A new taxonomic status for Darwin’s “ predicted ” pollinator: Xanthopan praedicta stat. nov. (Lepidoptera Sphingidae Sphinginae) Author Minet, Joël Author Basquin, Patrick Author Haxaire, Jean Author Lees, David C. Author Rougerie, Rodolphe text Antenor 2021 2021-12-31 8 1 69 86 journal article 10.5281/zenodo.5856141 960fb566-8c5d-4412-ac54-ed1679cf45f8 2275-9298 5856141 Morphological differences between Xanthopan m. morganii and X. m. praedicta Illustrations in ARDITTI & al . (2012) may give the impression that “ Xanthopan morganii morganii ” is a highly variable taxon, but some of the specimens pictured were erroneously referred as originating “from Gabon” ( ibid ., fig. 7 A-C) when actually being from Madagascar, and thus representing “ Xanthopan morganii praedicta ”. The main differences between X. m. morganii and X. m. praedicta are listed hereafter, following an anteroposterior order for the enumeration of the corresponding characters. Head . 1 , proximal section of the proboscis – morganii : reddish brown; praedicta : black. 2 , ventral vestiture of segment 1 of the labial palpus (in lateral view) – morganii : with numerous dark grey scales, at least distally, so that the coloration of segment 1 contrasts moderately with the brown region of segment 2; praedicta : relatively light-coloured, contrasting strongly with the dark brown region of segment 2. Foreleg . 3 , tarsus in dorsal view – morganii : basitarsus lightcoloured, contrasting strongly with tarsomeres 2-5, which are black or blackish (with some white at the extremities of tarsomere 2); praedicta : tarsomere 2 at least partially brown, with an intermediate coloration between that of the basitarsus (ochre or pale brown) and that of tarsomeres 3-5 (dark brown). Forewing shape . 4 , termen – morganii : angulate at M 3 (fig. 2 , a-d); praedicta : regularly arched between CuA 2 and the apex (fig. 2 , e-h). Forewing (dorsal surface) . 5 , dark subbasal costal mark – morganii : a trian - gular spot, extending distad along the costa (fig. 2 , a-c); praedicta : not distinctly triangular, usually in the form of a short zigzag, which is roughly parallel to vein CuA 2 and often unites two very small dark spots (the costal spot being neither clearly triangular nor markedly prolonged distad along the costa: fig. 2 , e and g). 6 , lower discal light-coloured spot (on discocellular m2-m3) – morganii : typically broad, greyish white, without well defined dark edges, and often slightly extended on the base of vein M 2 (fig. 2 , a-c); praedicta : thin, circumflexshaped, white or whitish, relatively well delimited, and not extended distad on M 2 (fig. 2 , e and g). 7 , region between veins M 1 and M 3 – morganii : greyish or beige (fig. 2 , a-c), sometimes (proximally) with a narrow pinkish band between M 2 and M 3 (fig. 2 , c); praedicta : typically, with a large chestnut-brown area, which is ill-delimited above M 2 and does not extend beyond the (often obsolete) submarginal line (fig. 2 , e and g). 8 , blackish subapical streak below vein Rs 4 – morganii : usually thin (fig. 2 , a-c), though sometimes fairly broad (plate IV, fig. 7 , in CARCASSON, 1976 ); praedicta : usually broader (fig. 2 , e and g; contrary to a statement in ROTHSCHILD & JORDAN (1903) , the broad streak is not a constant characteristic of the taxon praedicta ). Forewing (ventral surface) . 9 , apical region – morganii : uniformly brownish (fig. 2 , d); praedicta : with an upper light-coloured area that contrasts with the lower – brown – apical area (fig. 2 , f and h; the light-coloured area is occasionally faintly indicated, notably in “faded” specimens; it was found to be almost absent in a specimen in fairly good condition collected by one of us (PB) in Lakato forest, near Moramanga). FIG. 2. — Male (a-f) and female (g, h) adults of Xanthopan morganii (a-d) and X. praedicta stat. nov. (e-h), in dorsal (a-c, e, g) and ventral (d, f, h) views. – a , Adiopodoumé, Ivory Coast. – b , Ziéla, Dosso, Niger. – c and d , Grande Comore (= Ngazidja). – e and f , Andrangoloaka, east of Lake Mantasoa, Madagascar. – g and h , Ambatofitorahana, Betsileo territory, Madagascar. – Forewing lengths = 57 mm (a-d), 69 mm (e, f), and 79 mm (g, h). Hindwing (dorsal surface) . 10 , central and anal light-coloured patches – morganii : orange yellow, contrasting with the whitish colour of the discal cell and costal region (fig. 2 , a-c); praedicta : relatively pale yellow, having approximately the same colour as the cell and the costal region (fig. 2 , e and g). 11 , yellow central patch – morganii : well-developed and distinctly triangular (fig. 2 , a-c); praedicta : usually a bit smaller and subtriangular (fig. 2 , e and g). 12 , dark central region – morganii : uniformly brown (fig. 2 , a-c); praedicta : usually greyish brown, with dark brown longitudinal stripes between the veins (fig. 2, g). 13 , dark streak of the anal angle – morganii : usually well indicated, edged with whitish or pale grey (fig. 2 , a-c); praedicta : usually less conspicuous (fig. 2 , e and g). 14 , fringe of the termen – morganii : dark brown at vein apices, whitish or greyish between veins; praedicta : pale chestnut (at vein apices) and yellowish (between veins). Hindwing (ventral surface) . 15 , light-coloured patch below the discal cell – morganii : yellow, well indicated, extending from wing base to the median section of CuA 2 and the basal region of CuA 1 (fig. 2 , d); praedicta : rather inconspicuous and ill-delimited, never large, usually yellowish or pinkish (fig. 2 , f and h). Thorax and abdomen (coloration) . 16 , ventral region – morganii : white or yellowish white (fig. 2 , d); praedicta : always pink (fig. 2 , f and h), as already mentioned in the original description ( ROTHSCHILD & JORDAN, 1903: 32 ). Male genitalia . 17 , in the phallus, ratio between a (= distance between the anterior “elbow” and the posterior extremity of the bulbus ejaculatorius) and b (= length of the sclerotized phallus) – morganii : a long, varying from 48 to 57 percent of b (fig. 3 , b); praedicta : a usually shorter, varying from 41 to 48 percent of b (fig. 3 , f and h). 18 , saccus – morganii : elongate (fig. 3 , a and c; fig. 7, g and h); praedicta : shorter and more robust (fig. 3 , e and g; fig. 7 , e and f). 19 , valva – morganii : relatively narrow, its greatest breadth (c) being less than 1/3 of its total length (d) (fig. 3 , a and c); praedicta : broader, c being more than 1/3 of d (fig. 3 , e and g). These distances have been measured on spread genitalia, with c extending transversely from the most convex part of the costa to the ventral edge of the valva, below the praesacculus. FIG. 3. — Male genitalia of Xanthopan morganii (a-d) and X. praedicta stat. nov. (e-h); phalli detached, in ventral view (b, d, f, h). – a and b , specimen from Ivory Coast. – c and d , Tsoundzou, Mayotte. – e and f , Antongil Bay, Madagascar. – g and h , Amber Mountain (Montagne d’Ambre), Madagascar. – same magnification for a-d; for e-h. 20 , praesacculus (“harpe”) – morganii : usually with smooth dorsal edge (disregarding pilosity), except for a strong, undivided, dorso-distal tooth (fig. 3 , a and c; fig. 4 , f and h-l), occasionally with a supernumerary small tooth proximad of the dorso-distal tooth (fig. 4 , g); praedicta : dorsal edge of the praesacculus irregular, with several teeth proximad of the broad dorso-distal projection, which is not very prominent and usually has a crenulate edge (fig. 3 , e and g; fig. 4 , a-e). According to figure 53 in GRIVEAUD (1959) , the dorso-distal projection of the praesacculus of X. praedicta may occasionally resemble that of X. morganii but it should be noted that his figure is neither a photograph nor a detailed drawing. KITCHING (2002: 489 , character 65) had already noticed the difference between X. morganii and X. [“ m .”] praedicta with regard to the dorsal edge of the proximal region (“lobe”) of the “harpe”, described as “rather” truncate and smooth in the case of the former taxon, provided with “several blunt, irregular teeth” in the case of the latter. FIG. 4. — Praesacculus in different specimens of Xanthopan praedicta stat. nov. (a-e) and X. morganii (f-l). – a , Antongil Bay. – b , Perinet (Analamazaotra). – c , Fort-Dauphin (Tôlañaro). – d , Amber Mountain. – e , Tsarafidy, near Ambohimahasoa. – f , Tsoundzou, Mayotte. – g , Grande Comore. – h , Mikese, Tanzania. – i , Uganda. – j , Kisangani, DRC (Zaire). – k , Ipassa, Gabon. – l , Liberia. – same magnification for a-l. 21 , juxta – morganii : exhibiting on the left side a small lateroventral projection, which is usually elongate (fig. 3 , a and c; fig. 5 , a-c), although occasionally short and triangular (as in a specimen from Grande Comore: fig. 5 , d); praedicta : without well-developed lateroventral projection on the left side (fig. 3 , e and g; fig. 6 , a and b) or, at most, with a short, triangular lateroventral projection (fig. 6 , c and d). 22 , apex of the gnathos (observed ventrally between slide and coverslip) – morganii : distinctly notched mesally (fig. 3 , a and c; fig. 7 , c and d); praedicta : blunt or convex mesally, never notched (fig. 3 , e and g; fig. 7 , a and b). Female genitalia . 23 , posterior pouch of the corpus bursae (delimited ventrally, in lateral view, by the anterior, widened region of the ductus bursae) – morganii : fairly deep (fig. 10 , c-e), its depth (x) being greater than or equal to one fifth of the total length (y) of the corpus bursae; praedicta : apparently less deep (fig. 10 , a and b), its depth (x) being less than one fifth of y. This difference in the shape of the corpus bursae is more difficult to appreciate when the genitalia are observed in ventral view (figs. 8 and 9 ). 24 , signum – morganii : well developed, elliptical and more or less horseshoe-shaped (fig. 11 , d-h); praedicta : smaller (fig. 11 , a and b) to almost absent (fig. 11 , c). 25 , ostium bursae and sterigma – morganii : lateral sclerotized plates of the lamella antevaginalis posteriorly fused with the lateral edges of the ostium bursae (fig. 12 , d-f; also fig. 13 B in KITCHING, 2002 ) so that the plates and the ostium form a single pair of posterior projections in lateral view (fig. 12 , j-l); praedicta : posterior extremities of the lateral plates of the lamella antevaginalis not merged with the posterior extremities of the edges of the ostium bursae (fig. 12 , a-c) so that the plates and the ostium form two pairs of posterior projections in lateral view (fig. 12 , g-i). FIG. 5. — Juxta in four specimens of Xanthopan morganii . – a , Liberia. – b , Ipassa, Gabon. – c , Mikese, Tanzania. – d , Grande Comore. – same magnification as in fig. 6. FIG. 6. — Juxta in four specimens of Xanthopan praedicta stat. nov. – a , Antongil Bay. – b, Amber Mountain (Montagne d’Ambre). – c , Perinet (Analamazaotra). – d , Tsarafidy, near Ambohimahasoa. – same magnification for a-d. FIG. 7. — Uncus and gnathos in Xanthopan praedicta stat. nov. (a, b) and X. morganii (c, d) (ventral or lateroventral views); saccus in X. praedicta (e, f) and X. morganii (g, h) (ventral views). – a , Antongil Bay. – b , Amber Mountain. – c , Tsoundzou, Mayotte. – d , Grande Comore. – e , Perinet (Analamazaotra). – f , Tsarafidy, near Ambohimahasoa. – g , Grande Comore. – h , Liberia. – same magnification for a-h. FIGS 8 and 9. — 8 , Female genitalia of Xanthopan morganii in ventral view (specimen from Gabon). – scale bar = 5 mm. – 9 , Female genitalia of Xanthopan praedicta stat. nov. in ventral view (specimen from Antananarivo). – scale bar = 5 mm. In the above list, at least seven morphological characters provide clear-cut differences between the two taxa (viz. numbers 3, 4, 10, 16, 22, 25, and also 20 if one takes into account only the proximal region of the praesacculus). The differences corresponding to characters 1, 2, 5, 14, 15, and 18 may also be clear-cut ones, although some of them (such as 14) would need to be examined on fresh specimens (rather than on collection specimens, which often tend to be faded). By contrast, numbers 4, 17, 19 and 23 correspond to insufficiently studied quantitative differences, and numbers 6, 7, 8, 9, 11, 12, 13, and 21 provide differences that are observable in most, though not all, cases. FIG. 10. — Corpus bursae in specimens of Xanthopan praedicta stat. nov. (a, b) and X. morganii (c-e) (lateral views). – a , Antananarivo. – b , Ranomafana, west of Ifanadiana. – c , Tsoundzou, Mayotte. – d , La Maboke, Central African Republic. – e , Gabon. – x: depth of posterior pouch; y: total length of corpus bursae. – same magnification for a-e. On account of (1) a deep, ancient genetic divergence (dated to ca. 7.4 Mya by NETZ & RENNER, 2017 ) and (2) the above-mentioned clear-cut morphological differences (e. g. in the ostium bursae), Xanthopan morganii (Walker, 1856) and Xanthopan praedicta Rothschild & Jordan, 1903 stat. nov. should henceforth be regarded as distinct species (no longer as “subspecies”), the latter being endemic to Madagascar while the former inhabits sub-Saharan Africa and the Comoro Islands. FIG. 11. — Signa of specimens of Xanthopan praedicta stat. nov. (a-c) and X. morganii (d-h). – a , Amber Mountain. – b , Antananarivo. – c , Ranomafana, west of Ifanadiana. – d , Tsoundzou, Mayotte. – e , La Maboke, Central African Republic. – f , Gabon. – g , DRC (Zaire). – h , Ivory Coast. – same magnification for a-h. As emphasized in the next paragraph, the proboscis and forewing lengths are usually greater in Xanthopan praedicta than in Xanthopan morganii (e. g. fig. 13 ) but they vary in both species, with overlap zones in the measurements. Proboscis length variability in the two species of Xanthopan Although the first measure of the proboscis length (22.5 cm) of Xanthopan praedicta was given in the original description of this taxon ( ROTHSCHILD & JORDAN, 1903 ), very few studies have been devoted, up to now, to the precise knowledge of this length, with assessment of its variability ( ARDITTI & al. , 2012 : Table 4). Such data are even scarcer in scientific literature in the case of Xanthopan morganii . For the latter, the length of the proboscis would be “seven inches and a half” (19 cm) according to WALLACE (1891: 148) , 12 cm according to MARTINS & JOHNSON (2007 ; Table 3 – one male), and would have an average value of either 13.8 cm ( RYCKEWAERT & al. , 2011 – 6 individuals) or ca. 15 cm ( BALDUCCI & al. , 2019 – mean based on 12 individuals). In fact, after measuring the proboscides of 47 specimens from the MNHN (Additional data, Table 1 ), we find an average length of 14.18 cm for X. morganii , with the range being 8.9-21.8 cm (it should be noted that the mention “over 22 cm” in PINHEY (1962) concerns the taxon praedicta , not the true morganii ; Table 4 of ARDITTI & al. (2012) may be misleading on that point). We managed to measure a higher number of proboscides in the case of X. praedicta (Additional data, Table 2 : 77 individuals, most from MNHN and NHMUK collections, 24 released after measurements (Ranomafana National Park, DCL, March 2020)): we thus have for the proboscis of this Malagasy hawkmoth an average length of 20.8 cm – higher than the means proposed by NILSSON & al. (1985) and RYCKEWAERT & al. (2011) , namely, respectively, 19.6 cm (from 8 individuals) and 18.26 cm (from 5 individuals) – with the range being 15.2-28.5 cm (nevertheless RYCKEWAERT & al. found a shorter proboscis (14.5 cm long) in an individual of the Malagasy Xanthopan ). While the proboscis lengths of X. morganii and X. praedicta overlap partially (fig. 14 ), the commonest lengths are still more remote from one another (12-13 cm; 20 cm) than are their respective means (ca. 14.2 cm; 20.8 cm). Examining the length of the proboscis in terms of moth size (appraised via forewing length) and taking into account the sexes (fig. 15 ), females of X. morganii are routinely larger than the males of this taxon and usually have longer proboscides (16.2 cm on average; range: 12.3-21.8 cm – contra 13.4 cm (range: 8.9-18.3 cm) in the males), with a few values over 20 cm (three in our dataset). A 2 nd order polynomial trendline made with Excel (fig. 17 ) shows that proboscis length is roughly proportional to forewing length in the case of X. morganii . By contrast, in X. praedicta , the longest proboscides were found in males (fig. 16 based on the data in Additional data, Table 2 ), with one value (28.5 cm, from Ranomafana) seemingly representing an absolute record for a lepidopteran proboscis length (28 cm, in certain individuals of Amphimoea walkeri (Boisduval, [1875]) , had been previously regarded as the highest figure among insect haustellum lengths: MILLER, 1997 ). Nevertheless, in X. praedicta too, the average length of the proboscis is higher in females (21.97 cm; range: 18.6-25.4 cm) than in males (20.65 cm; range: 15.2-28.5 cm). For this species, the 2 nd order polynomial trendline ( fig. 18 ) is distinctly curved and becomes more or less horizontal for the larger specimens (females), which suggests that proboscides significantly exceeding 28.5 cm are unlikely to be found in Madagascar. FIG. 12. — Eighth abdominal segment (with ostium bursae) in females of Xanthopan praedicta stat. nov. (a-c, g-i) and X. morganii (d-f, j-l), in ventral (a-f) and lateral (g-l) views. – a and g , Amber Mountain. – b and h , Antananarivo. – c and i , Ranomafana, west of Ifanadiana. – d and j , Tsoundzou, Mayotte. – e and k , DRC (Zaire). – f and l , Ivory Coast. - same magnification for a-l. FIG. 13. — Two males of Xanthopan with unrolled proboscides. – small specimen: X. morganii , Tsoundzou , Mayotte (proboscis length: 9 cm – forewing length: 4.7 cm). – large specimen: X. praedicta stat. nov. , Ranomena, east of Ambositra (proboscis length: 28 cm – forewing length: 7 cm). Both specimens: RCPB. Discussion and conclusion Despite the absence of an absolute difference in proboscis length, the proboscis of Xanthopan praedicta is usually much longer than that of X. morganii , which assuredly reflects adaptations to different sets of visited flowers, especially among long-spurred orchids. Since most sphingophilous flowers are visited at night, their pollinators remain poorly known, notably in tropical regions. To date, only three African orchids have been recorded as sphingophilous flowers visited or pollinated by X. morganii , namely Aerangis kotschyana , Rangaeris amaniensis , and Bonatea steudneri ( MARTINS & JOHNSON, 2007 ; BALDUCCI & al. , 2019 ). None of these three Orchidaceae occurs in Madagascar, an island which nevertheless houses numerous species of long-spurred orchids ( BOSSER & LECOUFLE, 2011 ; NETZ & RENNER, 2017 ). The predicted pollination of Darwin’s Comet Orchid ( Angraecum sesquipedale , endemic to Madagascar) by X. praedicta was eventually confirmed by a viscidium found on a moth in 1992 and photographs of it visiting flowers of the orchid (see ARDITTI & al ., 2012: 419 ), also thanks to behavioural observations in captivity ( WASSERTHAL, 1993 , 1996 , 1997 ) and, most recently, in the wild, with a film (made by Philip DEVRIES in 2004) that can still be watched on the Web (https://www.youtube.com/watch?v=OMVN1EWxfAU). However X. praedicta may possibly not be the sole pollinator of A. sesquipedale , even though this orchid has an exceptionally long nectar tube (the “spur” ranging from 27 to 43 cm according to WASSERTHAL, 1997: 344 ). Indeed Agrius convolvuli (Linnaeus, 1758) was observed visiting flowers of A. sesquipedale in a large flight tent, although without removing the pollinaria of these flowers ( WASSERTHAL, 1997 ). It should be noted that, with an average length of 11 cm (range: ca. 8.5-15.5 cm – RYCKEWAERT & al. , 2011 ), the proboscides of Malagasy specimens of Agrius convolvuli are markedly longer than those of conspecifics from Europe or mainland Africa ( NILSSON & al. , 1985: 8 ). Moreover, although – quite surprisingly – reported only recently (e. g. NILSSON & al ., 1985 ), Madagascar possesses another Sphinginae equipped with a very long proboscis, namely Coelonia solani (Boisduval, 1833) , which has an average tongue length of 17.6 cm (range: 13-24 cm according to RYCKEWAERT & al. , 2011 ). Although C. solani is the main pollinator of Angraecum sororium ( WASSERTHAL, 1997 ) , the sister species of A. sesquipedale ( NETZ & RENNER, 2017 ) , we cannot completely rule out a possible role of this hawkmoth in the pollination of Angraecum sesquipedale (despite the fact that it did not visit two flowers of this orchid in a greenhouse in Germany: ibid ., p. 353). On the other hand, X. praedicta is known to visit flowers of different species, some in Orchidaceae ( Angraecum sesquipedale and Angraecum compactum ; perhaps also Aerangis fuscata : NILSSON & al. , 1987 ), some in other families: Malvaceae (the baobab Adansonia perrieri : BAUM, 1995 ), Lamiaceae ( Clerodendrum : WASSERTHAL, 1997 ), Verbenaceae ( Lantana ), Amaryllidaceae ( Crinum ), and probably several others. In fact, there are complex interactions between orchids and their pollinators ( HOULIHAN & al. , 2019 ), so that we can rule out KRITSKY’ s (1991) hypothesis that there would be in Madagascar “a gigantic moth with a proboscis even longer” than that of X. praedicta in view of the existence of Angraecum longicalcar , an orchid with a nectary “nearly 40 cm long, 10 cm longer than that of A. sesquipedale ”. It should be noted that, actually, the “spur” of A. sesquipedale ranges from 27 to 43 cm and contains a varying volume of nectar, a parameter that cannot be neglected in studies about pollination ( WASSERTHAL, 1997 : Fig. 4 ). According to NETZ & RENNER (2017) , the shape of the labellum should also be taken into account: it may form a landing platform for large hawkmoths, as in A. sesquipedale , or not, as in A. longicalcar (their Fig. 1B shows Xanthopan praedicta “trying to visit” flowers of this rare orchid). It should be noted that botanists from Royal Botanic Gardens, Kew, using camera traps, failed to observe any pollinator, not even X. praedicta , at the sole known wild population of A. longicalcar (https://www.theguardian.com/world/ 2017/may/13/madagascar-mass-extinction-plants-kewgardens). FIG. 14. — Histogram showing proboscis length variability in Xanthopan morganii and X. praedicta stat. nov. (orange and blue bars respectively). FIG. 15. — Proboscis length values in relation to size (estimated using forewing length), in males and females of Xanthopan morganii . FIG. 16. — Proboscis length values in relation to size (estimated using forewing length), in males and females of Xanthopan praedicta stat. nov. FIG. 17. — Xanthopan morganii : trend of increasing proboscis length with moth size (2 nd order polynomial trendline made with Excel). – x-axis: forewing length (cm); y-axis: proboscis length (cm). – data points correspond to average values; the standard deviation for a given point is represented by a vertical line. FIG. 18. — Xanthopan praedicta : trend of increasing proboscis length with moth size (2 nd order polynomial trendline made with Excel). – x-axis: forewing length (cm); y-axis: proboscis length (cm). – see fig. 17 for remarks about data points and vertical lines.

Sex	Forewing	Proboscis	Locality
M	6.8	21.4	Les Roussettes, Amber Mountain
M	6.85	23.35	Les Roussettes, Amber Mountain
M	6.8	19.0	Isaka forest, north of Tôlañaro
M	5.75	15.2	Les Roussettes, Amber Mountain
M	6.8	19.2	La Mandraka (near Manjakandriana)
M	6.3	16.95	Lokobe, Nosy Be
M	6.5	18.4	Les Roussettes, Amber Mountain
M	6.35	18.3	Perinet (Analamazaotra)
M	6.4	21.35	La Mandraka (near Manjakandriana)
M	6.55	18.2	Hiaraka,Antongil Bay
F	7.7	21.5	Antananarivo
M	6.6	19.7	Les Roussettes, Amber Mountain
M	6.6	19.4	La Mandraka (near Manjakandriana)
M	7.0	23.7	road Ambositra-Ambohimanga Atm.
F	7.6	22.6	La Mandraka (near Manjakandriana)
M	7.0	23.0	near Anosibe An’Ala
M	6.4	21.0	Perinet (Analamazaotra)
M	6.9	20.3	north of Lakato, near Moramanga
M	7.1	23.4	La Mandraka (near Manjakandriana)
F	7.0	20.3	Lokobe, Nosy Be
M	6.8	23.5	La Mandraka (near Manjakandriana)
M	6.5	16.9	Ankafina forest, near Tsarafidy
M	6.0	16.85	Tampolo, Masoala peninsula
M	7.1	20.2	Ambohitantely (near Ankazobe)
M	6.7	20.5	Moramanga
M	6.5	21.2	near Anosibe An’Ala
M	6.7	19.3	Moramanga
M	6.6	16.7	north of Lakato, near Moramanga
M	6.8	24.7	north of Lakato, near Moramanga
M	6.4	20.7	Ikoly, Itremo massif
F	7.5	24.4	Perinet (Analamazaotra)
M	6.9	21.3	Andrangoloaka, Eof Lake Mantasoa
M	7.0	22.5	Ambohitantely (near Ankazobe)
M	6.1	15.6	Ambohitantely (near Ankazobe)
M	6.45	18.5	NW of Manantenina
M	7.0	28.0	Ranomena, Eof Ambositra
M	6.4	20.7	north of Lakato, near Moramanga
M	6.6	20.4	north of Lakato, near Moramanga
M	6.7	19.8	north of Lakato, near Moramanga
M	6.7	18.0	north of Lakato, near Moramanga
M	6.8	20.7	Anjozorobe (NE of Antananarivo)
M	7.4	20.7	near Anosibe An’Ala
F	7.3	18.6	north of Lakato, near Moramanga
M	7.0	20.1	Ranomafana, Wof Ifanadiana
F	8.0	19.8	Antananarivo
M	6.9	19.2	Ranomafana, Wof Ifanadiana
F	7.4	21.0	Amber Mountain
M	6.6	18.3	Ranomafana, Wof Ifanadiana
M	7.1	21.5	Ranomafana, Wof Ifanadiana
M	7.1	20.8	Ranomafana, Wof Ifanadiana
M	7.2	23.1	Ranomafana, Wof Ifanadiana
M	6.8	19.3	Ranomafana, Wof Ifanadiana
M	6.7	21.3	Ranomafana, Wof Ifanadiana
M	6.5	18.8	Ranomafana, Wof Ifanadiana
F	7.8	25.4	Ranomafana, Wof Ifanadiana
M	7.1	21.0	Ranomafana, Wof Ifanadiana
M	6.7	20.1	Ranomafana, Wof Ifanadiana
M	6.8	20.2	Ranomafana, Wof Ifanadiana
M	6.3	22.1	Ranomafana, Wof Ifanadiana
M	7.1	21.6	Ranomafana, Wof Ifanadiana
M	6.7	20.6	Ranomafana, Wof Ifanadiana
M	6.3	18.6	Ranomafana, Wof Ifanadiana
M	6.7	18.7	Ranomafana, Wof Ifanadiana
M	6.6	19.3	Ranomafana, Wof Ifanadiana
M	7.1	20.7	Ranomafana, Wof Ifanadiana
M	7.0	25.0	Ranomafana, Wof Ifanadiana
M	6.8	19.2	Ranomafana, Wof Ifanadiana
M	6.9	18.5	Ranomafana, Wof Ifanadiana
M	7.2	27.3	Ranomafana, Wof Ifanadiana
F	8.0	24.1	Ranomafana, Wof Ifanadiana
M	6.7	20.8	Ranomafana, Wof Ifanadiana
M	6.8	26.2	Ranomafana, Wof Ifanadiana
M	6.3	22.3	Ranomafana, Wof Ifanadiana
M	6.9	28.5	Ranomafana, Wof Ifanadiana
M	6.8	26.5	Ranomafana, Wof Ifanadiana
M	6.7	18.6	Ranomafana, Wof Ifanadiana
M	6.1	17.5	Ambohitantely (near Ankazobe)

The taxonomic change we propose here not only for the first time gives appropriate weight to clear-cut morphological differences (including in female genitalia) and high mitochondrial divergence. It also gives long deserved recognition, at the species level, to one of the most celebrated of all Malagasy endemics.