eds), Trace fos sils: III. A large ra di at ing bur row-sys tem in Ju ras sic mica- Fos sils 2. Geo log i cal Jour nal, Spe cial Is sue, 9: 9–40. ceous sand stones of Jameson Land, East Greenland. Rap port Bandel, K., 1973. Trace fos sils from two Upper Penn syl va nian GrÝnlands Geologiske UndersÝgelse, 49: 23–30. CON STRUC TION OF ICHNOGENERIC NAMES 543

Bromley, R. G. & Ekdale, A. A., 1982. Ichnofabric and early Clarke, J. M., 1921. Or ganic de pend ence and disease: Their or i gin diagenesis: exam ples from Upper Creta ceous chalk, Den- and sig nif i cance. Bul le tin of the New York State Mu seum, mark. Geo log i cal So ci ety of Amer ica, Ab stracts with Pro- 221–222 [for 1919], 1–113. grams, 14(7): 452. Codez, J., 1957. Études de CirripÀdes acrothoraciques fossiles. Bromley, R. G. & Fürsich, F. T., 1980. Com ments on the pro posed DiplÛme d’Études supérieures, Poitiers. amend ments to the In ter na tional Code of Zoo log i cal No men- Colbert, E. H. & Merrilees, D., 1967. Cre ta ceous di nosaur foot - cla ture re gard ing ichnotaxa. ZN (S.) 1973. Bul le tin of Zoo log - prints from Western Aus tralia. Jour nal of the Royal So ci ety of i cal No men cla ture, 37: 6–10. West ern Aus tra lia, 50: 21–25. Bromley, R. G. & Surlyk, F., 1973. Bor ings pro duced by brachi o - Cox, E. T., 1874. Collettosaurus indianaensis Cox. In: Collett, J., pod pedi cles, fos sil and re cent. Lethaia, 6: 349–365. Ge ol ogy of War ren County. Fifth Annual Report of the Geo- Brongniart, A. T., 1822. Sur la clas sifi ca tion et la distri bu tion des log i cal Sur vey of In di ana, made dur ing the year 1873, 247– végétaux fossiles en général, et sur ceux des terrains de sédi- 248. ment, supérieur en particulier. Muséum d’Histoire Naturelle, Crimes, T. P., 1970. Tri lo bite tracks and other trace fos sils from Mémoires, 8: 203–240, 297–348. the Up per Cam brian of North Wales. Geo log i cal Jour nal, 7: Brongniart, A. T., 1849. Tab leau des gen res de végétaux fossiles 47–68. considérées sous le point de vue de leur dis tri bu tion géologi- Dahmer, G., 1937. Lebensspuren aus dem Taunusquarzit und den que. In: d’Orbigny, A. (ed.), Dictionnaire Universel d’His- Siegner Schichten (Unterdevon). Jahrbuch der Preussische toire Naturelle, 13: 52–173. Geologische Landesanstalt, 57 [for 1936]: 523–539. Bronn, H. G., 1837–1838. Lethaea Geognostica oder Abbildungen Davitashvili, L. Sh., 1945. Tsenozy zhivykh organizmov i organi- und Beschreibungen der für die Gebirgs-Formationen bezei- chieskikh ostatkov [Biocoenoses of liv ing or gan isms and of chnendsten Versteinerungen. E. Schweizerbart, Stuttgart, v. or ganic re mains]. Soobshcheniya Akademii Nauk Gruzhin- 1, p. 1–672 (1837); v. 2, p. 673–1350 (1838). skoy SSR, 6(7): 527–534. [Bu reau de Recherches Géologi- Brönnimann, P. & Norton, P., 1960. On the classi fi ca tion of fos sil ques, Géophysiques et MiniÀres, Traduction no. 2077.] fae cal pellets and de scrip tion of new forms from Cuba, Gua - Dawson, J. W., 1890. On bur rows and tracks of in ver te brate an i - temala and Libya. Eclogae Geologicae Helvetiae, 53: 832– mals in Palaeozoic rocks, and other mark ings. Quar terly 842. Jour nal of the Geo log i cal So ci ety of Lon don, 46: 595–617. Brown, L. (ed.), 1993. The New Shorter Ox ford Eng lish Dic tio - Delair, J. B., 1963. Notes on Purbeck fossil foot prints, with de - nary on His tor i cal Prin ci ples, 4th Edition . Clar en don Press, scrip tions of two hith erto un known forms from Dorset. Pro- Oxford, 2 v., 3801 pp. ceed ings of the Dorset Nat u ral His tory and Ar chae o log i cal Brown, R. W., 1956. Com po si tion of Sci en tific Words, 2nd Edition . So ci ety, 84: 92–100. Wash ing ton, DC, Smith so nian In sti tu tion Press, 882 pp. Demathieu, G., Ginsburg, L., Guérin, C. & Truc, G., 1984. Étude Buatois, L. A. & Mángano, M. G. 2013. Ichnodiversity and ichno- paléontologique, ichnologique et paléoécologique du gîse- dis par ity: sig nif i cance and ca ve ats. Lethaia, 46: 281–292. ment oligocÀne de Saignon (bassin d’Apt, Vau cluse). Bul le tin Buck land, W., 1829. A pa per by Dr. Buck land ... 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