Typus: Danais fragrans (Lam.) Pers.

= Alleizettea Dubard & Dop in J. Bot. (Morot) 3: 6. 1910. Typus: Alleizettea bracteata Dubard & Dop [= Danais volubilis Baker].

Scrambling, low to high-climbing, suffrutescent to woody climbing plants, climbing as high as 25 m, terrestrial, unarmed, with raphides, glabrous or variously pubescent, regularly branched, apparently not deciduous, sometimes with foetid odor. Leaves opposite, ternate, or 4-verticillate, decussate, subsessile to petiolate, sometimes with pubescent domatia in abaxial axils of secondary veins, with higher-order venation not lineolate, sometimes with higher-order venation laxly to closely reticulated and prominulous; stipules interpetiolar or infrequently fused around the stem (Danais coriacea C.M. Taylor sp. nov., D. longipedunculata Homolle), generally persistent or fragmenting with age, with interpetiolar portion triangular to broadly so, truncate, elliptic, or ovate, in its upper part variously acute to obtuse, rounded, subtruncate, serrate, bilobed, bidenticulate, laciniate, or multifid, erect or infrequently reflexed. Inflorescences terminal and/or axillary, 1- to many-flowered, fasciculate, cymose, thyrsiform, or racemiform, subsessile to pedunculate, bracts reduced to developed. Flowers subsessile to pedicellate, distylous, diurnal or perhaps sometimes nocturnal; calyx limb shortly to deeply (4–)5(–6)-lobed, lobes equal to unequal on an individual flower, without calycophylls; corolla salverform to funnelform, white to pale green, olive green, lavender, dull to dark purple, purple-black, yellow, orange, or dull to bright red with colors often differing on tube and lobes, externally generally glabrous, tube 2–22 mm long, usually slender with the distalmost portion swollen to accommodate anthers in long-styled form, internally usually puberulous to pilosulous, in throat usually pilose with pubescence often denser in long-styled form, sometimes at base of tube fenestrate with short preformed slits, these alternate to corolla lobes, 1–2 mm long, and with opening blocked internally by interlocking trichomes, lobes (4–)5(–6), valvate-reduplicate in bud, lanceolate to triangular or ligulate, acute to obtuse, at tip without appendages; stamens (4–)5(–6), inserted in upper part of corolla tube, anthers dorsifixed, ellipsoid to narrowly oblong, dehiscent by longitudinal slits, without connective appendages, in long-styled form included, subsessile, and positioned in corolla throat, in short-styled form exserted on well-developed filaments, pollen characterized by Buchner & Puff (1993) as spheroidal to subspheroidal, 10–26 µm, 3- to 5-colporate, and dimorphic, with supratectal verrucae present in grains of short-styled flowers but absent in grains of long-styled flowers; ovary 2-locular, with numerous seeds on peltate axile placentas, with papillate disk surrounding style at top, stigmas 2, linear, exserted in long-styled flowers, included and positioned in upper part of tube in short-styled flowers. Infructescences similar to inflorescences. Fruits capsular, deeply loculicidal from top, subglobose to ellipsoid or oblate, papery to chartaceous, coriaceous, or woody, 2–12 mm in diam., sometimes partially fused together, with calyx limb persistent or tardily deciduous, without beak or with weakly broadly angled beak portion up to 1 mm high; seeds numerous, 0.5–6 mm long, reticulated, flattened and circular to elliptic with circumferential wing, or infrequently angled to cylindrical and unwinged (D. aptera, D. baccata, D. sambiranensis), margin of wing when present entire to irregular or lacerate.

Notes. – At least 49 species distributed widely in humid to wet vegetation in the WIO region and eastern Africa, mainly on Madagascar but three species in the Mascarenes (Mauritius, Réunion, Rodrigues), two species in the Comores (one shared with Madagascar), and one species in East Africa (Tanzania).

Lectotypus (designated here): Madagascar: Domaine oriental, sur la côte, s.d., Chapelier s.n. (P [P00057135] image!; isolecto-: P [P00057136, P00057137] images!).

Distribution and ecology. – This species was collected once on the east coast of Madagascar, and nothing more is known of its ecology.

Notes. – Danais aptera is poorly known. It was described from one incomplete, fruiting collection and its type has not been matched to modern collections. This species is characterized by opposite, elliptic, subsessile leaves, narrowly triangular stipules, axillary flowers that are solitary or few and fascicled on slender peduncles or pedicels c. 2 cm long, narrowly triangular calyx lobes c. 1 mm long, subglobose loculicidal capsules c. 8 mm in diam., and cylindrical seeds that are apparently about 1 mm long (if the seeds on the type are mature).

Homolle (1936) classified this species in Danais apparently because it has the habit, fruit form, and general aspect of Danais. The original label on the lectotype describes the plant as a “liane grimpant”, so it has the diagnostic character of this genus. Buchner & Puff (1993: 73–74) excluded this species from Danais, however, based on its seeds that lack wings. As discussed above, other genera in Danaideae vary extensively in seed form and wing development (Buchner & Puff, 1993; Razafimandimbison et al., 2022), and a seed wing is not here considered diagnostic for Danais. Accordingly, this species is here reinstated in Danais.

The authorship of Danais aptera has been sometimes attributed solely to Homolle, but in the protologue she credited the coining of this name for the Chapelier type collection to “D. aptera Wedd. mss. in Herb. Mus. Par.”, which satisfies the criteria of ICN Art. 46.5 (Turland et al., 2018) for crediting Weddell for this epithet. Homolle provided the morphological description and taxon circumscription and thus remains the actual author of this name. Puff annotated two specimens of Chapelier's at P as the holotype and isotype of D. aptera, respectively, and Buchner & Puff (1993: 73) cited these two specimens as such without further detail to identify the corresponding specimens. Three duplicates of this collection are now curated at P, with the newly discovered one annotated by C.V. Naudin (1815–1899) but not Puff. None of these specimens were annotated or otherwise directly indicated as a holotype by Homolle. Buchner & Puff's incomplete published citation does not fulfill the requirements for an inadvertent lectotypification because it does not conclusively identify their intended holotype, so this name is typified here. The specimen annotated by Puff as the holotype was apparently originally chosen for having the most ample material including seeds, and that specimen has been consulted as such by later authors and is chosen as the lectotype.

Lectotypus (designated here): Madagascar. Reg. Ihorombe [Prov. Fianarantsoa]: alluvions des bords de l'Itomampy, VI.1919, Perrier de la Bâthie 12663 (P [P00057140] image!; isolecto-: MO-04613606!, P [P00057141, P00539961] images!).

Distribution and ecology. – This species is found in lowland moist evergreen forest, sometimes on sandy substrates, from sea level to 600 m along the eastern escarpment of Madagascar (Reg. Anosy [Prov. Toliara], Reg. Analanjirofo [Prov. Toamasina], Reg. Ihorombe [Prov. Fianarantsoa]).

Notes. – Danais baccata is not well known. It was described based on one incomplete collection, and its type has now been matched to two modern collections. This species is characterized by a branching pattern with the leaves and flowers borne on short lateral stems arising along an elongated, weakly flexuous main stem; opposite, elliptic, shortly petiolate leaves; triangular, deltate to sometimes acuminate, small stipules; axillary flowers that are solitary or few and fascicled on slender peduncles or pedicels 5–15 mm long; narrowly triangular calyx lobes c. 1 mm long; a slenderly salverform, brown-flushed corolla with the tube c. 13 mm long and fenestrate at the base; subglobose, loculicidal capsules 8–12 mm in diam.; and angled to ellipsoid, unwinged seeds about 1 mm long. The two modern specimens of D. baccata are from widely separated localities, but the distinctive branching pattern and wingless seeds indicate that these belong to the same species or represent closely related species that cannot currently be distinguished. The habit of this species appears to be variable, or at least difficult to characterize.

The branching arrangement of Danais baccata is uncommon in this genus, with the leaves and fruits borne on short lateral stems from elongated main stems. The type collection has regular, rather stiff branching with some of the long stems weakly curving, while the two modern specimens (Ludovic 1614, Manjato et al. 264, MO) have a similar growth form but markedly flexuous main stems. The labels of the type and one of the modern collections describe the plant as a shrub, while the label data of the other modern collection describe it as a liana. As Puff & Buchner (1994) noted, the habit of Danais is basically climbing but sometimes described by collectors as shrubby when the young plants develop a clumped form, or when the plants spread and clamber over a substrate such as rock. Thus, the conflicting label descriptions do not seem to confirm this species has an erect rather than supported habit.

Danais baccata is similar to D. aptera in its unwinged seeds, and it has a similar classification history. Homolle described this species as “Danais?”, apparently because it has the general aspect and fruit form of Danais but a supposed shrub habit and wingless seeds. Buchner & Puff (1993: 74) excluded this species from Danais based on these last two features. The seeds of D. baccata are ellipsoid, angled, and not clearly winged, although many of them have a weakly flattened margin that is often to flange-like along one side. As discussed above, winged seeds are not a diagnostic feature for Danais, and the habit of this species is at least sometimes climbing so it is here reinstated in Danais.

Buchner & Puff (1993: 74) cited the holotype specimen of Danais baccata as the single specimen they saw at P, without details to identify the individual sheet, and Puff annotated one specimen of the type collection there as the holotype [P00057140]. However, neither the protologue nor annotations by Homolle denote a single specimen that can be considered the holotype. Three duplicates of the type collection are deposited at P, and so must be considered syntypes. Two of these were annotated with this name and “n. sp.” by Homolle [P00057140, P00057141], and the third lacks any annotation by her [P00539961]. P00057140 has more and better plant material, a label with original collection data, an analysis in Homolle's hand, and Puff's annotation as holotype, and is therefore designated here as the lectotype. These last two specimens belonging to the same gathering are here considered isolectotypes.

Holotypus: Madagascar: “North Madagascar”, s.d., Baron 6607 (P [P00057143] image!; iso-: K [K000046178] image!).

Distribution and ecology. – This species is known from one collection from northern Madagascar, which lacks collection data.

Notes. – Danais baronii was incompletely treated by Puff & Buchner (1994: 62–63), and it is still incompletely known and problematic as to its identity. It was described based on one collection with immature fruits from an unspecified locality somewhere in northern Madagascar. Danais baronii is characterized by puberulous vegetative organs; opposite, petiolate leaves with the higher-order venation closely, densely reticulated and prominulous on both surfaces of the blade; interpetiolar, triangular stipules; cymose, pedunculate inflorescences borne in both axillary and terminal position on peduncles 0.5–2 cm long; flowers and fruits borne in regularly umbelliform groups on pedicels 1–2 mm long; and ellipsoid to subglobose fruits 4–5 mm in diam. The leaf blades in the type images are shiny on both surfaces and have dried with a coriaceous texture. The type of D. baronii has not yet been matched to any other collections, so is characterized here based on its few known features. The identity of this species is not only problematic in general, it is problematic for the taxonomy of Danais: some other species share its few known features but cannot be conclusively separated or synonymized with it because D. baronii is incompletely known.These similar species are separated and treated here because they can be characterized and analyzed, but one of these names may eventually be found to apply to the plants described as D. baronii.

Danais baronii is similar to D. rubra, which is also known from northern Madagascar. Danais rubra is also incompletely known, from only one flowering collection. Danais rubra differs from D. baronii in its longer petioles (20–25 mm long vs. 5–20 mm long in D. baronii); shorter calyx lobes (0.5–0.8 mm long vs. c. 1.5 mm long in D. baronii); and a different general aspect. Danais schatzii C.M. Taylor sp. nov. (described below) is also similar to D. baronii and is found in a different floristic region in east-central Madagascar. Danais schatzii differs from D. baronii in its glabrous stems and leaves; peduncles 4.5–10.7 cm long; pedicels 0.5–4 mm long and varying in length within a cyme so the flowers are less regularly arranged; and a shorter calyx limb, c. 0.5 mm long.

Holotypus: Madagascar: “forests of the Tanala country, chiefly in Betsileo-Land”, s.d., Baron 189 ex parte (K [K000046142] image!; iso-: P [P00057148 fragm.] image!).

Distribution and ecology. – This species is found widely in lowland and medium altitude moist evergreen forests from sea level to 1800 m, from northern to southern Madagascar (Reg. DIANA and SAVA [Prov. Antsiranana], Reg. Alaotra-Mangoro and Analanjirofo [Prov. Toamasina], Reg. Vatovavy-Fitovinany [Prov. Fianarantsoa], Reg. Anosy and Atsimo-Andrefana [Prov. Toliara]).

Notes. – Danais cernua is circumscribed more narrowly here than by Puff & Buchner (1994) and diagnosed in part by different characteristics. This species is widespread and commonly collected, and characterized here by an extensively branched habit with elongated, flexuous stems; glabrous to puberulous or hirtellous stems and abaxial leaf surfaces; opposite or rarely ternate, petiolate, elliptic to obovate or oblanceolate leaf blades 3–9 × 1.5–4.5 cm that dry with a papyraceous to chartaceous texture; interpetiolar stipules that are triangular in outline and bilobed; axillary cymose inflorescences about half as long as the leaves, with the flowers all pedicellate; a calyx limb divided nearly to the base into narrowly triangular to linear lobes 0.8–3 mm long that are often unequal in length on an individual flower; slenderly funnelform corollas with the lobes 2–4 mm long and generally orange to red; and subglobose fruits 4–9 mm in diameter. The calyx lobes characteristically become reflexed with age, which is unusual in Danais and distinctive for D. cernua when present. The corolla tubes are 5–9 mm long, and they apparently lack slits at the base but occasionally have corresponding lines of thinned tissue here. The corolla tubes expand regularly and notably (for Danais) in diameter from the base to the top and are wide enough at the top to accommodate the included anthers of the long-styled form without an abrupt swelling. Danais cernua is frequently collected in secondary vegetation.

Puff & Buchner (1994: 57–59) circumscribed Danais cernua more broadly than done here, to include wider variation in both vegetative and flowering characters, and they effectively broadened its circumscription even more in their specimen annotations. In particular, they included in D. cernua plants with leaf blades that dry variously membranaceous to papery, subcoriaceous, or coriaceous; stipules ranging from obtuse to acute and bilobed; and calyx limbs varying from deeply divided to tubular and only shallowly lobed, even though they characterized this structure in their commentary as “typically.... the calyx lacks a basal tubular part” (Puff & Buchner, 1994: 58). Puff & Buchner (1994: 58, as D. fragrans) noted accurately that D. cernua is similar to and easily confused with D. lyallii, but review of specimens identified by them finds no consistent use of their own diagnostic features and no evident morphological differences between their working concepts of these two species. Review here of specimens that correspond to Puff & Buchner's D. cernua and D. lyallii does find two morphologically distinguishable groups of plants, however, which generally correspond to their stated species concepts. One group of plants consistently has a papery or chartaceous leaf drying texture, distinctly bilobed stipules, and a calyx limb divided to the base into narrow, usually spreading to reflexed lobes; this group agrees with the type of D. cernua, and the circumscription of this species here. The other group of plants consistently has leaves with a chartaceous to coriaceous drying texture, stipules that are acute or infrequently denticulate, and a calyx limb divided for 1/3–2/3 of its length into triangular, erect lobes; this second group agrees with the type of D. lyallii, and this second species is diagnosed here by these features. Puff & Buchner also contrasted these two species as to flower size, with D. cernua characterized as having longer corolla tubes, but the measurements they gave for these two species were overlapping: “(4–)5–9 mm” (Puff & Buchner, 1994: 57) for D. cernua, vs. 2–4.5 mm long in D. lyallii (Puff & Buchner, 1994: 54, as D. fragrans). Review here with numerous additional collections finds no significant difference in corolla size between these two species, however: corollas of D. cernua have a tube 4–9 mm long, and those of D. lyallii have a tube 2.5–6 mm long. The corollas of D. cernua are indeed often longer than those of D. lyallii, as noted by them, but definitely not always; the corollas of D. cernua are often also relatively more slender and can appear longer in comparison due to this difference in form. In the taxonomy here, most of the plants Puff & Buchner included in D. fragrans and many of those they included in D. cernua are placed in D. lyallii. Several hundred collections corresponding to these two species were studied in this review, and only one specimen (De Block et al. 2335, MO) appears to be intermediate between the two species, with the tubular, shortly lobed calyx limb of D. lyallii, the bidentate stipules of D. cernua, and a corolla tube c. 5 mm long that agrees with both species. Given the frequency and sympatry in secondary vegetation of these two species and the hundreds of specimens that clearly correspond to one species or the other, this one collection is regarded as having a putative hybrid origin rather than as an intermediate plant that indicates that these species are not distinct.

Puff & Buchner (1994: 57) synonymized several other names under Danais cernua, which all agree with this species in the circumscription here and are cited again only in Tables 1 and 2.

Lectotypus (designated here): Madagascar. Reg. DIANA [Prov. Antsiranana]: Domaine du Centre, massif de Manongarivo, montagne du Sambirano, IV.1909, Perrier de la Bâthie 3823 (P [P00057145] image!; isolecto-: P [P00057146] image!).

Distribution and ecology. – This species is found in medium altitude moist evergreen forest at c. 850 m in northwestern Madagascar (Reg. DIANA [Prov. Antsiranana]).

Notes. – Danais coerulea is characterized by its combination of hirsute pubescence on stems, leaves, stipules, and inflorescences; opposite, petiolate leaves with the blade drying papyraceous; well-developed, ovate to lanceolate stipules 5–15 mm long that are often reflexed in their upper parts; axillary, pedunculate, subcapitate inflorescences; well-developed, linear calyx lobes 4–5.5 mm long; and purple or bluish black corollas with the tube 18–21 mm long.This species was described based on one flowering specimen, and its identity has not been clear until recently when a modern collection was matched to the type (Gautier 2988, G, MO, P).

Danais coerulea was treated as doubtfully placed in this genus by Puff & Buchner (1994: 39–40), and not keyed or given a full description by them so it has been overlooked. They excluded this from Danais based on its label data describing it as a shrub and because they could not confirm it has winged seeds. As discussed above, specimens of Danais can be described as shrubs or “scrambling shrubs” due to a habit confusion with lianas growing in dense vegetation or on rocks or young plants. The fruits of D. coerulea have apparently still not been documented, but this new collection confirms this plant is a climber and it is here reinstated in Danais.

Danais coerulea has been confused in herbarium identifications with D. sambiranensis, described and separated below, and D. capituliformis Homolle. Danais capituliformis can be recognized by its ligulate to narrowly triangular stipules 2–10 mm long that are erect and often prolonged at the tip, narrowly ligulate to triangular calyx lobes 1.5–3.5 mm long, and a corolla tube c. 12 mm long. Danais capituliformis was known to Puff & Buchner (1994: 39–40) from only three collections, and they noted that these three specimens showed notable variation in leaf size, inflorescence branching, and calyx lobe shape so further morphological variation was expected. However, the characters of D. coerulea are quite different and these two species are clearly distinct.

Two duplicates of the type collection of Danais coerulea are curated at P, and both were annotated with this name by Cavaco. There is no indication in the protologue or specimen annotations that one of these was to be considered the holotype, so they must be regarded as syntypes. Puff annotated one of these as the holotype and did not annotate the other. Puff & Buchner (1994: 63) cited the holotype as deposited at P but gave no more identifying information so this cannot be considered as an inadvertent lectotypification. A lectotype is designated here, and the specimen annotated as the holotype by Puff is chosen.

≡ Sabicea verticillata Wernham, Monogr. Sabicea: 74. 1914 [not Danais verticillata Baker, 1883].

Holotypus: Madagascar: “North Madagascar”, s.d., Humblot 213 (P [P00150784] image!).

Etymology. – The species epithet refers to the confused classification history of this species and competing informal names that have been used for it.

Distribution and ecology. – This species is known only from the type, which is from Madagascar but otherwise lacks collection data.

Notes. – Danais confusa is characterized by its densely hirsute stems, calyces, and hypanthia; ternate, subsessile, medium-sized, elliptic to obovate leaves; shallowly to deeply pectinate or laciniate stipules c. 8 mm long; flowers that are axillary at lower stem nodes, solitary, and subsessile or with a peduncle up to 5 mm long; notably well-developed calyx limbs, c. 9 mm long, that are divided nearly to the base into narrowly elliptic lobes; and slenderly salverform, externally hirsute corollas with the tube 17–20 mm long and the narrow lobes c. 9 mm long. This species is only known from one collection, which is in flower, but no color was noted.

This species was described in Sabicea Aubl., and it agrees with several species of that genus on Madagascar (Wernham, 1914) in its laciniate stipules, presumably climbing habit (no label data document this, but no author has questioned it), solitary axillary flowers with a well-developed calyx limb, and general corolla form. This plant is also similar to Danais in these same features, and was studied by Homolle, who provided a hand-written analysis of the type specimen that is mounted on it. She did not finally publish a name for this collection in Danais, similarly to others that she studied and annotated as new species for this genus but never finally described (Cavaco, 1965). Cavaco later regarded this plant as a species of Danais and annotated it as such, but did not formally transfer it taxonomically to this genus. In contrast, Puff & Buchner (1994: 64) regarded this as a species of Sabicea, based on the apparent lack of winged seeds. Some Sabicea species on Madagascar are similar to some species of Danais, but these genera differ in corolla aestivation, fruit type, and seed shape. In Sabicea, the corolla aestivation is valvate, the fruits are baccate and fleshy or juicy, and the seeds are angled and unwinged; in Danais the corollas are reduplicate-valvate, the fruit is capsular, and the seeds are angled to flattened and winged or not. The corolla lobes on the type specimen of S. verticillata are poorly preserved and their aestivation not entirely clear in the online image available, but these appear to be thinly valvate-reduplicate. The fruits are immature but not notably fleshy. Homolle's hand-written analysis on the type noted the details of a dissected ovary and the presence of wings already present on the developing seeds, but neither an ovary that has been dissected nor any seeds seem to be preserved on this specimen today (seen only as an image). Puff & Buchner concluded that the observation of winged seeds was inaccurate or at least not confirmable, and as discussed above, they regarded winged seeds as diagnostic for Danais. However, Homolle's drawing and statement that she found the developing seeds to be winged is considered credible here, and indicates this species is better classified in Danais. Additionally, some other features of S. verticillata are more similar to Danais than Sabicea. Wernham noted in the protologue that verticillate leaves were not otherwise known in Sabicea, and this apparently is still the case, but these are found in Danais. Also, the corollas of the type specimen are anomalous in Sabicea but agree with Danais in their very slender tubes that are abruptly enlarged at the top to accommodate the included anthers, and the short fenestrate slits at their bases are common in Danais but not known in Sabicea. Accordingly, this species is here transferred to Danais.

Puff & Buchner (1994) noted that Cavaco actually had regarded this species anecdotally as Danais twice, separately, once in a print citation and once in a herbarium annotation. He used two different names for it, proposed as replacement names in Danais, and these have been used by others and in curation, but neither is validly published. One of his unpublished names honored Herbert Fuller Wernham (1879–1941) and was cited in as a nomen nudum in a publication, while the other name honored Prof. Anne-Marie Homolle (1905–1988) and was only used in specimen annotations. The replacement name published here is newly coined in part to avoid confusion with those previous invalid names.

Danais confusa is similar to D. laciniata C.M. Taylor, which differs in its opposite leaves, branched inflorescences that are terminal and sometimes also produced in the uppermost leaf axils, and longer pedicels 15–26 mm long. Danais confusa is also similar to D. vestita Baker, which differs in its opposite leaves and pedunculate cymose inflorescences.

Holotypus: Madagascar. Reg. Atsinanana [Prov. Toamasina]: Distr. Brickaville, comm. Maroseranana, Fkt. Ambodilendemy, Ankerana, 18°25′40″S 48°47′17″E, 990 m, 24.III.2011, Antilahimena 7781 (MO-6407815!; iso-: P,TAN).

Danais coriacea C.M. Taylor is distinguished from D. longipedunculata Homolle by its shorter, white to yellow corollas.

Climbing plants, climbing to 16 m high, fleshy; stems glabrous, flattened to weakly quadrangular becoming terete. Leaves opposite; blade elliptic to elliptic-oblong or obovate, 7–14.5 × 3–6 cm, at base cuneate to acute, at apex obtuse to rounded or subtruncate and sometimes with acuminate tip 1–3 mm long, drying chartaceous to coriaceous, adaxially and abaxially glabrous; secondary veins 6–9 pairs, extending to near margins, without domatia, with 1–2 well-developed intersecondary veins present between pairs of secondary veins, with costa and secondary veins thickened to prominulous adaxially and prominent abaxially, with tertiary and quaternary venation laxly reticulated and prominulous adaxially and sometimes abaxially; petiole 1.2–3 cm long, glabrous; stipules shortly united around stem, persistent or caducous, glabrous, with interpetiolar portion triangular and sometimes becoming broadly triangular to subtruncate with age (as stipule is stretched due to stem diameter increase), 3–4 mm long, obtuse to acute or acuminate, abaxially glabrous and sometimes costate, with intrapetiolar portion 0.5–2 mm long and sometimes becoming narrower with age (as tissue is stretched around stem). Inflorescences axillary or sometimes terminal on short paired axillary stems, cymose to paniculiform, pedunculate, densely puberulous or strigillose to glabrescent; peduncle 1–5 cm long; branched portion 4.5–15 × 4–15 cm, branched to 3 orders, with 3–5 pairs of secondary axes, with flowers borne in subcapitate groups of 3–5, these groups borne at apices of axes; bracts deltate to ligulate, 0.5–1 mm long, strigillose to puberulous or glabrous. Flowers subsessile, distylous; hypanthium obconic, c. 1 mm long, densely puberulous to strigillose; calyx limb densely puberulous to strigillose, 1–1.2 mm long, 5-lobed for about half its length, lobes deltate; corolla funnelform, white or sometimes with yellow lobes, externally glabrous, tube 3–3.5 mm long, 1.2–1.8 mm in diam. near middle, without slits at base, internally glabrous in basal half and densely pilose or hirsute in upper half and throat, lobes 5, ligulate, 2–2.5 mm long, acute, plane at tip; anthers narrowly ellipsoid, in short-styled form 1–1.2 mm long and exserted on well-developed filaments, in long-styled form c. 1.5 mm long, exserted, and positioned just above top of corolla tube; stigmas linear, 1.2–1.5 mm long, in short-styled form exserted and positioned just above top of corolla tube, in long-styled form well exserted. Fruits ellipsoid, 3–3.5 × 2.5–3.5 mm, weakly laterally flattened, glabrous but with surface granular, woody, with smooth rounded beak portion c. 1 mm high; seeds rhombic to irregularly angled in outline, flattened, 0.5–1 mm long, central seed portion elliptic and smooth, marginally with irregular lacerate wing.

Etymology. – The species epithet refers to the relatively thick texture of the leaves that is an uncommon character in Danais.

Distribution and ecology. – This new species is found in lowland and medium altitude moist evergreen forests between 606–990 m in central eastern Madagascar (Reg. Atsinanana [Prov. Toamasina]).

Conservation status. – Danais coriacea is known from only three recent collections dating from 2006 to 2011. These collections represent three unique occurrences, two populations, and two locations (sensu IUCN, 2022) based on the most likely serious threat of small-scale, shifting agriculture. The EOO is estimated at 23 km^2, within the limits for “Critically Endangered” under IUCN Red List Criterion B1; and the AOO is estimated at 8 km^2, also within the limits for “Critically Endangered” under Criterion B2 (IUCN, 2012). Of the three known collections, two are from the area near the summit of Ankerana massif and are considered a single location (sensu IUCN, 2022) with regard to the most likely serious threat. Ankerana Massif is part of the Ankeniheny Zahamena Corridor (IUCN category VI, natural resources reserve), a large, protected area in eastern Madagascar. This site is threatened by slash-and-burn agriculture, illegal mining (quartz, pink beryl, gold), and illegal logging (for Dalbergia L. f. species, Fabaceae). The protected site was designated in 2015, but a loss of 6.6 % in forest cover has been reported from 1996–2016 (Goodman et al., 2018). The remaining collection is from the area of Ambinanindrano II, which is not a formally protected area and thus is considered a separate locality; it is one of the main areas of sugar cane plantations in the region. Only two documented locations for this species meet the IUCN criterion for “Endangered”; however, one locality is in an unprotected area subject to threat from plantation and shifting agriculture practices as well as human settlement, potentially decreasing suitable habitat. The species has not been collected at this locality since 2006 and may no longer be extant there. A continuing decline is inferred for the extent of occurrence, the area of occupancy, the area, extent, and/or quality of habitat, and the number of locations and D. coriacea is assessed as “Critically Endangered” [CR B1ab(i,ii,iii,iv)+2ab(i,ii,iii,iv)].

Notes. – Danais coriacea is characterized by opposite, petiolate, stiff-textured leaves; triangular to subtruncate stipules that are shortly fused around the stem; axillary, pedunculate, cymose inflorescences with the flowers subsessile in congested groups; a relatively small, funnelform, white to yellow corolla; and relatively small, subglobose capsules with relatively small seeds. This species is unusual in Danais in its relatively thick-textured leaves and white flowers. The woody capsules are closely clustered on the infructescence, but do not appear to become fused as in some Danais species.

Danais coriacea is similar to D. longipedunculata (Fig. 3B), which is the only other species of the genus with stipules fused around the stem. Danais longipedunculata can be separated by its corollas with the tube dull purple to brown or blackened and 5–6 mm long, and the lobes usually bright yellow to red. Danais coriacea is also similar to D. aurantiaca (Fig. 3A) which can be separated by its interpetiolar stipules, leaves with the higher-order venation closely densely reticulated and prominulous adaxially, and bright orange to red corollas with the tube 4–9 mm long.

Additional specimens examined. – Madagascar. Reg. Atsinanana [Prov. Toamasina]: Distr. Brickaville, Comm. Maroseranana, Fkt. Ambodilendemy, vers le sommet d'Ankerana, 18°25′42″S 48°46′57″E, 816 m, 19.III.2011, Ravelonarivo et al. 3715 (MO, P, TAN); Distr. Vatomandry, Ambalabe, Ambinanindrano II, à l'W de Toby Foara, 19°09′53″S 48°34′33″E, 606 m, 19.IX.2006, Razanatsima 167 (MO, P, TAN).

≡ Paederia fragrans Lam., Encycl. 2: 260. 1786.

– Danais denudata Pers., Syn. Pl. 1: 198. 1805 [nom. illeg. superfl.].

Lectotypus (first-step designated by Puff & Buchner, 1994: 54; second-step designated here): Mauritius: “Isle de France, dans les bois de Palma”, s.d., Commerson s.n. (P-LA [2-part specimen: P [P00308616, P00308619] microfiche 294/17-18!; isolecto-: B [B -W 05104 -01 0] image!, LINN [LINN-HS432-2, LINN-HS432-3] images!, MPU [MPU023584, MPU023587, MPU023588] images!, P [P00539954, P00539953, P00057156] images!).

Distribution and ecology. – This species is found in dry to humid lowland to highland forests on Mauritius and Réunion Islands (Verdcourt et al., 1989), and frequently today in or on the edge of forest remnants.

Notes. – Danais fragrans is circumscribed markedly more narrowly here than done by Verdcourt et al. (1989) and Puff & Buchner (1994: 53–56), who both included also in this species many Danais plants from Madagascar that we here ascribe to another taxon. In particular, the circumscription of Puff & Buchner included a notably wide range of variation, especially morphologically in inflorescence, flower, and fruit characters, and in ecological range. The morphological description presented by Puff & Buchner for D. fragrans was apparently based only on plants from Madagascar, where they studied this genus. Their treatment cited also names from the Mascarenes but did not apparently incorporate morphological features from those plants, except they noted (Puff & Buchner,1994: 55) in commentary that the plants from these two regions generally differ in flower and fruit size. The plants from Madagascar and the Mascarenes that were said to belong to the same species do not seem to have been directly studied together taxonomically until recently, when they were analyzed with molecular data by Krüger et al. (2012) and Razafimandimbison et al. (2022). Those studies found them separated into two lineages, with one lineage in each region. Morphological review here finds the plants of these two regions to be generally morphologically separable, although some individual features of some plants in each region are similar to those of some plants of the other region so the full descriptions of these two groups have some overlapping characteristics. The plants of these two regions are formally separated taxonomically here as D. fragrans, found in the Mascarenes, and D. lyallii, found on Madagascar (Razafimandimbison et al., 2022; Fig. 3C). A description and nomenclatural synonymy of D. fragrans corresponding to its circumscription here were presented by Verdcourt et al. (1989: 41).

Danais lyallii differs from D. fragrans by its leaf blades that dry with a chartaceous to subcoriaceous texture, salverform to broadly funnelform corolla tubes 2–6 mm long, and stiffly chartaceous to woody fruits, vs. leaf blades drying papyraceous to chartaceous, salverform corolla tubes 5–10 mm long, and chartaceous to papery fruits in D. fragrans. Danais fragrans is also similar to D. xanthorrhoea of Tanzania, in continental East Africa, which has a corolla tube 2.5–6 mm long. All three of these species are rather weakly distinguishable morphologically, in spite of their clear molecular separation (Razafimandimbison et al., 2022). Danais fragrans and D. xanthorrhoea are the most widely separated species geographically in the genus, and by far the most similar species pair in Danais.Their close but apparently homoplasious morphological similarity may be informative, when the genus is better understood, about diversification and adaptive potential in Danais.

The original material corresponding to this name comprises several specimens. The protologue treated this species under both a common name, “Danaïde odorante”, and the formal binomial Paederia fragrans, with only the common name attributed to Commerson. The name P. fragrans has been attributed to “Comm. ex Lam.” in various treatments based on this citation, but this does not meet the criteria for ascribing the published name in Paederia to Commerson (ICN Art. 46.4). The type collection is represented by a number of specimens, especially in the historical collections of MPU and P, but no individual specimen was denoted as the type in the protologue nor annotations by Lamarck so these must be considered syntypes. Puff & Buchner (1994: 54) cited the “holotype” of this name as deposited in P-LA along with isotype material in P, which constitutes a first-step lectotypification. There are six specimens from Mauritius in Lamarck's herbarium (P-LA), which are imaged online and also photographed in the microfiche of that herbarium (Burns-Balogh, 1987). One of these is attributed to Joseph Martin (1760?–1826) as “Jos. Mart.”, and three lack any collector information although one has the number “499”. Two other flowering specimens there are curated with one label, and so comprise a two-part specimen. This label has the locality information “Isle de Fr.” for Isle de France, now Mauritius, and “Com.on” for Commerson in Lamarck's hand. It is the only one in either Lamarck's herbarium or other general collections that is also annotated with “danaïde odorante”. This P-LA specimen agrees with the protologue and is therefore designated here as the lectotype. Two specimens at MPU originating from Cambessèdes's herbarium [MPU023588, MPU023584] and another in the general collection there [MPU023587], one in LINN [LINNHS432-2], and one in the P general collection [P00057156] also correspond to this gathering, and have flowers, original label data with the locality information as given in the protologue, Commerson as the collector, and no conflicting dates so these are apparently isolectotypes. Additional specimens of this species are attributed to Commerson in various collections, and some of these were also cited as isotypes by Puff & Buchner. Most of those are labelled only with information copied from the protologue of P. fragrans. Several of these match the lectotype and isolectotypes in condition and leaf size and lack conflicting information, and also seem to be isolectotypes. Some other specimens are curated as possible types of this name but are not here confirmed as isolectotypes. One at B-W [B -W 05104 -01 0] lacks any collector information and is in fruit while the confirmed Commerson collection comprises only flowering stems. One specimen at MPU [MPU023586] is curated as a type but lacks confirming label data. Two other specimens curated as possible types seem to have instead been collected after P. fragrans was described, one at BM [BM001024823] that is labelled as collected in 1810 and one at K [K000395000] that corresponds to Sieber's collecting trip to Mauritius in the 1820's.

Holotypus: Madagascar. Reg. Analanjirofo [Prov. Toamasina]: Distr. Maroantsetra, 5 km E of Sabavary village, 15°19′11″S 49°50′40″E, 180 m, 30.VIII.2009, Ralimanana, Andrianaivoravelona, Mananjandry & Jaovita 1231 (MO-6593981!; iso-: BR, G [G00415957]!, K, TAN).

Danais gracilis C.M. Taylor is distinguished from D. terminalis Boivin ex Drake by its leaf blades with the higher-order venation closely regularly reticulated and prominulous adaxially, and regularly axillary inflorescences with shorter peduncles and pedicels.

Climbing plants, to 3 m high; stems glabrous, weakly flattened becoming terete. Leaves opposite; blade ovate to elliptic, 4.2–10 × 2.2–5 cm, at base acute to usually obtuse, at apex acute to acuminate with tip 5–23 mm long, drying stiffy chartaceous, on both surfaces glabrous, abaxially with epidermis comprising relatively large cells; secondary veins 4–6 pairs, looping to interconnect, without domatia, with 1–2 weakly developed intersecondary veins present between pairs of secondary veins, with costa and secondary veins adaxially thinly sulcate, abaxially costa, secondary, and sometimes intersecondary veins prominent and remaining venation closely regularly reticulated and prominulous on both surfaces; petiole 0.6–1.4 cm long, glabrous; stipules interpetiolar, persistent, glabrous, broadly triangular, 1–1.5 mm long, obtuse and shortly apiculate. Inflorescences axillary, cymose or fasciculate, 3- or 5-flowered, pedunculate, glabrous; peduncle 0.2–0.5 cm long, articulated near middle, with 1–2 peduncles arising from an apparently persistent inflorescence base; branched portion umbelliform or branched to 1 order with 1 pair of secondary axes; bracts narrowly triangular, 0.3–0.5 mm long, glabrous; pedicels 3–7 mm long, sometimes articulated. Flowers pedicellate, whether distylous unknown; hypanthium ellipsoid, c. 1 mm long, glabrous; calyx limb glabrous, c. 0.5 mm long, deeply 5-lobed, lobes narrowly triangular; corolla salverform, pink to white, pale green, or orange, externally glabrous, tube 16–18 mm long, c. 0.5 mm in diam. near middle, with slits at base, internally glabrous except pilosulous in upper 1/4, lobes 5, narrowly ligulate, 5–5.5 mm long, obtuse to acute, fleshy, plane at tip; anthers in long-styled form narrowly oblong, c. 1.8 mm long, positioned at top of corolla tube; stigmas linear, in long-styled form 4–5 mm long and well exserted. Fruits not seen.

Etymology. – The species epithet refers to the relatively very slender corolla tubes.

Distribution and ecology. – This new species is found in lowland moist evergreen forest at c. 180 m in east central Madagascar (Reg. Analanjirofo [Prov. Toamasina]).

Conservation status. – Danais gracilis is known from only two recent collections dating from 2003 and 2009. These collections represent two unique occurrences, one population, and one location (sensu IUCN, 2022) based on the most likely serious threat of small-scale, shifting agriculture. The EOO is estimated at 0 km^2; however, if EOO is less than AOO, EOO should be changed to make it equal to AOO to ensure consistency with the definition of AOO as an area within EOO (IUCN, 2012). Thus, the EOO is adjusted here to equal 8 km^2, the estimated area of occupancy (AOO). Both the EOO and AOO are within the limits for “Critically Endangered” under IUCN Red List Criterion B1 and B2 (IUCN, 2012). Both of the known collections are in the Farankaraina Forest area, a community-managed forest without formal protection status. Forests without formal protection on the Masoala Peninsula are subject to degradation by small-scale slash-and-burn agriculture (tavy) and resource exploitation, including logging, hunting and mining (Goodman et al., 2018). A continuing decline is inferred for the extent of occurrence, the area of occupancy, the area, extent, and/or quality of habitat, the number of locations and the number of mature individuals, and D. gracilis is assessed as “Critically Endangered” [CR B1a b(i,ii,iii,iv,v)+B2ab(i,ii,iii,iv,v)].

Notes. – Danais gracilis is characterized by opposite, petiolate, medium-sized leaves, with the higher-order leaf venation closely regularly reticulated and prominulous on both surfaces and the secondary veins sulcate adaxially; axillary, umbelliform inflorescences borne on short articulated peduncles and with three to five pedicellate flowers; and medium-sized, notably slender corollas that are variously pale green or pink to white and/or orange. This species is known only from two collections, but these match each other and were made some years apart at different localities in the same region, which is not well explored botanically. Danais gracilis has very slender corolla tubes even for this genus that are distinctive. The label data on both specimens of this new species describe differing colors on the buds, tube, and lobes, as in most species of Danais, but vary in the colors noted. Such variation in corolla color has been reported in several Danais species with corollas of similar shape and size (Puff & Buchner, 1994), none of which have been studied in the field for this. The fruits of D. gracilis are not yet documented, but the flowers and inflorescences distinguish this new species within Danais.

Danais gracilis is similar to D. terminalis, which is found in the same general region and, in contrast, only known in fruit. Danais terminalis can be separated from D. gracilis by its narrower leaf blades, 1–2.8 cm wide, with the venation weakly and laxly reticulated and plane or only irregularly thickened, along with its usually terminal inflorescences with longer peduncles, 2–3.8 cm long.

Additional specimen examined. – Madagascar. Reg. Analanjirofo [Prov. Toamasina]: Distr. Maroantsetra, comm. Anjahana, Fkt. Andranofotsy, Farankaraina forest, 15°26′30″S 49°51′00″E, 11.VI.2003, Antilahimena 1985 (MO, P, TAN).

Lectotypus (designated here): Madagascar: “Central Madagascar, in forests of the province of Imerina”, s.d., Baron 1304 (K [K000046174] image!; isolecto-: K [K000046175] image!).

Distribution and ecology. – This species is found in medium altitude moist evergreen forest in the Central Plateau of Madagascar.

Notes. – Danais hispida is characterized by its combination of paired leaves, triangular stipules that are densely ciliate marginally, cymose inflorescences that range from shorter than to about as long as the leaves, a corolla tube 3.5–6 mm long, and fruits 5–6 mm in diameter with papery walls. Puff & Buchner (1994: 61) included in this species plants with the leaves and stems variously “covered with long multicellular hairs” and “hairy to more or less glabrous”, but the pubescence is characterized in more detail here as villosulous, hirtellous, or hispid and generally persistent. Puff & Buchner (1994: 62) reported that D. hispida was poorly documented and no modern material had been seen by them, and suggested it may be extinct due to the extensive habitat destruction of the Central Plateau of Madagascar. No modern specimens of D. hispida have been seen in this study. Puff & Buchner also noted that this species is similar to D. cernua, so it may be confused with that common species in some herbaria although the pubescence and stipules clearly separate it.

Puff & Buchner annotated two specimens of Baron 1304 at K as the “holotype” [K000046175] and “isotype” [K000046174] of Danais hispida, and cited these two specimens (PUFF & BUCHNER, 1994: 61) as such. No individual specimen was identified as the type of D. hispida in the protologue or annotations by Baker, so these must be considered syntypes. No conclusive identifying information for the individual type specimens was cited by Puff & Buchner in their published treatment either, so that citation cannot be considered as an inadvertent lectotypification. Accordingly, D. hispida is typified here. The specimen annotated by Puff & Buchner as the isotype bears an original label of Baron's with the number 1304 with an annotation with this species name, apparently by Baker, and a note that the specimen was sent to K in 1882; two stems with well-developed infructescences and fruits; and an annotation that this specimen is the type of this name with the protologue reference in an unidentified hand that may be Sandwith's. This specimen seems to effectively represent the holotype, because it was apparently available at K to Baker before D. hispida was described in 1883. The other specimen at K was annotated by Puff & Buchner as the holotype, but this may be problematic. This second specimen has one stem in good condition; a hand-written and hand-drawn morphological analysis of the plant; and a similar label to the other specimen identifying this as D. hispida and as Baron 1304 from Central Madagascar but noting that this was purchased by K in June 1889. This second specimen seems to be a duplicate of the other and part of the type gathering, but perhaps not available to Baker before this species was described. These two specimens seem to logically represent the holotype and an isotype of D. hispida, but this is reverse of the interpretation of Puff & Buchner, so this name is lectotypified here for clarity.

Lectotypus (designated here): Madagascar. Reg. Alaotra-Mangoro [Prov. Toamasina]: forêt d'Anamalazaotra, 800 m, s.d., Perrier de la Bâthie 6930 (P [P00057171] image!; isolecto-: P [P00057172] image!).

= Danais masoalana C.M. Taylor in Candollea 68: 175. 2013, syn. nov. Holotypus: Madagascar. Reg. Analanjirofo [Prov. Toamasina]: Masoala Peninsula, S of Ambanizana, Andranobe, Piste A, 15°41′S 49°58′E, 50 m, 12.II.1999, Hoffmann, Robinson, McPherson & Harimalala 36 (MO-5938276!; iso-: G [G00369979]!, K, TAN).

Distribution and ecology. – This species is found in lowland and medium altitude moist forest from sea level to 1200 m, from northeastern to southeastern Madagascar (Reg. SAVA [Prov. Antsiranana], Reg. Analamanga [Prov. Antananarivo], Reg. Analanjirofo and Alaotra-Mangoro [Prov. Toamasina], Reg. Atsimo-Atsinanana [Prov. Fianarantsoa], Reg. Anosy [Prov. Toliara]).

Notes. – Danais longipedunculata is characterized by opposite, petiolate, elliptic to usually obovate leaves; distinctive tubular stipules; lax, axillary, pedunculate, cymose inflorescences; and slender corollas with the tube dull- or dark-colored and 5–6 mm long and the lobes generally yellow to red. The stipules are subtruncate to triangular and 2–5 mm long, and are initially fused around the stem in a tubular sheath 0.5–1 mm long. With age the upper, interpetiolar portion of the stipule usually is deciduous, and the intrapetiolar portion of the sheath tardily splits as the stem enlarges so the stipule form can be mistaken if only older nodes are studied. This tubular stipule form is unusual in Danais and was considered unique in the genus by Puff & Buchner (1994) but is now also known in D. coriacea; see that species for their distinctions.

Danais longipedunculata was known to Puff & Buchner (1994: 31–32) from nine specimens that all have average-sized leaves for Danais, and it was characterized by them by its short stipules, 2–2.5 mm long, that are fused around the stem; peduncles they characterized as “long” and “up to 100 mm long” (p. 31); and habitat at 150–1200 m along the eastern escarpment. Danais masoalana originally appeared to differ from this in its triangular stipules 4–5 mm long, shorter peduncles 1.5–3.5 cm long, and range in the Masoala Peninsula at 0–400 m, and it was known from only three specimens with relatively large leaves. Additional and more recent specimens now show wide morphological variation in various features, including leaf size, stipule form and size, and peduncle length, and this variation is now documented as continuous, with the extreme forms of the variation sometimes both found on the same collection. Therefore, these species can no longer be separated. Danais longipedunculata is now documented across a relatively wide geographic range by more than 25 collections.

Two duplicates of the type collection of Danais longipedunculata are curated at P, and both were annotated with this name by Homolle. There is no indication in the protologue or by annotation that one of these was a holotype, so these must be considered syntypes. Puff annotated one of these as the holotype and did not annotate the other. Puff & Buchner (1994: 31) cited the “holotype” as deposited at P, but gave no more identifying information to distinguish which specimen that was so this cannot be considered as an inadvertent lectotypification. A lectotype is designated here, and the specimen annotated as the holotype by Puff is chosen.

Lectotypus (designated by Puff & Buchner, 1994: 54): Madagascar: “Central Madagascar”, s.d., Baron 3895 (K [K000046124] image!).

Danais obovata Drake in Bull. Soc. Bot. France 45: 348. 1899. Lectotypus (designated by Puff & Buchner, 1994: 54): Madagascar: sine loco, s.d., Boivin s.n. (P [P00057182] image!).

Distribution and ecology. – This species is found widely and frequently on Madagascar, in lowland and medium altitude moist evergreen forest and associated secondary vegetation from sea level to 2000 m, (Reg. Analamanga, Bongolava, Itasy, and Vakinankaratra [Prov. Antananarivo], Reg. DIANA and SAVA [Prov. Antsiranana], Reg. Amoron'i Mania, Atsimo-Atsinanana, Ihorombe, Matsiatra Ambony, and Vatovavy-Fitovinany [Prov. Fianarantsoa], Reg. Sofia [prov. Mahajanga], Reg. Alaotra-Mangoro, Analanjirofo, and Atsinanana [Prov. Toamasina], Reg. Anosy [Prov. Toliara]).

Notes. – Danais lyallii is circumscribed markedly differently here from the treatment of Puff & Buchner (1994). This species is characterized by glabrous to puberulous or pilosulous stems and leaves; opposite, petiolate, elliptic to oblanceolate or obovate leaves with the blades drying chartaceous to coriaceous; interpetiolar, triangular stipules that are apically acute to broadly rounded or infrequently bidenticulate; axillary, pedunculate, cymose inflorescences that are shorter than the leaves and often less than half as long as them; a calyx limb 0.3–1 mm long that is lobed for about 1/3–2/3 of its length; funnelform to salverform, relatively small, orange to red corollas; and stiffly chartaceous to woody capsules 5–8 mm in diameter. The leaf blades are 3–9 × 1.5–6 cm and vary in shape markedly from narrowly elliptic to broadly obovate, and apically in shape from acute to acuminate, obtuse, or sometimes rounded to subtruncate with a short (3–5 mm long), abruptly narrowed tip. The venation is laxly reticulated and prominulous adaxially, and the blade is usually shiny on both surfaces. The corollas also vary in size and form, as detailed below, but overall have a tube 2.5–6 mm long and lobes 1.5–4 mm long, with basal slits sometimes present on the tube.

As noted in the discussion above of Danais fragrans, the plants here treated as D. lyallii were included previously within the circumscription of that other species (VERDCOURT et al., 1989; Puff & Buchner, 1994); see the discussion there for their distinctions. And, as noted in the discussion of D. cernua, Puff & Buchner (1994: 53–56) also included in D. fragrans some plants but here separated into D. cernua; see the discussion of D. cernua for the separation of that other species from D. lyallii. Even with these taxonomic changes, D. lyallii is still the most morphologically variable, commonly collected, and widely distributed species of Danais on Madagascar.

Three groups of plants can be generally characterized within Danais lyallii and may represent diversifying forms although they are not fully distinguishable morphologically. These are noted here for further study:

– Most of the plants included here in Danais lyallii agree with the circumscription of Puff & Buchner (1994, as D. fragrans), and have leaf blades that are generally obovate to broadly elliptic, obtuse to subtruncate at the apex sometimes with a short acuminate tip, stiff-textured to coriaceous when dry, and relatively larger compared to the full size range of the species; stipule margins that are entire or more often densely ciliolate; and a funnelform corolla with the tube 2.5–5 mm long and the lobes usually half or more as long as the tube, 1.5–3 mm long. These corollas are rather slenderly funnelform, with the throat at most twice as wide as the base. These plants are found at 0–1770 m widely from northern to southern Madagascar and match the types of D. lyallii and D. obovata.

– Another group of plants included here in Danais lyallii has leaf blades that are obovate to oblanceolate, acute to rounded and acuminate at the apex, chartaceous when dry, and relatively smaller compared to the full size range of the species; stipule margins that are densely ciliolate; and a salverform corolla with the tube 4–6 mm long and the lobes 1.5–4 mm long. These corolla tubes are cylindrical, but wide enough in diameter to accommodate the included anthers without an abrupt distal swelling. These plants are found at 700–2000 m from northern to central-southern Madagascar (e.g. Anderberg et al. 18, Leeuwenberg 14209, Razanatsoa 26). These correspond to some of the plants that were included in D. cernua by Puff & Buchner (1994) that are here treated as D. lyallii (e.g. Croat 29469, Dorr et al. 4566).

– A last group of plants included here in Danais lyallii is similar to the preceding group in its leaves and stipules but has corollas with a stout funnelform tube 2.5–3.5 mm long and more than twice as wide at the throat as at the base, and corolla lobes that are always more than half as long as the tube and often almost as long. These plants are also found widely from northern to southern Madagascar and from 60–1600 m. This group includes plants that can easily be keyed incorrectly as D. cernua in Puff & Buchner ‘s (1994) treatment (e.g. Antilahimena & Félix 3160, Antilahimena et al. 3245, Leeuwenberg et al. 14480), and these are the plants that have contributed significantly in practice to the confusion over the identity and circumscription of D. lyallii.

Lectotypus (first-step designated by Puff & Buchner 1994: 45; second-step designated here): Madagascar. Reg. DIANA [Prov. Antsiranana]: Bas Sambirano, IV.1924, Perrier de la Bâthie 16277 (P [P00057178] image!; isolecto-: P [P00057179, P00057180] images!).

Climbing plants, climbing to 8 m high; stems glabrous or puberulous, weakly flattened becoming terete. Leaves opposite; blade ovate, 5–8 × 1.8–4 cm, at base obtuse to rounded, at apex acute to acuminate with tip 6–10 mm long, drying stiffy chartaceous to subcoriaceous, adaxially glabrous, abaxially glabrous to puberulous; secondary veins 5–10 pairs, weakly to well differentiated from higher-order venation, weakly looping to interconnect near margins, without domatia, sometimes with 1 weakly developed intersecondary vein present between pairs of secondary veins, adaxially with costa sulcate and secondary and tertiary venation laxly, sparsely, regularly reticulated and plane to thickened, abaxially costa and sometimes secondary veins prominent and remaining venation plane to thickened; petiole 0.3–0.7 cm, glabrous to puberulous; stipules interpetiolar, persistent or tardily deciduous, glabrous, broadly triangular, 1–2 mm long, acute or apiculate, sometimes becoming indurated with age. Inflorescences terminal and also axillary along lower parts of stems, laxly cymose, pedunculate, puberulous; peduncle 0.8–3.5 cm long, flexuous, often articulated in lower half, axillary peduncles sometimes borne from apparently persistent inflorescence bases; branched portion 2–4 × 2–5 cm, umbelliform or branched to 1 order with 1 pair of secondary axes, with 3–9 flowers in cymes of 3–5; bracts narrowly triangular to linear, 1–3.5 mm long, puberulous; pedicels 6–15 mm long. Flowers pedicellate, whether distylous unknown; hypanthium ellipsoid, c. 1.2 mm long, puberulous; calyx limb puberulous, 5-lobed to base, lobes narrowly triangular to linear, 0.5–3 mm long and unequal on an individual flower, at least some lobes 1.5 mm long or longer; corolla salverform, dull to dark red or dark purple, externally glabrous, tube 16–18 mm long, c. 0.8 mm in diam. near middle, with slits at base, internally glabrous except densely pilosulous in uppermost 2.5 mm, lobes 5, narrowly triangular, 7–8 mm long, acute, fleshy, plane at tip; anthers narrowly oblong, in long-styled form c. 3 mm long, included, positioned at top of corolla tube; stigmas linear, in long-styled form c. 3 mm long and well exserted on slender style. Fruits oblate to ellipsoid, c. 6 mm in diam., chartaceous, glabrous, with truncate top; seeds not seen.

Distribution and ecology. – This species is found in lowland moist evergreen forest from 40 to 310 m in northern Madagascar (Reg. DIANA [Prov. Antsiranana]).

Notes. – Danais nigra is characterized by a slender, laxly twining habit; opposite, petiolate leaves with ovate, stiff-textured blades with the higher-order venation adaxially plane to thickened; short interpetiolar stipules that are triangular and acute or sometimes apiculate; pedunculate, lax, terminal and axillary inflorescences with well-developed axes and flexuous pedicels; relatively well-developed, narrow calyx lobes; and slenderly salverform, dark purple to dark red corollas with a well-developed tube. The stems are notably flexuous and laxly branched. The few flowering specimens all agree with the long-styled form of Danais species, but whether this species is distylous cannot yet be confirmed.

Danais nigra is circumscribed differently here than done by Puff & Buchner (1994), and in particular, more narrowly. They included in this species plants from east-central to northwestern Madagascar and reported it from a wide range of habitats with a wide range of corolla colors. The type of D. nigra is from northwestern Madagascar, and this species is circumscribed here to include only plants from that region, which are morphologically consistent and distinct from those of the east-central region. This second, east-central group of plants does not match any other named species of Danais and is newly described here as D. puffiana Razafim. & C.M. Taylor (see below). That new species differs from D. nigra in its leaf blades adaxially with the costa prominulous and the higher-order venation prominulous and regularly reticulated; larger inflorescences, with 12 to 36 flowers, and shorter pedicels, half or less as long as those of D. nigra; and a yellow to pale purple, internally glabrous corolla tube. Puff & Buchner (1994: 46) tentatively also included D. perrieri Homolle in their circumscription of D. nigra, but this species is also distinct as outlined in its treatment below.

Danais nigra is also similar to D. rubra, which is known only in flower and can be separated by its leaf blades with the higher-order venation densely closely reticulated and prominulous on both surfaces, shorter calyx lobes 0.5–0.8 mm long, and shorter corollas, with the tube 7.5–8.5 mm long. Danais nigra is also similar to D. terminalis, which is only known in fruit; D. terminalis can be separated by its elliptic to elliptic-ovate leaves, calyx limb with deltate lobes 0.5–1 mm long, and allopatric range in central eastern Madagascar.

Two syntype collections were cited in the protologue of Danais nigra: Perrier de la Bâthie 3830 and 16277. Puff & Buchner (1994: 45) cited the latter as the lectotype. Three duplicates of the latter collection are deposited at P. Thus, a second step lectotypification is necessary and P00057178 annotated by Puff & Buchner as lectotype is here designated as such.

Holotypus: Madagascar. Reg. Alaotra-Mangoro [Prov. Toamasina]: Distr. Moramanga, Andasibe, Menalamba, Ambatovy forest, 18°49′38″S 48°18′48″E, 1205 m, III.2005, Antilahimena, Andriantiana & Félix 3725 (MO-6406412!; iso-: G [G00415958]!, P [P04541056] image!, TAN).

Danais ovata C.M. Taylor is distinguished from D. tsaratananensis Homolle by its regularly axillary as well as sometimes terminal inflorescences, longer pedicels, larger corollas, and flowers with the stamens and stigmas all partially or fully included.

Climbing plants, extensively twining, climbing to 7 m high; stems glabrous, weakly flattened becoming terete. Leaves opposite; blade ovate, 2.5–9 × 1–4 cm, at base rounded to usually truncate or cordulate, at apex acute to acuminate with tip 5–12 mm long, drying stiffly chartaceous to subcoriaceous, on both surfaces glabrous and shiny; secondary veins 5–6 pairs, weakly differentiated from higher-order venation, weakly looping to interconnect near margins, without domatia, with 1–2 weakly developed intersecondary veins present between pairs of secondary veins, with costa thinly prominulous adaxially and prominent abaxially, with remaining venation regularly, laxly reticulated and prominulous on both surfaces; petiole 0.4–1.5 cm long, glabrous; stipules interpetiolar, persistent, glabrous, broadly triangular to truncate, 0.2–1 mm long, obtuse and apiculate or reduced to a thickened line, perhaps glandular. Inflorescences axillary and sometimes terminal, cymose to paniculiform, pedunculate, glabrous; peduncle 0.5–2.5 cm long, flexuous, articulated at base and sometimes also near middle, axillary peduncles borne from an apparently persistent inflorescence bases; branched portion 1–5 × 1–3 cm, branched to 1 order, with 1–2 pairs of secondary axes, with flowers solitary on a peduncle or in lax cymes of 3 (or sometimes 5); bracts narrowly triangular, 0.5–1.5 mm long, glabrous; pedicels 6–25 mm long. Flowers pedicellate, distylous; hypanthium ellipsoid, c. 0.8 mm long, glabrous; calyx limb glabrous, 0.8–1 mm long, 5-lobed shallowly or for up to half its length, lobes deltate to narrowly triangular; corolla salverform, yellowish green to yellowish pink or yellow, externally glabrous, tube c. 18 mm long, c. 0.8 mm in diam. near middle, with slits at base, internally glabrous in basal 1/3 and pilosulous in upper 1/3, lobes 5, narrowly triangular, 10–12 mm long, acute, fleshy, plane at tip; anthers narrowly oblong, in short-styled form c. 3 mm long and partially exserted, in long-styled form c. 3 mm long, included, positioned at top of corolla tube; stigmas linear, in long-styled form c. 4 mm long and partially exserted, in short-styled form 1–2 mm long, included, and positioned near middle of corolla tube. Fruits ellipsoid to subglobose, 5–10 × 5–10 mm, glabrous, chartaceous to subcoriaceous, with smooth rounded beak portion 2 mm high, with pedicel becoming markedly thickened at top; seeds elliptic to circular, flattened, 2.5–3 mm in diam., central seed portion elliptic and smooth, marginally with narrow lacerate wing 0.2–0.8 mm wide.

Etymology. – The species epithet refers to the ovate shape of the leaf blades.

Distribution and ecology. – This species is found in lowland and medium altitude moist evergreen forest between 200 and 1200 m in eastern Madagascar (Reg. Alaotra-Mangoro, Analanjirofo, Atsinanana [Prov. Toamasina]).

Conservation status. – Danais ovata is known from only six recent collections dating from 2002 to 2012. These collections represent six unique occurrences, four populations, and three locations (sensu IUCN, 2022) based on the most likely serious threat of small-scale, shifting agriculture. The EOO is estimated at 1,325 km² and the AOO at 12 km^2. Both the EOO and AOO are within the limits for “Endangered” under IUCN Red List Criterion B1 and B2 (IUCN, 2012). Three of the six occurrences are within the mine footprint of the Ambatovy (Reg. Alaotra-Mangoro) nickel-mining project and have been removed from the dataset for the calculation of assessment parameters due to planned or actual destruction of this habitat. Of the remaining three occurrences, one is from Ankerana forest (Reg. Atsinanana), which has no known formal protection; one is from Miarinarivo Ifasina, along the Onibe River (Reg. Atsinanana), also with no known formal protection; and the northern-most occurrence is from Maroantsetra, within the Farankaraina Forest (Reg. Analanjirofo), a community-managed forest also without formal protection status. Forests without formal protection on the Masoala Peninsula are subject to degradation by small-scale slash-and-burn agriculture (tavy) and resource exploitation, including logging, hunting and mining (Goodman et al., 2018). Each of these three collection sites is considered an individual location, and this low number of locations is within the limits of “Endangered” under the subcriterion ‘a’ of criterion B2 (IUCN, 2012). A continuing decline is inferred for the extent of occurrence, the area of occupancy, the area, extent, and/or quality of habitat, the number of locations, and the number of mature individuals, and D. ovata is assessed as “Endangered” [EN B1ab(i,ii,iii,iv,v)+2ab(i,ii,iii,iv,v)].

Notes. – Danais ovata is characterized by an extensively twining habit; opposite, petiolate leaves with stiff-textured, ovate blades; interpetiolar, broadly triangular stipules that are relatively quite small; axillary and sometimes also terminal (Antilahimena et al. 1216), few- to several-flowered, pedunculate, cymose inflorescences with well-developed pedicels; slender, medium-sized, yellowish green to yellow or yellowish pink corollas; and fruiting pedicels that become strongly thickened at the top. The ovate shape of the leaf blades is distinctive and unusually consistent among the specimens of this species, and this feature aids identification of D. ovata. The leaf blades are also distinctive in being shiny on both surfaces, with the laxly reticulated tertiary venation usually thickened on both sides. Notably, D. ovata differs in its distylous morphology from most species of this genus: in most species the stigma or stamens are fully exserted and held well above the top of the corolla in the corresponding long-styled or short-styled from, with well-developed slender filaments or styles, and these structures are reciprocally positioned but only partially exserted in the corresponding flower forms in this new species.

Danais ovata is similar to D. tsaratananensis, which can be separated by its inflorescences that are all terminal and sometimes also produced in the uppermost leaf axils, shorter pedicels 2–3 mm long, smaller corollas, with the tube 8–10 mm long and the lobes 3–4 mm long, and flowers with either the stigmas or the stamens fully exserted and held well above the top of the corolla.These two species are both notable in Danais in their distinctive, consistently ovate leaf blades. Danais ovata is also similar to D. terminalis, detailed below, which is known only in fruit but can be separated by its elliptic to elliptic-ovate leaf blades; mostly shorter petioles, 4–7 mm long; triangular to acuminate stipules 1–1.5 mm long; fruiting pedicels that do not become strongly thickened at the top; and seeds with a broader marginal wing.

Additional specimens examined. – Madagascar. Reg. Alaotra-Mangoro [Prov. Toamasina]: Distr. Moramanga, Antsangimaso, Fkt. Ampitambe, comm. Ambohibary, Ambatovy, 18°51′32″S 48°19′29″E, 1115 m, 24.II.2005, Razanatsoa 293 (MO, P, TAN), 344 (MO, P, TAN); ibid., 18°50′43″S 48°17 ′22″E, 1044 m, 26.II.2005, Razanatsoa 344 (MO, P, TAN). Reg. Analanjirofo [Prov. Toamasina]: Distr. Maroantsetra, comm. Anjahana, Fkt. Ambodivangy, 15°25′47″S 49°50′07′E, 200 m, 19.VII. 2002, Antilahimena et al. 1216 (MO, P, TAN); Distr. Vavatenina, comm. Miarinarivo Ifasina, à 4 km au NO d'Ampasimaventy, le long de la rivière Onibe, 17°44′42″S 48°58′42″E, 300–350 m, 10.VII.2003, S. Randrianasolo et al. 392 (MO, P, TEF). Reg. Atsinanana [Prov. Toamasina]: Distr. Brickaville, Anjahamamy, Anivoranokely, Ankerana forest, 18°24′50″S 48°49′26″E, 768 m, 28.I.2012, Antilahimena 8103 (MO, P, TAN).

Lectotypus (designated here): Madagascar. Reg. Atsinanana [Prov. Toamasina]: Mont Andriantantely au Nord d'Anivorano, VI.1922, Perrier de la Bâthie 14752 (P [P00057185] image!; isolecto-: P [P00057184, P00057186] images!).

Distribution and ecology. – This species is found in lowland moist evergreen forest between 60 and 70 m in eastern Madagascar (Reg. Atsinanana [Prov. Toamasina], Reg. Vatovavy [Prov. Fianarantsoa]).

Notes. – The identity of Danais perrieri has not been entirely clear to recent authors (e.g. Puff & Buchner, 1994), and its seeds are still not known but the type collection can now be matched to another collection that significantly clarifies its identity (Capuron 28053-SF, MO). Danais perrieri is characterized by opposite leaves with relatively short petioles, 2–5 mm long; ovate to lanceolate leaf blades that dry papyraceous and have tertiary venation that is laxly reticulated and weakly thickened on both surfaces; interpetiolar, variably triangular to bilobed stipules c. 1 mm long that become subtruncate, indurated, and fragmented with age; terminal, pedunculate, cymose, lax inflorescences with 12–30 flowers borne on well-developed pedicels, 3–15 mm long; a calyx limb 0.8–1 mm long that is deeply divided into triangular lobes; slenderly salverform yellow corollas with the tube 13–19 mm long; and thin-textured fruits c. 4 mm in diameter.This newly identified second collection agrees with the first specimen of D. perrieri, except perhaps for its stipule morphology.The protologue described the stipules of D. perrieri as small and membranaceous, without detailing the size or shape. The stipules on the type collection are not well preserved and not entirely visible in the online scans but seem to vary from triangular to subtruncate and possibly are also bilobed. The stipules on the second specimen are well preserved, and those on younger nodes are variously triangular, bilobed, and subtruncate with well-developed colleters inserted adaxially, while those on older nodes are thickened and subtruncate with, apparently, their upper portion tardily deciduous. Puff & Buchner noted variation within several Danais species in stipule form, from acute to bilobed, and this is found in some species studied here (e.g. D. lyallii), so the apparent variation on this specimen does not disagree with the characterization of this species.

Puff & Buchner (1994: 46–47) included Danais perrieri in their circumscription of D. nigra, and noted some morphological differences between them but commented that the differences did not seem diagnostic. However, the circumscription here of D. nigra is substantially changed, and these are no longer similar. Danais nigra can now be separated from D. perrieri by its chartaceous to subcoriaceous leaf blades, inflorescences with 3 to 9 flowers, dark to dull red or purple corollas, and distribution in northeastern Madagascar. The plants that Puff & Buchner actually confused with D. perrieri and used to characterize it are those separated here from D. nigra into the new species D. puffiana. Danais puffiana can be separated by its chartaceous leaf blades with the higher-order venation regularly areolate and prominulous on the adaxial surface, shorter pedicels 0.5–3 mm long, linear calyx lobes, and a distribution centered further north, around the Masoala Peninsula and the escarpment near there.

Puff & Buchner (1994: 46) cited the holotype of Danais perrieri as deposited at P, without further specification. Three duplicates of the type collection are curated there, with one of them annotated by Puff as the “holotype”. However all of these are similarly annotated by Homolle as types of this name, and no indication was made on the specimens or in the protologue that one of them was the primary specimen so they must be considered syntypes. Two of these specimens have ample and complete material and full label data, while the third has a small stem and few flowers. The specimen with the most and best material is designated here as the lectotype. This is not the specimen that Puff annotated as the holotype, but it seems preferable as the reference specimen for this name.

Holotypus: Madagascar. Reg. Analanjirofo [Prov. Toamasina]: Distr. Maroantsetra, NW coast of Masoala Peninsula, trail along Ampanga River, c. 5 km S of village of Hiaraka, ESE of Maroantsetra, 300 m, 13.X.1986, Lowry, Rakotozafy & Nicoll 4100 (MO-3436425!; iso-: K, P [P04004803] image!).

Danais puffiana Razafim. & C.M. Taylor is distinguished from D. nigra Homolle by its leaf blades adaxially with the costa prominulous and the higher-order venation prominulous and regularly reticulated, larger inflorescences with 12–36 flowers and shorter pedicels half or less as long as those of D. nigra, and an internally glabrous corolla tube.

Climbing plants, climbing to 2 m high; stems glabrous, weakly flattened becoming terete. Leaves opposite; blade ovate, 6–13 × 3.5–7.5 cm, at base obtuse to subtruncate, at apex acute to acuminate with tip 5–10 mm long, drying stiffy chartaceous, glabrous on both surfaces, abaxially matte and perhaps pale; secondary veins 5–7 pairs, weakly to well differentiated from higher-order venation, weakly looping to interconnect near margins, without domatia, sometimes with 1 weakly developed intersecondary vein present between pairs of secondary veins, adaxially with costa and secondary veins prominulous and higher-order venation prominulous and reticulated, with areoles irregular in size and shape, 1–3 mm wide, abaxially costa prominent, secondary veins prominulous, and remaining venation thickened to prominulous; petiole 0.8–2.5 cm long, glabrous; stipules interpetiolar, glabrous, basal portion truncate, 0.1–0.2 mm long, persistent, upper portion narrowly triangular, 1–2 mm long, acute, deciduous. Inflorescences terminal, laxly cymose, subsessile to pedunculate, glabrous; peduncle 0.1–8.5 cm long, flexuous, not articulated; branched portion 2–8 × 2–16 cm (not including corollas), pyramidal, branched to 2–3 orders with 1–3 pairs of secondary axes, with 12–36 flowers in cymes of 3–5; bracts narrowly triangular to linear, 1–4 mm long, glabrous; pedicels 0.5–3 mm. Flowers pedicellate, whether distylous unknown; hypanthium ellipsoid, c. 1 mm long, glabrous; calyx limb glabrous, 5-lobed to base, lobes narrowly triangular to linear, 1–1.5 mm long, subequal on an individual flower; corolla salverform, externally pale purple and glabrous, tube 16–18 mm long, c. 0.8 mm in diam. near middle, with slits at base, internally glabrous, lobes 5, narrowly ligulate, 7–8 mm long, acute, orange-yellow, fleshy, plane at tip; anthers narrowly oblong, in short-styled form c. 1.5 mm long, well exserted; stigmas linear, in short-styled form c. 6 mm long and positioned in uppermost part of corolla tube. Fruits subglobose, c. 5 mm in diam., glabrous, weakly pustulose, subcoriaceous, with truncate top, with pedicel becoming thickened and up to 5 mm long; seeds elliptic, flattened, c. 1 mm in diam., central seed portion elliptic and smooth, marginally with narrow lacerate wing 0.2–0.5 mm wide.

Etymology. – The species epithet honors the first author of the previous treatment of Malagasy Danais, Prof. Dr. Christian Puff (1949–2013), for his valiant detailing of this genus with a limited specimen representation.

Distribution and ecology. – This new species is found in lowland moist evergreen forest between 205 and 550 m in eastern Madagascar (Reg. Analanjirofo [Prov. Toamasina]).

Conservation status. – Danais puffiana is known from only six collections dating from 1986 to 2013. These collections represent six unique occurrences, six populations, and four locations (sensu IUCN, 2022) based on the most likely serious threat of small-scale, shifting agriculture. The EOO is estimated at 522 km² and the AOO at 24 km^2. Both the EOO and AOO are within the limits for “Endangered” under IUCN Red List Criterion B1 and B2 (IUCN, 2012). The northwestern-most location is in the Makira Natural Park (Reg. Analanjirofo) (IUCN category II, natural park). Though a formally protected area since 2012, Makira is subject to threat from slash-and-burn agriculture (tavy), illegal mining (gold, quartz), and illegal logging for charcoal (Goodman et al., 2018). The remaining five collections are along the northwest coast of the Masoala Peninsula. The next two most northern occurrences constitute two locations, are not in Masoala National Park, and do not have protected status. The remaining three occurrences constitute a single location within the Masoala National Park (IUCN category II, national park); however, two of the occurrences are very close to human settlements and occupation (Benjana, Antalavia) near the boundary of the Park, and most likely are subject to threat from subsistence agriculture as well as uncontrolled fires and resource exploitation that includes logging, hunting, and mining (Goodman et al., 2018). The low number of locations corresponds to “Endangered” under the subcriterion ‘a’ of criterion B2 (sensu IUCN, 2022). A continuing decline is inferred for the extent of occurrence, the area of occupancy, the area, extent, and/or quality of habitat, the number of locations, and the number of mature individuals, and D. puffiana is assessed as “Endangered” [EN B1ab(i,ii,iii,iv,v)+2ab(i,ii,iii,iv,v)].

Notes. – Danais puffiana is characterized by opposite, ovate leaves with the higher-order venation regularly prominulous and reticulated on the upper surface; interpetiolar stipules with a short sheath portion and the small, triangular upper portion deciduous; terminal, cymose inflorescences with pedicellate flowers; medium-sized, yellow to purple or yellow-orange corollas; and medium-sized capsules. Only one specimen with mature flowers has been seen. Another specimen has buds, and both of these agree with the long-styled form of Danais flowers. Only one specimen with developing fruits has been seen, and its fruits are not fused together. Most of the seeds seen are clearly immature but one more mature one matches the elliptic, lacerate-winged seeds of most other Danais species. These plants were included within the circumscription of D. nigra by Puff & Buchner (1994), but they are separated here as discussed above under D. nigra.

Danais puffiana is similar to D. dauphinensis Cavaco, which is found in southeastern Madagascar and can be separated by its leaf blades with the higher-order venation plane or only irregularly thickened adaxially and the abaxial surface shiny; shorter calyx lobes, up to 1 mm long but mostly shorter; and white to olive green corollas with shorter lobes, 4–6 mm long. Danais puffiana is also similar to D. terminalis, which is also found in southeastern Madagascar and can be recognized by its leaves with the higher-order venation abaxially plane or only irregularly thickened; flexuous pedicels 5–20 mm long; and relatively large fruits and seeds, 8–12 mm in diameter and 2.5–4 mm in diameter, respectively. Danais puffiana is also similar to D. perrieri, see that last species for their distinctions.

Additional specimens examined. – Madagascar. Reg. Analanjirofo [Prov. Toamasina]: Distr. Maroantsetra, Masoala Peninsula, S of Ambanizana, Andranobe, 15°41′S 49°58′E, 205 m, 16.II.1999, Hoffmann et al. 55 (K, MO); ibid., E of village of Hiaraka, 550 m, 10.X.1986, Lowry et al. 4061 (MO); ibid., Ambanizana, 15°37′51″S 49°58′36″E, 233 m, 16.I.2013, Razafimandimbison et al. 1245 (MO, S); ibid., c. 3 km N of Antalavia, along Antalavia river, 15°47′S 50°02′E, 200–380 m, 13–16.XI.1989, Schatz et al. 2823 (K, P); Makira Protected Area, near Ankirindro, c. 9 km NW of Ambinanitelo, 15°17′44″S 49°33′12″E, 500 m, 22.V.2007, Briggs 183 (K, MO, P).

Holotypus: Madagascar. Reg. Anosy [Prov. Toliara]: Distr. Taolagnaro [Fort-Dauphin], Antsotso Avaratra, forêt Tsitongambarika, 24°34′16″S 47°12′05″E, 271 m, 1.IV.2008, Razakamalala, Rabenantoandro, Randriatafika, Roger & Mara 4132 (MO-6308607!; iso-: P, TAN).

Danais reticulata C.M. Taylor is distinguished from D. aurantiaca Homolle by its leaf blades that are not bullulate and have closely densely reticulated higher-order venation that is regularly prominulous on both surfaces, longer stigmas, capsules that do not become fused to each other, and narrowly rhombic seeds.

Climbing plants, climbing to 4 m high; stems glabrous or sometimes densely strigillose, flattened becoming terete. Leaves opposite; blade elliptic to oblanceolate-elliptic, 3–10 × 1.5–4.5 cm, at base cuneate to rounded, at apex obtuse then weakly acuminate to rounded, drying chartaceous to subcoriaceous, adaxially glabrous, abaxially glabrous or moderately to sparsely strigillose; secondary veins 6–8 pairs, weakly differentiated from higher-order venation, usually weakly looping to interconnect near margins, without domatia, sometimes with 1–2 weakly developed intersecondary veins present between pairs of secondary veins, with costa and secondary veins thickened adaxially and prominent abaxially, with tertiary and quaternary venation closely, regularly reticulated and prominulous on both surfaces; petiole 1–2.2 cm long, glabrous or densely strigillose; stipules interpetiolar, persistent, glabrous or densely strigillose, triangular to broadly triangular or subtruncate, 1–2 mm long, broadly obtuse, marginally erose and/ or densely ciliolate, abaxially glabrous or densely strigillose. Inflorescences axillary, cymose, pedunculate, glabrous; peduncle 1.5–6 cm long; branched portion 1.5–2 × 1.5–3 cm, branched to 2 orders, with 1–2 pairs of secondary axes, with flowers borne in subcapitate groups of 3–5, the groups borne at ends of axes; bracts narrowly triangular to narrowly lanceolate, 1–3 mm long, glabrous. Flowers subsessile, distylous; hypanthium obconic to ellipsoid, c. 1 mm long, glabrous; calyx limb glabrous, 0.5–1 mm long, 5-lobed shallowly or for up to half its length, lobes deltate; corolla salverform, orange, externally glabrous, tube c. 6 mm long, c. 1 mm in diam. near middle, with slits at base, internally glabrous in basal half and pilosulous in upper half, lobes 5, narrowly triangular, 4–5 mm long, acute, plane at tip; anthers narrowly oblong, in short-styled form c. 2 mm long and exserted on well-developed filaments, in long-styled form included, positioned just above top of corolla tube, and c. 1.8 mm long; stigmas linear, in short-styled form c. 5 mm long and partially exserted, in long-styled form 3.5–4 mm long and well exserted. Fruits subglobose to ovoid, 5–10 × 5–10 mm, glabrous, woody, with smooth rounded beak portion 1–2 mm high; seeds narrowly rhombic, flattened, 2–3 × 1 mm, central seed portion elliptic and smooth, marginally with lacerate wing prolonged on each end.

Etymology. – The species epithet refers to the closely reticulated leaf venation.

Distribution and ecology. – This new species is found in lowland moist evergreen forest, sometimes on basalt, between 116 and 662 m in southern Madagascar (Reg. Atsimo-Atsinanana [Prov. Fianarantsoa], Reg. Anosy [Prov. Toliara]).

Conservation status. – Danais reticulata is known from only six recent collections dating from 2007 to 2010. These collections represent six unique occurrences, three populations, and three locations (sensu IUCN, 2022) based on the most likely serious threat of small-scale, shifting agriculture. The EOO is estimated at 2,455 km² and the AOO at 16 km^2. Both the EOO and AOO are within the limits for “Endangered” under IUCN Red List Criterion B1 and B2 (IUCN, 2012). Four of the six occurrences are in the area of Tsitongambarika forest (Reg. Anosy) (IUCN category VI, natural resources reserve), which has protected status since 2015. However, the forest is subject to threats from illegal logging, slash-and-burn agriculture (tavy), hunting, and illegal mining (mica) (Goodman et al., 2018). These four occurrences are considered a single location (sensu IUCN, 2022). The fifth, western-most occurrence is within Andohahela National Park (Reg. Anosy) (IUCN category II, national park), a formally protected area. The lowland evergreen forest of Andohahela is subject to threat from slash-and-burn agriculture (tavy), uncontrolled fires, illegal logging and mining for mica (Goodman et al., 2018). The last remaining occurrence is to the north of these and located in Ankatriky forest (Reg. Atsimo-Atsinanana), which is part of Ankarabolava Natural Resources Reserve (IUCN category VI, natural resources reserve), a small area protected since 2015. There are threats to this reserve from illegal logging, slash-and-burn agriculture (tavy), clove plantations, and uncontrolled fires (Goodman et al., 2018). These last two occurrences constitute two locations, and along with the third location at Tsitongambarika place this species within the limits of “Endangered” under the subcriterion ‘a’ of criterion B2 (IUCN, 2012). A continuing decline is inferred for the extent of occurrence, the area of occupancy, the area, extent, and/or quality of habitat, the number of locations, and the number of mature individuals, and D. reticulata is assessed as “Endangered” [EN B1ab(i,ii,iii,iv,v)+2ab(i,ii,iii,iv,v)].

Notes. – Danais reticulata is characterized by its opposite, shiny, elliptic leaves with the higher-order venation closely densely reticulated and prominulous on both surfaces; interpetiolar, triangular to subtruncate stipules; axillary, pedunculate, cymose inflorescences with subsessile flowers; medium-sized orange corollas; woody, free capsules; and narrowly rhombic seeds. Similarly reticulated leaf venation is found in several Malagasy species of Danais and aids their identification. Unusually for Danais, the inflorescences of D. reticulata are glabrous even when the vegetative structures are pubescent (e.g. Birkinshaw et al. 1723, MO); in most species these are pubescent even when the vegetative structures are glabrous. The corollas are densely pubescent inside the throat and the upper part of the tube, but the pubescence does not protrude out of the tube in a “beard” as in many species of this genus.

Danais reticulata is similar to D. aurantiaca (Fig. 3A) which can be separated by its leaf blades that become at least weakly bullulate between the secondary veins with age; higher-order leaf venation that is regularly and moderately to densely reticulated and prominulous adaxially but only irregularly so abaxially; generally shorter inflorescences, 1–6 cm long, that are half or less as long as the leaves; shorter stigmas, c. 2 mm long; capsules that become partially fused as they mature; and seeds that are mostly elliptic to circular and 1.7–2 mm in diameter. Danais reticulata is also similar to D. baronii and D. schatzii, the latter newly described below. Both of these differ in their larger leaf blades, 9–15 × 4.5–8.8 cm, and larger inflorescences, with the branched portion 6–8.5 cm wide; and, both are allopatric, in northern and east-central Madagascar, respectively.

Additional specimens examined. – Madagascar. Reg. Atsimo-Atsinanana [Prov. Fianarantsoa]: Distr. Farafangana, Matanga, Mandritsara, Andranofasika forest, Ankatriky forest, 23°27 ′24″S 47 °30′17 ″E, 116 m, 2.VI.2007, Birkinshaw et al. 1723 (MO, P, TAN). Reg. Anosy [Prov. Toliara]: Distr. Fort-Dauphin [Taolagnaro], Andohahela National Park, 8.IV.2010, Krüger & Razafimandimbison 89 (MO, S); Antsotso Avaratra, 24°34′11″S 47°12′17″E, 160 m, 11.XII.2007, Razakamalala et al. 3913 (MO, P., TAN); ibid., 24°33′42″S 47°12′02″E, 662 m, 11.XII.2007, Razakamalala et al. 4003 (MO, P, S, TAN); ibid., forêt Tsitongambarika, 24°34′16″S 47 °12′05″E, 271 m, 1.IV.2008, Razakamalala et al. 4076 (MO, P, TAN).

Holotypus: Madagascar. Reg. DIANA [Prov. Antsiranana]: Distr. Ambanja, forêt classée d'Ambato, village de Bevoay, 40 km au N d'Ambanja, 13°27′30″S 48°33′01″E, 80–100 m, 17.V.1998, Randrianaivo, Birkinshaw, Antilahimena, Ranaivojaona, Randrianasolo, Rakotomamonjy & Rasolomanana 244 (MO-5206218!; iso-: P [P03794019] image!, TAN).

Danais sambiranensis C.M. Taylor is distinguished from D. longipedunculata Homolle by its leaves with more numerous secondary veins and the higher order venation laxly reticulated, interpetiolar stipules, longer calyx lobes, and longer corollas.

Climbing plants, height not noted; stems densely puberulous-strigillose, weakly flattened becoming terete. Leaves opposite; blade elliptic to obovate, 8–22 × 5–9 cm, at base acute to obtuse, at apex obtuse to acute and sometimes acuminate with tip 1–6 mm long, drying stiffly chartaceous, adaxially glabrous, abaxially densely puberulous-strigillose; secondary veins 14–16 pairs, looping to interconnect near margins, without domatia, with 1–2 weakly developed intersecondary veins present between pairs of secondary veins, with costa and secondary veins plane to thickened adaxially and prominent abaxially, with remaining venation laxly reticulated and plane on both surfaces or sometimes thickened adaxially; petiole 1.5–2 cm long, puberulous; stipules interpetiolar, persistent or tardily fragmenting, adaxially puberulous, triangular, 3–5 mm long, acute to obtuse, sometimes apparently resinous. Inflorescences axillary, paniculiform, pedunculate, densely puberulous; peduncle 8.5–12.5 cm long, flexuous; branched portion 5.5–10 × 8–18 cm, branched to 3 orders, with 2–3 pairs of secondary axes, with flowers closely set in umbelliform cymes of 5–15; bracts narrowly triangular, 0.5–4 mm long, puberulous; pedicels 0.5–5 mm long. Flowers pedicellate, distylous; hypanthium ellipsoid to ovoid, 1–1.2 mm long, densely puberulous; calyx limb densely puberulous, 1.2–1.5 mm long, 5-lobed nearly to base, lobes linear, margins sometimes involute; corolla salverform, pale green to orange on tube and orange to red on lobes, externally sparsely puberulous, tube c. 19 mm long, c. 0.3 mm in diam. near middle, with slits at base, internally glabrous except pilosulous at throat, lobes 5, narrowly ligulate, c. 3.5 mm long, acute, fleshy, plane at tip; anthers narrowly oblong, c. 2 mm long, in short-styled form well exserted, in long-styled form included and positioned in upper part of corolla tube; stigmas linear, in short-styled form c. 4 mm long, included, and positioned in upper part of corolla tube, in long-styled form c. 4 mm long and well exserted. Fruits ellipsoid, c. 4 mm in diam., puberulous, woody, with smooth rounded beak portion c. 0.3 mm high; seeds lenticular, c. 1 mm in diam., flattened to angled, sometimes with a very narrow ridge or wing on one side.

Etymology. – The species epithet refers to Sambirano biogeographic domain in northwestern Madagascar (Koechlin et al., 1974), where this species is found.

Distribution and ecology. – This new species is found in lowland moist evergreen forest between 80 and 425 m in northwestern Madagascar (Reg. DIANA [Prov. Antsiranana]).

Conservation status. – Danais sambiranensis is known from only three recent collections dating from 1998 to 2013. These collections represent three unique occurrences, two populations, and two locations (sensu IUCN, 2022) based on the most likely serious threat of small-scale, shifting agriculture. The EOO is estimated at 6 km² and the AOO at 12 km^2. The EOO is within the limits for “Critically Endangered” under IUCN Red List Criterion B1 and the AOO is within the limits for “Endangered” under IUCN Red List Criterion B2 (IUCN, 2012). Two of the occurrences are adjacent to the north of Tsaratanàna Strict Nature Reserve (Réserve Naturelle Intégrale de Tsaratanàna; Reg. DIANA) and constitute a single location. The remaining occurrence is to the northeast of the first location, in the Ambato Classified Forest (Reg. DIANA) and is near the town of Ambanja and other human settlements. Neither of these two locations are under formal protected status and the areas are subject to threat by slash-and-burn agriculture (tavy), uncontrolled fires, and resource exploitation, including logging, hunting and mining (Goodman et al., 2018). The restriction of this species to two locations falls within the limits of “Endangered” under the subcriterion ‘a’ of criterion B2 (IUCN, 2012). A continuing decline is inferred for the extent of occurrence, the area of occupancy, the area, extent, and/or quality of habitat, the number of locations, and the number of mature individuals, and D. sambiranensis is assessed as “Endangered” [EN B1ab(i, ii,iii,iv,v)+2ab(i,ii,iii,iv,v)].

Notes. – Danais sambiranensis is characterized by opposite, petiolate, well-developed leaves with relatively numerous secondary veins (for Danais); interpetiolar, triangular stipules; lax, well-developed, paniculiform inflorescences with elongated flexuous peduncles and pedicellate flowers; relatively well-developed calyx lobes; slender corollas with red lobes; and relatively small fruits and seeds. Also distinctive is the arrangement of the higher-order leaf venation, which is laxly reticulated and adaxially plane or sometimes weakly, sparsely thickened. The inflorescences are relatively large for Danais and appear to be pendulous. Few mature seeds are preserved on the fruiting paratype, and these appear to be to be variously unwinged or narrowly winged along one side. As discussed above, winged seeds were considered a characteristic feature for Danais by Puff & Buchner (1994) but are not diagnostic for this genus, and several species with unwinged seeds are included here in Danais. Danais sambiranensis is known only from the Sambirano biogeographic domain in northwestern Madagascar, which has a distinctive flora (Koechlin et al., 1974).

Danais sambiranensis is similar to D. longipedunculata (Fig. 3B), which can be separated by its leaf blades with fewer secondary veins, 5 to 11 pairs, and the higher-order venation regularly reticulated and prominulous adaxially; stipules that are at least shortly united around the stem; shorter calyx lobes 0.3–0.8 mm long; and shorter corollas, with the tube 5.5–6 mm long.

Additional specimens examined. – Madagascar. Reg. DIANA [Prov. Antsiranana]: Distr. Ambanja, lower slopes of Kalabenono, 13°38′11″S 48°39′56″E, 215 m, 20.III.2009, Briggs et al. 250 (K, MO, TAN); Distr. Ambilobe, Beramanja, Anketrabe Belinta, forêt de Manongarivo-Kalobinono, 13°38′16″S 48°40′21″E, 425 m, 29.IX.2013, Razakamalala et al. 7612 (MO, G, P, TAN).

Holotypus: Madagascar. Reg. Atsinanana [Prov. Toamasina]: Sahafina forest, W of Brickaville, 18°47′04″S 48°58′02″E, 200 m, 9.I.2017, Schatz, Lowry, Razakamalala & Randrianaivo 4430 (MO-6991769!; iso-: MO-6917297!, P, TAN).

Danais schatzii C.M. Taylor is distinguished from D. longipedunculata Homolle by its interpetiolar stipules and longer corollas.

Climbing plants to 10 m high; stems glabrous, weakly flattened becoming terete. Leaves opposite; blade lanceolate to ovate or elliptic-ovate, 8–15 × 3.5–8.8 cm, at base rounded to truncate or cordulate, at apex acute to acuminate with tip 1–6 mm long, drying stiffly chartaceous to subcoriaceous, on both surfaces glabrous and shiny; secondary veins 6–9 pairs, weakly differentiated from higher-order venation, weakly looping to interconnect near margins, without domatia, with 1–2 weakly developed intersecondary veins present between pairs of secondary veins, with costa thinly prominulous adaxially and prominent abaxially, secondary veins prominuluous on both surfaces, and remaining venation regularly, closely, densely reticulated and prominulous on both surfaces; petiole 1.8–4 cm long, glabrous; stipules interpetiolar, persistent, glabrous, triangular, 1–2 mm long, acute. Inflorescences axillary and/or terminal, cymose to paniculiform, pedunculate, glabrous or sparsely puberulous; peduncle 4.5–10.7 cm long, flexuous; branched portion 6–8.5 × 10–12 cm, branched to 3 orders, with 1–3 pairs of secondary axes, with flowers in umbelliform cymes of 3 or 5; bracts narrowly triangular, 0.1–4 mm, glabrous; pedicels 0.5–4 mm long. Flowers pedicellate, whether distylous unknown; hypanthium ellipsoid to subglobose, c. 1.2 mm long, glabrous; calyx limb glabrous, c. 0.5 mm long, 5-lobed for 1/2–3/4 its length, lobes deltate; corolla salverform, green on tube, white in throat, and orange on lobes, externally glabrous, tube c. 8 mm long, c. 0.8 mm in diam. near middle, with slits at base, internally glabrous except densely pilosulous in upper 1/4 and on bases of lobes, lobes 5, narrowly ligulate, 3.5–4 mm long, acute, fleshy, plane at tip; anthers narrowly oblong, in long-styled form c. 1.8 mm long, included, and positioned at top of corolla tube; stigmas linear, in long-styled form c. 3.5 mm long and well exserted. Fruits subglobose, c. 4 mm in diam., glabrous, woody, with smooth rounded beak portion c. 0.5 mm high; seeds not seen.

Etymology. – The species epithet honors Dr. George E. Schatz, who studied the Madagascar flora in depth for several decades and made the type collection.

Distribution and ecology. – This new species is found in lowland moist evergreen forest between 200 and 250 m in eastern Madagascar (Reg. Analanjirofo and Atsinanana [Prov. Toamasina]).

Conservation status. – Danais schatzii is known from only two collections dating from 1989 and 2017. These collections represent two unique occurrences, two populations, and two locations (sensu IUCN, 2022) based on the most likely serious threat of small-scale, shifting agriculture. The EOO is estimated at 0 km^2; however, if EOO is less than AOO, then procedurally EOO is changed to make it equal to AOO to ensure consistency with the definition of AOO as an area within EOO (IUCN, 2012). The AOO is estimated at 8 km^2, and the EOO is adjusted to equal 8 km^2. Both the EOO and AOO are within the limits for “Critically Endangered” under IUCN Red List Criterion B1 and B2 (IUCN, 2012). One of the occurrences is adjacent to Mananara-Nord National Park (Reg. Analanjirofo) but in an unprotected area of human settlements and agriculture and is considered a single location. This collection was made in 1989, and no new collections have been reported from this area; the population at this location may no longer be extant. The other occurrence is in the Harmonious Protected Landscape of Sahafina (Reg. Atsinanana) (IUCN category V, protected harmonious landscape), which is subject to threat from illegal logging of hardwoods and from invasive species (Goodman et al., 2018). Each of these collection sites is considered a location, and their small number is within the limits of “Endangered” under the subcriterion ‘a’ of criterion B2 (IUCN, 2012). However, one of these localities is in an unprotected area subject to threat from shifting agriculture practices as well as human settlement, potentially decreasing suitable habitat. A continuing decline is inferred for the extent of occurrence, the area of occupancy, the area, extent, and/or quality of habitat, the number of locations, and the number of mature individuals, and D. schatzii is assessed as “Critically Endangered” [EN B1ab(i,ii,iii,iv,v)+2ab(i,ii,iii,iv,v)].

Notes. – Danais schatzii is characterized by opposite, petiolate, well-developed leaves with the venation closely reticulated; interpetiolar triangular stipules; axillary and terminal, cymose, pedunculate, well-developed inflorescences with shortly pedicellate flowers; medium-sized, salverform corollas with a green tube and orange lobes; and rather small capsules. The color scheme of the corollas is the common one in Danais flowers. Several immature capsules and a few mature ones that are fully dehisced are preserved on the paratype specimen, but no fully developed seeds remain.

Danais schatzii is similar to D. longipedunculata (Fig. 3B), which is found in the same general region and can be separated by its stipules that are shortly united around the stem and its shorter corollas, with the tube 5.5–6 mm long. Danais schatzii is also similar to D. baronii; see the discussion of that species above for their distinctions.

Additional specimen examined. – Madagascar. Reg. Analanjirofo [Prov. Toamasina]: Distr. Mananara-Nord, Ivontaka-Sud, fort à 100 m de la mer, 16°20′S 49°48′E, 250 m, 10–11.XI.1989, Dumetz 866 (MO, P, TAN).

Lectotypus (designated here): Madagascar. Reg. Analanjirofo [Prov. Toamasina]: Distr. Sainte-Marie [Nosy Boraha], 1848, Boivin 1773 (P [P00057196] image!; isolecto-: P [P00057198] image!, W [W 1889-0147599] image!). Syntypus: ibid., 1849, Boivin 1773 (G [G00014460]!, TCD [TCD0017832] image!).

Distribution and ecology. – This species is found in lowland moist evergreen forest from sea level to 245 m in eastern Madagascar (Reg. Analanjirofo [Prov. Toamasina], Reg. Vatovavy [Prov. Fianarantsoa]), on Nosy Boraha and the coastal mainland.

Notes. – Danais terminalis was described from incomplete, fruiting material, and is not clearly diagnosed within the genus although it does not seem to match any other species. Danais terminalis is characterized by a combination of common characters in Danais: opposite, shortly petiolate leaves with rather small elliptic to oblanceolate blades, 4–6.5 × 1–2.8 cm, and laxly reticulated venation that is plane or only irregularly thickened; triangular interpetiolar stipules c. 1 mm long; few-to several-flowered, terminal and axillary inflorescences with lax peduncles 20–38 mm long and flexuous pedicels 5–20 mm long; and relatively large fruits and seeds, 8–12 mm in diam. and 2.5–4 mm in diam., respectively. Beyond this, the identity of D. terminalis is somewhat problematic, and was not fully clarified by Puff & Buchner (1994: 48). They distinguished D. terminalis by the combination of its relatively larger fruits and “characteristic inflorescence”, which they described as terminal but occasionally also axillary, 1- to 7-flowered, lax, and with a form they considered reduced for the genus (Buchner & Puff, 1993) but did not detail further. Buchner & Puff (1993: 33–36) interpreted the inflorescences of both D. terminalis and D. nigra as reduced thyrses (here interpreted as cymes) with partial inflorescence sections comprising 1 or 3 flowers and articulations where axes have been reduced, vs. 5 to 15 or more flowers in the unarticulated inflorescences they considered fully developed. Their “reduced” inflorescence form is found in D. terminalis, but also now documented in a number of other, similar Danais species, including some treated by Puff & Buchner without mention of the inflorescence arrangement (e.g. D. dauphinensis, D. coronata (Pers.) Steud.). Danais terminalis is indeed characterized in part by its lax and apparently reduced inflorescence structure, but this is not diagnostic alone for any Danais species. And, the relatively large fruits and seeds of D. terminalis seem to help identify it, but several of the similar species are known only in flower. Thus, as in the case of D. baronii, the identity of D. terminalis is problematic for the taxonomy of Danais because some other species share its few known features but are also poorly known and cannot be conclusively separated or synonymized with it. Danais terminalis is treated here based on the available information, so it eventually can be matched with additional specimens and its identity clarified. The four specimens included in this species by Puff & Buchner are from separated sites across a rather wide range, so it is possible that they represent more than one species.

Danais terminalis is similar in particular to D. gracilis, described above; see that species for their distinctions.

The holotype of Danais terminalis was cited by Puff & Buchner (1994: 48) as deposited at P, where Boivin and Drake worked, with no additional information to identify a particular specimen. There are two duplicates of the type collection curated there, both of them complete specimens and none labelled as part of a set. No primary specimen was designated in the protologue or denoted in annotations by Drake, so these must be considered syntypes. One of the specimens has a hand-written label with collection data and the name “Danais ? terminalis Boiv”, mature fruits and seeds, and the date 1848. This was annotated by Puff as the “holotype”, and it is the only duplicate with mature fruits and is here chosen as the lectotype. Another sheet at P has a date range, 1847–1852, that includes the collection date on the lectotype and is here considered a duplicate of the type collection. Other specimens at G, P, and TCD labelled as Boivin 1773 have a collection date of 1849. They belong to a different gathering and are here regarded as syntypes.