Saalmulleria Mabille, 1891

Lehmann, Ingo & Dalsgaard, Thure, 2023, Revision of Saalmulleria Mabille, 1891 (Lepidoptera, Metarbelidae) from Madagascar with the description of three new genera and fifteen new species, Evolutionary Systematics 7 (1), pp. 133-182 : 133

Saalmulleria Mabille, 1891

Type species of genus (cf. Lehmann 2019b).

Saalmulleria stumpffi ( Saalmüller, 1884).

Incorrect subsequent spelling (unavailable name, cf. ICZN 1999, Articles 33.3, 33.5): " Saalmülleria " in Gaede (1929) and in Viette (1974).

Note to the genus name.

Mabille (1891) wrote " Saalmulleria ", although “Saalmuelleria” should have been written, since diacritic marks cannot be put on scientific names (ICZN 1999, Article 27). Nevertheless, the original spelling mistake by Mabille has to be kept as "correct original spelling" (ICZN 1999, Article 32; also Fletcher and Nye 1995, p. 144).

Gaede (1929) transferred the genus " Saalmülleria " from the family Cossidae to the Metarbelidae . This was found to be correct by Lehmann (2019b) who included Saalmulleria in the Metarbelidae based on morphological characters and designated Saalmulleria stumpffi as type species of the genus.

Autapomorphies diagnosis.

The genus is defined by the following combination of characters: very large and pear-shaped lobes of papillae anales are in horizontal position; the size of one lobe is at least 45% the size of the papillae anales and both lobes have very long setae mainly ventrally and a narrow, long, oblique graben-like structure without setae in its center.

Differential diagnosis shared with Eberhardfischeria gen. nov. from Madagascar.

The basal point of the well visible fork of R 1+R 2 has the same distance to the anterior angle of median cell as the basal point of the fork of R 3+R 4 (in contrast cf. species of Morondavania gen. nov.). Both basal points are at ca. 30-50% of the length of R 3.

Diagnostic characters in females of Saalmulleria .

The head is unusually small if compared to the large size of females, e.g. in the holotype of S. stumpffi only 3.0 mm in diameter, and is among the smallest heads in Metarbelidae.

Labial palpi are three-segmented, all segments of almost equal length, basal segment broadest, also if compared to species of the other genera on Madagascar, namely ca. 1.5 × broader than other segments, rectangular in shape (cf. labial palpi of species of Morondavania gen. nov. and Eberhardfischeria gen. nov.).

The lower part of the fronto-clypeus of the head has no plate-like structure.

The discocellular cell of the hindwing has a shape like a fish-tail with the upper and lower tip in opposite position, and with the upper tip strongly pointed and bent; in contrast, cf. Morondavania gen. nov. and Eberhardfischeria gen. nov.

The hindwing has usually three anal veins present including a strongly sclerotized, broad 1A+2A (a very rare character in Metarbelidae).

The forewings and hindwings are not largely transparent (largely means herein that at least 50% of the forewing and at least 50% of hindwing are transparent).

A note to Gen. Nov. Eberhardfischeria dubiefi (Viette, 1974).

According to the autapomorphies and diagnostic characters of Saalmulleria presented above, the species " Saalmülleria Saalmulleria dubiefi " (Viette, 1974; published with an incorrect subsequent spelling of the genus name) is excluded here from the genus Saalmulleria . There are several reasons for this step that are based on the detailed picture of the male holotype as well as a drawing of its venation done by Professor Dr. Joël Minet and presented to I.L. at the MNHN (Paris) in 2009: first, the only male of Saalmulleria dubiefi (collected in Ambanja, 02nd August 1970 by M. P. Dubief, located at an altitude of 12-50 m in the Sambirano Region and "Madagascar Subhumid Forests" ecoregion) has triangular-shaped wings, largely transparent forewings and largely transparent hindwings as well as a termen that is bent inwards on the hindwings. This is a unique combination of characters among Metarbelidae . Largely transparent wings are extremely rare among Metarbelidae and only known from two species in the monotypic genera Janegoodallia Lehmann, 2014 and Dukearbela Mey, 2018, both occurring on the African mainland. Second, the wing venation of Dukearbela dubiefi is similar to the species of Morondavania gen. nov., comprising one possible synapomorphy in the forewing venation with the latter genus (cf. diagnostic characters of Morondavania gen. nov.). Third, based on the comprehensive morphological study of the Metarbelidae by I.L. it is unlikely that M. mineti sp. nov. and Morondavania dubiefi are congeneric. However, as long as the male genitalia of Morondavania dubiefi has not been studied (it is not allowed to dissect the holotype due to the regulations in the MNHN, J. Minet pers. comm. to I.L. in 2009) it should be treated as a species that belongs to an undescribed genus of Metarbelidae . Due to the latter fact, Morondavania dubiefi is not presented herein with any detailed description.

Description of Saalmulleria .

Female Head (Figs 4e-g , 5c , 7b , 14a, b , 15 , 16C, E, F ): Unusually small; rough-scaled; long hair-like scales of light brownish-olive mixed with deep olive-buff and sepia scales on fronto-clypeus, some with a light lilac and golden glint, or without glint; eyes olive or brownish-olive with small black patches or sepia or dark olive spots; a pair of pits is present, rudimentary or entirely absent (cf. S. stumpffi ) on lower fronto-clypeus, a pair of conical projections always absent, pits behind labial palpi absent; lower fronto-clypeus is broad, as broad as half of eye-diameter (viewed anteriorly) and smooth, without any structures; labial palpi dark olive-buff, very short, less than half of eye- diameter, narrow, consisting of three segments, all segments are of almost equal length, 1st (basal) segment is very broad, at least 1.5 × broader than central segment, rectangular. Antennae bipectinate, narrow and long branches up to 4.0 × longer than width of shaft, branches are widely separated at base with up to 2.0 × width of branch, dorsal and lateral sides of branches not scaled, but with many setae in pairs ventrally and laterally, dorsal and lateral sides of flagellum scaled deep olive-buff mixed with brownish-olive.

Thorax: Densely covered with hair-like scales and broader scales of olive-brown with dark olive tips on patagia and on tegulae, some scales on tegulae with a light lilac glint; scale crest on metathorax is pronounced with long olive-brown hair-like scales with a slightly broader tip of dark olive. Fore and mid legs olive-brown, dorsally sepia, with long dense hair-like structures and a light lilac glint. Epiphyses present, up to 2.5 mm long, medium broad and flat. Hind legs with two pairs of tibial spurs, upper pair more narrow and longer, up to 2.1 mm long, spurs in lower pair broader, up to 1.4 mm long. Wingspan is the longest among Metarbelidae in a worldwide context (cf. S. stumpffi ), namely between 55.0 mm up to 70.5 mm. Forewing large, broad with a rounded apex, upperside without any geometric design, deep olive-buff or greyish-olive and towards termen with a light golden glint, a simple scale pattern is present, usually with a lunule-like sepia patch (sometimes absent or reduced) at center of forewing from base of M2 to base of CuA1, the patch is edged inwards by a small transparent spot (cf. S. stumpffi ), sometimes this spot is absent, various dark olive terminal, sub-terminal and post-medial patches and bands, sometimes broadly V-shaped or slightly Y-shaped, from near costa to CuA2, termen without lunules or with weak dark olive lunules, a dark chestnut or sepia patch is present below base of 1A+2A, up to 30% length of 1A+2A. Hindwing rounded with a pointed apex, termen not bent inwards, largely with short scales of deep olive-buff or greyish-olive, with a light golden glint, with a sepia patch (sometimes absent or reduced) at center of hindwing from base of M2 to base of CuA1, but the patch is not edged inwards by a small transparent spot as in forewing. Underside with scales of deep olive-buff with a light golden glint. Cilia short with up to 1.1 mm length, deep olive-buff with a glint. Forewing venation (Fig. 5c ) with strongly sclerotized and broad veins, 1A+2A deeply forked at base, fork is up to 25% the length of 1A+2A; CuP absent, sometimes represented by a continuous weak fold that is not sclerotized, or CuP is slightly sclerotized on first 2/3 of its length and disappears on last third completely; CuA2 originating from near hind margin of posterior cell; CuA1, M3 and M2 separated and initiating from apical angle of posterior cell; M1 originating from distal margin of median cell and is broadly separated from its anterior angle; areole absent; R1+R2 originating from a long stalk (the stalk has the length of ca. 30-40% of R3) and initiating from near anterior angle of median cell; R3+R4+R5 are long stalked and originating from anterior angle of median cell, the basal point of this stalk is exactly opposite of the basal point of the stalk of R1+R2; Sc more or less parallel to R1. Hindwing venation with 3A present, 1A+2A present or represented by a sclerotized fold, with a small fork at base, CuP present; CuA2 originating from near hind margin of posterior cell; CuA1, M3 and M2 originating from apical angle of posterior cell, separated, sometimes M2 has a tiny fork at base (cf. S. stumpffi ); M1 and Rs originating from anterior cell, broadly separated, with M1 at center of distal margin of anterior cell (cf. different position of basal point of M1 in hindwing of Morondavania gen. nov.); a short bar from Rs to Sc+R1 is absent or weak (Fig. 5c ), a strongly sclerotized vein in discocellular cell on both fore- and hindwing is present and sometimes long forked distally in forewing (a very rare character in Metarbelidae , cf. S. stumpffi ). The discocellular cell on forewing and hindwing is small, only up to 20% of wing size. The discocellular cell on the hindwing is similar in shape like a fish-tail, with the upper and lower tip in opposite position, and the upper tip is strongly pointed. Fringe scales short if compared to the large wing size, up to 2.0 mm, deep olive-buff with a glint. Retinaculum and frenulum absent.

Abdomen: Very long, up to 32.0 mm ( S. stumpffi ) with hair-like scales of deep olive-buff or greyish-olive with a strong light golden glint, mixed with dark olive, upper part of abdomen sepia or dark chestnut, end of abdomen sometimes sepia, abdominal tuft short, up to 5.0 mm long, or ca. 15% of abdomen length, sepia or dark chestnut.

Female postabdominal structure and genitalia with very large, pear-shaped lobes of papillae anales in horizontal position, one lobe at least 45% the size of papillae anales (cf. species of Eberhardfischeria gen. nov. with much smaller lobes), lobes ventrally with long setae and few long setae along the edge, each lobe with a long oblique graben-like structure that has no setae at center; papillae anales covered with many short and many long setae. Segment 8 represents a medium broad rectangular sclerotized band, more narrow ventrally, setose along its whole posterior margin with long setae, or without any setae on dorsal part of posterior margin, and without any setae on segment 8, with a very narrow band attached ventrally extending to the base of anterior apophysis, where the band is weakly attached and has a very broad rectangular-shaped base that fits like a segment into segment 8 (Fig. 7b , only visible in fresh preparations); anterior apophysis slightly or strongly bent downwards, up to 3.0 × as long as segment 8 dorsally, on their basal half of length up to 2 × as broad as at tip, within the first 30% of their length strongly knee-like shaped (cf. Eberhardfischeria with anterior apophysis knee-like shaped at middle), on almost the whole length with a deep horizontal graben-like structure; posterior apophysis narrow with four times broader base, up to 60% the length of anterior apophysis, with large sclerotized base up to 40% the size of papillae anales in lateral view, posterior apophysis longer or equal in length of dorsal part of segment 8; ductus bursae and corpus bursae are both membranous (cf. S. ampandrandavaensis sp. nov.), or ductus bursae is thickly membranous (cf. S. analameranaensis sp. nov.), ductus bursae broad, short (up to 50% the length of corpus bursae) with or without a small thickly membranous, pear-shaped structure at base, the base is not broader and not sclerotized, corpus bursae very large, up to 4 × as large as segment 8 in lateral view, membranous, not sclerotized at any part, without any structures, elongated, oval (cf. S. ampandrandavaensis sp. nov.).

Male: unknown.

Species richness.

Currently, Saalmulleria comprises four species, including three species new to science of which two new species are described here. One species that is represented by a female (deposited in MNHN) was not studied and is treated here as undescribed - it is labelled as follows: "Madagascar Est [East] 42 km N. de Sambava [North of Sambava / Eastern Region] forêt d’ Analabe 50 m [Analabe forest altitude 50 m / "Madagascar Humid Forests" ecoregion], 15/20-XI-1958 P. Griveaud, A. Peyrieras et P. Viette" [leg.].

Distribution.

Species of Saalmulleria occur in the "West Malagasy regional centre of endemism" as well as "East Malagasy regional centre of endemism" sensu White (1983) and might be still present on the whole island of Madagascar if natural woody habitats are still available (cf. distribution map in Lehmann 2019b, fig. 47). In northern Madagascar, the distribution range covers the "Sambirano Region", "Western Region" as well as "Eastern Region" sensu Humbert (1955, 1965), extending from the Réserve Naturelle de Lokobe off the northwestern coast of Madagascar towards the Réserve Naturelle de la Analamerana in the Northeast of the main island and from there southeastwards to north of Sambava close to the eastern coast with the Indian Ocean (cf. note above). Within this area, species of Saalmulleria are known from different lowland forest types within the "Madagascar Dry Deciduous Forests" ecoregion, the "Madagascar Subhumid Forests" ecoregion and the "Madagascar Humid Forests" ecoregion. The altitude range is 5-419 m and comprises forests with an average annual rainfall of 1500 mm to more than 2000 mm (cf. Eberhardfischeria gen. nov. with a similar habitat range). In southern Madagascar, species of Saalmulleria occur in the submontane and subarid area of Ampandrandava in the "Madagascar Succulent Woodlands" ecoregion sensu Crowley (2004) and close to the southwestern limit of the central highlands, "Central Region", with an average annual rainfall of only 575-1330 mm.

Biological traits.

The biology of species of Saalmulleria is unknown at present. However, lowland tropical Metarbelidae species are strongly associated to habitats with a dominance of woody legumes ( Lehmann 2008, 2019b). Leguminosae on Madagascar are in general diverse in the northern, western and southern parts, with particular high densities in areas with a marked dry season and deciduous vegetation ( Labat and Moat 2003). Two species of Saalmulleria presented here from northern and southern Madagascar are linked to the latter areas.

Key to the species of Saalmulleria

The key is based primarily on characters of the genitalia; hence, it cannot serve as a field identification key. For all species, only one female specimen is available, so identifications obtained from this key should be cross-checked carefully with the description, distribution, and figures presented in this paper.

1	Female	2
2(1)	Lobes of papillae anales in horizontal position, very large, one lobe is at least 45% the size of papillae anales	3
3(2)	Lobes of papillae anales not longer than segment 8 dorsally	4
-	Lobes of papillae anales much longer than segment 8 dorsally	5
4(3)	Anterior apophyses strongly bent downwards, 2.0 × longer than dorsal width of segment 8, large extension of ductus bursae near its base	S. analameranaensis sp. nov.
5(3)	Lobes of papillae anales largest in Metarbelidae , one lobe is 50% the size of papillae anales, anterior apophyses 3.0 × longer than dorsal width of segment 8, posterior apophyses long, 50% length of anterior apophyses	S. stumpffi
-	Not as above, one lobe is 45% the size of papillae anales, posterior apophyses short with 30% length of anterior apophyses	S. ampandrandavaensis sp. nov.

Figures

Figure 4. a. Morondavania mineti sp. nov., holotype, male, Madagascar, Western Region, north of Morondava, Marofandilia Forest; b. M. mineti sp. nov., paratype, male, Madagascar, Western Region, north of Morondava, western part of Marofandilia Forest; c. Eberhardfischeria husemanni sp. nov., holotype, female, Madagascar, Western Region, Diego Suarez (today Antsira ṅana) or areas nearby; d. E. husemanni sp. nov., paratype, female, Madagascar, Western Region, Diego Suarez (today Antsira ṅana) or areas nearby; e. Saalmulleria stumpffi (Saalmueller, 1884), " Type ", female, Madagascar, Sambirano Region, Nosy Be Island, Lokobe; f. S. analameranaensis sp. nov., holotype, female, Madagascar, Western Region, Analamerana Forest, ca. 10 km to 40 km west of the Indian Ocean coastline; g. S. ampandrandavaensis sp. nov., holotype, female, Madagascar, Central Region, Ampandrandava, ca. 50 km northeast of Bekily.

Figure 5. Wing venation: a. Eberhardfischeria husemanni sp. nov., paratype, female; b. Morondavania mineti sp. nov., holotype, male; c. Saalmulleria stumpffi (Saalmueller, 1884), " Type ", female.

Figure 7. Female postabdominal structures and male genitalia in a not pressed condition: a. Eberhardfischeria husemanni sp. nov., paratype, female; b. Saalmulleria stumpffi (Saalmueller, 1884), " Type ", female; c. Shimbania kerstinhempae sp. nov., holotype, female; d. Morondavania mineti sp. nov., paratype, male, with aedeagus below; e. S. wichgrafi (Gruenberg, 1910), " Type ", male, with aedeagus below (all drawings by I. L.).

Figure 14. Female postabdominal structures of a. Saalmulleria stumpffi (Saalmueller, 1884), " Type ", female, Madagascar, Sambirano Region, Nosy Be Island, Lokobe; b. S. analameranaensis sp. nov., holotype, female, Madagascar, Western Region, Analamerana Forest, ca. 10 km to 40 km west of the Indian Ocean coastline.

Figure 15. Genitalia of Saalmulleria ampandrandavaensis sp. nov., holotype, female, Madagascar, Central Region, Ampandrandava, ca. 50 km northeast of Bekily, with a thinly membranous and long ductus bursae (a) and a thinly membranous elongated oval-shaped corpus bursae (b), both without any structures. The two lobes of the papillae anales (c) are in horizontal position, an autapomorphy of species of Saalmulleria, and represent the largest lobes among Metarbelidae, with one lobe at least 45 % the size of the papillae anales.

Figure 16. SEM pictures of diagnostic head structures of Metarbelidae from Madagascar: A. The lower fronto-clypeus of Morondavania mineti sp. nov., holotype, male, is narrow and without a pair of pits as well as without a pair of projections; B, D. The lower fronto-clypeus of Eberhardfischeria husemanni sp. nov., paratype, female, is broad with a strongly sclerotized plate-like structure (a) with a well-defined dorsal ridge (b) and a pair of rudimentary pits (c); C, E, F. The very broad and smooth lower fronto-clypeus of Saalmulleria ampandrandavaensis sp. nov., holotype, female, is without any plate-like structure, but with a pair of well-developed pits (d), a variable character in species of Saalmulleria that is sometimes entirely absent. A pair of pits on the lower fronto-clypeus is a homoplasy among Metarbelidae in both sexes (cf. Lehmann 2019 b) and it is in species of various genera a variable character (SEM pictures taken by Karin Ulmen, ZFMK, Bonn 2013).