Schismatoglottis calyptrata (Roxb.) Zoll. & Moritzi

This species is distinguished from others in this group by its hapaxanthic, hypogeal stems and stoloniferous, often colony-forming terrestrial habit, the clustered inflorescences on rather short peduncles which elongate and become declinate in fruit, the strongly hourglass-shaped spadix with a poorly defined interstice positioned level with the top of the lower spathe chamber, the scattered (sometimes very few, occasionally absent) interpistillar staminodes about equalling to more usually somewhat exceeding the pistils, the dome-shaped appendix slightly and abruptly ider than the top of the male zone. Typically, local ‘populations’ are strikingly uniform, and frequently seem rather sharply differentiated from adjacent ones. This may well be a result of the colonial habit of this species enabling the formation of very extensive, long-lived clonal ‘superindividuals’.
Overall, this species is extremely variable in leaf shape and size and in inflorescence size - nowhere more so than in New Guinea (extreme dimensions in parentheses in the description), where it is very common and the only species of this genus represented. There, the most diminutive forms have been called S. klossii, S. dorensis,  S. hellwigiana, S. hellwigiana var. subcordata and S. parvifolia. Though all those are described from Irian Jaya, similarly small plants are recorded from localities scattered throughout New Guinea at a wide variety of altitudes. In these the leaf is elliptic with an obtuse to very slightly cordate base, and as little as 5.5 cm long. Robust forms with dramatically variegated, spattered leaves have been called S. neoguineensis (a form extending to the Bismarck Archipelago, Solomon Islands and Vanuatu). The most robust forms have ovato-sagittate leaves as much as 40 cm long. The very small forms tend to have reduced numbers of interpistillar staminodes which often do not exceed the pistils in height, unlike in the more robust forms. There is a complete set of intermediates including the types of all other names applied to Schismatoglottis in New Guinea, and no evidence has been found from amongst the herbarium material examined that there is ecological differentiation (e.g. to substrate or vegetation type) of any of them. Whether there is biological differentiation by pollinators remains to be discovered. Schismatoglottis calyptrata is common and variable, but not to the same extent, throughout most of the rest of Malesia. However, this species is surprisingly uncommon in Borneo, though there are a number of geographically and ecologically (where known) restricted segregate species there, such as Schismatoglottis ahmadii, S. lingua, S. venusta and S. viridissima.
A few specimens occur in the Malay Peninsula which are intermediate between this species and S. wallichii (e.g. Hay et al. 9159). In the latter species, the sterile and male zones of the spadix are in reverse order to that in S. calyptrata, so that there is no appendix (though there may be a very small terminal cluster of staminodes at the spadix apex). In the intermediate specimens, the male zone is longer than in S. calyptrata s. str. and is separated from the female by a narrow zone of staminodes of the type found in the appendix of S. calyptrata and the sterile zone in S. wallichii s. str.
The appendix is much shorter, and the distal portion of the spadix is rather narrowly clavate: i.e. intermediate between the more or less cylidrical spadix of S. wallichii and the abruptly clavate one of S. calyptrata and without the slight but abrupt difference in width between the male zone and appendix of the latter species. Occasional specimens are also found in which the structure of the spadix is typical for S. calyptrata except for a very short sterile zone between the male and female zones; these have been determined as S. calyptrata, but if hybridization has taken place with S. wallichii, they may suggest that some introgression is occurring. Cytological investigation and experimental crossing are required to confirm whether or not these species are genetically isolated. The descriptions of both S. calyptrata and S. wallichii presented here are based on material judged to be broadly ‘typical’. Calla calyptrata Roxb. was based on material cultivated at Calcutta and obtained from Ambon. It was lectotypified by Hay (1996), where it was noted that an epitype was required, as both the elements from which the lectotype could be selected are somewhat ambiguous. The designated specimen is the only complete material seen from Ambon and it conforms with the sense in which the name Schismatoglottis calyptrata is widely used.
Hasskarl’s publications in Flora (1842a) and Tijdschr. Nat. Gesch. & Physiol. (1842b), cited above, appear to have been published in reverse order to that in which they seem to have been written. C. humilis var. major, designated variety a, appears in Hasskarl (1842 b; Aug/Sept), but not in Hasskarl (1842a; July), where only var. minor appears, designated variety ß. It is evident that Hasskarl considered var. major the ‘type’ form from which var. minor was segregated, and not another variety in addition to an implicit var. humilis. We conclude therefore that var. major is a superfluous name for the nomenclaturally typical variety. That, in turn, is a superfluous name for Calla calyptrata, as they are both based on the above-cited plate in Rumphius’ Herbarium Amboinense.
Although C. humilis Hassk. is a synonym of Schismatoglottis calyptrata, it cannot be deemed completely clear if var. minor is actually Schismatoglottis too. No element was cited that could serve as type, the description is very sparse and it is not indicated from where the plant is supposed to have originated. No herbarium material bearing this name has been located at Leiden. Hasskarl (1842a & b), in discussing the varieties of Colocasia humilis, was clear about the specific character of the clavate appendix which does not accord with Colocasia in the current sense at all. It does however, accord with Schismatoglottis calyptrata, though also with other Schismatoglottis species (though not in Java). Hasskarl (1842a) also mentions capitate interpistillar staminodes as a character of C. humilis var. minor — another characteristic in accord with S. calyptrata, particularly when combined with the additional specific character of being stoloniferous (cf. Hay 1996). It therefore seems highly likely, though not certain, that Colocasia humilis var. minor is Schismatoglottis calyptrata, and we have found nothing to suggest it might be anything else. It is not a name in use. Besides the issue of its identity, Colocasia humilis var. minor Hassk., though erected under an illegitimate species name, is valid (Tokyo Code Art. 55.2). A new type is therefore needed. Smith 930 is designated neotype here; it conforms to Hasskarl’s description in its small size and somewhat hastate, rather narrowly lanceolate and acuminate leaf blade (though it is not variegated).
No material was cited in the protologue of Colocasia neoguineensis and so the accompanying plate is selected as lectoype. It is not detailed however, and in interpretive epitype is required. Material studied and preserved by Brown at Kew in 1885 when he made the combination in Schismatoglottis, which was obtained from Linden and matches the description of C. neoguineensis, is therefore designated as the epitype.
Schismatoglottis calyptrata var. albidomaculata was first published by Hallier (loc. cit.) as a nomen nudum. A year later he reduced it to the first of two validly published forms of a new variety maculata. The second, f. flavidomaculata, included Schismatoglottis neoguineensis. One of these forms has to be regarded as the nomenclaturally typical (and hence superfluous) one, and the first is taken here. Ridley, apparently unaware of Hallier’s 1898 reduction of var. albidomaculata to a forma in var. maculata, validated S. calyptrata var. albidomaculata, citing a specimen of his own. Schismatoglottis calyptrata var. albidomaculata Ridl. could be regarded as an altogether new, valid and legitimate variety. However, since Ridley clearly cites Hallier as the originating authority, the name can be regarded as a superfluous renaming of S. calyptrata var. maculata Hallier f. Engler (1912) attributed Schismatoglottis calyptrata var. bivittata to Hallier, and considered his own S. picta f. bivittata to be based on it. However, the former was not valid; there being no description, and is thus not the basionym. Schismatoglottis picta f. bivittata is therefore typified with the material cited by Engler and not with material linked to Hallier’s trinomial.
Engler cited two syntypes of S. emarginata in the protologue, Nieuwenhuis 393 (BO) from Bochang and Amdjah 420 (BO) - Borneo, without specific locality. Alderwerelt (loc. cit.) pointed out that the protologue and the illustration do not agree with the specimens Engler cited. Both specimens are sterile (Amdjah 420 with a fruiting spathe), while the description and illustration include flowering parts. Both the description and illustration are of a stoloniferous plant with a central band of variegation along the midrib. The cited specimens do not show these features either. Engler (loc. cit.) indicated that he had seen cultivated material in addition to the cited specimens, though he did not specify what, and apparently none was preserved and annotated with the name at the time. Years later, Alderwerelt found at Bogor two living plants cultivated under this name, one of which (XI.B.X.15) matched the illustration and original description of S. emarginata. On the basis of this living plant and its resemblance to the protologue, he provided a more exhaustive description (preserving a specimen in the Gardens Herbarium here designated the epitype). He also indicated that Nieuwenhuis 393 & Amdjah 420, though inadequate for certain identification, were probably referable to S. irrorata Engl. (= S. motleyana Schott s. lat.). From this it is concluded that he effectively lectotypified S. emarginata with Engler’s illustration.
Unfortunately, the plant on which the illustration was allegedly based is of unknown original provenance (even as to island). Schismatoglottis emarginata belongs in the group around the widespread S. calyptrata which includes a number of narrow segregates whose definition sometimes relies on geographic location and substrate preference to support slight morphological differences. Schismatoglottis emarginata differs only slightly from typical S. calyptrata (in the rather somewhat more numerous primary lateral veins relative to the size of the leaf blade — cf. the Sulawesi elements S. bitaeniata, S. sublaxiflora and S. vanvuurenii here also placed in synonymy with  S. calyptrata). It is at present impossible to evaluate as no exactly matching material exists of known provenance. It is therefore placed provisionally in the synonymy of S. calyptrata.
The holotype of Schismatoglottis acutangula, a species described from sterile material cultivated at Bogor and collected from there by Engler in 1906, is not at B. It presumably was destroyed during the war. Alderwerelt (loc. cit.) expressed some doubt about whether the plant he subsequently described in full under this name was the same cultivated plant that Engler had seen. He appears to have been right to doubt it, as Alderwerelt’s plant was cited as originally collected from Mt Alkmaar by Djibdja, who accompanied Versteeg there in 1907 (van Steenis-Kruseman, 1950), the year after Engler had visited Bogor. That material is therefore not used as a neotype. The specimen so-designated is one determined with this name by Engler, made from a plant growing at the Berlin Botanic Garden in 1911. It is not clear whether this plant was obtained from Bogor during Engler’s visit. However, it matches the protologue quite closely.
The type of Schismatoglottis bitaeniata, a Sulawesi element, does not seem to be distinguishable from S. calyptrata s. lat. in inflorescence, though the leaf is somewhat distinctive, combining a cuspidate tip, finely crispate margin, rather reduced posterior lobes and unusually closely parallel primary lateral veins. Bornean Schismatoglottis ahmadii and other species are also extremely similar to S. calyptrata in inflorescence but have rather minor differences in leaf and are recognised here as segregate species.
However, in those instances there is evidence of ecological differentiation. This may, of course, transpire to be the case with S. bitaeniata, in which case it would reasonably be pulled out of the synonymy of S. calyptrata, but there are no ecological data available at present. This particular form is quite widely cultivated (usually as Schismatoglottis picta, under which name it is illustrated in Graf (1978: 320)). Schismatoglottis sublaxiflora and S. vanvuurenii, also from Sulawesi and provisionally regarded as synonyms of S. calyptrata, are elements rather similar to S. bitaeniata.
The identity of Schismatoglottis nieuwenhuisii appears to have been somewhat confused. It was discussed by Alderwerelt (loc. cit.) who noted that of the two specimens cited by Engler in the protologue, one, Nieuwenhuis 1485, collected by Jaheri (who was cited as the collecting locality) and grown at Bogor, was a Homalomena. The second was a Schismatoglottis preserved directly from the wild (Winkler 2129).
Alderwerelt was also of the opinion that Engler had used, though did not cite, other material cultivated at Bogor to form the basis of his diagnosis of S. nieuwenhuisii and the figure which appeared in Pflanzenreich. Alderwerelt therefore decided to redefine S. nieuwenhuisii on this basis. He not only rightly excluded the Homalomena element from S. nieuwenhuisii, but also, unseen, the only other element Engler cited (the Berlin Hay and Yuzammi, Schismatoglottideae (Araceae) in Malesia I 113 duplicate of Winkler 2129), basing the redefined S. nieuwenhuisii on a cultivated plant XI.B.X.160. Having rearranged things to his liking, Alderwerelt used the Bogor duplicate of Winkler 2129 (incorrectly cited as 2199), an isosyntype of S. nieuwenhuisii, as the holotype of a new species Schismatoglottis maculata Alderw. In order to establish an identity for S. nieuwenhuisii, it does not seem useful or acceptable, for the purpose of lectotypification, to go down Alderwerelt’s very speculative path, after over a decade and a half had elapsed (Engler visited Bogor in 1906), given all the opportunities that exist for the misplacement of plant labels, of specifying what other material growing in the Bogor gardens Engler might have seen and used to formulate his concept. Where there is this much confusion, and in a case where the name can hardly be deemed to be in current use, it seems preferable to take a mechanical approach rather than to try to fathom and make a tenuous case for what the author may have really meant in the event that the cited materials do not agree well with the description. In that case S. nieuwenhuisii is to be lectotypified by the Berlin duplicate of Winkler 2129 [evidently Engler’s inclusion of material of a Homalomena was a mistake; besides which the epithet nieuwenhuisii is already occupied in that genus]. However, the Berlin duplicate, which was the only one cited by Engler, appears to have been destroyed in WWII. The other duplicates of Winkler 2129 are not syntypes and therefore the lectotype itself cannot be selected from among them, but as duplicates they have a strong, if mechanical, link with the name S. nieuwenhuisii, and they are therefore here designated as isolectotypes. The specimens concerned conform uncontroversially with S. calyptrata. As Schismatoglottis niewenhuisii and S. maculata were based on different duplicates of Winkler 2129, they are not homotypic and S. maculata is to be regarded as legitimate.
The holotype of Schismatoglottis klossii is mounted on a sheet which includes a second, virtually identical specimen labelled as having been collected on Mt Carstenz at Camp III in Dec 1912. Ridley’s citation states ‘Camps III to IV…’, which agrees exactly with the label associated with the specimen cited as holotype here. Alderwerelt made two specimens from the type plant of Schismatoglottis ruttenii Alderw., either of which could be selected as lectotype. The second, Alderwerelt 323, is in BO (+ BO spirit). Alderwerelt also made two specimens from the type plant of Schismatoglottis acutangula f. staminodiifera Alderw., either of which could be selected as lectotype. The second, Alderwerelt 295, is in BOKR (+ BO spirit).