The month of April 2016 starts off with April Fools' Day. In addition, the 12th marks 119 years since the death of American paleontologist and herpetologist Edward Drinker Cope (1840-1897). Cope was known for a sense of humor, so on April first it may be appropriate to took at one of his generic names that has sometimes been seen as a joke: *Nimravus*. Cope's derivation and intended meaning of the name *Nimravus* are commonly misunderstood.
The idea that the name *Nimravus* was meant as a joke appears in a number of sources. Charles Lewis Abbott (1862-1927) was a writer, editor, sometime Greek scholar and translator, and amateur naturalist who lived in South Dakota. He self-published a number of short works on evolution, including "Evolution: True or False" in 1890 (said to have been favorably noted by Thomas Huxley) and "What Comes from What; Or, The Relationships of Animals and Plants" in 1922.
His 1922 pamphlet contains this passage:
"The saber-toothed tiger and its entire family are extinct. Nimrod 'was a mighty hunter before the Lord.' Gen. x. 8. Some palaeontologist has humorously named one of the ancestors of the cat *Nimravus* (Nimrod's grandfather)!" (page 43)
The "paleontologist" in question was, of course, Edward Drinker Cope. Unfortunately, Cope himself never explained the name *Nimravus* nor supplied an etymology, so the idea that his generic name *Nimravus* was meant as some sort of a joke based on Nimrod from the Book of Genesis can't be confirmed directly.
Other sources also have assumed that Cope derived the name *Nimravus* from the biblical Nimrod, a great hunter in his youth, who later became a warrior and a powerful ruler, and who, according to tradition, is said to have built the Tower of Babel. The term "nimrod" has come to mean a "hunter" in English.
T. S. Palmer's 1908 Index Generum Mammalium gave this etymology:
*Nimravus*: Nimr(od), hunter; Lat. *avus*, ancestor. (Palmer 1908: 460)
Older dictionaries also have given an etymology from Nimrod as an allusion to a great hunter:
A version of this supposed meaning currently appears in a number of places, including the National Parks guide book "Agate Fossil Beds: Agate Fossil Beds National Monument Nebraska":
"*Nimravus* (ancestral hunter)..." (page 32)
It has even carried over to other languages. In Chinese, *Nimravus* is called *liehu* "hunting tiger," for example.
There is an obvious question: If Cope derived his name *Nimravus* from the biblical name Nimrod as supposed, why didn't he spell it "Nimrodavus" rather than dropping the last part of the name? (For the record, the Greek and Latin forms of Nimrod are *Nebrod*.)
What has apparently been overlooked with the proposed "Nimrod" etymology of Cope's *Nimravus* is that *nimr* itself exists as a word with a relevant meaning. In fact, the current scientific name for the critically endangered subspecies the Arabian leopard is *Panthera pardus nimr* (Hemprich & Ehrenberg, 1833).
http: // www.iucnredlist.org/details/15958/0
So, conceivably, Cope may have derived *Nimravus* from *nimr* (a scientific name used for a type of leopard) with no spelling change, instead of from a chopped off version of the biblical name Nimrod.
A bit of linguistic and zoological history is unavoidable.
Between 1820 and 1825, the German naturalist Christian Gottfried Ehrenberg (1795--1876) conducted a scientific expedition through parts of the Middle East, accompanied by the German zoologist Wilhelm Friedrich Hemprich (1796-- 1825), who tragically died of a fever in Massawa (now in Eritrea), bringing an end to their joint exploration. The Prussian Academy of Sciences in Berlin was the major funder of the expedition.
For more on the expedition, see:
After Ehrenberg returned to Germany, he published a series of illustrated books in Latin (under the title "Symbolæ physicæ") on the wide range of fauna and flora collected. (Confusingly, the text pages were not numbered, although the lithograph illustrations were numbered as plates.) Ehrenberg kept the late Hemprich as his co-author.
The volumes can be viewed at the Biodiveristy Heritage website:
Of note for the name *Nimravus* was Ehrenberg's description and illustration of a species of big cat he named *Felis Nimr*, establishing the Neo-Latin usage of *nimr* for a type of leopard (now the endangered Arabian subspecies *Pathera pardus nimr* (Hemprich & Ehrenberg, 1833) as noted above).
Symbolae physicae seu icones et descriptiones, Mammalium., II: 17 (F. nimr). 1833. (text pages unnumbered)
According to the original description in Latin of *Felis, Pardus?, Nimr*: "today is called Nimr or el Nimr by the Arabs of Syria, Arabia and Nubia, i.e, Spotted, Mottled":
For a hand-tinted lithographic image of *Felis Nimr* in the original publication (image can be downloaded as jpeg) :
The term *nimr* is found in modern Arabic, including as a man's given name (Nimr) or as an honorific epithet to suggest power (Al-Nimr "the Tiger"; "the Panther"), often meaning a tiger, but also a leopard or a panther. Alternate forms and Latin alphabet transcriptions of the word from Arabic dialects include *nimar* and *numair*.
There are many cognates of *nimr* in other Semitic languages in the Middle East that also contain the consonants n + m + r but with differnet vowels: *namer* in Hebrew, etc.
The 18th-century French naturalist Count Buffon (1707-1788) cited "Nemer" as a name used by the Arabs for a large panther:
In English translation:
In 1868 the Austrian zoologist Leopold Fitzinger (1802--1884) erected the species *Panthera Nimr* (pg. 461) as <<der ostafrikanische Panther>> [the East African panther], contrasting with *Panthera pardus* (pg. 457), as the West African panther:
Fitzinger, L. F. J. 1868. Revision der zur natürlichen Familie der Katzen (Feles) gehörigen Formen. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften 58:421–519.
In 19th century sources, the term "nimr" was used for a type of "panther" found in the Middle East:
"....[W]e think that a distinct variety of Panther has not received due consideration by naturalists, except by Ehrenberg. It is the Nimr or Syrian Panther (not Leopard)....," (pg. 180)
Smith, Charles Hamilton (1858). Introduction to Mammalia. Edinburgh,W. H. Lizars.
The "Panther" Puzzle
What is a panther? The term "panther" has a long history in zoology, but with shifting and often confusing meanings.
In modern English usage, the term "panther" is commonly applied to leopards with a melanistic color mutation ("black panthers"), a variation that also occurs in jaguars. Spotted leopards and black panthers are thus individual variations within the same species, and not biologically or taxonomically distinct.
In older literature, however, the term "panther" was commonly used for a supposed species of spotted big cat found in Africa, the Middle East, and Asia that was thought to be distinct from a leopard, based on certain details its physical appearance and build--NOT for a uniform black coloration. Confusingly, the exact distinctions between a panther and a leopard differed according to the author and the geographical region where the cats were found. In addition, the cheetah (Acinonyx jubatus) was sometimes referred to as a "hunting leopard" or a "hunting panther," and even as a *nimr.*
A thorough history of the term "panther" in zoology is not possible here. However, a few major points may be useful.
Buffon distinguished between <<la panthère>> and <<le léopard>> based on size (the panther was generally larger) and on differences in the patterns of spots.
Cuvier distinguished between the panther (*Felis pardus*) and the leopard (*Felis leopardus*) based mainly on their patterns of spots:
For a more detailed discussion of the complicated older nomenclature for spotted big cats (as of 1839 at least), see the Penny Cyclopaedia entry for LEOPARDS (page 430):
The distinction between panthers and leopards (and pards for some authors) was long debated, especially by British naturalists in India. For some authors, the panther was the larger, for others, the smaller, of the two.
"Panther" for larger form, "leopard" for smaller form (pg. 97)
Jerdon, T. C. 1867. The mammals of India.
"Panther" for smaller form, "pard" for larger (pg. 183)
Sterndale, R. A. 1884. Natural History of the Mammalia of India and Ceylon.
Blanford (1891) suggested that the physical differences likely resulted from growth stages and local color variations, and there was only one species (pg. 68).
Blanford, W. T. 1891. Mammalia. Fauna of British India.
In current taxonomy, there is no distinction recognized between what were once called panthers and what were called leopards. As noted, the idea that "panther" is the term for a large cat with black coloration is not the historical distinction.
To add to the linguistic muddle, the term "panther" has also been used for the *Puma concolor*, the mountain lion, cougar, or puma found in the Americas ("Florida panther," "Eastern panther," "American panther," etc.), as well as for the jaguar (*Panthera onca*) ("American panther," "South American panther," "Brazilian panther," etc. ). The ocelot is sometimes called the "Mexican panther."
The term "panther" was used for the puma in the British colonies of Eastern North America dating back to the 17th Century. For example, 18th century colonial laws establishing animal bounties refer to "panthers," along with wolves.
Note that the now widely used term "cougar" is, in fact, a word invented by the 18th century French naturalist Count Buffon as a contracted form "couguar" of the reported Brazilian name *cuguacu ara*:
O.C. Marsh and the Latin "suffix" *avus* "ancestor"
E. D. Cope was noted for coining imaginative and unusual generic names, although the derivations and meanings of some of them have become obscure or misunderstood thanks to his linguistic invention and erudition. By contrast, his rival O.C. Marsh came up with a few famously evocative names such as *Brontosaurus*, but sometimes opted for duller fare in the likes of *Allosaurus* and *Apatosaurus*, or repetitive combinations as with *Limnocyon*, *Limnofelis*, *Limnohyus*, *Limnotherium*, *Limnophis*, and *Limnosaurus*, all for animals that lived around an ancient Eocene lake in Wyoming.
However, Marsh deserves credit for one notable nomenclatural innovation--turning Latin *avus* "grandfather, ancestor* into a kind of suffix to suggest evolutionary relationships in fossil animals.
*Boavus* Marsh, 1871
"... the fossil specimens represent a separate genus, for which the name Boavus* is proposed, in allusion to the not improbable relationship of the two types.
*Boa and avus, grandfather"
His other examples include: *Graculavus* Marsh, 1872; *Iguanavus* Marsh, 1872; *Lemuravus* Marsh, 1875; *Sciuravus* Marsh, 1871; *Talpavus* Marsh, 1872; *Tapiravus* Marsh, 1877; *Viverravus* Marsh, 1872; *Vulpavus* Marsh, 1871.
These so-named taxa were described as being related in some way to the living genus used as the basis for the name, or at least resembling that genus.
In post-Darwinian nomenclature, Marsh's suffix -avus "ancestor" joined the repurposed prefixes pro- "before" and proto- "first" to indicate evolutionary ancestry, either in a general sense or sometimes to suggest a direct link.
Apparently Cope (professional and personal rivalry aside) appreciated Marsh's nomenclatural inspiration and used -avus in the name *Nimravus* to suggest ancestry--the first example of another researcher other than Marsh using -avus in the name of a fossil animal.
What is a less clear is exactly what kind of "ancestry" Cope had in mind.
Cope's Classification of Felids
In 1879 Cope reviewed the living and extinct genera of felids and canids in a short paper, which introduced the generic name *Nimravus* (pg. 169) with the species *Nimravus brachyops* (pg. 170).
He used the generic name *Uncia* for most of the big cats now included in *Panthera*:
"Thus I combine [Gray's] *Uncia*, *Tigris*, *Leo* and *Leopardus* into a genus for which I employ his name *Uncia*, as the 'least objectionable,' after having confirmed by autopsy the circular character of the pupil." (pg. 169)
Cope's list of large living felids included the following (with his usual common name for each species added):
*Uncia pardus* ("leopard"; Cope evidently did not distinguish between an Old World "panther" and a leopard) [*Panthera pardus*]
*Uncia concolor* ("American panther" or "panther") [*Puma concolor*]
*Uncia onca* ("jaguar") [*Panthera onca*]
*Nimravus* and the Nimravidae
The Nimravidae Cope, 1880 are now recognized as an important clade of extinct predators (Eocene to Oligocene) within the Feliformia, but not particularly close to the Felidae in phylogenetic terms. The catlike appearance, and probably catlike behavior, of the nimravids is a notable example of convergent evolution.
For the nomenclatural and taxonomic history of *Nimravus* (with *Nimravus brachyops* Cope, 1878 as the type species), see:
Toohey, L. (1959). The species of *Nimravus* (Carnivora, Felidae). Bulletin of the American Museum of Natural History, 118, 71–112.
For recent discussions of the Nimravidae:
Peigné, S. (2003). Systematic review of European Nimravinae (Mammalia, Carnivora, Nimravidae) and the phylogenetic relationships of Palaeogene Nimravidae. —Zoologica Scripta, 32, 199–229.
Cope published two detailed descriptions of *Nimravus* (1880 and 1884), based primarily on the species he called *Nimravus gomphodus* ("nail tooth", from Greek *gomphos*, here meaning a large wedge-shaped nail, alluding to the straight, pointed shape of the enlarged canines). Both descriptions were preceded by a description of the genus *Archaelurus* that compared that animal to the "American panther" (1880: 842) and to the "panther (*Uncia concolor*)" (1884: 953), establishing that Cope used "panther" to refer to the puma, and not to the Old World "panther."
His 1880 description of *Nimravus* compares it to a "panther" in size, noting that the "panther" (puma) had elongated proportions, contrasting with the "more robust" leopard and jaguar:
"The *Nimravus* gomphodus is as large as the full-grown panther of the large varieties. It probably stood as high above the ground, but whether the body had the elongate proportions of that animal, or the more robust form of the leopard and jaguar, cannot be ascertained in the absence of necessary material." (Cope 1880 pg. 844)
Cope, E. D. 1880. On the extinct cats of America. American Naturalist. XIV 833-858, 15 figs.
His 1884 description notes numerous other points of resemblance to a "panther" (puma), including the general proportions of the skull and the convex frontal and nasal region.
Report of the United States Geological Survey of the Territories, Volume 3
Cope, E. D. 1884. The Vertebrata of the Tertiary formations of the West. Book I. Report of the United States geological survey of the Territories. F. V. Hayden, United States geologist-incharge. Washington (Department of the interior) III xxxv + 1009 pp., 38 figs., pls. I-LXXVa.
In his 1880 discussion of the relationship between the families Nimravidae and Felidae, Cope proposed that some nimravids were the direct ancestors to felid sabertooths:
"There can be no reasonable doubt that the genera *Drepanodon* and *Smilodon* in the Felidae are the descendants of *Hoplophoneus* and allied genera. In fact, the Nimravidae and Felidae are 'homologous groups,' having corresponding terms in the manner I foreshadowed as a general principle in 1868 (Origin of Genera)." (1880: 840)
The "false sabre-tooth" *Nimravus*, however, is not presented as a direct ancestor to later felids. Cope's use of *avus* as "ancestor" in *Nimravus* was evidently meant to suggest an ancient catlike predator "homologous" to a living puma ("panther"), similar in some of its anatomy and in its size.
*Nimravus*: a Multilingual Joke?
The etymology for *Nimravus* that makes the most sense based on the available evidence would appear to be:
*Nimravus* Cope, 1879 "panther [= puma] ancestor" from Arabic *nimr* as "panther" + Latin *avus* "ancestor"
Cope took the Arabic word *nimr* (already in use as a specific name) as "panther" (for an Old World big cat) to allude to "panther" as used in American English for the New World puma, in a combination with Latin *avus* "ancestor" to suggest it was a figurative ancestor of the modern puma, distantly (not directly) related, but similar in size and some points of anatomy.
The multilingual formation of the name *Nimravus* would be similar to the name of another mammal Cope named in 1879:
*Paciculus* Cope, 1879 "little paca" for a small fossil rodent, from *paca*, a Brazilian Tupi name for a type of rodent (used as the species name *Cuniculus paca* (*Agouti paca*)) + Latin suffix -iculus "little"
A proposed derivation of *Nimravus* from the biblical name Nimrod to mean "hunter ancestor" or "Nimrod's grandfather" may be a bit TOO clever and erudite--and requires arbitrary shortening of the name Nimrod.
[As a side note, some older sources suggested that the name Nimrod itself was somehow derived from *nimr* "panther"--supposedly to mean "panther subduer," "son of the panther," or "panthers," depending on the putative derivation of -od in the name. However, such etymologies are implausible for many reasons. As noted above, languages change and evolve over time, and there is no evidence that the modern Arabic *nimr* with a meaning "panther" was the form of the word in use thousands of years ago when any historical figure resembling Nimrod could be said to have lived or later when the Hebrew Bible was first set down in writing.]
Cope DID create "joke" generic names after famous names in ancient history, but with descriptive meanings based on alternate etymologies.
*Empedocles* Cope, 1878. Empedocles was the name of an ancient Greek philosopher (4th Century BC). Cope used the name for a diadectid with interlocking processes on its vertebrae, helping hold the spine rigid, from Greek *empedos* "firm-set, steadfast" + *klees* "glory, fame." Cope later replaced the preoccupied name Empedocles with *Empedias* Cope, 1881, the name of a Spartan during the Peloponnesian War, again derived from Greek *empedos* "firm-set, steadfast" + -ias "characterized by."
Cope's generic names *Claudius*, *Hadrianus*, *Thorius*, and *Tuditanus* look like the names of Roman leaders, but are, in fact, jokes: *Claudius* (Roman Emperor Claudius) for a turtle (from Latin *claudo* "lock up," after its shel); *Hadrianus* (Roman Emperor Hadrian (Latin Hadrianus), named for his birthplace Hadria) for an extinct giant tortoise (so as if from Greek *hadros* "robust, bulky"); *Thorius* Cope, 1869 (Thorius was an Ancient Roman name), as if from Greek *thoroeis* "embryonic," alluding to the poorly ossified, larval-like adult skull in a living salamander; *Tuditanus* (a Roman leader) for a fossil amphibian with a "broad, flat" head (from Latin *tudis* "mallet"--supposedly the Ancient Roman Tuditanus was nicknamed for the broad, mallet-shape of his head).
Cope's name *Nimravus* appears to be a bit of linguistic mischief, or at least linguistic misdirection, using an Arabic word *nimr* to mean "panther"--but to refer to the American "panther" (*Puma concolor*). The joke may not be as elaborate as a supposed reference to the biblical great hunter Nimrod, but it explains the spelling better and fits with Cope's original descriptions of *Nimravus* comparing the ancient predator to a "panther."