Zootaxa 4147 (4): 433–442
http://www.mapress.com/j/zt/
Copyright © 2016 Magnolia Press
Article
ISSN 1175-5326 (print edition)
ZOOTAXA
ISSN 1175-5334 (online edition)
http://doi.org/10.11646/zootaxa.4147.4.5
http://zoobank.org/urn:lsid:zoobank.org:pub:86E9C17F-F09C-49C2-BB33-4F8FFF2381A2
A new small-sized Theloderma (Anura: Rhacophoridae) from Laos
NIANE SIVONGXAY1, MONEKHAM DAVANKHAM1, SOMPHOUTHONE PHIMMACHAK1,2,
KEOCHAY PHOUMIXAY1 & BRYAN L. STUART3,4
1
National University of Laos, Faculty of Natural Science, Department of Biology, P.O. Box 2273, Dong Dok Campus, Vientiane, Laos
Kasetsart University, Faculty of Science, Department of Zoology, Chatuchak, Bangkok, 10900, Thailand
3
North Carolina Museum of Natural Sciences, 11 West Jones Street, Raleigh NC 27601, USA
4
Corresponding author. E-mail: [email protected]; Telephone: 1-919-707-8861
2
Abstract
A new species of the rhacophorid frog genus Theloderma is described from the forested shoreline of the Nam Lik Reservoir, Vientiane Province, Laos. The new species differs from its congeners by having the combination of males with SVL
17.0–20.6; pearly asperities on dorsum; no vomerine teeth; disc diameter of finger III ca. 40% of tympanum diameter;
uniformly gray venter; light brown dorsum with darker brown and black markings; and a uniformly bronze iris with small
black reticulations. Molecular phylogenetic analysis of mitochondrial DNA sequence data infers that the new species is
most closely related to T. lateriticum from northern Vietnam. Evidence for the monophyly of Theloderma is reviewed.
Key words: Nam Lik Reservoir, Frankixalus, Nasutixalus, Theloderma lateriticum, Theloderma moloch
Introduction
The genus Theloderma Tschudi 1838 is a group of Asian rhacophorid treefrogs that has long captured the interests
of herpetologists and hobbyists alike. Many of the species are charismatically ornamented with spines, cryptic
color patterns resembling moss or bird droppings, and/or bright or highly contrasting color patterns (Orlov 1997;
Orlov et al. 2006; Dever et al. 2015; Li et al. 2016). Some species perform a balled defensive posture, and those
with known modes of reproduction breed in rain-filled tree or bamboo cavities (Orlov 1997; Orlov et al. 2006;
Grosjean et al. 2008; Nguyen et al. 2009; Rowley et al. 2011; Vassilieva et al. 2016). Several of the larger
ornamented species are regularly found in the international hobbyist trade (Orlov & Rybaltovksy 1999; Arinin &
Ryabov 2006; Rowley et al. 2007; Ziegler & Nguyen 2008; Gawor et al. 2012).
The taxonomic content of the genus is currently in a state of flux (Poyarkov et al. 2015; Frost 2016; Li et al.
2016). Twenty-four species are recognized here, following recent synynomies of two species from southern
Vietnam (Poyarkov et al. 2015), but excluding three species that were transferred from Nyctixalus Boulenger 1882
to Theloderma (Poyarkov et al. 2015; below). Molecular phylogenetic analyses have clarified species boundaries
in the genus (e.g. Rowley et al. 2011; Poyarkov et al. 2015) and provided evidence to justify the transfer of several
species of the genus Philautus Gistel 1848 to Theloderma (e.g. Yu et al. 2008; Rowley et al. 2011; Nguyen et al.
2014).
The Nam Lik Reservoir in Laos was formed by the Nam Lik 1-2 Hydroelectric Power Project’s damming of
the Nam Lik River in 2010 (Global Energy Observatory 2016). At full capacity, the reservoir covers approximately
24.4 km2 (Global Energy Observatory 2016). During fieldwork conducted in forested shoreline of the Nam Lik
Reservoir during 2014–2015, we obtained a series of very small-sized Theloderma that differed morphologically
and genetically from all other known species. Herein, we describe this species as new.
Accepted by J. Rowley: 16 Jun. 2016; published: 5 Aug. 2016
433
Materials and methods
Sampling. Specimens were collected in the field and fixed in 10% buffered formalin after preserving liver in 20%
DMSO-salt saturated storage buffer. Specimens were later transferred to 70% ethanol for permanent storage.
Specimens and tissue samples were deposited at the National University of Laos, Faculty of Natural Science,
Department of Biology (NUOL) and the North Carolina Museum of Natural Sciences (NCSM).
Morphology. Measurements were taken to the nearest 0.1 mm with dial calipers: snout-vent length (SVL);
head length from tip of snout to rear of jaws (HDL); maximum head width (HDW); snout length from tip of snout
to anterior corner of eye (SNT); eye diameter (EYE); horizontal diameter of tympanum (TMP); interorbital
distance (IOD); internasal distance (IND); shank (= tibia and fibula) length (SHK); thigh (=femur) length (TGH);
forearm length, from elbow to base of thenar tubercle (FAL); manus length from tip of third digit to base of thenar
tubercle (HND); and pes length from tip of fourth toe to base of inner metatarsal tubercle (FTL). Morphological
data for described species were taken from their original descriptions and Poyarkov et al. (2015). Specimens were
sexed by internal examination of gonads.
Molecules. Total genomic DNA was extracted using the ArchivePure DNA Cell/Tissue Kit (5 Prime). An
1,044–1,045 nucleotide basepair (bp) fragment of mitochondrial (mt) DNA that encodes part of the 16S rRNA
gene (16S) was amplified, sequenced and edited from two individuals (NCSM 84682 and NCSM 84683) following
Stuart et al. (2013). Sequences were deposited in GenBank under accession numbers KX095245 (NCSM 84682)
and KX095246 (NCSM 84683).
All available 16S sequences of Theloderma and Nyctixalus, and one species each of the outgroup taxa
Rhacophorus, Kurixalus, Nasutixalus, and Gracixalus (following Pyron & Wiens 2011; Poyarkov et al. 2015;
Jiang et al. 2016) were downloaded from GenBank on 28 March 2016. GenBank sequences were aligned to the
newly generated sequences using the default parameters in MAFFT v. 7 (Katoh & Standley 2013). Two alignments
were analyzed. In the first (hereafter as “complete” alignment), the entire sequence contained in each 16S GenBank
accession was used, including flanking regions of the 12S rRNA gene (12S) or transfer RNA (tRNA) if present. In
the second (hereafter as “trimmed” alignment), GenBank sequences were trimmed to match the length of the newly
generated sequences (and GenBank accession KU170015 of Theloderma moloch was excluded from analysis
owing to lack of overlap with the newly generated sequences). The complete alignment contained 137 terminal
taxa and 2,595 characters, and the trimmed alignment contained 136 terminal taxa and 1,105 characters.
The model of sequence evolution that best described the data (TIM2+I+G for both alignments) was inferred
using the Akaike Information Criterion as implemented in jModelTest 2 (Darriba et al. 2012). Four independent
Bayesian analyses were performed using MrBayes 3.2 (Ronquist et al. 2012). In each analysis, four chains were
run for 10 million generations using the default priors, trees were sampled every 2,000 generations, and the first
25% of trees were discarded as ‘burn-in.’ A 50% majority-rule consensus of the sampled trees was constructed to
calculate the posterior probabilities of tree nodes. Uncorrected pairwise (p) distances were calculated using PAUP*
4.0b10 (Swofford 2002).
Results
Theloderma lacustrinum sp. nov.
Holotype. NCSM 84682 (field tag SP 00299), adult male (Fig. 1), Laos, Vientiane Province, Feuang District, Ban
Naxang Village, bank of Nam Lik Reservoir, Houay Poungfan Stream, 18º48’59.5”N 102º08’06.1”E, 292 m elev.,
coll. 16 March 2015 by Niane Sivongxay, Monekham Davankham, Somphouthone Phimmachak, and Keochay
Phoumixay.
Paratypes. NUOL 00043 (one juvenile), NUOL 00044 (one male), same data as holotype. NUOL 00970 (one
male), same data as holotype except coll. 16 December 2014 by Niane Sivongxay, Monekham Davankham, and
Keochay Phoumixay. NCSM 84683, NUOL 00045 (two males), same data as holotype except Houay Kengthang
Stream, 18º51’58.2”N 102º07’20.6”E, 361 m elev., coll. 17 March 2015.
Etymology. The specific epithet taken from lacustrinus L., of lakes, in reference to Nam Lik Reservoir, a
large, man-made reservoir formed by a hydropower dam that has inundated the vicinity of the type locality.
Diagnosis. Assigned to the genus Theloderma on the basis of molecular data (Fig. 2). A very small-sized
(second smallest known) species of Theloderma having the combination of males with SVL 17.0–20.6; pearly
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SIVONGXAY ET AL.
asperities on dorsum; no vomerine teeth; disc diameter of finger III ca. 40% of tympanum diameter; uniformly gray
venter; light brown dorsum with darker brown and black markings; and a uniformly bronze iris with small black
reticulations.
Description of holotype. Habitus slender. Head longer than wide. Snout slightly rounded in dorsal and lateral
views; nostril oval, slightly oblique, much closer to tip of snout than to eye, internarial shorter than interorbital
distance; canthus rostralis indistinct, rounded, slightly constricted behind nares; lores oblique, concave; interorbital
region slightly convex, interorbital distance greater than upper eyelid width; pupil diamond-shaped, horizontal; eye
diameter and snout length subequal; tympanum distinct, round, 70% of eye diameter, tympanic rim elevated
relative to skin of temporal region; pineal ocellus absent; vomerine teeth absent; choanae oval, at margins of roof of
mouth; tongue wide, attached anteriorly, deeply notched posteriorly; supratympanic fold from posterior margin of
eye to level slightly posterior to axilla.
Forelimb slender. Finger tips with slightly expanded discs having weakly visible circummarginal grooves,
finger III disc width 40% tympanum diameter; fingers slender; relative finger lengths I < II < IV < III; webbing on
fingers absent; subarticular tubercles conspicuous, surfaces rounded, formula 1, 1, 2, 2; two indistinct, oval, palmar
tubercles in contact, subequal in size, surfaces flat; oval, thenar tubercle, surface flat; round, indistinct
supernumerary tubercles.
Hindlimb slender. Toe tips with slightly expanded discs having weakly visible circummarginal grooves,
diameter of discs subequal to those of fingers; toes slender; relative toe lengths I<II<III<V<IV; rudimentary web
on toe I to level of center of subarticular tubercle, continuing as fringe on preaxial side of toe II to base of tip, on
postaxial side of toe II to level of distal margin of subarticular tubercle, continuing as fringe on preaxial side of toe
III, on postaxial side of toe III to level midway between subarticular tubercles, on preaxial side of toe IV to level of
distal margin of proximal subarticular tubercle, continuing as fringe to base of disc, postaxial side of toe IV to level
of proximal margin of penultimate subarticular tubercle and continuing as a fringe to base of tip, and on toe V to
level of proximal margin of subarticular tubercle; subarticular tubercles conspicuous, surfaces rounded, formula 1,
1, 2, 3, 2; inner metatarsal tubercle oval, surface flat; outer metatarsal tubercle weakly visible.
Skin smooth, with pearly asperities on side of head and dorsal surfaces of head, back, limbs, and outer margin
of foot; venter coursely granular; very weak dermal fringe on outer margin of toe V from base of foot to base of tip.
Velvety, ovoid nuptial pad on dorsal and posterior margin of finger I.
Color of holotype in life. Iris uniformly bronze with small black reticulations; dorsum light brown with brown
hourglass-shaped marking on top of head, continuing as stripe on back to sacrum; dorsal surfaces of limbs orangebrown with irregular, black crossbands, dorsal surfaces of digits bronze (matching iris); flank gray; irregular black
band from tip of snout to axilla, as narrow band below canthus and eye but as enlarged spot over nostril and
tympanum; irregular, large, black spots and reticulations on flank; ventral surfaces uniformly gray.
Color of holotype in preservative. Black coloration faded to brown; brown marking on top of head and back
less distinct; bronze coloration on digits faded to brown.
Variation. Paratypes closely resemble holotype. NUOL 00043 lacks a nuptial pad and has the smallest body
size in the type series (SVL 15.5), and so is deemed immature. Females, eggs, larvae, and vocalizations are
unknown. Measurements are summarized in Table 1.
Molecules. The new species was recovered in both analyses as sister to T. lateriticum Bain, Nguyen & Doan
2009, with a Bayesian posterior probability of 1.00 (Fig. 2). The holotype and sequenced paratype (NCSM 84683)
differed by only a single insertion-deletion in the 16S gene fragment (and so had a p-distance of 0%). Four
sequences of T. lateriticum (GenBank accession numbers LC012848–51) had p-distances of 0.33–4.40%. The new
species and T. lateriticum (n=4) had p-distances of 10.54–10.81%. Bayesian inference of the trimmed alignment
differed from that of the complete alignment only by failing to recover statistical support for the monophyly of
Theloderma laeve and a sister relationship between the outgroup genera Kurixalus and Nasutixalus.
Distribution, natural history, and conservation. Theloderma lacustrinum sp. nov. is currently known only
from the type and paratype localities (Fig. 3), where it was collected at night (1820–2215 h) on leaves 30–80 cm
above the ground near two rocky streams (292–361 m elev.) in semi-evergreen forest that flow into the Nam Lik
Reservoir (Fig. 1). The two known localities are approximately 5.7 air-km apart. We did not observe the new
species at these localities during visits in April, July, and September 2015, suggesting that it may be seasonally
active or most detectable during cooler, drier months (e.g., at lower heights in the forest canopy). The vicinity of
the two known localities had not been surveyed for amphibians prior to construction of the hydroelectric dam, and
thus it is not known how inundation from the reservoir may have impacted the range of the species.
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TABLE 1. Measurements (mm) of types of Theloderma lacustrinum sp. nov. Abbreviations defined in the text.
Measurement
Holotype male
n=1
All males
n=5
Range; Mean ± SD
Juvenile
n=1
SVL
20.6
17.0–20.6; 18.8 ± 1.7
15.5
HDL
9.7
7.3–9.7; 8.3 ± 1.1
6.9
HDW
6.6
5.5–6.7; 6.2 ± 0.6
5.3
SNT
3.0
2.8–3.1; 3.0 ± 0.1
2.9
EYE
2.8
2.0–3.0; 2.5 ± 0.4
2.6
TMP
1.9
1.4–1.9; 1.7 ± 0.3
1.3
IOD
3.8
2.5–3.8; 3.2 ± 0.5
2.9
IND
1.8
1.6–1.8; 1.7 ± 0.1
1.6
SHK
12.0
9.7–12.0; 11.0 ± 0.9
8.5
TGH
10.9
9.3–11.1; 10.3 ± 0.7
8.2
FAL
10.2
8.4–10.2; 9.2 ± 0.8
7.3
HND
5.2
4.5–5.3; 4.9 ± 0.4
4.0
FTL
8.1
6.2–8.1; 7.1 ± 0.8
6.0
Comparisons. Theloderma lacustrinum sp. nov. is most closely related to T. lateriticum (Fig. 2) from Lao Cai,
Bac Giang, and Son La Provinces, Vietnam (Bain et al. 2009, Nguyen et al. 2015), but is readily distinguished from
T. lateriticum by having a light brown dorsum with dark markings (nearly uniformly red in lateriticum), by having
a uniformly gray venter (distinctly spotted in lateriticum), and by having a bronze iris (deep brick-red in
lateriticum).
Theloderma lacustrinum sp. nov. differs from all other Theloderma by having adult males with SVL <21 mm,
except T. baibengense (Jiang, Fei & Huang 2009) that has adult males with SVL 15.0–16.2, n=2 (Jiang et al. 2009).
However, T. lacustrinum sp. nov. is readily distinguished from T. baibengense by having a light brown dorsum
with dark markings (dark with conspicuous white pattern in baibengense), by having a uniformly gray venter
(distinctly contrasting marbled pattern in baibengense), and by having a bronze iris (reddish-brown in
baibengense).
Twelve additional species of Theloderma have small (sensu Rowley et al. 2011) adult body sizes, with SVL
<35 mm: T. albopunctatum (Liu & Hu 1962), T. andersoni (Ahl, 1927), T. asperum (Boulenger 1886), T. laeve
(Smith 1924), T. licin McLeod & Norhayati 2007, T. nebulosum Rowley, Le, Hoang, Dau & Cao 2011, T. palliatum
Rowley, Le, Hoang, Dau & Cao 2011, T. petilum (Stuart & Heatwole 2004), T. rhododiscum (Liu & Hu 1962), T.
stellatum Taylor 1962, T. truongsonense (Orlov & Ho 2005), and T. vietnamense Poyarkov, Orlov, Moiseeva,
Pawangkhanant, Ruangsuwan, Vassilieva, Galoyan, Nguyen & Gogoleva, 2015. Theloderma lacustrinum sp. nov.
differs from T. laeve and T. truongsonense by having dorsal asperities (absent in laeve and truongsonense).
Theloderma lacustrinum sp. nov. differs from T. petilum by lacking vomerine teeth (present in petilum) and by
having relatively smaller finger discs, with the disc diameter of finger III ca. 40% of tympanum diameter (≥65% in
petilum). Theloderma lacustrinum sp. nov. differs from T. albopunctatum, T. asperum, T. baibengense, T. licin, and
T. stellatum by having a light brown dorsum with dark markings (dorsum dark with conspicuous white pattern in
albopunctatum, asperum, baibengense, and stellatum, nearly uniformly brown or white in licin). Theloderma
lacustrinum sp. nov. differs from T. albopunctatum, T. asperum, T. licin, T. nebulosum, T. palliatum, T.
rhododiscum, T. stellatum, T. truongsonense, and T. vietnamense by having a uniformly gray venter (distinctly
contrasting spotted, marbled or reticulated ventral pattern in albopunctatum, asperum, baibengense,
chuyangsinense, lateriticum, licin, nebulosum, palliatum, rhododiscum, stellatum, truongsonense, and
vietnamense). Theloderma lacustrinum sp. nov. differs from T. andersoni by lacking two yellow spots on flank
(present in andersoni). Theloderma lacustrinum sp. nov. differs from T. laeve, T. nebulosum, T. palliatum, T.
petilum, and T. truongsonense by having a uniformly colored iris (distinctly bicolored in laeve, nebulosum,
palliatum, petilum, and truongsonense).
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SIVONGXAY ET AL.
FIGURE 1. Holotype NCSM 84682 (A) and paratype NUOL 00970 (B) of Theloderma lacustrinum sp. nov. in life.
Microhabitat (C) and macrohabitat (D) of type locality of T. lacustrinum sp. nov. Dorsal view (E) and ventral view (F) of
holotype NCSM 84682 of T. lacustrinum sp. nov. immediately prior to preservation. Dorsal view (G) and ventral view (H) of
holotype NCSM 84682 of T. lacustrinum sp. nov. in preservation. Scale bar = ca. 10 mm.
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FIGURE 2. Fifty percent majority-rule consensus phylogram resulting from Bayesian analysis of 2,595 aligned sequence
characters (“complete” alignment) of the mitochondrial 16S gene and portions of flanking 12S and tRNA genes from
rhacophorid frogs. Black circles at nodes indicate Bayesian posterior probabilities ≥ 0.99, and open circles at nodes indicate
Bayesian posterior probabilities ≥ 0.95. Numbers at terminal tips are GenBank accession numbers. An asterisk (*) indicates the
species name was modified from that provided in the GenBank accession at the time of download.
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FIGURE 3. Map illustrating the type locality (white star) of Theloderma lacustrinum sp. nov. in Vientiane Province, Laos, and
the type locality (black star) of its closest known relative, T. lateriticum, in Lao Cai Province, Vietnam.
Discussion
The monophyly of Theloderma has been questioned, but primarily on the bases of an erroneously identified
voucher that has been perpetuated in the literature and misinterpretation of unresolved nodes in phylogenetic trees.
Sequences attributed to Theloderma moloch (voucher specimen “6255Rao,” GenBank accessions GQ285679,
GQ285690, GQ285721, GQ285753, GQ285782, GQ285800) by Li et al. (2009), and that were re-incorporated into
numerous other phylogenetic studies (e.g., Rowley et al. 2011, Hertwig et al. 2013; Li et al. 2013; Nguyen et al.
2015; Poyarkov et al. 2015; Biju et al. 2016; Jiang et al. 2016; Li et al. 2016), resulted in the disparate placement
of T. moloch within the Rhacophoridae, and consequently, the non-monophyly of Theloderma. Voucher “6255Rao”
(and its associated sequences) has been re-assigned to the newly-described genus and species Nasutixalus
medogensis Jiang, Yan, Wang, Zou, Li & Che 2016, and, almost concurrently, to an unnamed species in the newlydescribed genus Frankixalus Biju, Senevirathne, Garg, Mahony, Kamei, Thomas, Shouche, Raxworthy,
Meegaskumbura & Van Bocxlaer 2016. After excluding the misidentified sequences associated with voucher
“6255Rao,” numerous phylogenetic studies have actually recovered Theloderma to be a monophyletic clade, and
sister to a monophyletic clade containing species of Nyctixalus (e.g., Wilkinson et al. 2002; Frost et al. 2006;
Grosjean et al. 2008; Li et al. 2008, 2009, 2013; Yu et al. 2009; Hertwig et al. 2011; Pyron & Wiens 2011; Dever et
al. 2015; Meegaskumbura et al. 2015; Nguyen et al. 2015; Biju et al. 2016; Jiang et al. 2016; Li et al. 2016; this
study). The monophyly of Theloderma is further corroborated in studies that incorporated new, and presumably
correctly identified, samples of T. moloch (Jiang et al. 2016; Biju et al. 2016; Li et al. 2016; this study). The date of
publication printed on the hard cover of Zoological Research Volume 37, Issue 1, was 18 January 2016, and that
provided on the first page of PLoS ONE 11(1), e0145727, was 20 January 2016, giving publication priority to
Nasutixalus by two days. As such, we treat Frankixalus Biju, Senevirathne, Garg, Mahony, Kamei, Thomas,
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Shouche, Raxworthy, Meegaskumbura & Van Bocxlaer 2016 as a subjective junior synonym of Nasutixalus Jiang,
Yan, Wang, Zou, Li & Che 2016. We also maintain T. moloch (Annandale 1912) in Theloderma on the basis of the
newly available (and hopefully correctly identified) samples of this species (Biju et al. 2016; Jiang et al. 2016; Li et
al. 2016; this study).
The Bayesian inference tree generated by Poyarkov et al. (2015:253) placed Nyctixalus within Theloderma
(other than the erroneously identified T. moloch sample), and consequently these authors synonymized the two
genera, treating Nyctixalus as a subgenus of Theloderma. However, the node supporting that placement lacked
statistical support (Bayesian posterior probability of 0.86), and the relationships among (i) a clade containing T.
horridum and the T. stellatum group, (ii) a clade containing Nyctixalus, and (iii) a clade containing remaining
species of Theloderma (other than the erroneously identified T. moloch sample) were unresolved in Poyarkov et al.
(2015). Rowley et al. (2011:8) also found statistical support for the phylogenetic placement of Nyctixalus pictus
within Theloderma in their Bayesian inference tree, but not in their maximum likelihood tree. Their analyses were
based on short sequence fragments of 16S and minimal taxon sampling (Nguyen et al. 2015), as the authors only
intended to “support the placement of our newly collected specimens within the genus Theloderma, rather than to
resolve phylogenetic relationships within the group” (Rowley et al. 2011: 6). Owing to the statistical polytomy in
Poyarkov et al. (2015), the limitations in the phylogenetic analyses of Rowley et al. (2011), our results (Fig. 2), and
those of numerous other studies (cited above), we reject the synonymy proposed by Poyarkov et al. (2015), and
maintain Theloderma and Nyctixalus as separate genera.
Acknowledgements
The Feuang District Office of Natural Resources and Environment, Laos, provided permission and participated in
the field research. Fongfany Lybounyasao and Phousavanh Inthapanya assisted with fieldwork. The Wildlife
Conservation Society Laos Program provided logistical support. The manuscript benefited from discussion with
Darrel Frost. The U.S. National Science Foundation (grant DEB-1145922), and the Partnerships for Enhanced
Engagement in Research (PEER) Science program (grant PGA-2000003545), which is a partnership between the
U.S. Agency for International Development (USAID) and the U.S. National Science Foundation, funded this
research.
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