taxonID	type	description	language	source
03E98786FFB82410FF64FE7FFE96326A.taxon	description	Backer (1940) conducted a further investigation into Gynura in Java. At herbarium U, he located a specimen collected by Zollinger, without any number or precise locality (except for Java) but named by Miquel himself as G. densiflora. He deduced that it was most probably Zollinger H. 2592 (as “ Z. M. 2592 ”; see below for the meaning of “ Z. M. ”) and concluded that G. densiflora was conspecific with G. ajakensis. He thus synonymized G. ajakensis with G. densiflora. In addition, he suspected that the record of Bandung for the type locality of G. densiflora was probably an error, because the species was only known from East Java, and given that Bandung was a well explored region, it was hard to believe that a very conspicuous plant had been overlooked by all later collectors. Regarding G. mollis, Backer (1940) also agreed that it would be conspecific with G. densiflora, because they were both based on Zollinger H. 2592. However, he adopted G. densiflora over G. mollis with a preference for certainty, despite the latter name having priority. Backer’s (1940) treatment, however, was not accepted by two major worldwide revisions of Gynura (Davies 1980; Vanijajiva & Kadereit 2011). Both Davies (1980) and Vanijajiva & Kadereit (2011) treated G. densiflora and G. mollis as synonyms of G. aurantiaca and further considered G. densiflora to be a superfluous name of G. aurantiaca. Davies (1980) indicated that the holotype of G. densiflora was kept at G, but stated that he had not seen the type of G. mollis. Vanijajiva & Kadereit (2011), however, treated G. densiflora and G. mollis as homotypic names, citing their holotype at G. At G, we located three specimens of Zollinger H. 2592 (G 00604939, G 00630048 (with two sheets), and G 00630050; Fig. 2). One of these (G 00604939; Fig. 2 A) bears annotations from both Hochreutiner and Davies (Fig. 2 B). This specimen is likely the type indicated by Hochreutiner (1934) and Davies (1980) for G. densiflora, and it may also be the type suggested by Vanijajiva & Kadereit (2011) for G. mollis. However, neither Schultz nor Miquel appears to have seen these specimens, because no annotations have been made by them on these sheets. At U, we saw the specimen (U 0001285; Fig. 3 A) that Backer (1940) suggested as the type material of Gynura densiflora. It indeed bears Miquel’s determination and is a potential original material for G. densiflora. However, it does not completely match the original description and type information of G. densiflora. First, G. densiflora is described as having shortly petiolate lower leaves, but U 0001285 only contains three sessile leaves. Second, Miquel cited the type locality as Bandung, but no precise locality has been given on U 0001285, except for Java (Fig. 3 B). Third, U 0001285 does not bear any collection number. The annotation of “ probably Z. M. 2592 ” on the sheet (Fig. 3 B) was actually made by Backer, as indicated in his paper (Backer 1940), with “ Z. M. ” standing for Zollinger and Moritzi. Moritzi was a close friend of Zollinger and, before his death, responsible for the summary determination and distribution of the specimens collected by Zollinger from Java (Moritzi 1845 – 1846; Briquet 1940). Therefore, there should be some other materials used by Miquel in describing G. densiflora. We know that, during the preparation of Flora van Nederlandsch Indie, Miquel had access to Zollinger’s private herbarium via a loan from Comte de Franqueville (Briquet 1940; Steenis 1950; Stafleu 1970), who bought Zollinger’s herbarium after his death. The herbarium then went to Emmanuel Drake del Castillo (the Drake herbarium) and ultimately incorporated into P (Stafleu 1970). At P, we sorted out two specimens (P 02562905 and P 02562907; Fig. 4) of Zollinger H. 2592 from the Drake herbarium (with the annotation of “ HERBIER E. DRAKE ”). The P 02562907 sheet (Fig. 4 A) is indicated to be collected from Bandung and bears the annotations of “ Gynura aurantiaca (non DC.) Schultz ” and “ Gynura densiflora Miq. ” in Miquel’s hand (Fig. 4 B), perfectly matching the information provided in the protologue of G. densiflora (“ Gynura aurantiaca (haud DC.) Schultz Bip. in Herb. Zoll. ad n. 2592 ”). The morphology also agrees with the original description of G. densiflora, especially in having shortly petiolate lower leaves. As for P 02562905 (Fig. 4 C, D), it is in significant conflict with P 02562907 (Fig. 4 A, B) and the protologue of G. densiflora in both morphology and collection locality. It has lower leaves three to five times larger than those of P 02562907 and is indicated to be collected from Mt. Ardjuno, which is located in East Java and far from Bandung. There is also no annotation made by Miquel on P 02562905 (Fig. 4 C). Therefore, Miquel probably did not see P 02562905; otherwise, he would not have overlooked such obvious incongruences. Since Zollinger H. 2592 is the only gathering cited in the protologue of G. densiflora, and P 02562907 (Fig. 4 A) is the only specimen of this gathering seen by Miquel, even though the aforementioned U 0001285 sheet (Fig. 3) is a potential original material, P 02562907 (Fig. 4 A) should be recognized as the holotype of G. densiflora, and it is confirmed to be conspecific with G. ajakensis. Apart from Gynura densiflora, we also located the original material of G. mollis at P. We know that Schultz’s herbarium was sold to Ernest Saint-Charles Cosson after his death, and later presented to P by Cosson’s grandson (Bonifacino et al. 2009). At P, we sorted out three sheets (P 02562892, P 02562893, and P 02562909; Figs. 5, 6) of Zollinger H. 2592 from Schultz’s herbarium (annotated with “ Herb. Schultz Bip. ” and “ HERB. E. COSSON ”). However, as with G. densiflora, these three specimens belong to two species. P 02562892 (Fig. 5 A) and P 02562893 (Fig. 5 C) are similar in morphology and match the original description of G. mollis. In addition, P 02562893 bears Schultz’s determination of “ Gynura mollis ” (Fig. 5 B) and P 02562892 is annotated with the localities of Mt. Tankuban and Mt. Ardjuno (Fig. 5 D). Together, they perfectly align with the information provided in the protologue of G. mollis. In contrast, P 02562909 (Fig. 6) has sessile lower leaves semiamplexicaul at the base, conflicting with the petiolate leaves described for G. mollis (Miquel 1856) and possessed by both P 02562892 (Fig. 5 A) and P 02562893 (Fig. 5 C). Although P 02562909 also bears a determination of “ Gynura mollis Schultz ” (Fig. 6 B), it appears to have been written by Zollinger rather than Schultz. Instead, under this determination and on a separate slip (Fig. 6 B), there is the same determination of “ Gynura aurantiaca DC. ” clearly in Schultz’s hand. Therefore, Schultz should have only used P 02562892 (Fig. 5 A) and P 02562893 (Fig. 5 C) to describe G. mollis, while P 02562909 (Fig. 6 A) was probably distributed independently, after Zollinger received the determination of G. mollis by Schultz, who was one of the experts helping identify Zollinger’s collections. Based on P 02562892 (Fig. 5 A) and P 02562893 (Fig. 5 C), G. mollis is actually conspecific with G. densiflora and G. ajakensis, and, with the priority, G. mollis turns out to be the correct name for this species.	en	Zhu, Li-Cui, Hariri, Muhammad Rifqi, Li, Cheng-Sheng, Ren, Chen (2025): Clarification of the taxonomic and nomenclatural confusions with three Gynura names (Asteraceae: Senecioneae) based on Zollinger’s collections from Java, Indonesia. Phytotaxa 697 (3): 219-233, DOI: 10.11646/phytotaxa.697.3.1, URL: https://doi.org/10.11646/phytotaxa.697.3.1
03E98786FFB1241CFF64FE38FC32311B.taxon	description	We have been able to sorted out six specimens of Zollinger HZ. 442, viz., BR 0000029078073 (Fig. 9 A), P 02562892 (Fig. 5 A), P 02562893 (Fig. 5 C), P 02562894 (Fig. 9 C), P 02562896 (Fig. 10 A), and P 02562909 (Fig. 6 A). Among these, P 02562892 (Fig. 5 A), P 02562893 (Fig. 5 C), and P 02562909 (Fig. 6 A), which are further numbered as H. 2592, have been discussed above. They are from Schultz’s herbarium but not the type material of Gynura lyrata. Instead, P 02562892 (Fig. 5 A) and P 02562893 (Fig. 5 C) are the syntypes of G. mollis. P 02562909 (Fig. 6 A) was likely distributed only after Zollinger received Schultz’s determination on G. mollis, and probably not used for describing G. lyrata either, because the two species were published simultaneously. Moreover, P 02562909 (Fig. 6 A) is in conflict with the original description of G. lyrata, with elliptic and undivided rather than lyrate leaves as described by Schultz (in Zollinger 1854) for G. lyrata. The other two specimens, BR 0000029078073 (Fig. 9 A) and P 02562894 (Fig. 9 C), are not from the Schultz Herbarium and probably not seen by Schultz. In fact, they represent the additional cases of cross-labelling between HZ. 442 and H. 2592. Both specimens are G. mollis and should belong to H. 2592 but are mislabeled as HZ. 442. The remaining specimen, P 02562896 (Fig. 10 A), matches the protologue of G. mollis perfectly. It is from Schultz’s herbarium (annotated with “ Herb. Schultz Bip. ” and “ HERB. E. COSSON ”; see above) and bears the determination of “ Gynura lyrata ” in Schultz’s hand (Fig. 10 B). This specimen should be the holotype of G. lyrata. Morphologically, it well conforms with G. auriculata and is accepted as a synonym of the latter.	en	Zhu, Li-Cui, Hariri, Muhammad Rifqi, Li, Cheng-Sheng, Ren, Chen (2025): Clarification of the taxonomic and nomenclatural confusions with three Gynura names (Asteraceae: Senecioneae) based on Zollinger’s collections from Java, Indonesia. Phytotaxa 697 (3): 219-233, DOI: 10.11646/phytotaxa.697.3.1, URL: https://doi.org/10.11646/phytotaxa.697.3.1
03E98786FFB1241DFF64F886FF3C3077.taxon	materials_examined	Type: — INDONESIA. West Java, Mt. Gede, 1 June, Blume 521 [lectotype designated by Davies (1980: 730): L 0001986!; isolectotype: L 0001987!].	en	Zhu, Li-Cui, Hariri, Muhammad Rifqi, Li, Cheng-Sheng, Ren, Chen (2025): Clarification of the taxonomic and nomenclatural confusions with three Gynura names (Asteraceae: Senecioneae) based on Zollinger’s collections from Java, Indonesia. Phytotaxa 697 (3): 219-233, DOI: 10.11646/phytotaxa.697.3.1, URL: https://doi.org/10.11646/phytotaxa.697.3.1
03E98786FFB1241DFF64F886FF3C3077.taxon	materials_examined	Type: — INDONESIA. West Java, Mt. Tankuban, March 1944, Zollinger HZ. 442 [holotype: P 02562896!; isotype: P 02562905! (misnumbered as H. 2592)] (Figs. 4 C, 10 A). Notes: — Gynura aurantiaca was originally described by Blume (1826) based on his collections from Mt. Gede and Mt. Burangrang. Davies (1980) designated Blume 521 (mistakenly as “ 520 ”) at L as the lectotype (incorrectly as “ holotype ”, correctable under Art. 9.10 of the International Code of Nomenclature for algae, fungi, and plants; Turland et al. 2018). We located two sheets (L 0001986 and L 0001987) of Blume 521 at L. Both specimens bear Davies’ determinations, but only L 0001986 is further annotated by Davies with “ Holotype ” and should be accepted as the effective lectotype. However, it should be noted that although both are numbered as “ 521 ”, L 0001986 and L 0001987 are indicated with different localities: Mt. Gede and Mt. Burangrang, respectively. They may represent separate gatherings but were tentatively treated as one here, pending further evidence. The issue of cross-labelling among Zollinger HZ. 442, HZ. 606, and H. 2592 has been discussed above. Based on the available evidence, we reassign the mislabeled specimens to their correct gatherings. For Zollinger HZ. 442, only P 02562896 (Fig. 10 A) and P 02562905 (Fig. 4 C) are assigned to this gathering. P 02562896 (Fig. 10 A) is the holotype of G. lyrata. P 02562905 (Fig. 4 C) is numbered H. 2592, but has same morphology as P 02562896 (Fig. 10 A). It is highly probably a duplicate of HZ. 442 and recognized here as an isotype. Among the other specimens labeled “ HZ. 442 ”, BR 0000029078073 (Fig. 9 A), P 02562892 (Fig. 5 A), P 02562893 (Fig. 5 C), and P 02562894 (Fig. 9 C) are G. mollis and assigned to the gathering of Zollinger H. 2592, while P 02562909 (Fig. 6 A) represents another species, G. daviesiae. It is noteworthy that Schultz actually noticed the mixture of Zollinger HZ. 442 with H. 2592. When describing G. lyrata, he (in Zollinger 1854) explicitly cited the type as “ Mixta cum No. 3 sub No. 442 HZ ”, restricting the type to the “ No. 3 ” specimen of Zollinger HZ. 442. In addition, P 02562892 (Fig. 5 A) and P 02562893 (Fig. 5 C), which are the types of G. mollis, were annotated with both HZ. 442 and H. 2592. However, Schultz exclusively assigned them to H. 2592, by only citing this number in the protologue of G. mollis. In fact, Schultz also left tangible evidence on the specimens. He made a “ wrong ” mark on the number of “ HZ. 442 ” on P 02562893 (Fig. 5 D) and specifically annotated the single individual on P 02562892 as “ 2592 ” (Fig. 5 B), aligning with the citation in the protologue of G. mollis. However, Schultz’s citation of “ No. 3 ” appears to be a mistake. The “ No. 3 ” should refer to the notation immediately following the collection number “ 442 ” on P 02562896 (Fig. 10 B). While it resembles the number “ 3 ”, it is actually the letter “ Z ”, indicating that the specimen belonged to Zollinger’s private herbarium (Moritzi 1845 – 1846; Zollinger 1854).	en	Zhu, Li-Cui, Hariri, Muhammad Rifqi, Li, Cheng-Sheng, Ren, Chen (2025): Clarification of the taxonomic and nomenclatural confusions with three Gynura names (Asteraceae: Senecioneae) based on Zollinger’s collections from Java, Indonesia. Phytotaxa 697 (3): 219-233, DOI: 10.11646/phytotaxa.697.3.1, URL: https://doi.org/10.11646/phytotaxa.697.3.1
03E98786FFB02419FF64FB25FD0F379E.taxon	materials_examined	Type: — INDONESIA. East Java, Mt. Ardjuno, 1200 – 1800 m, September 1944, Zollinger H. 2592 (lectotype designated here: P 02562893!; isolectotypes: BR 0000029078073!, G 00604939!, G 00630048!, G 00630050!, P 02562892!, and P 02562894!) (Figs. 2, 5, 9).	en	Zhu, Li-Cui, Hariri, Muhammad Rifqi, Li, Cheng-Sheng, Ren, Chen (2025): Clarification of the taxonomic and nomenclatural confusions with three Gynura names (Asteraceae: Senecioneae) based on Zollinger’s collections from Java, Indonesia. Phytotaxa 697 (3): 219-233, DOI: 10.11646/phytotaxa.697.3.1, URL: https://doi.org/10.11646/phytotaxa.697.3.1
03E98786FFB02419FF64FB25FD0F379E.taxon	materials_examined	Type: — INDONESIA. East Java, Mt. Tengger, 2300 m, November 1944, Zollinger HZ. 606 (lectotype here designated: P 02562906!; isolectotypes: BO 1281963!, P 02562907!, and P 04389013!) (Figs. 4 A, 7, 8).	en	Zhu, Li-Cui, Hariri, Muhammad Rifqi, Li, Cheng-Sheng, Ren, Chen (2025): Clarification of the taxonomic and nomenclatural confusions with three Gynura names (Asteraceae: Senecioneae) based on Zollinger’s collections from Java, Indonesia. Phytotaxa 697 (3): 219-233, DOI: 10.11646/phytotaxa.697.3.1, URL: https://doi.org/10.11646/phytotaxa.697.3.1
03E98786FFB02419FF64FB25FD0F379E.taxon	materials_examined	Type: — INDONESIA. East Java, Mt. Tengger, 2300 m, November 1944, Zollinger HZ. 606 [cited as “ H. 2592 ” by Miquel (1856); see below] (holotype: P 02562907!; isotypes: BO 1281963!, P 02562906!, and P 04389013!) (Figs. 4 A, 7, 8).	en	Zhu, Li-Cui, Hariri, Muhammad Rifqi, Li, Cheng-Sheng, Ren, Chen (2025): Clarification of the taxonomic and nomenclatural confusions with three Gynura names (Asteraceae: Senecioneae) based on Zollinger’s collections from Java, Indonesia. Phytotaxa 697 (3): 219-233, DOI: 10.11646/phytotaxa.697.3.1, URL: https://doi.org/10.11646/phytotaxa.697.3.1
03E98786FFB02419FF64FB25FD0F379E.taxon	materials_examined	Type: — INDONESIA. East Java, Mt. Tengger, Ajak-ajak, 2600 m, 20 January, 1905, Hochreutiner 2736 (holotype: G 00035628!) (Fig. 1). Notes: — We suggest that the gathering Zollinger H. 2592 should include BR 0000029078073 (Fig. 9 A), G 00604939 (Fig. 2 A), G 00630048 (Fig. 2 C), G 00630050 (Fig. 2 D), P 02562892 (Fig. 5 A), P 02562893 (Fig. 5 C), and P 02562894 (Fig. 9 C), while Zollinger HZ. 606 should contain BO 1281963 (Fig. 7 A), P 02562906 (Fig. 7 C), P 02562907 (Fig. 4 A), and P 04389013 (Fig. 8 A). In particular, P 02562907 (Fig. 4 A) needs some notes. It is the holotype of Gynura densiflora and cited as H. 2592 by Miquel in the protologue. However, the specimen is actually annotated with both HZ. 606 and H. 2592 (Fig. 4 B). Despite both gatherings undoubtedly representing the same species, P 02562907, with coarser leaves more densely arranged along the stem and a more robust synflorescence, aligns more closely with the specimens of Zollinger HZ. 606 (BO 1281963, P 04389013, and P 02562906; Figs. 7, 8). Consequently, we consider it better to assign P 02562907 (Fig. 4 A) to Zollinger HZ. 606. P 02562892 (Fig. 5 A) and P 02562893 (Fig. 5 C) are the two specimens Schultz relied on to describe G. mollis. Among them, P 02562893 (Fig. 5 C) is better preserved and bears Schultz’s handwritten determination, and is selected as the lectotype of G. mollis. Gynura aurantiaca var. robusta is also based on Zollinger HZ. 606 (Zollinger 1845). Among the four sheets of Zollinger HZ. 606, P 02562906 (Fig. 7 C) and BO 1281963 (Fig. 7 A) were annotated on the determination slips as a potential variety of G. aurantiaca, although no varietal epithet was specified for P 02562906 (Fig. 7 D) and a different epithet “ rugosa ” given for BO 1281963 (Fig. 7 B). These should be the basis of var. robusta. P 02562906 (Fig. 7 C) is here chosen as the lectotype because it is better preserved with an intact synflorescence. Davies (1980) and Vanijajiva & Kadereit (2011) regarded G. densiflora as a superfluous name for G. aurantiaca. This is probably due to a misinterpretation of Miquel’s (1856) citation in the protologue of G. densiflora, where Miquel (1856) cited “ Gynura aurantiaca (haud DC.) Schultz Bip. in Herb. Zoll. ad n. 2592 ” following the morphological description. Davies (1980) may have assumed that Miquel cited legitimate G. aurantiaca as a synonym, thereby rendering G. densiflora superfluous. However, Miquel (1856) actually intended to indicate a misidentification of Zollinger H. 2592. Therefore, Gynura densiflora is a legitimate name. Backer (1940) once considered that Hochreutiner (1934), when describing G. ajakensis, mistakenly compared it to G. densiflora by erroneously adopting the original description of G. densifolia Miquel (1856: 99) as that of G. densiflora. Gynura densifolia is a completely different species, but simultaneously described with G. densiflora with a confusingly similar epithet, and it is listed immediately after G. densiflora. We, however, consider that this confusion is actually caused by two typographical errors in Hochreutiner’s (1934) text that mistook G. densifolia as G. densiflora (lines 25 and 30 on page 331). In the first paragraph of his discussion, Hochreutiner (1934: 331) clearly stated that he saw the type of G. densiflora (“ Nous avons a Geneve, le type du G. densiflora …… ”; Hochreutiner 1934: 331), and as a matter of fact, he annotated one of the sheets (G 00604939; Fig. 2 B). In addition, he compared G. densiflora with G. ajakensis in the second paragraph. However, in the third paragraph, he paradoxically reclaimed not to have seen G. densiflora (“ Nous n'avons pas vu le G. densiflora …… ”; Hochreutiner 1934: 331) and re-compared G. densiflora with G. ajakensis. This inconsistency is difficult to reconcile without attributing it to typographical errors. Notably, Hochreutiner likely never examined the type of G. densifolia, which was absent from G, further supporting our hypothesis about the errors. Additional evidence comes from Hochreutiner’s (1934) reference to Koorders, who noted that the species was also found by Kuntze in Guedeh near Sindaglaja. This actually referred to the gathering Kuntze 4462. It was described by Kuntze (1891) as Crassocephalum densifolium var. atropurpureum Kuntze, but later synonymized by Koorders (1912) just with G. densifolia. Therefore, Hochreutiner (1934) actually intended to compare G. ajakensis with G. densifolia in the third paragraph of the discussion. Unfortunately, the typographical errors mistook G. densifolia as G. densiflora and led to the confusion. Despite Backer’s (1940) misinterpretation of Hochreutiner’s (1934) discussion, we agree with Backer’s (1940) conclusion that G. ajakensis and G. densiflora are conspecific. Hochreutiner’s (1934) considered G. ajakensis different from G. densiflora in having denser synflorescences and different types of leaves and indument. However. we do not observe significant differences in leaf shape or indument. While the synflorescence on the type of G. ajakensis is indeed denser compared to those on the specimens of Zollinger H. 2592 kept at G (Fig. 2), which Hochreutiner (1934) regarded as the “ type ” of G. densifolia, these variations fall well within the normal range of this species. The capitula are generally densely arranged, especially in comparison to G. aurantiaca, but they also exhibit some variability, with peduncles elongating to a certain extent. Gynura mollis is clearly most similar to G. aurantiaca. Their differences have been well summarized by Backer (1940). Gynura aurantiaca is a climbing or creeping plant characterized by purple hairs on young parts. The leaves are largely lyrate and often have conspicuous auricles at the base, and the capitula are generally loosely arranged. In contrast, G. mollis is an erect plant with white hairs on young parts, undivided leaves lacking auricles at the base, and generally dense synflorescences. In addition, G. mollis is restricted to East Java, whereas G. aurantiaca is a common species in Sumatra, West and Central Java, only slightly overlapping the distribution range of G. mollis in the westernmost part of East Java (Mts. Lawu and Wilis).	en	Zhu, Li-Cui, Hariri, Muhammad Rifqi, Li, Cheng-Sheng, Ren, Chen (2025): Clarification of the taxonomic and nomenclatural confusions with three Gynura names (Asteraceae: Senecioneae) based on Zollinger’s collections from Java, Indonesia. Phytotaxa 697 (3): 219-233, DOI: 10.11646/phytotaxa.697.3.1, URL: https://doi.org/10.11646/phytotaxa.697.3.1
