Stumpffia analamaina is a very small frog, with a snout-vent length range of 10 - 13.4 mm in males and females (Klages et al. 2013, Rakotoarison et al. 2017). The head is marginally longer than it is wide and is narrower than the slender body. The snout is round or slightly pointed in both the dorsal and lateral view and protrudes slightly. The nostrils are barely protuberant, oriented laterally, and are positioned equally between the snout and the eyes. The canthus rostralis is rounded and faintly marked and the loreal region can vary from straight to slightly convex. Compared to overall body size, the eyes are rather large, with round pupils, and an interorbital distance almost twice that of the internarial distance. Both the tympanum and supratympanic fold are obscured. The forelimbs are slender and the hands have three flattened, palmar tubercles with faint markings on them, of which the rounded inner tubercle is largest. The hands and feet have slightly reduced first digits. The hand to snout-vent ratio length ranges from 0.22 - 0.26. The short fingers are generally round, unwebbed, and lack obvious terminal discs. Relative finger lengths from least to greatest are I < IV < II < III. Small, flat, rounded subarticular tubercles can be found on finger III (2 tubercles) and finger II (1 tubercle), and the prepollex is absent. The hands of the holotype lack nuptial pads. The hindlimbs are slender as well and the tibiotarsal articulation extends to the nostril when pressed adjacent to the body. The tibia and foot are approximately the same size. The feet each have an inner metatarsal tubercle that is an ambiguous flattened ovoid shape. There is no outer metatarsal tubercle. The ratio of foot plus tarsus to the snout-vent length is 0.68 - 0.81. The toe tips are marginally wider than the rest of the toe with circummarginal grooves, but only the IV toe has a faintly distinguishable toe disc. The toes have rounded subarticular tubercles. Like the fingers, the toes are unwebbed. They have relative toe lengths of I < II < V < III < IV with the second toe ranging from slightly to distinctly longer than the fourth, and the third ranging from slightly to distinctly longer than the fifth. The body shape is externally very similar to other species of Stumpffia. The dorsum may have small tubercles but both the dorsum and ventrum become smooth in preservative (Klages et al. 2013).
Stumpffia analamaina is differentiated from other Stumpffia by several morphological and vocal characters. It is distinguished from both S. mamitika and S. larinki by slightly smaller body size (10 - 12 mm vs. 12.3 – 15.0 mm and 12.5 – 12.8 mm, respectively), and a longer call duration than S. mamitika. It is distinguished from S. helenae by slightly smaller body size (10 - 12 mm rather than 14 - 15 mm) and the absence of expanded toe discs present in the digits of both the hind and forelimbs of S. helenae. Stumpffia analamaina is drastically smaller than S. be and S. staffordi (10 - 12 mm vs. ~25 mm and 27 - 28 mm, respectively) and both species have expanded toe discs (vs. lack of toe discs). It is also distinctly smaller than S. megsoni and S. hara (10 - 12 mm vs. 21 - 22 mm and 22 - 25 mm, respectively) and again both species have expanded toe discs (vs. lack of toe discs). It is distinct from S. roseifemoralis by smaller body size (10 - 12 mm vs. 18 - 19 mm), and lack of red pigmentation on the ventral side of limbs. Reduced digit number in S. tridactyla, and S. tetradactyla distinguish these species from the more typical digit morphology of S. analamaina. The markings of S. analamaina are much less distinct browns and grays compared to the clearly defined brown patches of S. grandis (also with black markings on blue background across the venter), and S. grandis is considerably larger (19 - 24 mm vs. 10 - 12 mm). Stumpffia analamaina is marginally smaller than S. psologlossa (10 - 12 mm vs. 14 - 16 mm), has a more obviously reduced first toe, and an advertisement call characterized more by a series of short chirps, rather than the short trills of S. psoglossa. The dorsum of S. analamaina is much less tubercular, or completely smooth, compared to the plainly tubercular dorsum of S. gimmeli; in addition, S. analamaina has a slightly shorter first toe and first finger (well developed in S. gimmeli), a smaller body size (10 – 12 mm vs. 15 - 18 mm), and lacks terminal discs on all digits (at least slightly developed toe discs in S. gimmeli, particularly in larger individuals) (Klages et al. 2013). Stumpffia analamaina has a significantly smaller body than S. kibomena (10 - 12 mm vs. 17 – 21 mm) (Glaw et al. 2015).
Stumpffia analamaina is most likely to be confused with S. pygmaea, S. madagascariensis, and its sister taxon S. yanniki., However, the distinct border between the light dorsum and darker flanks in S. madagascarensis (Köhler et al. 2010) is absent in S. analamaina; in addition, the hands and feet of S. analamaina are distinctly longer relative to their snout-vent lengths than S. madagascareiensis (0.22 - 0.26 and 0.68 - 0.81 respectively vs. 0.16 mm and 0.61) and S. pygmaea (0.16 - 0.18 and 0.53–0.63) (Vences and Glaw 1991; Klages et al. 2013). Stumpffia analamaina and S. yanniki only differ marginally in terms of morphology: S. analamaina has slightly larger hands and feet relative to their snout-vent lengths (0.22 - 0.26 and 0.68 - 0.81 respectively vs. 0.18 - 0.19 and 0.62 - 0.65) than S. yanniki (Rakotoarison et al. 2017).
In life, S. analamaina has a grey to brownish-red background color consistent throughout the dorsum with no contrast to the ventrum. On the dorsum there is generally an ambiguous X-shaped brown patterning that varies amongst individuals. The flanks are densely covered with small whitish spots. Distinct boundaries in coloration between the dorsum and flanks may be present. The inguinal region has a distinctive dark patch and faded brown bands and patches can be seen on both forelimbs and hindlimbs. These brown patches often extend along the flanks and the coloration continues along the supratympanic fold into the snout. The eyes are distinct with copper brown irises. In preservative, colors remain relatively unchanged after two years, with slightly more prominent contrast between the patterning and background dorsal color (Klages et al. 2013).
Morphology remains consistent across individuals, with no sexual dimorphism or dichromatism. The red-brown background color prevalent in some specimens is substituted with a lighter beige coloration in other specimens. The brown dorsal patterning of certain individuals extends into a general X-shaped pattern, reaching anteriorly towards the eyes and posteriorly towards the base of the hindlimbs; whereas, in other individuals, this patterning forms more of the classic “gummi-bear” marking seen in other Stumpffia, such as S. psologlosa (Glaw and Vences 2007; Klages et al. 2013).
Distribution and Habitat
Country distribution from AmphibiaWeb's database: Madagascar
Stumpffia analamaina, like the rest of Stumpffia, is endemic to Madagascar and, more specifically, is found in the northwestern region of Madagascar. Stumpffia analamaina’s primary habitat is in dense leaf litter or shrubbery of dry deciduous forest fragments, and it can be found alongside streams in high concentrations during the dry austral winter when humidity is low (Klages et al. 2013, Rakotoarison et al. 2017).
The current distribution is based off of two separate populations, one from where the type specimen was collected and another population much further south that is marginally genetically differentiated and thus only recognized as S. analamaina in 2017 on the basis of morphology and molecular genetics (Rakotoarison et al. 2017). The type specimen was found in the forest habitat 27 km northwest of Antsohihy at 140 m asl and is suspected to inhabit the regions north and northeast of Antsohihy as well (Klages et al. 2013). The secondary locality is in Ankarafantsika National Park, extending much further south of Antsohihy at 139 m asl (Rakotoarison et al. 2017, IUCN 2016).
Life History, Abundance, Activity, and Special Behaviors
Stumpffia analamaina is a terrestrial leaf litter inhabitant and can be found in dry, deciduous, gallery forest habitats near small streams; occurring in sympatry with several other frogs: Mantidactylus ulcerosus, Mantella ebenaui, Boophis tampoka, and B. brachychir (Klages et al. 2013).
In captivity, S. analamaina are mainly active at night and generally moved slowly, only jumping when disturbed; however, jumps have been recorded as long as 60 cm outside of a terrarium (Klages et al. 2013).
Calls seem to be relatively independent of rainfall and air temperature, and most prevalent during evening/nighttime (although they have been observed calling on rainy mornings). Peak calling time appears to be at 17:00, and individuals make their calls either from cover underneath the leaf litter, or on top of the leaf litter.
The advertisement call of S. analamaina males is characterized by a noisy display of small chirping notes (21 - 27 per minute) that are less tonal than most other Stumpffia species (Klages et al. 2013). The calls at the species two locations varied slightly. Individuals from the type locality had calls recorded in terrariums with air temperatures of 23°C, and emitted calls from underneath dead leaves. The duration of calls is 228 - 286 ms (260 ± 23.6 ms) and inter-call intervals are 1974 - 2260 ms (2242 ± 136 ms). The dominant frequency of the calls range from 6130 - 6330 Hz (6210 ± 80.4 Hz) (Rakotoarison et al. 2017). The maximum call energy ranges from 6000 – 6400 Hz (Klages et al. 2013), and the distinct variables of S. analamaina’s call is quite similar sounding to that of S. pygmaea. Vocalizations that have been recorded in captivity are arranged in series of 3 – 8 notes, but it is speculated that S. analamaina might be able to emit notes in a much longer series for many minutes on end. Across five recorded note series in captivity, notes were produced at constant intervals and all sounded quite similar to one another. Consequently, the authors are confident that it represents the true advertisement call, despite a possibility that calls were shorter in duration than what was actually recorded due to echoing in the captive terraria. Yet, if that is the case it would still be a diagnostic taxonomic character as the measured notes as a whole were a shorter length than most other Stumpffia (Klages et al. 2013).
At the secondary locality (Ampijoroa, Ankarafantsika National Park) call duration is 134 - 175 ms (153 ± 13.5 ms) and inter-call intervals are 1055 - 1160 ms (1120 ± 30 ms). The dominant frequency of the calls range from 5343 - 5531 Hz (5399 ± 91 Hz). The Ampijoroa population appears to call twice as frequently, and the calls are more tonal in nature; however, these could both be possible artifacts as calls were not only recorded at different temperatures, but different environments at the different localities: in captivity in terrariums for the specimens from the type locality, and in the field for specimens in the Ampijoroa population (Rakotoarison et al. 2017).
Information on reproductive mode in all of Stumpffia is data deficient, however, some reproductive activity has been observed in S. analamaina in captivity by Klages et al. (2013). Stumpffia analamaina construct foam nests (~50 mm in diameter) and in captivity one individual was found guarding a nest for several days after a period of high intensity calling. The foam nest was approximately 50 mm in diameter (Klages et al. 2013). Foam nests produced in the wild are expected to be made amongst the leaf litter and likely yield non-feeding tadpoles based on field observations of S. pygmaea’s reproductive mode (Rakotoarison et al. 2017). Despite finding the foam next in captivity, no S. analmaina egg laying has observed (Klages et al. 2013).
Stumpffia analamaina has been observed exhibiting lateral tongue protrusion during prey capture. This trait is not known in any other member of the Cophylinae subfamily, but is common in the rest Microhylidae (Klages et al. 2013, Meyers et al. 2004)
Trends and Threats
This species is very rare and not well studied. Adequate microhabitat (damp leaf litter adjacent to streams with intact gallery forest) for S. analamaina is scarce and further threat is posed by rapid habitat destruction due to logging and burning for agricultural clearing purposes in the northwestern part of Madagascar (Klages et al. 2013). This species is not confirmed to live within the boundaries of a protected area but may possibly occurs within the Bora Special Reserve. Until this is confirmed, the species has no formal protection. Protection and management of remaining forest fragments is needed to protect S. analamaina, however, further research into life history, population size and trends, taxonomy and threats is also necessary to develop an effect management plan for the species (IUCN 2016).
Possible reasons for amphibian decline
General habitat alteration and loss
Habitat modification from deforestation, or logging related activities
Subtle changes to necessary specialized habitat
The species authority is: Klages, J., Glaw, F., Köhler, J., Müller, J., Hipsley, C. A., Vences, M. (2013). “Molecular, morphological and osteological differentiation of a new species of microhylid frog of the genus Stumpffia from northwestern Madagascar.” Zootaxa, 3717, 280-300.
Stumffia analamaina is characterized by strong mitochondrial differentiation from other species, with more than 10% uncorrected pairwise distance within the 16S rRNA gene. Additionally, it shows uniqueness in the RAG1 nuclear gene (Klages et al. 2013).
Rakotoarison et al. (2017) has analyzed the 12S, the two 16S fragments, Cox-1, and Rag-1 genes in Stumpffia sp. using Partitioned Bayesian Inference. The analysis provided a well-resolved tree where four main clades were distinguished. Stumpffia analamaina is placed within the northern clade, with a total of 17 species. It is further partitioned into the northwestern clade, which includes small to miniaturized species, such as S. yanniki, S. madagascariensis, and S. pygmaea. Based on these analyses, S. analamaina is most closely related to S. yanniki (Rakotoarison et al. 2017).
The origin of this species epithet, “analamaina”, comes from its unique habitat. The name is composed of two Malagasy words, “ala” meaning “of the forest” and “maina” meaning “dry” and is in reference to the dry northwestern forest habitat of Madagascar (Klages et al. 2013).
Glaw, F., and Vences, M. (2007). Field Guide to the Amphibians and Reptiles of Madagascar. Third Edition. Vences and Glaw Verlag, Köln.
IUCN SSC Amphibian Specialist Group. (2016). ''Rhombophryne analamaina''. The IUCN Red List of Threatened Species 2016: e.T79680099A84185500. https://dx.doi.org/10.2305/IUCN.UK.2016-1.RLTS.T79680099A84185500.en. Downloaded on 01 May 2020.
Klages, J., Glaw, F., Köhler, J., Müller, J., Hipsley, C.A., Vences, M. (2013). ''Molecular, morphological and osteological differentiation of a new species of microhylid frog of the genus Stumpffia from northwestern Madagascar.'' Zootaxa, 3717(2), 280-300. [link]
Köhler, J., Vences, M., D'Cruze, N., and Glaw, F. (2010). ''Giant Dwargs: discovery of a radiation of large-bodied 'stump-toed frogs' from karstic cave environments of northern Madagascar.'' Journal of Zoology, 282, 21-38.
Meyers, J.J., O'Reilly, J.C., Monroy, J.A., Nishikawa, K.C. (2004). ''Mechanism of tongue protraction in microhylid frogs.'' Journal of experimental biology, 207, 21-31. [link]
Rakotoarison, A., Scherz, M.D., Glaw, F., Köhler, J., Andreone, F., Franzen, M., Glos, J., Hawlitschek, O., Jono, T., Mori, A., Ndriantsoa, S.H., Raminosoa, N.R., Riemann, J.C., Rödel, M., Rosa, G.M., Vieites, D.R., Crottini, A., Vences, M. (2017). ''Describing the smaller majority: integrative taxonomy reveals twenty-six new species of tiny microhylid frogs (genus Stumpffia) from Madagascar.'' Vertebrate Zoology, 67(3), 271-398.
Written by Jimmy W. Wehsener (jwehsener AT berkeley.edu), UC Berkeley
First submitted 2020-08-27
Edited by Ann T. Chang (2020-08-27)
Species Account Citation: AmphibiaWeb 2020 Stumpffia analamaina <http://amphibiaweb.org/species/8074> University of California, Berkeley, CA, USA. Accessed Nov 25, 2020.
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Citation: AmphibiaWeb. 2020. <http://amphibiaweb.org> University of California, Berkeley, CA, USA. Accessed 25 Nov 2020.
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