Lobed bats, butterfly bats, particoloured bats, thick-thumbed bats, Dormer's bats, bats, bats, BATS... did I mention the bats? (vesper bats part XVII)


Here we are, so close to the very end. I am pleased and surprised to find that we're now looking at the vesper bats within Vespertilionini - the clade that (in the topology I'm using here: that of Roehrs et al. (2010)) includes the pipistrelles and noctules and their closest relatives. We'll get to those extremely familiar bats soon. As usual, they have a bunch of far less familiar close relatives... or alleged close relatives, or possible close relatives... and it's those bats that we'll be looking at here. After, that is, that we deal with lobed bats and butterfly bats. They aren't members of Vespertilionini: as we'll see, they might be hypsugines or serotines.


Australasian lobed bats and African butterfly bats: probably not close relatives after all

Australia, New Guinea, New Caledonia and New Zealand are inhabited by the seven species of pied bats, lobe-lipped bats, groove-lipped bats or wattled bats (Chalinolobus) [the composite illustration above shows Gould's wattled bat C. gouldii on the left (by Neville W. Cayley, from wikipedia). The skull on the right belongs to an Asian particoloured bat Vespertilio sinensis (from HorácÄk (1997))]. Both a population of lobed bats from New Caledonia - regarded by some authors as the distinct species C. neocaledonicus - and the Long-tailed wattled bat C. tuberculatus from New Zealand are critically endangered. All of the lobe-lipped bats have dark brown or even blackish, fine, velvety fur, though a fringe of white hairs in C. picatus has led to the common name Pied wattled bat for this species.

As suggested by some of the common names (and the generic name: it means 'lobe mouthed'), small fleshy lobes (termed lip-lobules) project outwards from near the corners of the mouth, though these are quite small in some species (Chruszcz & Barclay 2002). The antitragus descends down the side of the face, extends beneath the eye, and terminates in another lobe close to the lip-lobule. The function of these facial lobes (if they have one) seems not to be known. Glands are present on the surface of the short, relatively wide snout. The diagram below - from Parnaby (1992) - shows the range of lip-lobules, tragi and antitragi present in lobed bat species.



The Chalinolobus species have sometimes been regarded as congeneric with the African butterfly bats or silvered bats (the nine species grouped together in Glauconycteris). Butterfly bats get their common name for their superficially moth- or butterfly-like flight style, made particularly noticeable by the fact that they often fly well before sunset. Their wings are often marked with patches of cream, brown and black, and their wing membranes have a particularly prominent venation: it's been suggested that these features might help the bats look like dead leaves and hence act as camouflage. The adjacent illustration (by Fiona Reid, from Gary Graham's 2002 Bats of the World) shows a Variegated butterfly bat G. variegata in flight and a Superb butterfly bat G. superba below it.

Chalinolobus and Glauconycteris are highly similar (butterfly bats even have those lip-lobules), and (apparently) only differ in minor features like the proportions of the metacarpals, the position of the outer incisor relative to the canine and the other incisors, and the relative size of the third upper molar (Koopman 1971). This strong similarity means that some authors have regarded them as congeneric: when this has been done, Chalinolobus gets priority since it was named in 1866 (Glauconycteris was named in 1875). Somewhat surprisingly, molecular data does not support close affinity of the two: Roehrs et al. (2010) found Chalinolobus to be part of their hypsugine group while Glauconycteris belonged in the serotine clade. Australasian lobed bats and African butterfly bats are thus not close at all if these results are correct - they're substantially separated, and their strong similarity must be the result of convergent evolution. Shockingly detailed convergent evolution, actually.

Vespertilio: the particoloured bats


Three short-eared, short-snouted Eurasian bats that possess concave regions on either side of the face are united in Vespertilio [adjacent photo of V. murinus by Mnolf, from wikipedia]. They're known as the frosted or particoloured bats due to silver-tipped hairs in the otherwise brownish pelage. A unique cartilaginous 'pseudobaculum' is present in addition to the baculum proper and makes the whole element long and slender, and thus superficially like the baculi of pipistrelles and noctules (Heller & Volleth 1984, HorácÄk 1997). Particoloured bats have long, slender wings and have a fast, direct style of flight. They will eat beetles and moths but mostly seem to prefer flies, aphids and other small insects (Rydell & Baagøe 1994).

Vespertilio murinus was the very first scientific name ever created for a bat. Linnaeus's very vague description of this species meant that later authors ended up applying the name to a whole list of European bats now known to be quite distinct from V. murinus as we understand it today, and an extraordinary amount of complexity was involved in sorting out the ensuing mess. As Wallin (1969) said "The road to the present nomenclature ran through an unparalleled chaos in vertebrate systematics" (p. 300). In truth, Linnaeus didn't definitely associate the name Vespertilio murinus with the Particoloured bat anyway: it has been argued that the bat he had in mind was more likely Myotis myotis (see discussion in Rydell & Baagøe (1994)). Even in recent decades, there are Vespertilio specimens and species that have been confused with serotines and noctules (see HorácÄk 1997).


Particoloured bat have typically been classified close to serotines, but some molecular data indicates instead that they form a clade with pipistrelles and noctules (Roehrs et al. 2010). However, Agnarsson et al. (2011) found particoloured bats to be most closely related to 'hypsugines' like Neoromicia and Hypsugo. Obviously, the name Vespertilionini has to go with whatever clade includes the particoloured bats. In accordance with the trees generated by Roehrs et al. (2010), I'm here using this name for the clade that also includes pipistrelles and noctules.

Until recently, fossil representatives of Vespertilio were only known from the Pleistocene and Holocene. A lower jaw from Russia has now extended their fossil record back to the Late Miocene (Rossina et al. 2006) [that jaw - identified as Vespertilio cf. villanyiensis - is shown here (the diagrams labelled a-c), with a lower jaw of V. murinus shown below. Both from Rossina et al. (2006). Scale bars = 3 mm].

Thick-thumbed bats and Dormer's bat

Two little known, superficially pipistrelle-like vesper bats with weird thickened pads on their thumbs and feet are united in Glischropus and sometimes known as the thick-thumbed bats (G. tylopus and G. javanus). Their affinities are uncertain and few phylogenetic studies have incorporated them, but in some recent classifications and phylogenies (Volleth & Heller 1984) they've been listed close to true pipistrelles and noctules.

A controversial species originally described as G. rosseti was argued to actually be a species of Myotis by Hill & Topál (1973) [the Myotis bats - the mouse-eared and little brown bats - were covered here]. Today it's sometimes called the Thick-thumbed myotis. The fact that vesper bats as phylogenetically distant as pipistrelle-like forms and mouse-eared bats might be confused with other another seems odd and surprising today, but prior to the 1970s it was fairly usual for bat experts to think that such taxa as Myotis, Pipistrellus (sensu lato) and Eptesicus were close relatives, and morphologically similar but for differences in premolar configuration.


Thick-thumbed bats are very small (head and body length is about 40 mm, weight is between 3.5 and 4.5 g) and they look as if they have particularly small eyes. The ears are long, reaching the tip of the snout. The upper surface of the snout slopes downwards, giving them a vaguely shrew-like profile. Thick-thumbed bats inhabit south-east Asia, Indonesia, the Philippines, Java, Sumatra and the Moluccas, and are often associated with bamboo, rock crevices and banana leaves. Like other bats with modified pads on the thumb and/or feet [adjacent diagram shows foot and thumb of G. tylopus], it seems likely that these features have evolved under selection pressure for an increased grip on smooth roosting surfaces. The pads in thick-thumbed bats are mobile (thanks to four sets of associated muscles) but they don't have an adhesive function: Thewissen & Etnier (1995, p. 933) concluded that the pads most likely are "involved in mechanical adhesion. It is possible that they interlock with the substrate or, more likely, they may act as a friction pad or wedge".

In the previous article we looked at both the bamboo bats or club-footed bat (Tylonycteris) - notable for their flattened skulls and flattened pads on their thumbs and feet - and at Eudiscopus denticulus, another flat-skulled little vesper bat that possesses 'gripping pads' (this time just on its feet). There's no clear indication that Tylonycteris, Eudiscopus denticulus or Glischropus are closely related (though affinities between some or all of them have been suggested), so it seems that these 'gripping' adaptations evolved independently on three separate occasions (actually more since the hypsugine Neoromicia nanus - popularly known as the Banana bat and until recently classified as Pipistrellus nanus (P. africanus is a synonym) - also has small pads on the wrist).


Dormer's bat Scotozous dormeri [shown here] is from India and Pakistan and has silver-tipped hairs scattered throughout its dark brown dorsal pelage. It has a reduced upper incisor dentition compared to pipistrelles, with the outer pair of incisors being either relictual or absent. You might remember from the nycticein article that the possession of a single pair of upper incisors was formerly regarded as a nycticein character. However, based mostly on its superficial resemblance to pipistrelles, authors have generally put it close to the pipistrelle assemblage (though remember that the large assemblage of vesper bats traditionally grouped together as pipistrelles or pipistrelle relatives are now known to be non-monophyletic). So far as I can tell, modern phylogenetic studies haven't incorporated it, and it remains little studied and poorly known.

And... we're not done yet. More soon. For previous Tet Zoo articles in the vesper bats series, see...

And for previous Tet Zoo articles on bats, see...

Refs - -

Agnarsson, I., Zambrana-Torrelio, C. M., Flores-Saldana, N. P. & May-Collado, L. J. 2011. A time-calibrated species-level phylogeny of bats (Chiroptera, Mammalia). PLoS Currents 011 February 4; 3: RRN1212. doi: 10.1371/currents.RRN1212.

Chruszcz, B. & Barclay, R. M. R. 2002. Chalinolobus gouldii. Mammalian Species 690, 1-4.

Heller, K. G. & Volleth, M. 1984. Taxonomic position of "Pipistrellus societatis" Hill, 1972 and the karyological characteristics of the genus Eptesicus (Chiroptera: Vespertilionidae). Zietschrift für zoologische Systematik und Evolutionsforshung 22, 65-77.

Hill, J. E. & Topál, G. 1973. The affinities of Pipistrellus ridleyi Thomas, 1898 and Glischropus rosseti Oey, 1951 (Chiroptera: Vespertilionidae). Bulletin of the British Museum (Natural History), Zoology Series 24, 447-454.

HorácÄk, I. 1997. Status of Vesperus sinensis Peters, 1880 and remarks on the genus Vespertilio. Vespertilio 2, 59-72.

Koopman, K. F. (1971). Taxonomic notes on Chalinolobus and Glauconycteris (Chiroptera, Vespertilionidae) American Museum Novitates, 2451, 1-10

Parnaby, H. 1992. An interim guide to identification of insectivorous bats of south-eastern Australia. Technical Reports of the Australian Museum 8, 1-33.

Roehrs, Z. P., Lack, J. B. & Van Den Bussche, R. A. 2010. Tribal phylogenetic relationships within Vespertilioninae (Chiroptera: Vespertilionidae) based on mitochondrial and nuclear sequence data. Journal of Mammalogy 91, 1073-1092.

Rossina, V. V., Kruskop, S. V., Tesakov, A. S. & Titov, V. V. 2006. The first record of Late Miocene bat from European Russia. Acta Zoologica Cracoviensia 49A (1-2), 125-133.

Rydell, J. & Baagøe, H. J. 1994. Vespertilio murinus. Mammalian Species 467, 1-6.

Thewissen, J. G. M. & Etnier, S. A. 1995. Adhesive devices on the thumb of vespertilionid bats (Chiroptera). Journal of Mammalogy 76, 925-936.

Wallin, L. 1969. The Japanese bat fauna. Zoologiska Bidrag från Uppsala 37, 223-440.


More like this

Glad to see our local Gould's Wattled Bat finally made it to your bat-list. This species seems to be commonest small bat in Melbourne. Some years ago, bat-boxes were put on trees in a local river-side park. It took more than a year for them to be accepted and the GWBs frequently shift about between the boxes or natural roosts.
They are checked every month - bats are weighed and sex ratios, pregnancy, and young noted. They are released again at dusk. Females gather together, males are sometimes found solo. Recently they have been joined in the box by one or two Broad-nosed Bats.
We once had one in the house. It flew around a large room and quite failed to notice when doors and windows were opened - saving energy with no sonar. We had to climb on a chair and flip it gently with a tea-towel before it switched on again and found its way out.

By Anthea Fleming (not verified) on 18 Apr 2011 #permalink

Darren, during the course of this vespertilionid series, you've implied that you're unhappy with certain vernacular bat names. Here, I couldn't help noticing that you did not mention Chalinolobus tuberculatus by its commonly used vernacular name, i.e., (New Zealand) long-tailed bat. Was that omission intentional?

Linnaeus didn't definitely associate the name Vespertilio murinus with the Particoloured bat anyway: it has been argued that the bat he had in mind was more likely Myotis myotis

For Linnaeaus to have chosen Vespertilio murinus as 'the' bat is surprising also in the light of the fact that the particoloured bat is not especially common in Linnaeus' native Sweden*. The most common and widespread Swedish bat species is the northern bat Eptesicus nilssonii. As for Myotis myotis, that species is not native to Sweden; as far as I know, only one individual has ever been found there (in 1985).

* At least not today; there is, of course, the theoretical possibility that the relative abundancies and distributions of Swedish bats have drastically changed since the 18th century.

In truth, Linnaeus didn't definitely associate the name Vespertilio murinus with the Particoloured bat anyway: it has been argued that the bat he had in mind was more likely Myotis myotis

Was he concerned with such details at all? We're talking about the man who didn't distinguish chimpanzee and orang-utan, lumped all crocodylians into Lacerta crocodilus, lumped all salamanders (except Siren lacertina which he and Ãsterdam described in 1677) into Lacerta salamandra, lumped all bacteria into Volvox chaos, and then believed he had described all species!

By David MarjanoviÄ (not verified) on 19 Apr 2011 #permalink


the man who didn't distinguish chimpanzee and orang-utan

Yeah, but neither did anyone else at the time. I think we can cut Linnaeus some slack here.

Thanks for comments. Regarding comment 2, the only reason I sometimes don't use the correct common names is because I tend to be looking at the technical literature at the time. So, omissions are not intentional (I will go and correct, thanks).

Chalinolobus neocaledonicus: the claim that it might be extinct comes from Chruszcz & Barclay (2002), cited above. However, I made a mistake, as I think this refers to a population of C. gouldii that formerly occurred on Norfolk Island. I'll correct.

Linnaeus (1758) and bats: Linnaeus recognised seven bat species, all classified within Vespertilio. Only two of these were European: V. auritus (thought to refer to Plecotus auritus) and V. murinus. His vague description of the latter means that it's hard/impossible to work out exactly which bat he was referring to. It seems that some 19th century authors assumed that he must really have been referring to Myotis myotis since it was "the most common species" (Keyserling & Blasius 1839), but this was then contested when it was pointed out that M. myotis doesn't occur in Sweden (the type area for V. murinus). I can't follow the whole story as I haven't seen Hinkel (1992): it seems, however, that the ICZN was petitioned in 1958 in order that the name V. murinus be linked to the Particoloured bat.

Ref - -

Hinkel, A. 1992. Die Zweifarbfledermaus(Vespertilio murinus) wurde von J. Natterer entdeckt. Nyctalus 4, 302-306.

Keyserling, A. & Blasius, I. H., 1839. Die Ãbersicht der Gattungs- u. Artcharaktere d. europaeische Fledermäuse. Archiv für Naturgeschichte 5(1).

Translation of the title of Hinkel (1992): "The particoloured bat (Vespertilio murinus) was discovered by J. Natterer".

The particoloured bat occurs in southern Sweden according to text and map of the Swedish Wikipedia article.

By David MarjanoviÄ (not verified) on 20 Apr 2011 #permalink


I can't follow the whole story as I haven't seen Hinkel (1992)

I looked up that reference. This nomenclatural issue is complicated indeed, but to make a long story short: Hinkel thinks that Linnaeus' description of 'Vespertilio murinus' is based on an illustration of a bat in a contemporary book* by a German naturalist, and that the bat in that illustration is most likely Myotis myotis (which was, and is, native to Germany).

* That book was 'Vorstellung der Vögel in Teutschland**' by J.L. Frisch. That's right; the bats were included in a book about birds (although Frisch did admit that bats can't really be considered birds).

**Sic; I'll leave it to David M. to explain how and when 'Teutschland' became 'Deutschland'.

the ICZN was petitioned in 1958 in order that the name V. murinus be linked to the Particoloured bat

Correct; it's now too late to try to change this (and doing that would hardly serve any useful purpose anyway).


The particoloured bat occurs in southern Sweden

Indeed it does but, as I said, it's rather uncommon and thus not the 'default bat' there.


Oh, then I give up.

Nah, don't be discouraged; you were doing fine.

"**Sic; I'll leave it to David M. to explain how and when 'Teutschland' became 'Deutschland'."

Wrong David, perhaps, but many languages were spelled phonetically, and, to our eyes, idiosyncratically. IIRC there are some ?47? Shakespeare signatures surviving - every one of them using a slightly different spelling.

In particular, some German dialects seem to have de-voiced (some) initial consonants. (or just confused voiced and unvoiced)

The Nuremberg Chronicle, published in 1493, for instance, regularly spells 'Bischof' (Bishop) with a initial 'P'. Similarly, as can be seen http://upload.wikimedia.org/wikipedia/commons/0/00/Schedel_konstantinop… from this Wiki image, Greek is spelled 'Kriechische' (with a 'k' instead of a 'g').

By David Houston (not verified) on 20 Apr 2011 #permalink

I'll leave it to David M. to explain how and when 'Teutschland' became 'Deutschland'."

Well, one possibility is that the T was just an attempt to graphically link the Germans to the Teutons. (That would likely be correct, but only in that there's an extinct word for "people" in both. "German" is "how normal people, as opposed to clergy, speak" -- "not Latin".)

The other...

In particular, some German dialects seem to have de-voiced (some) initial consonants. (or just confused voiced and unvoiced)

All* High German** dialects, for instance mine, have completely devoiced /b d g/. Because /p t k/ are not aspirated, this leaves a very small acoustic difference between /b d g/ and /p t k/; not only does this lead to occasional confusion, but most of these dialects have gone on to completely merge the two series. A largely unified spelling system for German only developped in the 19th century (and wasn't codified before 1901).

Spellings with p and k for /b/ and /g/ were apparently very common in what little was written in Old Bavarian 1000 years ago. p for what is elsewhere /b/ survives in various surnames and place names.

* Except the extreme southern ones, which have reinterpreted the entire sound system in Slovene terms.
** This is a geographical term, not a social one, for the dialects that are spoken where the country is high. Low German is (or was) spoken in the flatland of northern Germany.

By David MarjanoviÄ (not verified) on 26 Apr 2011 #permalink

I had forgotten...

Last summer I found a document from 1643 on exhibit in a museum. It's the first kitchen supply list from Upper Austria to mention turkeys and potatoes. The latter are ertöpfl, which would be Erdäpfel in modern Standard German. Here we have one of those elusive spellings of t for /d/.

(Ã is not a typo. It's a dialect feature caused by the l that forms the next syllable.)

By David MarjanoviÄ (not verified) on 26 Apr 2011 #permalink

Thanks, Davids, for the lessons in linguistics!


A belated comment on this:

Translation of the title of Hinkel (1992): "The particoloured bat (Vespertilio murinus) was discovered by J. Natterer".

Hinkel says that the particoloured bat was really discovered by Johann Natterer (the type locality was, incidentally, near Vienna, Austria), who named it Vespertilio discolor*. That binomen would have been more suitable for the particoloured bat, but, as mentioned, it was suppressed by the ICZN in 1958.

* The species was described in Heinrich Kuhl's monograph Die deutschen Fledermäuse in 1817.