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In 2006, the collaboration of 19 leading authorities led to a new estimate of amphibian phyologeny. It was published in The Bulletin of the American Museum of Natural History(Volume 297, Issue 1, March, 2006). The title of the publication is The Amphibian Tree of Life. In this section of the website, I planned to place scanning electron microscope images of APs from a fairly wide variety of anurans (frogs, toads and their allies). They were part of a large study I carried out, beginning in the mid-1970s, to provide inferences about AP evolution. The species were selected strategically on the basis of the generally accepted estimates of phylogeny at that time. They were published largely in a journal (Scanning Electron Microscopy) that is not widely available, so I thought it appropriate to place representative examples here.

In the process of doing so, I learned of the 2006 work (through Wikipedia), and, on studying the new tree of life realized that there were new tales to tell about the evolution of the anuran amphibian papilla. The new estimate of phylogeny is based not only on morphological characters, but also on analyses of mitochondrial and nuclear DNA. The base of the new tree, representing the anura with the most ancient pedigrees, is similar to that of the old estimates of phylogeny, and is reasonably well covered by my original selection of species. For the most ancient branch, the Leiopelmatidae, I have APs from two of the four living species, including both genera (Ascaphus and Leiopelma). For the next branch, the Xenoanura, I have APs from four species (two genera) from the Pipidae, one of two families in the branch. For the third branch, the Costata, I have APs for two species from one family (Bombinatoridae) and APs for one species from the other family (Alytidae). For the fourth branch, Anomocoela, I have APs for species from two of the four families (from Pelobatidae and Scaphiopodidae).

At this point the new tree proceeds into the neobatrachia, the anurans of more modern origins. I have no APs from species in the first neobatrachian branch, the Heleophrynidae- - the ghost frogs of Southern Africa. If I were continuing the study today, that would be the first group of frogs I would go for. The remaining neobatrachians have been grouped as the Phthanobatrachia, comprising two major limbs-- each of which needs to be climbed separately in our comparative morphology. For each limb, the Hyloides and the Ranoides, I have APs from species at the first branch (Sooglossidae for Hyloides, Allodapanura for Ranoides). For the Hyloides, I also have representative APs from an early middle branch, Brachycephalidae, a later middle branch, Hylidae, and terminal, or near terminal branches, Dendrobatidae and Bufonidae. For the Ranoides, I have no representatives from the middle branches, but I do have APs for species from terminal or near-terminal branches, Rhacophoridae and Ranidae. If I were continuing the study today, I would go for species in the mid-branches of the Ranoides limb (Ptychadenidae, Ceratobatrachidae, Micrixalidae, Phrynobatrachia); but most excitedly, I would go for frogs in the Terrarana of Hedges et al., see APs on an Eleutherodactylus Tree



 
 
Ascaphus truei
Tailed frog
Leiopelmatidae
Xenopus tropicalis
an African-clawed-frog species
Pipidae
Alytes obstetricans
Common midwife toad
Alytidae
Pelobates fuscus
Common spadefoot
Pelobatidae
 
 
Sooglossus thomasetti
Thomasett's frog
Sooglossidae
Gastrophryne olivacea
Plains narrowmouth toad
Microhylidae

The Amphibian Papilla from Ancient to Modern Frogs and Toads

The AP in the upper left belongs to a member of the most ancient family (Ascaphidae or Leiopelmatidae) of living anurans-- the tailed frog (Ascaphus truei). It consists of a single patch of sensory epithelium. There are three other living members of this family. In the one that we examined, Leiopelma hochstetteri, the AP also consisted of one patch. Next, in the upper row, is the AP from a species (Xenopus tropicalis) in the branch nearest the Leiopelmatidae. This Xenopus AP consists of two patches, which was true of all of the other anuran APs that we examined. In X. laevis, Cheuk Li observed the APs of tadploes at various stages and found that the two patches were widely separated (by a field of squamous epithelial cells) in early stages.

Next is the AP from the common midwife toad (Alytes obstetricans) a species from the third branch of the tree. Immediately to the right of the Alytes AP is one from the common spadefoot (Pelobates fuscus), a species from the fourth branch (Anomocoela), which is the branch nearest the Neobatrachia.

On the lower left is an AP from Sooglossus thomasetti , a species in the first branch (Sooglissidae) of the Hyloides limb. The animal, caught on Mahe in the Seychelle Islands in 1985, was given to us by R.A. Nussbaum. On the lower right is an AP from Gastrophryne olivacea, a species in the first branch (the Allodapanura) of the Ranoides limb.

The story that will emerge from these web pages is one of remarkable plasticity in the caudal patch of the Lalagobatrachia (Figure), with considerable evolutionary experimentation among pre-neobatrachian APs, and remarkable convergence among APs in the Hyloides and Ranoides limbs.

  More APs from
ancient anurans
APs from mid-branch
Hyloides
APs from hylids The puzzling bufonids
part I
The puzzling bufonids
part II
APs from Ranoides  




  APs on the
Tree of Life
APs on a
Rana Tree
APs on a
Bufo Tree
APs on an
Eleutherodactylus
Tree
What does the
caudal extension do?
Discussion  




Calibrated Images


The following index provides links to micrographs from a 1984 paper (Lewis ER, On the frog amphibian papilla, Scanning Electron Microscopy/1984/IV, pp. 1899-1913). They are shown with 600-pixel vertical resolution and with scale bars to indicate the magnification. Each AP either was taken from the left ear, or its image was mirrored to make its orientation consistent with the rest. The numerical order of the presentation reflects my thinking, in 1984, of the general pattern of evolution of AP shape. In light of the new tree of life, that thinking has changed.

  2. The AP and environs
in the Colorado-
River toad
3. Light micrograph
of a bullfrog
AP chamber
4. AP from the
tailed frog
5. AP from
Hochstetter's frog
6. AP from the
oriental fire-bellied
toad
7. AP from the
Surinam toad
 


  8. AP from the
midwife toad
9. AP from a
clawed frog
10. AP from the
Plains spadefoot
(New World)
11. AP from the
common spadefoot
(Old World)
12. AP from the
red-bellied toad
13. AP from
Bocourt's toad
 


  14. AP from the
Texas toad
15. AP from the
Plains narrow-mouth
toad
16. AP from
Strecker's
chorus frog
17. AP from the
green treefrog
18. AP from the
wood frog
19. AP from the
Cascades frog
 


  20. AP from the
green frog
21. AP from the striped
running frog
22. AP tectorium from
the tailed frog
23. AP tectorium from the
African clawed frog
24. AP tectorium from the
western toad
25. AP tectorium from the
cane toad
 


Last updated 03/02/09