Yesterday we looked at the smallest of the fruit bats (mega bats). Earlier we looked at several micro bats.
Here, at nearly the same size,
Notopteris (Figs. 1, 2( nests in the large reptile tree (LRT, 1671+ taxa, subset Fig. 5) as the most primitive extant megabat due to its long tail and a few other primitive traits.
Figure 1. Notopteris in vivo. Note the microbat proportions and relatively long tail. The wing membrane begins along a dorsal margin, not laterally as in other bats.
Notopteris macdonaldi (Gray 1859) is the long-tailed fruit bat or Fijian blossom bat. This is the most primitive megabat in the LRT and the only one that retains a long tail. It roosts in large cave colonies only on South Pacific islands. Note the mid-dorsal attachment of the proximal wing membranes, rather than a more lateral attachment. This is a derived trait not shared with other bats.
Figure 2. Notopteris skull and mandible. Note the primitive skull and derived simple cusp teeth.
The most primitive extant microbat
in the LRT (Figs. 3, 4) is the newly added Rhinopoma, the lesser mouse-tailed bat (Fig. 3). It is similar, to Notopteris (Figs. 1, 2), but with a shorter rostrum and retains primitive multiple cusps on its teeth. Both are cave dwellers.
Figure 3. Rhinopoma is the most primitive extant micro bat in the LRT. Note the long tail, long legs and small feet, all Chriacus-like and Onychonycteris-like primitive traits. Note the lateral insertion of the wing membrane on the torso, distinct from Notopteris (Fig. 1).
Rhinopoma hardwickei (Gray 1831) is the extant lesser mouse-tailed bat, an insectivore found from North Africa to India. The tail is 3/4 free and no calcar is present on the heel. The legs are long and the feet are small.
Figure 4. Rhinopoma skull from Digimorph.org and used with permission. Note the prominent ear bones (yellow) in this echolocating microbat.
Simmons et al. 1984 looked at echolocation in Rhinopoma.
They concluded, “Except for duration these signals are relatively inflexible and suggestive of a primitive kind of echolocation in which only one dimension is changed to achieve qualities which most other species of bats obtain by changing a variety of signal dimensions simultaneously.”
Nelson and Hamilton Smith 1982 looked at echolocation in Notopteris.
They concluded, “Some field experiments… showed these flying foxes were unable to avoid obstacles in complete darkness or when blindfolded, but were able to do so in very dim light. No audible or ultrasonic sounds that could be used in echolocation were detected during their flight.”
Holland et al. 2004 looked at echolocation in the megabat Rousettus.
They reported, “Rousettus aegyptiacus Geoffroy 1810 is a member of the only genus of Megachiropteran bats to use vocal echolocation, but the structure of its brief, click-like signal is poorly described.Rousettus aegyptiacus Geoffroy 1810 is a member of the only genus of Megachiropteran bats to use vocal echolocation, but the structure of its brief, click-like signal is poorly described. However, the low energy content of the signals and short duration should make returning echoes difficult to detect. The performance of R. aegyptiacus in obstacle avoidance experiments using echolocation therefore remains something of a conundrum.”
Simmons and Geisler 1998 looked at echolocation in Icaronycteris.
They reported, “We propose that flight evolved before echolocation, and that the first bats used vision for orientation in their arboreal/aerial environment. The evolution of flight was followed by the origin of low-duty-cycle laryngeal echolocation in early members of the microchiropteran lineage. This system was most likely simple at first, permitting orientation and obstacle detection but not detection or tracking of airborne prey.”
Veselka et al. 2010 concluded that Onychonycteris finneyi may have been capable of echolocation. in reply, Simmons et al. 2010 argued that Onychonycteris finneyi was probably not an echolocating bat.
Echolocation seems to have been convergently acquired
in microbats and Rousettus.
Figure 5. Subset of the LRT focusing on the resurrected clade Volitantia, including dermopterans, pangolins, bats and their extinct kin.
Basal bats in the LRT have more plesiomorphic traits overall,
like small ears, simple nose, long legs, long tail and small feet, all Chriacus-like (Fig. 6) traits. This is what we should expect when any cladogram models micro-evolutionary changes.
Figure 6. Chriacus and Onychonycteris nest as a sister to the undiscovered bat ancestor and a basal bat. Miniaturization was part of the transition. So was enlargement of the manus. It is still a mystery why the transitional form decided to start flapping.
We looked at the origin of bats from Chriacus-like ancestors
earlier here, here and at earlier links therein. These posts are –by far– the most popular posts at this PterosaurHeresies.
To summarize one of those posts
hanging pre-bats simply listened for the sounds of prey in leaf litter below, then pounced from above. Parachuting with flapping evolved into helicoptering then that evolved into flight to return to the branch the bat fell from. Larger hands and extradermal membranes would have increasingly aided entrapment at the moment of impact. Even larger hands and extradermal membranes would have increasingly helped helicoptering while falling. Smaller size and weight (Fig. 6) was co-opted to aid these behaviors. Echolocation seems to have evolved in bats seeking aerial prey and co-opted to live in caves in complete darkness.
Gray JE 1831. Description of some new genera and species of bats. The Zoological Miscellany, 1: 37-38.
Gray GR 1859. The annals and magazine of Natural History, Zoology, Botany and Geology 3. Series IV: 4859.
Holland RA, Waters DA and Rayner JMV 2004. Echolocation signal structure in the megachiropteran bat Rousettus aegyptiacus Geoffroy 1810. Journal of Experimental Biology 207:4361–4369.
Nelson JE and Hamilton-Smith E 1982. Some observations on Notopteris macdonaldi (Chiroptera: Pteropodidae) in Australian Mammal Society 5: 247–252.
Simmons JA, Kick SA and Lawrence BD 1984. Echolocation and hearing in the mouse-tailed bat, Rhinopoma hardwickei: acoustic evolution of echolocation in bats. Journal of Comparative Physiology A 154: 347–356.
Simmons NB and Geisler JH 1998. Phylogenetic relationships of Icaronycteris, Archaeonycteris, Hassianycteris, and Palaeochiropteryx to extant bat lineages, with comments on the evolution of echolocation. Bulletin of the American Museum of Natural History 235.
Simmons NB, Seymour KL, Habersetzer J and Gunnell GF 2010. Inferring echolation in ancient bats. Nature 466: E8.
Veselka et al. (8 co-authors) 2010. A bony connection signals larygenal echolocation in bats.Nature 463: 939–942.