I was recently reading a paper in which the phrase “anoline lizards” was frequently used, and it got me thinking: Just what does “anoline” mean? Is it synonymous with “anole”? And is it correct usage? Needless to say, the word is not in standard dictionaries.
Part of the potential confusion comes from the way it was used in the past, at least by some. Back in the old days, pre-molecular data, the standard wisdom was that the genera Chamaeleolis, Chamaelinorops, and Phenacosaurus were early offshoots of the anole radiation that had evolved prior to Anolis. Hence, Ernest Williams used the term “anoline” to refer to them, as well as Anolis species. In this sense, “anoline” might be construed to apply more broadly than just to members of the genus Anolis (and, of course, should not be confused with “anoloid” which referred to an even broader phylogenetic grouping that included other, more distantly related lizards). In this sense, “anoline” had a technical meaning beyond just Anolis. But…some workers, even back then, referred to these three other genera as “anoles” as well. Moreover, we now are confident that the three genera arose within Anolis, which leads most modern workers to no longer recognize those genera, but rather subsume the species within Anolis (though it is still perfectly appropriate to recognize them as subclades of Anolis).
In any case, what’s the history of the term? I searched on JSTOR and found three references from 1900 and before. Cope twice used the term in the 19th century, referring to “anoline iguanidae” in a paper (click on “show full citation”) in the Proceedings of the Academy of Natural Sciences of Philadelphia in 1861 and then again in the same journal in 1896 in which he also referred to the Anolinae. In addition, JSTOR reported another use of the term in the backmatter of the British Medical Journal in 1900. However, inspection of this journal discovers that it was, in fact, an ad for Lanoline, a type of toilet preparation (whatever that is*), and as you can see from the picture, the ornate script fooled whatever automated system JSTOR used, such that the florid L was not recognized as part of the word. JSTOR then reported an “anoline” void for half a century, until seven hits in the 1950s. The number then increased greatly, with 51 in 1960s and 210 in 1970s.
Who better to ask about this than Kevin de Queiroz, Curator of Herpetology at the Smithsonian’s National Museum of Natural History and expert on all matters systematic and nomenclatural. He said this in response to my question: What does ‘anoline’ mean and is it a correct term? I think it’s normally used as an adjective referring to anything about anoles. If so, is it correct, as opposed to say ‘anole ecology,’ etc.?”:
“The term “anoline” has normally been used (at least in the community of anologists, especially those following Ernest Williams) as a noun or adjective for an informal taxonomic group (anolines) that I infer to be (loosely, since it’s informal) associated with the rank of subfamily or tribe (the formal equivalents of which would be “subfamily Anolinae” or “tribe Anolini”) and composed of Anolis, Chamaeleolis, Chamaelinorops, and Phenacosaurus. However, it’s also been used as an adjectival form of “Anolis.” I do think that it is correct to say “anole ecology,” for example, and that this is probably less ambiguous than saying “anoline ecology,” as the latter could be interpreted as endorsing the recognition of Chamaeleolis, Chamaelinorops, and Phenacosaurus as separate genera (unless, of course, the user did wish to endorse that taxonomic scheme).
“Anoloid” has traditionally been used for a more inclusive informal taxon, though in this case the associated rank is less clear, given that the suffix “-oidea” is normally associated with the rank of superfamily in zoology (not in botany), but the people who used this term presumably considered the taxon to be at a lower rank than family, given that they considered anoloids to be in the family Iguanidae. The “-oid” ending also means “like” or “resembling”, and I believe that the term “anoloid” was used in that sense—that is, for anole-like or anole-related lizards (the group also included the “para-anoles” [Anisolepis, etc.], the “leiosaurs” [Leiosaurus, etc.] and Polychrus).
Currently accepted phylogenetic hypotheses call into question the distinction between anoles and anolines. If Chamaeleolis, Chamaelinorops, and Phenocosaurus are nested within Anolis, then anoles and anolines are the same. Moreover, this situation also has a precedent in the older literature, as Etheridge seems to have considered these other taxa to be anoles even when he treated them as separate genera (note that the title of his dissertation is “The Relationships of the Anoles”). Note that Etheridge and de Queiroz (1988) also called the group composed of all four genera “the Anoles,” even though they considered Chamaeleolis not to be derived from within Anolis. Given this situation, it seems reasonable to treat the terms “anole” and “anoline” as synonyms. Nevertheless, I would still recommend using “anole” as an adjectival modifier (as in “anole ecology”) rather than “anoline” because the latter term might be misinterpreted as referring to some larger taxon that includes more than just Anolis. (In fact, I used “Anolines” when I probably should have used “Anoloids” in my first publication.)
Looking into this a bit more objectively (Google Scholar), it seems that “anoline” has been used much more commonly as an adjective than as an informal taxon name. I say this because a GS search on “anoline” yielded about 2000 hits, while one on “anolines” yielded only 85, and the former are mostly in the phrase “anoline lizards.” On other other hand, one could say that “anoline lizards” is itself an informal taxon name (or a common name for a formal taxon, see below). Uses as an adjective not modifying “lizard” (“anoline display behavior” “anoline faunas”) appear to be less common.
I also searched on “Anolinae” and got 33 hits, one as recent as 2007. I think Cope (1896) may have been the first person to use this name, though he earlier (e.g., 1864) recognized a family Anolidae. Cope’s (1900) Anolinae included Chamaeleolis, Xiphocercus (a polyphyletic group of anoles with prehensile tails), Anolis, Scytomycterus (proboscis anoles), Norops, and Polychrus. Most of the more recent uses of this taxon name are not in taxonomic papers, so that the exact composition of the taxon can’t be determined. Many of them are in papers that use the name to indicate a higher taxon to which a study species is referred (e.g., “Anolis equestris (Anolinae)”) or that list several species included in a broad survey (e.g., of sex determining mechanisms in squamates). Given that the formal name “Anolinae” is still used occasionally, one could also interpret “anoline” as a common name equivalent of a formal taxon name (i.e., not only as an informal taxon name).
*Kevin de Queiroz remarked upon reading this: “’It’s advertised to be ‘An emollient, soothing application for irritable skins, useful for chapped lips and hands, and eruptions of many kinds.’ The name is likely a modification of ‘lanolin,’ which refers to a waxy substance found in sheep’s wool that’s used in various lip balms and ointments and was likely a component of lanoline. It’s derived from Greek lana (wool) plus oleum (oil).”
So, can we assume that the upcoming Phylocode companion won’t have a definition & diagnosis for Anolinae?
In addition to the para-anoles (which are southern South American in distribution), we also used to talk about the Peron-anoles, and also (since we were at the MCZ) the Pere-anoles.