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Chromosome Numbers of Hawaiian Angiosperms: New Records and Comments1

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Chromosome Numbers of Hawaiian Angiosperms: New Records and Comments1 View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by ScholarSpace at University of Hawai'i at Manoa Chromosome Numbers of Hawaiian Angiosperms: New Records and Comments1 Michael Kiehn2 Abstract: In this paper chromosome counts for 90 collections representing 67 native Hawaiian angiosperm species and eight hybrids in 22 families are presented and discussed. Included are the first records for 26 species, two sub- specific taxa, eight natural hybrids, and the endemic genus Pteralyxia (Apocyna- ceae). In four families Hawaiian representatives have been investigated cytologically for the first time. For three species the investigations are the first on Hawaiian material. Seven counts differ from earlier reports in the literature. Implications of the results are discussed in the context of autochthonous chro- mosomal evolution and of colonization events for the Hawaiian Islands. Chromosome numbers are available for growing root tips, shoot apices, or young nearly 40% of the native Hawaiian species flowers for mitotic counts, or young flower (Carr 1998). In most cases these numbers buds for meiotic investigations of plants cul- were obtained from counts of single or very tivated at the National Tropical Botanical few individuals. In this paper it is aimed Garden, La¯wa‘i, Kaua‘i, Hawai‘i (NTBG), to improve the knowledge on chromosome and of plants collected in the wild were made numbers for Hawaiian angiosperms, both in a freshly mixed 3:1 solution of ethanol with regard to taxa not previously investi- (96%) :glacial acetic acid. In each field fixa- gated and for additional collections of taxa tion, only one individual was sampled (except already counted. Data for 19 species pre- for Labordia degeneri, DL6453, collected and sented here were shared with G. Carr (Uni- fixed by D. Lorence, where several individuals versity of Hawai‘i at Ma¯noa, Honolulu) and may have been sampled in the same fixation). were included in his earlier survey (Carr Additional fixations of root tips were taken 1998, table 1.1) as ‘‘Kiehn unpubl.’’ These from plants cultivated at the Botanical Garden counts are documented and discussed further of the University of Vienna, Austria (HBV ), here and are footnoted ‘‘c’’ in Table 1. from seeds collected in Hawai‘i; the root tips were pretreated in 8-hydroxyquinoline materials and methods (0.002 m, ca. 6 hr at 8–10C). Each root tip fixation represented one individual. The In the course of eight visits to Hawai‘i be- fixed material was stored at 8–10C before tween 1989 and 2003, fixations of actively staining with Giemsa (after hydrolysis in 5 N HCl for 50 min. at 20C) or with Feulgen 1 Parts of this study were supported financially by the (Kiehn 1995). Hot aceto-carmine (2% solu- Austrian Science Foundation (FWF-proj. P-13107-Bio), tion in 45% acetic acid) was used to intensify the O¨ sterr. Forschungsgemeinschaft (proj. 06/3699), the staining if needed. Somatic chromosome Hochschuljubila¨umsstiftung der Stadt Wien (proj. Nr. numbers (2n) were established from any of H-21/94), and the National Tropical Botanical Garden, several tissue sources, including premeiotic La¯wa‘i, Kaua‘i, Hawai‘i. Manuscript accepted 20 August 2004. anthers, young flower buds, young ovaries, 2 Institute of Botany, University of Vienna, Rennweg ovary wall, and root tips. Gametic number 14, A-1030 Vienna, Austria (e-mail: michael.kiehn@ determinations (n) were derived from meiotic univie.ac.at). divisions of microsporocytes. Collection data for each studied sample are listed in Table Pacific Science (2005), vol. 59, no. 3:363–377 1. Voucher specimens have been deposited : 2005 by University of Hawai‘i Press in the herbaria at the University of Vienna All rights reserved (wu), at the National Tropical Botanical Gar- 363 364 PACIFIC SCIENCE . July 2005 den (ptbg), and/or in other herbaria as stated dra have been counted so far; both are tetra- in Table 1. Permanent slides for most of the ploids with n ¼ 24=2n ¼ 48 (Carr 1978, 1998, counts are in the collection of the author. Kiehn and Lorence 1996). The data reported here for Tetraplasandra bisattenuata (2n ¼ 46– results 48) fit well into the picture for this group. campanulaceae: The chromosome num- Chromosome counts for 67 species (90 ber determinations for seven species, five collections) of native Hawaiian angiosperms of which are new counts, are in accor- from 22 families are presented in Table dance with the earlier data for Hawaiian 1. They include the first chromosome num- species summarized by Lammers (1988) ber reports for 26 species, two subspecific and Carr (1998). All Hawaiian taxa of the taxa, eight natural hybrids, and the endemic family cytologically investigated to date ex- genus Pteralyxia (Apocynaceae). In four fami- hibit 2n ¼ 28; Carr (1998) considered single lies (Droseraceae, Myrsinaceae, Orchidaceae, deviating reports to be erroneous, a view cor- and Viscaceae) Hawaiian representatives have roborated by the results presented here. caryophyllaceae been investigated cytologically for the first : Of the seven Hawai- time. For three indigenous Hawaiian species ian species of Silene, S. alexanderi from Molo- the investigations are the first ones on Hawai- ka‘i reported here is the third to be counted ian material. Counts for seven species differ (see Carr 1998). All three species exhibit from earlier reports; 18 counts are in accor- 2n ¼ 24, in agreement with x ¼ 12 as the pre- dance with earlier published data. Some chro- dominant basic number for Silene (cf., e.g., mosome numbers are reported as a range. Goldblatt and Johnson 2000). cyperaceae This is the case in taxa with high chromo- : The chromosome number some numbers (e.g., in Hedyotis spp. or Cop- for Eleocharis obtusa is established for the rosma ernodeoides) where different counts on first time on Hawaiian material. The result several individuals led to differing results, or of 2n ¼ 10 is in accordance with earlier was caused by chromosomes sticking together counts for this species (e.g., from northeast- during fixation (e.g., in Alyxia, Labordia, Lipa- ern America [Gervais and Cayouette 1985]). droseraceae ris, or Korthalsella). : The first count for Dro- sera anglica on material from Hawai‘i revealed 2n ¼ 40. This result is in accordance with discussion earlier reports for this, by Murı´n and Ma´jov- sky´ (1987) from Europe and by Kondo Discussion of the new data in a taxonomic and Segawa (1988) from Asia. Thus there is context no apparent variation in the chromosome apocynaceae: The results reported number of this species across its whole range here for Alyxia stellata (¼ A. oliviformis, cf. of distribution. In contrast, the genus Drosera Middleton 2000) and for Pteralyxia kauaiensis in general is extremely variable in basic num- (first count for the genus) are in line with pre- bers (ranging from x ¼ 3tox ¼ 9, with addi- vious reports for the genus Alyxia (Skottsberg tional reports of x ¼ 13 and x ¼ 15 [Kondo 1955: 2n ¼ ca. 36, 2n ¼ ca. 39 for A. stellata; and Lavarack 1984, Kondo and Segawa Van der Laan and Arends 1985: 2n ¼ 36 for 1988, Sheikh and Kondo 1995]) and in ploidy two non-Hawaiian Alyxia species). The re- levels (from diploidy to dodecaploidy [Kondo ported range of 2n ¼ 34–36 is caused by the and Lavarack 1984, Sheikh and Kondo relatively large chromosomes being twisted 1995]). around each other in the available fixations. ericaceae: Two of the three native Ha- araliaceae: According to molecular data, waiian Vaccinium species (V. calycinum, V. re- the Hawaiian genera Munroidendron, Reynold- ticulatum) have already been counted; both sia, and Tetraplasandra are closely related to exhibit 2n ¼ 24 (Carr 1998). The result of each other (Wen et al. 2001). The monotypic 2n ¼ 48 obtained here for Vaccinium cf. den- Munroidendron and one species of Tetraplasan- tatum from Moloka‘i indicates the presence Chromosome Numbers of Hawaiian Angiosperms . Kiehn 365 of tetraploidy in Hawaiian representatives of reports by Borgmann 1964 for taxa from the genus. New Guinea [see Kiehn and Weber 1998 for The three native Hawaiian Vaccinium discussion]). The genus proves to be very uni- species, together with the southern Polyne- form karyologically. Some variation in chro- sian species V. cereum, form the monophyletic mosome length and size (as discussed by sect. Macropelma of Vaccinium (Vander Kloet Storey 1966) can be seen in Hawaiian mate- 1996). Vander Kloet (1996) hypothesized rial, but this is (at least partly) caused by that the Hawaiian taxa compose a coherent differences in fixation and pretreatments, be- evolutionary unit of occasionally anastomos- cause such variation was also observed in dif- ing selfing lineages. The production of such ferent fixations of the same taxon (e.g., of C. hybrid lineages could explain the presence of hawaiensis and of C. paludosa). two ploidy levels among the Hawaiian taxa. Ranges of numbers given for C. grayi, C. The difficulties in identifying the investigated kalihii, C. lessoniana, and C. platyphylla are Vaccinium collection from Moloka‘i could also due to the poor quality of the respective fixa- be due to a hybrid origin of the investigated tions and most probably do not reflect a real collection. It would be desirable to study mei- range of chromosome numbers in these spe- otic divisions in this group or to use molecu- cies. In the case of the C. lessoniana collection, lar fingerprint methods to further evaluate this assumption is supported by the obser- the hypothesis of Vander Kloet (1996). vation of regular meiotic divisions. Regular geraniaceae: The genus Geranium is meiotic divisions are also present in collec- represented in Hawai‘i by six native species. tions of C. hawaiensis, C. kauaiensis, C. paludosa The chromosome number established here var. paludosa, C. sandwicensis, and in two puta- for Geranium cuneatum subsp.
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