Identity of Three New Sea Turtles Named by J. Friedrich Eschscholtz
Abstract
Johann Friedrich Eschscholtz published two herpetological titles, the first in a journal named Quatember. Until now, this paper, which contains descriptions of three sea turtle taxa, including Chelonia olivacea, has been overlooked, and the nomenclatural implications have been ignored. The Quatember paper preceded, and has precedence over, Eschscholtz's better known Zoologischer Atlas, and therefore is the earliest description of C. olivacea. However, Plate III of the Atlas remains the earliest illustration for what is today called Lepidochelys olivacea.
Johann Friedrich Eschscholtz (1793–1831; for biography, see Adler 2007) was director of the Zoological Cabinet at the Imperial University of Dorpat, Russia (today Tartu, Estonia). Most of his publications are entomological, but he authored two titles in herpetology, both in the year 1829 (Eschscholtz 1829, 1829–1833). Of the two herpetological titles by Eschscholtz, the only one that has been remembered is his book, Zoologischer Atlas, which reports on the zoological collections made during the circumnavigation of the Russian warship, Predpriaetië, from 1823 to 1826. It includes descriptions of 4 Californian salamanders, Salamandrina attenuate (= Batrachoseps attenuatus), Triton ensatus (= Dicamptodon ensatus), Triton torosus (= Taricha torosa, attributed to Rathke), and Triton tereticauda (= Aneides lugubris), and 2 reptiles from the Philippines, an agamid lizard, Istiurus pustulatus (= Hydrosaurus pustulatus), and a sea turtle, Chelonia olivacea (= Lepidochelys olivacea) (Adler 2007). The original description of the turtle has long been credited to Eschscholtz's Atlas, in which publication of this taxon is based on 2 specimens. However, we have discovered that the original description of L. olivacea did not occur in the Atlas.
Several months before the publication of the first installment of the Atlas, in the January 1829 issue of Quatember (published in Mitau, Russia; today Jelgava, Latvia), Eschscholtz published descriptions of 3 new species of sea turtles, including C. olivacea. In the introduction to this article, Eschscholtz stated that of the 7 species of sea turtles contained in the Zoological Cabinet, 4 had been named, all belonging to the genus Chelonia, namely, Chelonia cephalo (= Caretta caretta), Chelonia mydas or Chelonia esculenta (= C. mydas), Chelonia virgata (= C. mydas), and Chelonia imbricata (= Eretmochelys imbricata). Because he considered that there were 3 additional undescribed species, he decided to name them Chelonia castanea from Surinam, Chelonia grisea (= E. imbricata) from the Caspian Sea (locality in error, see below), and C. olivacea (= L. olivacea [Eschscholtz 1829]) from Manila Bay, Philippines.
This article provides a detailed evaluation of these 3 new species erected by Eschscholtz. Unfortunately, the specimens (syntypes) on which these descriptions were based seem to be lost (see below); hence, the evaluations are based solely on the descriptions in Eschscholtz (1829). No other taxa are described in the Quatember article. They were translated from German to English by one of us (T.G.E.).
Chelonia castanea
This species is described on the basis of 3 specimens preserved in spirits (Eschscholtz 1829, p. 11), but Eschscholtz only gave measurements for one of them (carapace length [CL] 6.35 cm and carapace width [CW] 5 cm). The diagnostic characters used by Eschscholtz are 13 large dorsal scutes, thick and non-transparent brown coloration; middle vertebral scutes three times wider than long, with 6 sides, the anterior and posterior sides parallel, those on each side with 2 edges that come together at a sharp angle; 25 marginals, acute along the outer edges giving the appearance of saw teeth. Plastron is faintly yellow, with 13 middle scutes (excluding inframarginals), a small scute (intergular) inserted between the first pair of middle pectoral scutes. Head thick and rounded, and the snout protruding; head shields dark brown; frontoparietal (“the anterior rear main shield”) very large with 7 sides, the 2 parietals (“the two posterior ones”) slender and oddly arranged. The lower jaw terminates in a hook, and the cutting edges of the beak are not saw-toothed. The flippers are shorter than they are in other species; they possess a single claw per flipper. The measurements indicate a recently hatched animal, and the description is easily assignable to C. mydas. The specimens came from Surinam where the species is well known and documented (Shultz 1975) (Fig. 1). The number of dorsal scutes, excluding the marginals (13), the shape and number of the parietals, and having only one claw on each flipper differentiate it from other genera such as Lepidochelys and Caretta. This is not a specimen of Eretmochelys, by the coloration of the plastral scutes that are cream in C. mydas (including some population from the East Pacific) but brown to black in E. imbricata hatchlings (Fig. 2). The name C. castanea Eschscholtz (1829) should, therefore, be added to the list of synonyms of C. mydas. Unfortunately the specimens (syntypes) on which the description is based seem to be lost, as are other specimens that Eschscholtz described (see below).



Citation: Chelonian Conservation and Biology 15, 1; 10.2744/CCB-1190.1



Citation: Chelonian Conservation and Biology 15, 1; 10.2744/CCB-1190.1
Chelonia grisea
The description of C. grisea is based on 1 juvenile specimen (CL 13.9 cm). The description includes 13 large and very thin scutes on the carapace that are yellowish gray in coloration and from which the ribs are visible; along the vertebral scutes, there is a black ridge; the central scutes are almost twice as long as wide, the 4 most anterior ones have 6 sides; and there are 25 marginals that do not form a saw-toothed edge. The plastron has 2 weakly pronounced ridges that run lengthwise; the entire plastron is whitish. The number of middle scutes is 13 (excluding inframarginals); as in the previous species, a small scute (intergular) is inserted between the first anterior pair. The head is slender and pointed, covered with brownish shields; the anterior main shield (frontoparietal) has an 8-sided appearance; there are 3 equally sized parietals, arranged in a row, as C. cephalo (= C. caretta). The neck and eyelids are smooth, throat finely scaled; the lower jaw is hook-shaped and contains saw-tooth–shaped margins. Each flipper has a pointed claw; anterior limbs are very long and narrow.
In the last paragraph of his description, Eschscholtz compares this taxon with Testudo japonica (= C. mydas). He states that these 2 taxa are similar in the number of dorsal scutes; the presence of a ridge on the backbone (vertebral zone); the thin scutes through which the ribs are visible; and the marginals that do not form a saw-toothed edge. The differences between these taxa are the black coloration of the dorsal scutess (in T. japonica); the vertebral scutes that are nearly as wide as long (in T. japonica); high and protruding longitudinal dorsal ridge (in T. japonica); the presence of only 2 parietals (in T. japonica); a wide and long snout (in T. japonica); and shorter and wider front feet (in T. japonica).
According to Eschscholtz, a note attached to the specimen stated its origin as the Caspian Sea; he remarked that, if this information was correct, this would represent a record of a marine animal from a lake (Eschscholtz 1829, p. 13) (the Caspian Sea is saline and the largest inland body of water in the world). Clearly, this is an erroneous locality, or the animal was introduced by human agency.
Chelonia grisea is listed in the synonym list of E. imbricata by Fritz and Havaš (2007), who give credit for the name to Eschscholtz in the journal, Quatember. They also noted the erroneous type locality, but there is no mention of the type material, one juvenile. Fritz and Havaš (2007) did not remark on the other new sea turtle names mentioned in Eschscholtz's article.
Based on the description given above and our examinations of hatchlings and juveniles, as well of photographic material of C. mydas and E. imbricata from the Caribbean and western Atlantic region, we conclude that this taxon cannot represent the latter species. We base our conclusion on the following characters: hatchlings and juveniles of E. imbricata have dorsal ridges on the vertebrals and costals, giving the appearance of 3 dorsal ridges on the carapace (Fig. 2), not just the vertebral ridge; the marginals in adult E. imbricata have the appearance of a saw-tooth edge, whereas in a C. mydas (CL 36.5 cm), the appearance of saw teeth on the marginals is almost absent; hatchlings of E imbricata have a brown plastron (Fig. 2), not whitish, although this coloration becomes lighter with age, juveniles still show dark patches on the ventral scutes (Fig. 3); the head is slender and pointed in C. mydas, the snout is not pointed in small E. imbricata; the lower jaw of E. imbricata hatchlings and juveniles do not have saw-tooth–shaped teeth; the anterior flippers are proportionally longer in C. mydas compared with those of E. imbricata. Other characters are either similar between the 2 species (such as 13 large scutes on the carapace [excluding marginals], the anterior ones with 6 sides; 25 marginals; 13 middle [excluding the inframarginals] ventral scutes, with a small scute [intergular] inserted between the first anterior pair; head covered with brownish shields; lower jaw terminates in a hook) or are variable in these and other taxa (form and number of sides in the frontoparietal; number of parietals). Other characters are difficult to assess; these are thin dorsal scutes through which the ribs are visible (protruding ribs); center dorsal scutess are almost twice as long as wide; neck and eyelids are smooth; and the throat is finely shielded. Unfortunately, there is no mention of the condition of the prefrontals in C. grisea, which are divided in E. imbricata and undivided in C. mydas.



Citation: Chelonian Conservation and Biology 15, 1; 10.2744/CCB-1190.1
We believe that Eschscholtz's description actually refers to an anomalous C. mydas, but certainly not an E. imbricata. Therefore, the name Chelonia grisea Eschscholtz (1829) should be added to the synonym list of C. mydas and removed from that of E. imbricata.
Chelonia olivacea
The description of this taxon is not problematic because it was illustrated in Eschscholtz's Zoologischer Atlas. Comparing the written description in Quatember with the published plate, it is easy to follow. It is worth noting that in his Quatember paper, Eschscholtz also described a juvenile of L. olivacea (CL 17.7 cm, CW 16.51 cm, head length [HL] 5.7 cm, head width [HW] 3.1 cm) and an adult (CL 72.3 cm, CW 60.9 cm; HL 15.2 cm, HW14.6 cm). The juvenile specimen is depicted in plate III of the Atlas at about 72% its original size.
From the date stated on the initial page of volume 1 of the journal Quatember, it is clear that this version of the description takes precedence over that in the Atlas (see below); permission for publication was granted on 9 January 1829. Permission for publication was granted for issue 2 of that same volume on 29 May 1829; it is logical to think that issue 1 was already published by then. Eschscholtz's preface in the Atlas, which was published together with part I (containing the description of L. olivacea), is dated “Dorpat im Mai 1829” (May 1829). If we take this statement literally, then publication of part I of the Atlas had to have occurred no earlier than May 1829, and probably in June or later, given that the book was printed in Berlin. Therefore, the original publication of the name, C. olivacea, was in the journal and not in the Atlas.
Concerning the 2 specimens of C. olivacea described by Eschscholtz, Zug et al. (1998) mentioned that Eschscholtz examined 2 specimens (syntypes) “but not explicitly designated any, possibly deposited in Museum of Tartu but never reported, thus considered lost or destroyed . . .”. They credited this information to Smith and Smith (1979). Apparently, Zug et al. (1998) confused Eschscholtz's specimens with that cited by Kuhl (1820), the last one was part of “. . . the collection of Mr. Kuypers at Groningen, apparently . . . lost . . .” (Brongersma 1961; Smith and Smith 1979). There is no statement that the type used by Eschscholtz was lost or destroyed in Smith and Smith (1979). According to A. Miljutin (pers. comm., September 2009), Curator of Vertebrates at the Tartu Museum, Estonia, Eschscholtz's specimens were sold by his widow to the Zoological Museum of Moscow State University (ZMMU) in 1837. There are 2 catalogs of specimens and types deposited at the Moscow State University Museum (Kulagin 1888; Dunayev and Orlova 2001), neither of which list any Eschscholtz specimens. V. Orlova (pers. comm., October 2009), Curator of Reptiles at the Zoological Museum of Moscow State University, informs us that there are no records of transfers from the Tartu Museum, and apparently only entomological specimens from Eschscholtz exist on record. Therefore, we conclude that the type material of L. olivacea Eschscholtz is probably lost.
Additionally, Fritz and Havaš (2007) listed the type locality as “Manila Bay, Philippine Islands and Sumatra”; there is no mention in the original description of the specimens coming from Sumatra. Therefore, the type locality should be as given in the original description “Manila Bay” [Philippine Islands].
There has been a long history of confusion regarding the name, identity, and authorship of species of sea turtles (see for example; Brongersma 1961; Rhodin and Smith 1982; Bour and Dubois 1984; Frazier 1985; Smith and Rhodin 1986; Parham and Zug 1996). In particular, L. olivacea was confused for a long time with the loggerhead (Deraniyagala 1933, 1939a, 1939b, 1952; Parker 1939); see discussion in Smith and Smith (1979). Part of the problem has been the lack of type material (see Parker 1939). Although today it would be difficult to mistake the 7 recognized species of turtles, it seems appropriate to reproduce the plate III of the Zoologischer Atlas depicting the type specimen for L. olivacea that is lost (Fig. 4). In fact, the description of the small specimen is given first by Eschscholtz in both publications (Quatember and Atlas).



Citation: Chelonian Conservation and Biology 15, 1; 10.2744/CCB-1190.1
Finally, a note about the reprint of the herpetological sections of the Zoologischer Atlas that were issued in 1966 by the Ohio Herpetological Society (today, Society for the Study of Amphibians and Reptiles). This reprint was made from an original copy of the atlas in the library of the University of Michigan Museum of Zoology, in which the text of part I (only, but including the description of L. olivacea) is duplicated. The two versions are practically identical in content, but one is set in smaller type. This one appears to have been a preliminary version that was later discarded, because the version in larger type is the format followed in the rest of the work and is the version we have seen in other copies of the book. The Michigan copy is the only one that we can locate that includes this alternative version of part I.

Dorsal and ventral views of a hatchling of Chelonia mydas USNM 293594, from Surinam. Photo taken by J. Poindexter, reproduced here with permission of the Smithsonian Institution. Diagnostic characters used by Eschscholtz are the number of dorsal and marginal scutes, as well as the shape of the middle vertebral scutes, and the edges of marginals that seem like a saw teeth. There is only 1 dorsal ridge. The light plastron and dark dorsal color of head with 2 parietals. See text for more details.

Dorsal and ventral views of a hatchling of Eretmochelys imbricata USNM 163069, from Surinam. Photo taken by J. Poindexter, reproduced here with permission of the Smithsonian Institution. The number of dorsal and marginal scutes are the same as those on the specimen from Fig. 1. This species has 3 dorsal ridges on the carapace, and the plastron is not light in color (whitish). The snout is not pointed. See text for more details.

Ventral view of juveniles of Eretmochelys imbricata: (A) from Chenkán, Campeche, Mexico, in captivity in Ciudad del Carmen, Campeche, Mexico (photo by V. Matos Mendez, reproduced with permission); (B) from Isla Mona, Puerto Rico (photo by R. Van Dam, reproduced with permission). Notice the black marks on the ventral scutes of juveniles. See text for more details.

Plate III from Eschscholtz's “Zoologischer Atlas” depicting the juvenile of Chelonia olivacea (= Lepidochelys olivacea. Enhanced from the facsimile.
Contributor Notes
Handling Editor: Jeffrey A. Seminoff