Putting the 'perimyotines' well away from pips proper (vesper bats part XII)


Once upon a time, a huge variety of small to very small vesper bats - basically all of those that possess a simple tragus, a shortish face, two pairs of upper incisors and two upper and two lower premolars - were lumped together as the pipistrelles. You don't have to have a detailed or expert knowledge of vesper bat diversity or morphology to realise that at least some of these characters are primitive across Vespertilionidae, or have evolved repeatedly in disparate lineages. When these observations are combined with the morphological and molecular differences present among the many species concerned, you can see why bat experts have often doubted the monophyly of 'pipistrelles' as traditionally conceived. Indeed, it now turns out that many 'pipistrelles' aren't pipistrelles at all [composite below shows 'Eastern pipistrelle' at left (from here) and Common pipistrelle Pipistrellus pipistrellus (by Mnolf, from wikipedia) at right].


Here in Europe - where the history comes from - we're ok: our pipistrelles are pipistrelles, since they're the ones first given this name. But if you're North American - sorry - your pipistrelles turn out not to be pipistrelles. I'm referring of course to the familiar little bats once known as the Western pipistrelle Pipistrellus hesperus and Eastern pipistrelle P. subflavus.

Goodbye 'Western pipistrelle'... hello Canyon bat?


The Western pipistrelle [shown here, from wikipedia] is often white-bellied, yellowish-brown dorsally, and with a very dark face and ears. It's the smallest of North America's bats, with a mass of 3-6 g and wingspan of 19-23 cm. It occurs from Mexico north to Washington and is strongly associated across its range with rock faces, canyons and boulders. It roosts in cracks, crevices and among rocks and may also use rodent burrows. For these reasons some people have suggested that it should be known as the Canyon bat. Variation across its range has resulted in the recognition of seven 'subspecies' (see map below, from Hatfield (1936). It shows five, but a sixth - P. h. potosinus - was named in 1951 and a seventh - P. h. oklahomae - in 1959). How many of these might actually be valid phylogenetic entities is in doubt and I'm not sure if recent studies have recovered any distinct structure within the species.


It's adaptable, able to forage in woodland, shrubland and desert habitats, and there are indications that it can fly even when at low body temperature (O'Farrell & Bradley 1977). In fact there are records of these bats foraging when the air temperature was -8°C; some researchers think that (in the warmer parts of their range) these bats are probably active year-round (O'Farrell & Bradley 1970, Geluso 2007). Remember: there are bats that hibernate when it gets cool, bats that migrate when it gets cool, but also bats that can tolerate the cool and just carry on as usual. At the other extreme, this bat is also happy to be out foraging when air temperatures exceed 30°C.

And goodbye 'Eastern pipistrelle'... hello Tricolored bat?


The Eastern pipistrelle is typically yellowish-brown and with black wing membranes. Its forearms are reddish-orange [obvious in the adjacent image: photo from here]. Its hairs are distinctly tri-coloured, with black bases, light brown shafts and dark brown tips. This has led to the alternative name Tricolored bat (or Tri-colored bat), and some bat workers argue that this name should be used instead of the technically inaccurate 'Eastern pipistrelle'. It's one of North American's commonest and most widespread bats, yet comparatively little is known about its ecology and natural history (a familiar theme with North American mammals: do not assume that your wildlife is well understood!*). High-aspect wings suggest a preference for foraging in open habitats and there are some reports where foraging has been observed over fields and above woodland canopies. It may well be an ecotone species, frequently eating harmful insects associated with agricultural habitats.

* That wasn't meant to be a dig directed at North Americans - it's true of everywhere.

Unlike the Western pipistrelle, it seems to be an obligate hibernator, even in the southern parts of its range. Like many other temperate-zone vesper bats, females store sperm during hibernation and don't 'allow' their eggs to be fertilized until spring (they produce one or two babies). In terms of roost and hibernation sites, the Eastern pipistrelle is flexible, with the choice of sites including buildings, rock crevices, beneath Spanish moss on low branches, and also beneath branches in tree canopies. It occurs as far north as Nova Scotia and might persist here because it's warm compared to the rest of eastern Canada.


There are some indications that the species is expanded westwards across the USA, with new records occurring across the whole western boundary of its range (from South Dakota to Texas) as well as in the southern limits of its range (New Mexico) (White et al. 2006). Range expansion has also occurred around the Great Lakes: apparently this is the result of the modification of Bear Cave (Michigan) that happened in 1939-40 (Kurta et al. 2007). In Europe, there's been some suggestion that increasing numbers of Common pipistrelles have put pressure on rare populations of horseshoe bats. It's therefore conceivable that an increase in the Eastern pipistrelle might, similarly, have affected populations of other, rarer bats in eastern North America - though I'm only throwing this out there as a possibility and haven't heard of any evidence for it. A preference for east-facing caves as hibernating sites has been demonstrated (Briggler & Prarther 2003). [Adjacent Eastern pip image by Hammbeen, from wikipedia]

Of parastrelles and perimyotines

As noted at the start of this article, it's often been suggested that a great many of the vesper bats conventionally lumped together as pipistrelles are not really close relatives. Rather, they seem to be superficially similar, short-faced, small vesper bats with no close relationship. Indeed, ever since the 1940s morphologists and geneticists have been noting how various of the species involved probably warrant separation. The two North American 'pipistrelles' are among the offenders, differing from undoubted, Old World species of Pipistrellus in bacular and dental anatomy as well as karyotype.


Hoofer & Van Den Bussche (2003) found both species to be substantially removed from pipistrelles proper, and in fact outside the clade that included serotines, pipistrelles and vespertilionins. Neither species grouped closely to any other taxon, so it was decided that both should be given their own generic names. P. subflavus - the 'Eastern pipistrelle' - became Perimyotis subflavus, incorporating a generic name proposed for this species in 1984.

The situation with P. hesperus is a little more complex. HorácÄk & Hanák (1985, 1985-86) proposed the new name Parastrellus for this species, but never did so in formal fashion. Hoofer et al. (2006) argued that this rendered the name a nomen nudum and therefore established the name formally. The Western pipistrelle is therefore now Parastrellus hesperus. Given that bats in Pipistrellus are known as pipistrelles and bats in Falsistrellus are known as falsistrelles, it would be neat if bats in Parastrellus became known as parastrelles. Some people started doing this unofficially as soon as Hoofer et al. (2006) was published, and in 2008 at least one annotated checklist of American mammals did use 'American parastrelle' as the official name for what used to be called the Western pipistrelle (Manning et al. 2008).

We therefore now have the undesirable situation of there being three common names in use for this bat: the name 'Western pipistrelle' isn't going to go away, and there's also 'Canyon bat' and now 'American parastrelle'. The new name 'American perimyotis' has been suggested for what used to be called the Eastern pipistrelle (Manning et al. 2008). And this is in addition to 'Tricolored bat'.

Incidentally, the Western pipistrelle/Canyon bat/American parastrelle was (when first described in 1864) originally named as a new species of Scotophilus. This name is, as we saw recently, only used today for the house bats of the Old World, but back then it was also used for various North American bats and also for European pipistrelles. The historical nomenclature of vesper bats is a total nightmare and I'd like to mostly avoid it if that's ok with you.


Whatever they're called, are these two interesting little bats close relatives, or are they separate on the vesper bat tree? Opinions on this have differed but Roehrs et al. (2010) found them to group together in a clade close to the large (unnamed) clade that includes serotines, hypsugines, pipistrelles and noctules. There isn't (yet) a formal name for the 'American pipistrelle' clade, but Roehrs et al. (2010) referred to it as "the perimyotine group". Google indicates that 'Perimyotini' hasn't yet been used in the literature, and we await further studies (for a discussion of vesper bat phylogeny see the article on vesper bat phylogeny) [Eastern pipistrelle/Tricolored bat/American perimyotis shown here].

Incidentally, Hoofer et al. (2006) noted that the Western pipistrelle/Canyon bat/American parastrelle is very similar to the African hypsugine Hypsugo musciculus (this had previously been noted by Karl Koopman). Whether these similarities reflect affinity or convergence remains interesting, for one because it would demonstrate that perimyotines are not exclusively American. I don't think the idea has been properly tested since phylogenies haven't included both species; like so many obscure bats, H. musciculus is very poorly known and molecular data is apparently unavailable. I haven't covered hypsugines yet... patience, patience.

Anyway, that was rather more than I intended to say about just two species. Nevertheless, they have - historically - been 'important' and, as we'll see later, they've been harbingers of doom in terms of their implications for the content of Pipistrellus sensu lato.

For previous Tet Zoo articles in the vesper bats series, see...

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

Refs - -

BRIGGLER, J., & PRATHER, J. (2003). Seasonal Use and Selection of Caves by the Eastern Pipistrelle Bat (Pipistrellus subflavus) The American Midland Naturalist, 149 (2), 406-412 DOI: 10.1674/0003-0031(2003)149[0406:SUASOC]2.0.CO;2

Geluso, K. 2007. Winter activity of bats over water and along flyways in New Mexico. The Southwestern Naturalist 52, 482-492.

Hatfield, D. M. 1936. A revision of the Pipistrellus hesperus group of bats. Journal of Mammalogy 17, 257-262.

Hoofer, S. R. & Van Den Bussche, R. A. 2003. Molecular phylogenetics of the chiropteran family Vespertilionidae. Acta Chiropterologica 5, supplement, 1-63.

- ., Van Den Bussche, R. A. & HoráÄek, I. 2006. Generic status of the American pipistrelles (Vespertilionidae) with description of a new genus. Journal of Mammalogy 87, 981-992.

HoráÄek, I. & Hanák, V. 1985-1986. Generic status of Pipistrellus savii and comments on classification of the genus Pipistrellus (Chiroptera, Vespertilionidae). Myotis 23-24, 9-16.

Kurta, A., Winhold, L., Whitaker, J. O. & Foster, R. 2007. Range expansion and changing abundance of the Eastern pipistrelle (Chiroptera: Vespertilionidae) in the Central Great Lakes region. The American Midland Naturalist 157, 404-411.

Manning, R. W., Jones, C. & Yancey, F. D. 2008. Annotated checklist of recent land mammals of Texas, 2008. Occasional Papers, Museum of Texas Tech University 278, 1-18.

O'Farrell, M. J. & Bradley, W. G. 1970. Activity patterns of bats over a desert spring. Journal of Mammalogy 51, 18-26.

- . & Bradley, W. G. 1977. Comparative thermal relationships of flight for some bats in the Southwestern United States. Comparative Biochemistry and Physiology Part A: Physiology 58, 223-227.

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.

White, J. A., Moosman, P. R., Kilgore, C. H. & Best, T. L. 2006. First record of the Eastern pipistrelle (Pipistrellus subflavus) from southern New Mexico. The Southwestern Naturalist 51, 420-422.


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Among the best known, most widespread and most familiar of vesper bats are the pipistrelles. All bats conventionally regarded as pipistrelles are small (ranging from 3-20 g and 35-62 mm in head-body length), typically with proportionally short, broad-based ears and a jerky, rather erratic flying…
A group of mostly mid-sized pipistrelle-like bats of Africa and the northern continents are known as the serotines (Eptesicus) [species shown here is the one generally known simply as the Serotine E. serotinus: photo by Mnolf, from wikipedia]. Here in Europe this is - along with pipistrelles,…
As we've seen throughout this series (see links below for previous parts), recent phylogenetic studies have found a number of 'pipistrelle-like non-pipistrelles' to form a novel clade previously unsuspected from morphological studies [composite above shows - l to r - Hypsugo cf. joffrei (from…
Yay, more vesper bats! The groups we've looked at so far have - in anatomical terms - been pretty conservative. This time round we're looking at a really remarkable group; as is so often the case, their familiarity (relative to so many others of the world's bats) means that we tend to forget or…

The new name 'American perimyotis' has been suggested for what used to be called the Eastern pipistrelle

But if it's the only species that has been placed in the genus, why would you need the 'American'? Why not just 'perimyotis'?

On common names, I'm leaning toward retaining "pipistrelle". After all, American robins aren't robins, and American blackbirds aren't blackbirds, and American warblers aren't warblers, etc.* And nobody is in any hurry to fix those. Then again, it having been found that North American tanagers aren't really tanagers, I feel like changing those names: it's "Scarlet piranga" for me. Emerson.

*Just kidding. Actually, European warblers don't deserve to keep their names. Drab little things.

By John Harshman (not verified) on 12 Apr 2011 #permalink

Q: Why 'American perimyotis'?
A: Because it makes for better movie titles.

And... "European warblers: drab little things". My Hippolais icterina begs to differ.

Since I'm a botanist who had to do a bit of bat (paper) research, one thing I can add is that bat experts really need to study botany.

Botanists have (and still are) getting used to lots of name changes, particularly in the American Asteraceae. The problem is that many genera named in Europe actually aren't over here, we've just got the covergent forms...

In any event, there's a sustained push amongst the herbaria, activist groups, and state and federal science bodies to make plant names inter-translatable, even if different bodies don't totally agree on the taxonomy.

Considering that we've got a big push on for wind power, and bats are the #1 group getting pulped by wind turbines (not to mention white nose syndrome) , I'd strongly suggest that the bat systematists take a few days off arguing over the names and get on with figuring out how to translate the name changes and make them easy to follow. That way, the local planners, environmental consultants, and activists who are commenting on the wind projects won't totally get lost in the maze.

The end goal should be saving as many bats as possible, and (at least in my fantasy land) the taxonomists would realize this. Naming recently extinct species isn't much fun. I already know one taxonomist who named a "ssp. extinctus" from a decades-old museum collection, and it would be nice to keep the numbers of these low.

By heteromeles (not verified) on 12 Apr 2011 #permalink

Historical nomenclature! Historical nomenclature!

Don't mess with me, Naish. I'll see your Hippolais and raise you a dozen Dendroica, just for a start. Hippolais icterina barely rises even to the level of an orange-crowned or Nashville warbler, and they're among the least colorful parulids. By the way, if you're ever in a Chicago lake shore park in mid-May, you can find a good 30 species, maybe even 35, in an hour. Now that's what I call warblers.

By John Harshman (not verified) on 12 Apr 2011 #permalink

John: think of Parulidae as "beginner birder's warblers", and of Acrocephalidae (or whatever they are called nowadays) as "real expert's warblers". You can't seriously consider yourself knowledgeable of birds unless you can ID any Phylloscopus by sight or by sound at 100 m without knowing the geographic location or habitat.

Anyway, what I wanted to say is that "tricolored bat" totally sucks: there is Myotis tricolor, and lots of bats with tricolored hair.

Vlad: We have something similar, but we call them Empidonax. Of course they aren't warblers, but that was my point anyway.

By John Harshman (not verified) on 12 Apr 2011 #permalink

I just wanted to say:

You are the bat master, son!

By Stevo Darkly (not verified) on 12 Apr 2011 #permalink


the alternative name Tricolored bat (or Tri-colored bat)

Pip pip, what's all this, then? I say, old chap, I am jolly surprised that you didn't spell it 'coloured'.

Given that bats in Pipistrellus are known as pipistrelles and bats in Falsistrellus are known as falsistrelles, it would be neat if bats in Parastrellus became known as parastrelles.

There doesn't seem to be any bat genus called 'Kestrellus', which is probably just as well; otherwise we might have to worry about people confusing 'kestrelles' with 'kestrels'...


American warblers aren't warblers

Aren't they basically just modified, insect-eating sparrows (or something)?

Regarding Phylloscopus, by the way... John knows this, of course, but for the record: there is a native 'Old World warbler' species in the New World too. The Arctic warbler Phylloscopus borealis breeds in Alaska.

Parulids: I started writing articles on that group, shame I haven't finished them (one of them is called something like 'Parulids that aren't'... I'm sure you can see where I was going with that one). But when it comes to testing your skills, I agree with Vlad - distinguishing all those sylviids and acros and locustellas takes skill and experience. New World warblers are child's play :)

I went with the spelling 'Tricolored bat' because, obviously, it's mostly been used by Americans. It is indeed not ideal - I can see people easily confusing it with 'Tricolored big-eared bat', a name sometimes used for the phyllostomid Glyphonycteris sylvestris. Regardless, google shows that the name is being used quite a lot, often in connection with rabies (this bat is a confirmed vector).

You didn't mention the false pipistrelles of Australia -- genus Falsistrellus. There's supposed to be five species, the Western False Pipistrelle (F. mackenziei), the Eastern False Pipistrelle (F. tasmaniensis), the Chocolate Pipistrelle (F. affinus), the Pungent Pipistrelle (F. mordax), and Peters's Pipistrelle (F. petersi). How do they fit into the Vespertilionidae family tree, do you know?

Of course Australia also has a number of (still) true pipistrelles... though one less since the Christmas Island pip went extinct a year or so ago.

By John Atkinson (not verified) on 13 Apr 2011 #permalink

Actually, I did mention the falsistrelles, but only in passing since they're not (so far as we can tell) part of the perimyotine clade discussed here: rather, they appear to be hypsugines, a clade I haven't yet gotten to. Patience! :)

I love the name Falsistrellus.

I just wanted to say:

You are the bat master, son!


By David MarjanoviÄ (not verified) on 13 Apr 2011 #permalink

...and as for Batman; do you mean the Gotham City crimefighter, or the founder of the city of Melbourne, Australia?

American Periomyotis is a HORRIBLE name.

I don't see an issue with retaining pipistrelle as part of the common names of the American taxa, since they do at least look like old world pipistrelles and presumably it is a good enough name for this particular "morphotype"

Just like "chat", "flycatcher", and "finch" refer to a morphotype, and not a particular clade

By Morgan Churchill (not verified) on 16 Apr 2011 #permalink

the founder of the city of Melbourne, Australia

<hovering over link>

:-D :-D :-D :-D :-D

Day saved!

By David MarjanoviÄ (not verified) on 17 Apr 2011 #permalink