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L_spiritus
L_natalens
L_ocellatu
L_susanae
L_palmatus
L_flavomac
L_viridis
L_aubriodi
L_rufus
L_modestus
L_cf_ragaz
L_kivuensi
L_argenteu
L_vermicul
L_fiziensi
L_macrotis
L_vannutel
L_notatus
L_aubryi
L_yaldeni
L_gramineu
L_breviros
L_boulenge
Astylo_spp
L_parbocag
L_barbouri
L_bocagii
L_calcarat
L_parkeri
L_millsoni
!!!!!!!!!!!!!
L. parbocagii L. barbouri L. parkeri L. argenteus L. natalensis L. yaldeni L. vannutellii L. ragazzii L. susanae L. gramineus L. fiziensis L. vermiculatus L. kivuensis L. flavomaculatus L. bocagii L. modestus L. palmatus! L. aubryioides L. notatus L. brevirostris L. calcaratus L. boulengeri L. ocellatus L. aubryi L. viridis L. spiritusnoctis L. macrotis! L. millsoni! L. rufus! Astylosternus spp.
(58mm)! (43mm)! (56mm)! (52mm)! (65mm)! (43mm)! (46mm)! (50mm)! (53mm)! (63mm)! (38mm)! (85mm)! (36mm)! (70mm)! (58mm)! (41mm)! (110mm)! (43mm)! (74mm)! (64mm)! (57mm)! (81mm)! (58mm)! (54mm)! (48mm)! (49mm)! (84mm)! (87mm)! (87mm)! (NA)
Size
Morphological Analyses
Among other characters, the extent of webbing in feet has been a consistently distinguishable tool for our specimens. We have also found that the ratio of disc (DSC) to Tympanum (TMP) and TMP to head width (HDW) or head length (HDL) is useful in our samples as well. We continue to explore new features that may be incorporated into the literature for future aid in identification.
Deciphering Evolutionary Ancestry in African Tree Frogs: Genus Leptopelis Kyle E. Jaynes1,2 and Rayna C. Bell1
1Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution 2Department of Biology and Environmental Sciences, Adrian College
West and Central African Leptopelis species (L. rufus, L. macrotis, and L. millsoni) were previously thought to be the closest ancestors of L. palmatus due to both their large size and proximity to the island chain. According to our most recent data, along with more extensive sampling from our collaborators, we find that L. palmatus is not closely related to any of the species mentioned above. Our results indicate that L. palmatus is within a group of Central African species that are smaller in body size, suggesting that there may be a convergence in gigantism within this genus.
Moving forward, we are going to (1) increase our species and genetic sampling to determine which species of Leptopelis L. palmatus is closely related to, and (2) collect ddRADseq data for the L. macrotis/ L. millsoni/ L. rufus complex and look at more specimens to assess the morphological variation in these populations. These data will provide greater insight into the biogeography of land bridge and oceanic islands in the Gulf of Guinea archipelago and diversification in the Guineo-Congolian forests. In addition, we hope to find additional morphological characteristics to improve field identifications in this challenging species complex.
Conclusion
We collected 16s mitochondrial gene sequences for 33 species of Leptopelis and 146 individuals for our four target species. To determine whether L. palmatus is closely related to the L. rufus/ L. millsoni/ L. macrotis complex we generated a phylogeny using BEAST v 1.8.4 including one representative sequence for each species, with Astylosternus as an out-group. We assessed phylogeographic structure across the L. rufus/ L. millsoni/ L. macrotis complex with more than 100 specimens from 19 localities amongst the three species. We took 14 standard morphological measurements on more than 30 specimens to evaluate field identifications and search for more reliable morphological characters for species identification.
Methods
Results In figure 1 (a), Leptopelis palmatus is clearly separated from the L. macrotis/ L. millsoni/ L. rufus complex with strong
support (>95% CI). There is not strong support for anything beyond the divergence of these groups, although it appears that there is a closer relationship with species of the Guinean-Congolian distribution.
In figure 2 (a), we see clear phylogeographic structure across the species complex with support for distinct lineages of L. macrotis in West Africa and L. millsoni in Central Africa, distinct lineages of L. millsoni on Bioko island, and pronounced phylogeographic structure in L. rufus across Cameroon.
Results
Frost, Darrel R. 2017. Amphibian Species of the World: an Online Reference. Version 6.0 (07.24.2017). Electronic Database accessible at http://research.amnh.org/herpetology/amphibia/index.html. American Museum of Natural History, New York, USA. Jones, P.J. 1994. Biodiversity in the Gulf of Guinea: an overview. Biodiversity and Conservation. 3:772-784. Linder, H.P., K.M. de Klerk, J. Born, N.D. Burgess, J. Fjeldsa, and C. Rahbek. 2012. Journal of Biogeography. 10:1365-2699. Schiotz A. 1999. Treefrogs of Africa. Chimaira. 1:251-258.
References
We would like to thank our collaborators on the project: Adam Leaché, David Blackburn, Eli Greenbaum, Dan Portik, Greg Jongsma, Matt Fujita, and Mark Oliver-Rödel. We thank Gracia Gonzalez Porter and the L.A.B. for assistance with sequences and facilities. We thank the NHRE program coordinators at the NMNH, Smithsonian Institution for their contributions to the REU program. This research was funded by NSF OCE: 1560088 and an ADS research grant.
Acknowledgments
Photo credit: A. Stanbridge Leptopelis palmatus
Leptopelis palmatus
Photo credit: A. Stanbridge
Introduction
The genus Leptopelis is a group of African Tree Frogs that have a poorly understood taxonomic history, with 53 currently accepted species (Frost 2017). These species are closely related and difficult to distinguish by morphological features alone, often times being misidentified in the field. Leptopelis palmatus is an endemic species to Príncipe Island, one of four islands in the Gulf of Guinea archipelago (Jones 1994). Reconstructing its colonization history and subsequent speciation is key to understanding the biogeography of this archipelago.
L. macrotis, L. millsoni, and L. rufus form a species complex that occurs across West and Central Africa, as well as on Bioko Island, a continental land-bridge system. These species have been hypothesized to be the closest ancestors of L. palmatus due to their large size and proximity to Príncipe (Schiotz 1999). In addition, their distribution across continental Africa and Bioko Island make them an ideal study system for investigating speciation across a range of environments. In this project, we investigate questions of phylogeography and ancestral history in order to learn more about the evolutionary relationships amongst the organisms in this island system.
Introduction
Photo credit: A. Stanbridge Leptopelis palmatus
Figure 1 – (a) Phylogenetic tree of 16s mitochondrial gene. Includes 30 Leptopelis species with geographic distribution by color and maximum female size. Closed circles at nodes represent (>95% CI). Female size and distribution from (Schiotz et al 1999). (b) Biogeographical map of African ecosystems partitioned into 8 areas; Partitioned areas from (Linder et al. 2012).
Biogeographical Regions of Africa Phylogeny of 30 Leptopelis Species
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!!!!
L. rufus! !!!
!!! Cameroon -
Littoral Region !!!!
Cameroon !! Bioko
Gabon
! Ghana
!! Liberia- Guinea !
! Cameroon - East Region !
!!!
L. millsoni! !!!!!!!!!!!!!!!!!!!
L. macrotis! !!!!!!!!
Phylogeographic Tree of West-Central African Leptopelis Species Complex
Leptopelis millsoni Photo credit: Rayna Bell
Variation in Leptopelis Species Complex
Photo credit: Rayna Bell Leptopelis palmatus
Photo credit: B. Freiermuth Leptopelis rufus
Leptopelis rufus Leptopelis millsoni
Figure 3– Hand sketch of webbing extent in L. millsoni and L. rufus.
a)
a)
Sampling Map of West-Central African Leptopelis Species Complex
b)
b)
L. Rufus Type Locality
L. Millsoni Type Locality
L. Macrotis Type Localtiy
Figure 2 – (a) Phylogeographic tree of the L. rufus (red)/ L. macrotis (blue)/ L. millsoni (green) species complex, with geographically sampled and coded lineages. (b) Distribution map of L.macrotis (blue), L.millsoni (green), and L. rufus (red). Circle size is determined by the number of specimens sampled at that location. Triangles represent the type locality of that species.
Background Photo Credit: http://wallpapercave.com/rain-forest-background Figures 1 and 2 B map outline credit: https://www.pinterest.com/pin/394627986069379882/