Nomenclatural and Taxonomic Changes in Hawaiian Alectryon (Sapindaceae)!
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Pacific Science (1987), vol. 41, nos. 1-4 © 1988 by the University of Hawaii Press. All right s reserved Nomenclatural and Taxonomic Changes in Hawaiian Alectryon (Sapindaceae)! G EORGE L INNEy 2 Alectryon is endemic to the southwestern Pa proposal, some problems with the existmg cific, from Indonesia to Australia, the Philip nomenclature and taxonomy of the two Ha pines, Hawaii, and New Zealand. The species waiian species were found. are either shrubs or trees; all have pinnately Based on non-flowering specimens col compound leaves. Flowers, borne in panic lected around 1870 on Oahu (Makaleha Val ulate inflorescences, are less than 5 mm in ley) and Molokai (Pali of Kalaupapa), Hille diameter, apetalous, and are androdioe brand (1888) published the provisional genus cious, perhaps monoecious in A . grandifolius , "Mahoe, gen. nov. ?" Specimens have been though Radlkofer (1933) called them "falsely deposited in Berlin (B), Kew (K) , and Bishop polygamous." The fruit is 1-3 coccate, on the Museum , Honolulu,(BISH) herbaria. In No order of 10 mm in diameter in most species, vember 1889, Radlkofer saw and annotated at and lacks any obvious suture. At maturity it least one ofthe two specimens at K (Figure 1). ruptures irregularly to expose a bright red He also saw and annotated at least one of the "aril." The seed is borne distal to the "aril" and specimens at B (Figure 2). There was no date partially imbedded in it. Its exposed surface is of annotation on the sheet at B, and the sheet highly polished. Thirty-two names have been is no longer extant. An additional annotation published, including four since Alectryon on the label , probably in Radlkofer's hand was last monographed by Radlkofer in 1933. writing, reads, " Genus imperfecte cognitum The type species of the genus is A. excelsus Sapindacearum." Apparently based solely on J. Gaertner, 1788. Hillebrand's material, as no other collec Two species have been recognized from the tions are known to have existed at that time , Hawaiian Islands: A . macrococcus Radlk. Radlkofer (1890) published the new species A . (1890) and A. mahoe St. John and Frederick macrococcus in a footnote, along with A. (1949), both endemic. These are small trees strigosus and A . reticulatus, both New Guinea 3-8 m tall, rarely twice as tall, ofmesic forests. species. Alectryon strigosus and A. reticulatus Among the five major islands ofthe Hawaiian were given full Latin diagnoses, while A . ma chain, they are absent only from the island of crococcus was presented very casually, with a Hawaii. They differ from extra-Hawaiian spe reference to Hillebrand (1888), repeating in cies, as far as known, by the presence of a formation on fruit and leaflet sizes, and an hypodermis just below the upper epidermis of original contribution on the presence of the the leaf, by the very large size ofthe fruits, up hypodermis, all in German. It is possible that to 60 mm in diameter, and in having patelli he was not actually intending to publish the form as opposed to spheroid seeds. The two name at that time, only anticipating future Hawaiian species are supposedly separated by publication. Nevertheless, Radlkofer, in his the indument of the undersurface of mature later treatments, and all subsequent workers leaves: glabrate in A. mahoe, and densely to have accepted A. macrococcus as validly pub mentose in A. macrococcus. Young and unex lished in the 1890 paper. No type was des panded leaves ofboth species are more or less ignated. Indeed, no specimen at all was cited, densely sericeo-tomentose. Alectryon macro except by reference to Hillebrand, who had coccus is being proposed as rare and en cited collections. Because of this reference, dangered. During the preparation of the Hillebrand's text is to be considered part of the protologue. Upon the discovery of an additional popu I Manuscript accepted June 1985. lation at Auwahi, Maui, by Joseph Rock in 2 Department of Botan y. 1910, Radlkofer (Radlkofer and Rock, 1911) 68 Changes in Hawaiian Alectryon-LINNEY 69 FIGURE I. Sheet of A . macrococcus in Kew, annotated FIGURE 2. Sheet of A. macrococcus, formerl y in Berlin, by Radlk ofer, here selected as new lectotype. Annotation annotated by Radlk ofer. Annotation reads: "Alectryon reads: "Alectryon macrocarpus [sic] m. sp. nov., mox macrococus [sic] m. Radlk ." Photograph by H. St. John , edenda (Mahoe Hillebr. Fl. Hawiens. p. 86 XI. 89. used by permission. Determ. Radlkofer." Kew photograph , printed by permission. may have been responsible for all the sub sequent confusion. St. John and Frederick provided a full Latin description for A . macro (1949, here and throughout) were in error coccus. He cited the Hillebrand collections in in generally attributing dense tomentum to Band K as specimens seen, and Rock's newly plants of Mo lokai . discovered Maui material as not seen. Details After additional collections had been made of floral morphology, still unknown to Radl on Oahu and Kauai by several collectors, St. kofer , were provided by Rock. In his mono John and Frederick recognized two discrete graph of the Sapindaceae, Radlkofer (1933) entities among the collections. They described repeated his description of A . macrococcus the new species A . mahoe, comprising all of without substantial change. Neither Radlko the Oahu collections, and probably tho se fer and Rock (1911) nor Radlkofer (1933) from Kauai. At the same time, St. John and mentioned dense golden brown tomentum on Frederick undertook to lectotypify A . macro leaf undersurfaces. This is noteworthy as the coccus. The y ignored the K specimens and tomentum is the most salient feature of the perhaps were unaware oftheir existence, or at Auwahi population. On the other hand, Rock least ofthe Radlkofer annotation on the one. (1913) ascribed the dense tomentum to the They selected the sheet in B annotated by species in general, ignoring the fact that spec Radlkofer, which St. John had seen and pho imens other than those collected at Auwahi tographed in 1935. Curiously, they made their were glabrate. I believe this bias or oversight selection in the belief that the sheet had since 70 PACIFIC SCIENCE, Volume 41, 1987 been destroyed in 1943. Its destruction has and Rock (1911) and Radlkofer (1933) noted been confirmed by Dr. Paul Hiepko, Curator that the leaves were "glabra (vel subtus t. of Phanerogams, Berlin Dahlem (in litt.) Hillebr. laxe tomentosa)." If St. John and Hillebrand's original material was puta Frederick realized the contradiction, it may be tively from mixed collections (Makaleha Val that they gave weight to the annotated speci ley, Oahu, and Pali of Kalaupapa, Molokai). men over the protologue, believing the pro The leaf indument character was believed to tologue in error. This again would require that be different on the two islands (glabrate on the now lost B specimen in fact had been Oahu, tomentose on Molokai), and St. John tomentose. However, the ICBN "Guide for and Frederick recognized the two entities as the Determination of Types" (GDT) TA.(d) distinct species. Therefore, the identities ofthe indicates the protologue is given precedence two species might have been reversed, depend to annotations and other indications of au ing on the choice of lectotype. St. John and thor's intent when there is a contradiction. Frederick chose a Molokai specimen. Though Article 8.1 states that a lectotype designation they do not say specifically, for their taxono may be superseded if shown to be in serious my to be consistent, it must be presumed that conflict with the protologue. Second, as to the their lectotype possessed leaves densely to choice of the annotated B sheet over the an mentose beneath. Certainly, in their discus notated K sheet, it must be repeated that the B sion they attributed this character to A . ma sheet bore no date of annotation. The sheet crococcus S.s. However, the extant Hillebrand mayor may not have been annotated prior to material in K is glabrate, regardless of the the date of publication. Surely, Radlkofer had collection locality written on the sheet, and annotated it prior to his 1911 paper, as he might be believed to be from Oahu. Locality specifically cited it there. The K sheet bears information to the contrary might be con both a prepublication date and the note "sp. sidered to be in error. Rather, I am fully per nov., mox edenda." The purport of the note is suaded the now lost B material also was extremely powerful, as no other sheet bore glabrate, not densely tomentose. Collections anything comparable. Furthermore, this sheet unquestionably from Molokai (Degener 9534 agrees with the protologue. In view of GDT and 9536; Steve Anderson 545: all in BISH), T.4.(a) which states: "A lectotype must be are glabrate. Unfortunately, St. John's photo chosen from among the elements definitely graph of the B sheet, though excellent, pro studied by the author up to the time the name vides no assistance. In the photo the leaves of the taxon was published," it is argued appear to be quite clearly glabrate, but it is not that the K sheet was the proper choice for certain that a leaf undersurface is represented. lectotype. At any rate, A . macrococcus came to be cir The third irregularity is St. John and Frede cumscribed as having densely tomentose leaf ricks having chosen a non-existent specimen. undersurfaces, and A. mahoe glabrate. The With neither a tangible specimen, nor even a other diagnostic features noted by St. John description ofit to support their claim contra and Frederick will be considered below in the dictory to the protologue, the name is no more taxonomic treatment. fixed by their lectotypification than without There are four irregularities in St.