The fate and status of the supposed tertiarius Scudder, 1885 J.A. Dunlop

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J.A. Dunlop. The fate and status of the supposed fossil tick Ixodes tertiarius Scudder, 1885. Acarolo- gia, Acarologia, 2011, 51 (4), pp.399-404. ￿10.1051/acarologia/20112021￿. ￿hal-01600228￿

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Acarologia is under free license and distributed under the terms of the Creative Commons-BY-NC-ND which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original author and source are credited. Acarologia 51(4): 399–404 (2011) DOI: 10.1051/acarologia/20112021

THE FATE AND STATUS OF THE SUPPOSED FOSSIL TICK IXODES TERTIARIUS SCUDDER, 1885

Jason A. DUNLOP

(Received 19 August 2011; accepted 27 September 2011; published online 20 December 2011)

Museum für Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity at the Humboldt University Berlin Invalidenstrasse 43, D-10115 Berlin, Germany. [email protected]

ABSTRACT — A fossil described as Ixodes tertiarius Scudder, 1885 from Wyoming, USA represents the only non- record of a tick (Acari: Ixodida) from the Eocene. The original illustration does not include characters indicative of any particular modern tick ; or even Ixodida in general. Unfortunately, efforts to track down the type specimen were unsuccessful and it is almost certainly lost. Confusion about the original place of publication is addressed, and given that there is an illustration associated with the original mention of the name it is not a nomen nudum. However, since it is impossible to verify the accuracy of this illustration against a type specimen – and given the absence of convincing taxonomic features in the figure – Ixodes tertiarius is considered here a nomen dubium which should be excluded from the tick fossil record. KEYWORDS — Acari; Ixodida; fossil; Wyoming; Eocene; nomen dubium

INTRODUCTION or genera found as subfossils in, for exam- ple, the ear canal of a fossil rhinocerous (Schille Fossil parasitiform are surprisingly rare. 1916), coprolites of cats (Guerra et al. 2003), or a fos- While a few opilioacarid and gamasid mites are silised owl pellet (Sanchez et al. 2010). Reviews of known (e.g. Witali´nski2000; Dunlop et al. 2010), the tick fossil record – and their wider evolutionary nine of the fifteen curently valid fossil species in origins – can be found in, e.g., Klompen et al. (1996), the literature are (Ixodida). These incude Cre- Fuente (2003) and Mans and Neitz (2004). taceous records from the ca. 100 Ma Myanmar (Burmese) amber (Poinar and Brown 2003; Poinar There is another enigmatic fossil assigned to and Buckley 2008) and the slighty younger, 90 – 94 a living genus of ticks: Ixodes tertiarius Scudder, Ma, Raritan (or ) amber (Klompen and 1885 from the Eocene Green River Formation of Grimaldi 2001). From the Eocene there are records Wyoming, USA. For a recent account of this well from the 45 – 49 Ma Baltic amber (Weidner 1964; known (ca. 50 Ma) lake deposit, which has yielded Fuente 2003) and from the Miocene from the ca. 16 insects and – as well as vertebrates in- Ma Dominican Republic amber (Poinar 1985; Lane cluding fish, and bats – see Smith et al. (2008) and Poinar 1986; Keirans et al. 2002; Fuente 2003). and references therein. Note that citations to an The dataset also includes essentially modern tick Oligocene (ca. 30 Ma) age are erroneous. Ixodes ter- http://www1.montpellier.inra.fr/CBGP/acarologia/ 399 ISSN 0044-586-X (print). ISSN 2107-7207 (electronic) Dunlop J.A. tiarius was originally named and figured in the Ger- ple – derived from the A. S. Packard collection. man paleontological textbook Handbuch der Palaeon- Alpheus Spring Packard (1839-1905) was a well- tologie. However this consists of a drawing (repro- known entomologist and paleontologist, who held duced here as Fig. 1) and a name only, and lacks a professorship in zoology and geology at Brown a formal description in the text. The isolated fig- University, Rhode Island, USA from 1878 onwards. ure was reproduced (Scudder 1887) in the French Enquires at the geological museum at Brown Uni- volume Traité de Paléontologie. Scudder (1890: 47) versity did not turn up any record of Packard’s ma- again reproduced his earlier drawing, but this time terial (Douglas Morse, pers. comm.). However, it included a formal description, in which he stated was suggested that the specimen could have found ". . . its general appearance and size make it tolera- its way to the Museum of Comparative Zoology bly evident that it belongs to the or Ricini (MCZ) at Harvard University; the largest major mu- and probably to Ixodes proper." The fossil was de- seum in the vicinity. scribed as having a length of 3.5 mm and a width Indeed Laura Leibensperger (MCZ) confirmed of 1.75 mm; although significantly the hypostome that there is an empty box in their collections with [as "rostrum"] was believed to be missing while the a label which reads "Ixodes tertiarius Scudder. Zit- shape and disposition of the legs appear to have tel, Handb. 1885, 1, th 2, /733, fig. 906". There been the main features which led Scudder to pro- is also an entry in the database – probably from pose ixodid affinities. A weakly-defined scutum [as the retired curatorial associate Ardis Johnston (L. "frontal shield"] was alluded to in the text, but was Leibensperger, pers. comm.) – which reads: "Type ". . . (not represented in the plate and perhaps illu- was A. S. Packard specimen no. 258. MCZ only sory) . . . ". has label for specimen, in Henshaw’s handwrit- Scudder’s species was briefly mentioned in a ing. Prof. Frank Carpenter found that the Packard surprisingly large number of later works (see syn- specimens of Scudder’s types were in the introduc- onmy list below) but, apart from some hints that tory paleontology teaching collection at Brown Uni- its affinities might be questionable (Schulze 1929; versity. When he checked specimens remaining at Vitzthum 1942), generally without detailed com- Brown, most of the Scudder types were missing. He ment on its validity. Given that oldest part of the took the insect types. It is possible that other types Green River formation marginally predates Baltic are still at Brown. Fossil insects of North America, amber, Ixodes tertiarius has occasionally been explic- vol. 2: 47." itly listed as the oldest putative record of the fam- ily Ixodidae (cf. Petrunkevitch 1955; Selden 1993). A further note from Ardis in the empty specimen Most recently, in their discussion of problematic tick box reads: "Ixodes tertiarius Scudder, 1885: The type names Guglielmone et al. (2009: 307) suggested specimen was part of the Packard fossil collection. that I. tertiarius could be a nomen nudum – see also At the time, Packard was the state entomologist for Guglielmone and Nava (2011) – but concluded that , but at a later date he went to Brown they had not seen the original material and that: University as a professor, taking his fossil collection "It would be worthwhile to locate this fossil, whose with him. Regrettably, after his death his collection, study may shed light on tick evolution." In light of including type specimens, was used for about 40 this – and as part of a wider project to stabilise fos- years as student material in Brown’s introductory sil nomenclature – an attempt was made paleontology labs. In the 1930’s Dr. Carpenter dis- to track down Scudder’s type specimen; unfortu- covered what had happened and he rescued what nately without success as documented below. was left of the fossil insect type specimens. It is possible that Scudder’s tick is still at Brown. The type label is in Henshaw’s handwriting. Note: The MATERIALS AND METHODS above information was obtained in a conversation Scudder (1890) stated that the original specimen – with Dr. Frank Carpenter, Sept. 1988. A. Johnston". presumably the holotype and only known exam- Attempts to track the material down to another de-

400 Acarologia 51(4): 399–404 (2011)

FIGURE 1: Facsimile copy of a drawing of Ixodes tertiarius Scudder, 1885 from the Eocene of Green River, Wyoming, USA; reproduced from Scudder (1890, Pl. VI, fig. 12). While possibly a , the illustration does not reveal any obvious characters of Ixodes, or of Ixodida in general. Since the type is missing, presumed lost (see text), the name is treated here as a nomen dubium. partment of the MCZ were unsuccessful. Given that thorship incorrectly assigned to Weidner]; Gugliel- Brown University cannot trace it from their former mone et al., 2009: 316-317; Dunlop et al., 2010: 67; teaching collection either, it seems highly likely that Guglielmone and Nava, 2011: 60. Scudder’s type specimen has, unfortunately, been Material — Fish Cut, Green River, Wyoming, lost. USA. Paleogene (Eocene). Originally part of the A. Ixodes tertiarius Scudder, 1885 (nomen dubium) S. Packard collection, specimen no. 258; missing, (Figure 1) presumed lost. Ixodes tertiarius Scudder, 1885: 733, fig. 906; Scudder, 1887: 732, fig. 922; Scudder, 1890: 47, pl. DISCUSSION VI, fig. 12; Scudder, 1891: 269; Vitzthum, 1924: 732; Chodziesner, 1924: 527; Schulze, 1929: 745; Scudder’s (1885, 1887, 1890) illustration (Figure 1) Petrunkevitch, 1955: 97; Weidner, 1964: 144; Doss does not reveal any obvious characters of hard ticks et al. 1974: 542; Doss and Anastos, 1977: 87; Lane (Ixodidae), such as a characteristic capitulum (i.e. and Poinar, 1986: 75; Sonenshine, 1991: 23; Selden, hypostome + pedipalps), a well defined scutum 1993: 310; Klompen et al., 1996: 146; Camicas et al., and/or marginal festoons. At least on the right side 1998: 167; Walter and Proctor, 1999: 16 [erroneously five or even six appendages are preserved which as an amber specimen]; Witali´nski,2000: 620; Klom- raises questions about which of them are walking pen and Grimaldi, 2001: 10; Keirens et al. 2002: 61; legs; e.g. are the anteriormost limbs the pedipalps, Fuente, 2003: 333; Guerra et al., 2003: 181; Mans and in which case they seem to be unusually large and Neitz, 2004: 2; Robbins and Robbins, 2004: 13 [au- robust for a mite? The overall habitus could per-

401 Dunlop J.A. haps fit better to the more amorphous dorsal body being named . . . " For this reason, Scudder’s (1885) surface of a soft tick () – and as noted inclusion of a drawing of the fossil does make his above, both bats and birds (as potential hosts) have name available and it need not be treated as a nomen been recorded from the Green River locality (Sim- nudum. Nevertheless, the drawing is equivocal in a mons et al. 2008; Weidig 2010) – but this is spec- number of key features, and even the subsequent ulative, and again lacks explicit character support. accompanying description by Scudder (1890) does Looking at the inflated anteriormost appendages, if not enable the fossil to be placed convincingly in these were leg 1, a case could be made for the fos- any particular modern genus or even higher group sil being something like a large velvet mite (Trom- of mites. In the interests of stability, and given the bidiidae), but again this would be based on the out- absence of a type specimen, I. tertiarius is proba- line appearance of the rather than diagnostic bly best treated as a nomen dubium and should be characters. While Ixodes tertiarius could conceivably excluded from further discussions of the tick fossil be a mite of some description, in the absence of a record and their evolution. type (see above), and in light of the equivocal na- ture of the drawing (see also comments in Vitzthum ACKNOWLEDGEMENTS 1942), an exact assignment to any particular group now seems to be impossible. I am especially grateful to Patricia Davey and Dou- glas Morse (both Brown University) and Laura With respect to the status of the name, Vitzthum Leibensperger (MCZ) for their kind assistance in (1942) inadvertently assigned authorship of Ixodes trying to track down Packard’s specimens, and to tertiarius to Scudder (1884) [as 1885] – a paper care- the reviewers for helpful comments. fully checked by the present author and which is clearly restricted to Palaeozoic with no men- tion of I. tertiarius, or any other Acari. Vitzthum’s REFERENCES error was followed by other authors (e.g. Weidner Camicas J.-L., Hervey J.-P., Adam F., Morel P. C. 1998 — 1964; Lane and Poinar 1986; Klompen et al 1996; Les Tiques du monde. Paris: Orstom. pp. 233. Guglielmone et al. 2009), all of whom cite the wrong Chodziesner, M. 1924 — Beitrag zur Kenntnis der paper as the original description. Guglielmone et Zecken mit besonderer Berücksichtigung der Gattung al. did, in fact, cite the oldest mention of the name Koch. — Zool. Jb., Syst., 47: 505-572. by Scudder (1885) in the Handbuch der Palaeontologie. Doss M. A., Farr, M. M., Roach K. F., Anastos G. 1974 — This is confirmed by Scudder (1891: 269) who cata- Index catalogue of medical and veterinary zoology. — logued all fossil arachnid names up until that time Special publication No. 3. Ticks and tickborne dis- eases I. Genera and species of ticks. Part 2. Genera and cited the Handbuch entry from 1885 as the oldest H-N. Washington: University of & United occurrence of his own name, followed by Scudder States Department of Agriculture. pp. 593. (1887) [erroneously as 1886] and Scudder (1890). Doss M. A., Anastos G. 1977 — Index catalogue of medi- cal and veterinary zoology. — Special publication No. Yet as Guglielmone et al. (2009) noted, Scudder 3. Ticks and tickborne diseases III. Checklist of fami- (1885) consists of a named drawing without an ac- lies, genera, species and subspecies of ticks. Washing- companying diagnosis or text description and thus ton: University of Maryland & United States Depart- these authors suggested that the name is a nomen ment of Agriculture. pp. 97. nudum. For this to be the case Ixodes tertiarius – pub- Dunlop J. A., Sempf C., Wunderlich J. 2010 — A new opil- lished before 1931 – must fail to conform to Article ioacarid in Baltic amber. — In: Nentwig W., Entling M., Kropf C. (Eds) European Arachnology 2008. Bern: 12 of the International Code of Zoological Nomen- Natural History Museum. p. 59-70. clature which states that the name must be accom- Fuente J. de la 2003 — The fossil record and origin of ticks panied by an "indication". Among the permissi- (Acari: : Ixodida). — Exp. Appl. Ac- ble indications, Article 12.2.7 lists "the proposal of arol., 29: 331-344. doi:10.1023/A:1025824702816 a new genus-group name or of a new species group Guerra R. de M. S. N. de C., Gazêta G. S., Amorim M., name in association with an illustration of the taxon Duarte A. N., Serra-Freire N. M. 2003 — Ecological

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Weidig I. 2010 — New birds from the Lower Eocene COPYRIGHT Green River Formation, North America. — Rec. Aust. Mus., 62: 29-44. doi:10.3853/j.0067-1975.62.2010.1544 Weidner H. 1964. — Eine Zecke, Ixodes succineus sp. n. im Dunlop. Acarologia is under free license. Batischen Bernstein. — Veröff. Überseemus. Bremen This open-access article is distributed under the terms of A, 3: 143-151. the Creative Commons-BY-NC-ND which permits unre- Witali´nskiW. 2000 — Aclerogamasus stenocornis sp. n., a stricted non-commercial use, distribution, and reproduc- fossil mite from the Baltic amber (Acari: Gamasida: tion in any medium, provided the original author and ). — Genus, 11: 619-626. source are credited.

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