Pacific Science, vol. 54, no. 2: 159-167 © 2000 by University of Hawai'i Press. All rights reserved

Phylogenetic and Biogeographic Aspects of Sect. (Papilionaceae) 1

R. C. PENA,2 L. ITURRIAGA,2 G. MONTENEGRO,z AND B. K. CASSELS 3

ABSTRACT: Sophora comprises 45-50 of worldwide distribution, but no general proposal as to the evolution of this group has been put forth. We used cladistic relationships of the quinolizidine alkaloids (matrine, sparteine, methylcytisine, anagyrine, and sophoranol) with morphological and palynolog­ ical characters to suggest a hypothesis of evolutionary and biogeographic rela­ tionships. The mainland Chilean species of Sophora appear to have been de­ rived from' ancestors phylogenetically near the extant Argentinean species S. linearifolia and S. rhynchocarpa and the psammophyte S. tomentosa, growing at tropical coastal sites around the world. The Boreotropic hypothesis of Lavin and Luckow is incorporated in our model as the most parsimonious explana­ tion of the evolution of the species of Edwardsia. Sophora is a taxonomic group that meets the following criteria: a center of diversity in North America, an early Tertiary record in North America, and a pantropical distribution. Styph­ nolobium and Sophora (including Calia) are representatives of Sophora s.l. in the United States, suggesting a migration of the latter from the Northern Hemi­ sphere to South America. Consistent with the Boreotropic hypothesis, a pri­ mary diversification center in South America and subsequent migration to the Indian Ocean and New Zealand, the Juan Fernandez Archipelago, , and possibly the Hawaiian Islands is the simplest explanation for the evolution of the Edwardsia species.

VALID SPECIES OF Sophora following revisions The Sophora is a taxonomic group of Allan (1961), Yakovlev (1967), Green of pantropical distribution but with its center (1970), Godley (1979), Isely (1981), Tsoong of diversity and an early Tertiary record in and Ma (1981), and Heenan (1998) are listed North America. Sousa and Rudd (1993) dis­ in Table 1. Although Tsoong and Ma (1981) tinguished Sophora as perennials, , or merged Calia and , it is better without bracteoles and with a dehiscent to consider them as two distinctive taxa , and Calia and Styphnolobium as taking into account first the cytogenetic trees with with bracteoles and non­ differences, chromosome number, and karyo­ dehiscent fruit. Styphnolobium differs from types, and second the distinctive Calia in having with stipels and le­ mode of Styphnolobium and Sophora sect. gume with fleshy meso- and endocarp. Calia (Palomino et al. 1993). Styphnolobium affine (Torr. and A. Gray) Walp grows well in deciduous forests and grasslands. is a deciduous endeniic to'China.- Sophora 1 This study was made possible by the financial sup­ port of the Fondo Nacional de Ciencia y Tecnologia macrocarpa is a characteristic endemic (FONDECYT) Chile, Grant No. 1980967 to G.M. and from Chile that grows in open places or NIH Grant 2 UOI TW 00316-06 to B. Timmermann. under the forest canopy. , a Manuscript accepted I August 1999. divaricate suckering shrub from New Zea­ 2Ecology Department, Pontificia Universidad Cato­ lica de Chile, Casilla 114-D, Santiago, Chile. land, grows well in grassland and rocky 3Faculty of Sciences, Universidad de Chile, P.O. Box places. In contrast, the majority ofthe species 653, Santiago, Chile. of Edwardsia prefer lowland or lower mon- 159 160 PACIFIC SCIENCE, Volume 54, April 2000

TABLE 1

NOMENCLATURE OF STUDIED SPECIES, INCLUDING SECTIONS

SPECIES NAME SECTION

Sophora secundiflora Ortega ex DC. Calia (Salisb.) Seem. Edwardsia Sophora denudata Bory Edwardsia Sophorajernandeziana (phil.) Skottb. Edwardsia Sophora howinsula (Oliv.) Green Edwardsia Sophora longicarinata G. Simpson & J. S. Thompson Edwardsia Sophora macnabiana (Grah.) Skottb. Edwardsia J. E. Sm. Edwardsia Sophora masajuerana (phil.) Skottb. Edwardsia Ait. Edwardsia Sophora prostrata Buchan. Edwardsia J. Mill. Edwardsia L. Edwardsia Sophora rhynchocarpa Griseb. Sophora Sophora linearifolia Griseb. Sophora L. Sophora Styphnolobiumjaponicum (L.) Schott Styphnolobium

tane areas along rivers and forest margins or that S. macrocarpa is derived from S. micro­ outskirts. Sophora longicarinata (a recently phylla of New Zealand on the basis of the reinstated species close to S. microphylla) is a similarity of the polypeptide and protein shrub or tree of stressed habitats; it is known compositions of both taxa. It is surprising from the Takaka area of the northern South that those authors also recognized the strong Island of New Zealand, restricted to marble similarity between these taxa and Pisum sati­ or limestone sites (Heenan 1998). Sophora vum in this regard, which seems to indicate denudata, S. chrysophylla, and Sophora from that their method does not even discriminate Juan Fernandez are found in uplands-So at the suprageneric level. Similarly, Mark­ Jernandeziana appears on hills; S. denudata, a ham and Godley (1972) could not segregate small tree from La Reunion, grows over the Chilean, Gough Island, or Chatham Is­ Acacia heterophylla Willd. and is frequently land taxa from New Zealand material from associated with the ericaceous Philippia mon­ both North and South Islands using their tana Klotsch.; similarly S. chrysophylla is the phenolic constituent patterns. Additional cir­ major tree in the treeline ecosystem on both cumstantial evidence cited by Murray (1986) (Hawai'i) and Haleakala () considered the hydrochoric character of S. and is the successional replacer of Metrosi­ microphylla . Ramirez and Romero deros polymorpha Gaud. (Mueller-Dombois (1978) also observed the potential for hydro­ 1987). choric seed dispersal in S. macrocarpa. There The first hypothesis put forth to explain is no difficulty, however, in considering S. the origin of Edwardsia suggests that their macrocarpa as the founding species of the disttibution center lies in the-West Pacific section as a- result of its possible relationship with subsequent migration and differentia­ to American species in the section Sophora tion from west to east, reaching South and because of its potential for hydrochoric America, Hawai'i, Juan Fernandez and seed dispersal. Two species from Argentina Easter Islands, Gough Island in the Atlantic with a large, unwinged legume and dark Ocean, and La Reunion in the Indian Ocean brown seeds (cf. Burkart 1952), S. linearifolia (Sykes and Godley 1968). Recently, Murray and S. rhynchocarpa, restricted to Cordoba (1986) and Murray and Porter (1980) stated and San Luis Provinces, and to Salta Prov-

, - ...... '" , .....'" • "..,.-..-" .~, ... , ,I, ... ,~... ,.,.,r .or '....~_...... ,,_ Phylogenetic and Biogeographic Aspects of Sophora Sect. Edwardsia-"PENA ET AL. 161 ince, respectively, are closest to Chilean S. B.P. and Masafuera between 1.0 and 2.4 mil­ macrocarpa. In the same vein, Skottsberg lion yr RP. (Stuessy et al. 1984). Lava from (1956) noted that the Chilean insular taxa, S. the Poike crater of Easter Island is 2.5 mil­ jernandeziana, S. masajuerana, and S. toro­ lion yr old (Zizka 1988). Thus, an American miro, are related and might be united in S. origin for this section becomes an attractive tetraptera. In the Juan Fernandez Islands alternative contrasting with the Gondwanic there is a high degree of endemism, and the origin suggested for the Edwardsia species, as principal source of its flowering is the may deduced from Sykes and Godley's continental flora of South America, repre­ (1968) and Murray's (1986) reasoning (M. senting about 47% (Hoffmann and Marti­ Kalin Arroyo, pers. comm.). corena 1987). We have incorporated Lavin and The link between sections Sophora and Luckow's hypothesis in our model, because Edwardsia is S. macrocarpa. In Edwardsia, Sophora is a taxonomic group that meets the change in morphology can be re­ the following three criteria: a center of lated to the presence of a new pollination diversity in North America, an early Tertiary vector. Calia, Sophora, and Styphnolobium record in North America, and a have an erect flag and are melittophylic, pantropical distribution (Lavin and Luckow but Edwardsia has an extended one and is 1993). Styphnolobium, Sophora, and Calia visited by birds. Such a change has been are representatives of Sophora s.1. in the demonstrated in Harpalyce (Galegeae­ United States, suggesting migration of the Brongniartieae), whose resupinate flowers are latter from the Northern Hemisphere to South inadequate to ensure successful pollination America. Another two cases demonstrated in by (Arroyo 1976). Drepanididae visit the Papilionaceae are Robinia (Lavin and Sophora chrysophylla in Hawai'i and the tro­ Luckow 1993) and the Psoraleeae (Grimes chilid Patagona gigas visits S. macrocarpa in 1990). The fossil record of Styphnolobium is Chile (Arroyo 1981); other present in Middle Eocene deposits in North pollinate S. tetraptera and S. microphylla in America (Sousa and Rudd 1993). New Zealand. It is interesting that the Both palynological and chemical evidence of the ornithophilous S. microphylla kills might corroborate Or weaken the hypothesis bees, but is apparently not harmful to birds of a closer relationship between the Sophora (Arroyo 1981). taxa from Juan Fernandez Islands and South Flowering plants were present in New American stock close to S. macnabiana (s. Zealand by the Cenomanian, some 100 mil­ microphylla from South America) than with lion yr B.P. (before the present) when it was in their previously postulated Antarcto-Pacific direct contact with Australia, New Caledo­ relatives. nia, Antarctica, India, and South America No recent micromorphological studies are (Raven 1973). Nevertheless, there is no paly­ available, with the exception of a partial nological evidence for the existence of Soph­ analysis of the cuticles of some Sophora spe­ ora before Quaternary times (Mindelhall cies belonging to the section Edwardsia 1980). That there are only two species of (Lobin and Barthlott 1988, Alden and Zizka Sophora in Australia, S. jraseri Benth. and S. 1989), all of which are very similar. Many tomentosa subsp. australis Yakovl., both years earlier, Heusser (1971) had described from Queensland (fide Yakovlev 1967), sug­ the pollen grain~ of the continental Edward­ gests that it is a recent immigrant. Alfthe sia species as tricolporate, subprolate, with a volcanic islands of the Pacific Ocean appear suprareticulated exine I J..lm thick. Pefia et al. to be at most a few million years old. In the (1993) studied the architecture of pollen of Hawaiian Islands, Kaua'i is dated about 4.4­ various Edwardsia compared with Styphno­ 5.7 million yr B.P. and Hawai'i between 0.1 lobium. and 0.45 million yr RP. (Macdonald et al. The aim of this work was to put forth an 1983). Juan Fernandez Masatierra Island has alternate hypothesis to explain the origin of been dated to between 3.7 and 4.3 million yr Edwardsia by using comparative chemistry 162 PACIFIC SCIENCE, Volume 54, April 2000

TABLE 2

DATA MATRIX FOR Styphnolobium, Calia, AND THE Sophora SPECIES STUDIED; 9 INDICATES INDETERMINATE OR INAPPLICABLE CHARACTERS

TAXON"

STYPH 10002 00000 00100 00000 0000 CALIA 10000 00000 30000 10002 0191 TOMEN 00000 00001 01009 1 1210 1 110 LINEA 01000 00000 00000 10002 0000 RHYNC 01100 00000 00109 19999 9000 MACRO 00000 10100 00001 10022 0001 CHRYS 1 1000 1 1 1 1 1 2 1 1 1 1 10022 1 111 DENUD 10 100 1 1 1 1 1 0001 1 10121 1011 MACNA 1 1 1 1 1 01111 1 1 1 1 1 10220 0011 FERN 10101 1 0 1 0 1 00001 1 1022 1 101 MASAF 1 0 1 1 1 1 0 1 0 1 01101 19999 9011 TaRO 10101 1 1 1 1 1 1 1 1 1 1 1 1220 1001 HOW 10001 1 1 1 0 1 1 1 1 1 1 99999 9091 TETRA 1 1000 1 2 1 0 1 1 1 1 1 1 10220 0011 MICRO 1 1 1 1 1 1 1 1 1 1 11111 1 1220 1011 PROST 001 1 1 1 0 1 1 1 00110 10022 0091

"STYPH, Styphnolobium; TOMEN, Sophora tomentosa; LINEA, S. linearifolia; RHYNC, S. rhynchocarpa; MACRO, S. macrocarpa; CHRYS, S. chrysophylla; DENUD, S. denudata; MACNA, S. macnabiana; FERN, S. fernandeziana; MASAF, S. masafuerana; TaRO, s. toromiro; HOW, S. howinsula; TETRA, S. tetraptera; MICRO, S. microphylla; PROST, S. prostrata.

and morphological and palynological evi­ most parsimonious trees option (mhennig*) dence. with global branch swapping (bb*), implicit enumeration search option (ie*) with non- additive characters, and nelsen option (nel- MATERIALS AND METHODS sen*) for consensus resolutions of Hennig86 (Farris 1988). Character states are shown in The exines of pollen grains of Sophora Appendix 1. Indeterminate characters coded species were examined by scanning electron as 9 in Table 2 were processed as 0 (synapo­ microscopy. The pollen grains were metal- morphic). lized with a gold-palladium layer 30-40 nm The selection of an operative out-group thick and then photographed with an Auto- for Edwardsia lies in other sections of Soph­ scan Siemens microscope according to the ora, mainly Calia and the genus Styphno­ method of Lynch and Webster (1975). Pollen lobium. Many characters show that these two measurements were determined from low- genera are closely related. A number of resolution micrographs and exine ornamen- characteristics of Styphnolobium Schott. tation was described from plates taken at allow it to be considered as an out-group of 10,000 x magnification. The scanty herbar- Sophora: the presence of stipulae, of reserve ium materials do not allow the assignment of albumen in the seeds, the distinctive chro­ mean or modarvalues:orirymeasurement-mosome niliii5ef (n = l'1-),-tne-·presence of ranges are provided for pollen besides the seed galactomannans, hypogeal germination, ornamentation of exine. and the heterobrochate exine (Bailey 1974, Alkaloid extraction and GLC analysis Goldblatt 1981, Polhill 1981, Pefia et al. have been described and discussed in a pre- 1993) (Table 2). Both Calia and Edwardsia vious paper (Pefia and Cassels 1996). have cotyledons with epigeal germination, The characters were polarized by the out- contrasting with sections Styphnolobium and group technique and processed by multiple Sophora in which germination is hypogeal. Phylogenetic and Biogeographic Aspects of Sophora Sect. Edwardsia-PENA ET AL. 163

r---- S. microphylla '----- S. toromiro r---- S. macnabiana '----- S. tetraptera '------S. chrysophylla L..- S. denudata Section Edwardsia

L s. masafuerana ------1C=== S. howinsula '------S. prostrata L..- S. femandeziana L- S. macrocarpa '------S. secundiflora Section Calia L..- S.linearifolia '------S. tomentosa Section Sophora '------S. rhynchocarpa L- Styphnolobiumjaponicum

FIGURE 1. Cladogram of Sophora sect. Edwardsia, using 22 characters: 1-16, morphology; 17-23, chemistry; 23, palynology; and 24, germination.

RESULTS ships that were consistently resolved in five trees, depicted in Figure 1. Edwardsia is Incomplete data used in previous studies monophyletic by 6, 8, 15, and 19. Sophora did not allow a hypothesis to be elaborated microphylla, S. toromiro, S. tetraptera, mac­ to explain the origin and evolution of Soph­ nabiana, S. masaJuerana, and S. howinsula ora species, especially for section Edwardsia. are sister groups. Calia (s. secundiflora) ap­ We have now been able to reconcile geologi­ pears as a paraphyletic assemblage of Soph­ cal, palynological, morphological, germina­ ora sect. Edwardsia. Both Calia and Edward­ tion, and chemical evidence related to Chil­ sia have cotyledons with epigeal germination, ean mainland and oceanic island (5 species) contrasting with sections Styphnolobium and material and from another collection of Ed­ Sophora in which germination is hypogeal. wardsia (La Reunion [1], New Zealand [4], Here this condition is viewed as symplesio­ and Hawai'i [1]). morphic. Otherwise, three resolutions from The cladogram of Sophora sect. Edwardsia 19 remnant trees show four polyphyletic trees group (Figure 1) is one of the 43 most parsi­ for sections of Sophora, and five gave S. monious cladograms generated by Hennig86 prostrata, a species with many symplesio­ using multiple most parsimonious trees op­ morphic characters, near the base of the Ed­ tion (mhennig*) with global swapping (bb*). wardsia group. In three cases linkage of S. The cladogram has 84 steps, a consistency tetraptera with S. denudata and S. chrys­ index of 0.36, and a retention index of 0.60. ophylla was found; these tree species are The relationships discussed below are those considered closely allied by many specialists consistently resolved in 25 cladograms. Im­ (Green 1970). plicit enumeration search option (ie*) with all The following chemical compound char­ characters nonadditive gives similar results. acters are informative: replacement of tetra­ The other 16 cladograms are variants of the by tricycljc quin9lizidine alkaloids in .Calia - one presented and have no bearing on a dis­ [sect. Calia (Berland) Rudd.], which has a cussion of the origin of the Edwardsia group. high content, like Edwardsia, in con­ Only four represent a nonmonophyletic ar­ trast with section Sophora, which has a rangement of sect. Edwardsia, partly due to higher sparteine content. Hoeneisen et al. chemical homoplasious characters; therefore (1993) isolated formylcytisine from two S. tomentosa sometimes converges with the Chilean species, S. Jernandeziana and S. more advanced Edwardsia. macnabiana. This is probably the only chem­ The cladogram represents the relation- ical linkage found between these two species 164 PACIFIC SCIENCE, Volume 54, April 2000

to the apparent exclusion of others, although migration to the Pacific and Indian oceanic it seems only to represent an intermediate islands. biosynthetic step between methylcytisine and The Papilionaceae are usually discussed as cytisine, common to all Edwardsia. Mark­ of Gondwanic origin, because their major ham and Godley (1972) merged S. macnabi­ diversity is in tropical Africa and South ana (Chile) with S. microphylla (New Zea­ America (Raven 1973, Kamp 1980, Raven land) populations because of the similarity of and Polhill 1981). In this vein, it is assumed the flavonoid patterns. Our chemical re­ that North American tropical representatives sults with the seed alkaloids contradict a re­ evolved from South American ancestors lationship between these species, particularly (Taylor 1990). considering the methylcytisinejcytisine ratio: Sophora macnabiana, a Chilean species a New Zealand population preferentially ac­ close to S. microphylla and putatively derived cumulates the methyl derivative, whereas from it, represents the most advanced taxon Chilean populations contain higher propor­ in section Edwardsia. Sophora fernandeziana tions of the nonmethylated compound. and S. macrocarpa share ancestral, morpho­ Sophora denudata and S. chrysophylla logical, chemical, or palynological charac­ show some similarities in their morphological ters. Considering the flora of the Juan Fer­ (I, 6-10, 14-16), palynological (23), germi­ mindez Islands, S. fernandeziana appears nation (24), and some chemical (19, 21) closely related to S. macrocarpa (6-8, IS, 16, characters. Sophora microphylla and S. mac­ 19, 20, and 24), but parallel evolution seems nabiana show many synapomorphic charac­ to be the simplest explanation. Similarly, S. ters (1-5, 7-16, 18-19,23-24). Sophora toro­ masafuerana appears closer to the micro­ miro only differs in ornamentation of the phylla group. The palynological characters of exine (23), two chemical characters (17 and the former two, S. fernandeziana and S. 21), and three morphological characters (2, 4, macrocarpa (exine heterobrochate), contrast and 6). Chemically, these two species may be with the homobrochate exine of S. masa­ linked with S. tetraptera and s.. macnabiana fuerana, S. macnabiana, and most insular Ed­ by cytisine and sparteine contents (18, 19). wardsia species (pefia et al. 1993). Therefore, The relationship between S. masafuerana derivation of the insular species, particularly and S. howinsula deserves further comment. of S. fernandeziana, from continental species Many indeterminate characters here were cannot be dismissed. Sophora macrocarpa coded as °or symplesiomorphic, 17-21. In and S. fernandeziana share a number of addition, characters 16 and 23 in S. howin­ characters (15,19, and 20: absence ofauricles sula make this relationship dubious. on the , percentage of matrine, and methylcytisinejcytisine ratio). The relation­ ship of S. toromiro remains unclear: chemis­ try characters coincide with S. microphylla DISCUSSION but palynologically it is close to the South We previously proposed a South Ameri­ American group, with exine heterobrochate. can origin for section Edwardsia of Sophora (pefia et al. 1993) based upon palynological and chemical data. Many lines of evidence CONCLUSIONS were. not c.ongruentwith~Ne~~ealand center oforigin for Edwardsia as a whole. We . . Consistent wltfithe' BoreoiropiC hypothe­ later confirmed (pefia and Cassels 1996) this sis, a primary diversification center in South hypothesis by incorporating phytochemical America and subsequent migration to the evidence. Although the chemical characters Indian Ocean, New Zealand, the Juan Fer­ are poor discriminators, they do not falsify mindez Islands, Easter Island, and possibly our proposal of a South American primary the Hawaiian Islands is the simplest expla­ center of diversification and ultramarine nation for the evolution of Edwardsia species

. ., _ ....N.,~'~""_,..,. .... 'f'-""",." "'~" •• .,. ./ ...... ,.-., •• _ .." ",... , •..,,,,,. Phylogenetic and Biogeographic Aspects of Sophora Sect. Edwardsia-PENA ET AL. 165

...... ;... )

<:> Gough I.

.. '

FIGURE 2. Area cladogram of some representatives of Sophora depicting South American elements ultimately de­ rived from North American lineages Styphnolobium and Calia.

(two mainland and nine ultramarine) taking carpa and the psammophyte S. tomentosa into account comparative morphological, growing at tropical coastal sites around the palynological, and chemical evidence (Figure world. 2). Our results are consistent with Lavin and Luckow's (1993) hypothesis. First, Sophora is ACKNOWLEDGMENTS a taxonomic group that meets the following three criteria: a center of diversity in North We are grateful to Mary Kalin Arroyo for America, an early Tertiary fossil record in suggesting an American origin for Edwardsia North America, and a pantropical distribu­ and a number of tests of this hypothesis. We tion. Styphnolobium, Sophora, and Calia are also thank the generous sources of representatives of Sophora s.l. in the United material (SI, BISH, SSUC, SGo) and Carolina States, suggesting migration of the latter Villagran for her helpful comments. A copy --fn511l--fhe--Nortliern Hemispliere --to --South ofFarris' program Hennig86 was generously ­ America. Styphnolobium is present in Middle borrowed by Charlotte Taylor. Eocene deposits (Sousa and Rudd 1993). On phylogenetic grounds, the mainland Chilean LITERATURE CITED species of Sophora appear to have been de­ rived from ancestors near the extant Argen­ ALDEN, B., and G. ZIZKA. 1989. Der Toro­ tinean species S. linearifolia and S. rhyncho- miro (Sophora toromiro) eine ausges- 166 PACIFIC SCIENCE, Volume 54, April 2000

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Sophora L. y Styphnolobium Schott. Ann. relationships of angiosperms from Creta­ Mo. Bot. Gard. 80: 284-290. ceous and early Tertiary of the North PENA, R. c., and B. K. CASSELS. 1996. Phy­ American area. Bot. Rev. 56: 279-379. logenetic relationships between Chilean TSOONG, P.-c., and C.-Y. MA. 1981. A study Sophora species (Papilionaceae). Biochem. on the genus Sophora L. Acta Phytotaxon. Ecol. Syst. 24 (7-8): 725-733. Sin. 19: 1-22, 143-167. PENA, R. c., L. ITURRIAGA, A. M. MUJICA, YAKOVLEV, G. L. 1967. Zametki po sistema­ and G. MONTENEGRO. 1993. Amilisis rni­ tike i geographii roda Sophora L. i bliz­ cromorfologico de polen de Sophora kikh rodov. Proc. Leningr. Chern. Pharm. (papilionaceae). Gayana Bot. 50 (2): 57­ Inst. 21 (4): 42-62. 65. ZIZKA, G. 1988. Naturgeschichte der Oster­ POLIllLL, R. H. 1981. Sophoreae. Pages 213­ insel. Pages 21-38 in 1500 Jahre Kultur 230 in R. M. Polhill and P. H. Raven, eds. der Osterinsel. Schiitze aus dem Land des Advances in legume systematics. Royal Hotu Matua. Verlag Philipp von Zabern, Botanical Gardens, Kew. Mainz am Rhein. RAMIREZ, c., and M. ROMERO. 1978. El Pacifico como agente diseminante en el litoral chileno. Ecologia 3 (1): 19-30. RAVEN, P. H. 1973. Evolution of subalpine APPENDIX I and alpine plant groups in New Zealand. CHARACTER STATES: CODING NUMBERS, CHARACTERS, N. Z. J. Bot. 11: 177-200. AND STATES RAVEN, P. H., and R. M. POLIllLL. 1981. I: Life form: shrub, 0; tree, 1. Biogeography of Leguminosae. Pages 27­ 2: Number of leaflets: few, 0; 15-25, 1. 34 in R. M. Polhill and P. H. Raven, eds. 3: Length of leaflets: over I cm, 0; less than 1 cm, 1. Advances in legume systematics. Royal 4: Length/width of leaflets ratio: more than 2, 0; less than 2, 1. Botanical Gardens, Kew. 5: Pubescence of the leaflets: face and back, 0; back SKOTTSBERG, C. 1956. Derivation of the flora only, 1. and fauna of Juan Fernandez and Easter 6: Direction of flag: erect, 0; extended, 1. Islands. Pages 193-439 in The Natural 7: Flag/wings ratio: same, 0; much longer, I and 2. History of Juan Fernandez and Easter Is­ 8: Exertion of stamens: no, 0; yes, 1. 9: Strangulation of the lomentum: no, 0; yes, I. land. Almqvist Wiksells, Uppsala. 10: Length of seed: large (I cm or more), 0; small, 1. SOUSA, M., and V. E. RUDD. 1993. Revision II: Seed color: brown, 0; ocher, I; orange, 2; reddish, 3. del genero Styphnolobium (Leguminosae: 12: Number ofseeds per fruit: few (2-5), 0; 6 or more, I. Papilionoideae: Sophoreae). Ann. Mo. 13: Pubescence of the lomentum: marked, 0; slight, 1. 14: Presence of wings on the fruit: yes, I; no, O. Bot. Gard. 80: 270-283. IS: Presence of auricules on the petals: yes, 0; no, 1. STUESSY, T. F., K. A. FOLAND, J. F. SUTTER, 16: Presence of stipulae: yes, 0; no, 1. R. W. SANDERS, and M. O. SILVA. 1984. 17: Methylcytisine percentage: below 30%, 0; over 30%, Botanical and geological significance of 1. potassium-argon dates from the Juan Fer­ 18: Cytisine percentage: below 31%, 0; over 31%, 1. 19: Sparteine percentage: none detected, 2; below 1%, I; nandez Islands. Science (Washington, over 1%, O. D.C.) 225: 49-51. 20: Matrine percentage: below 31%, I; over 31%, O. SYKES, W. R., and E. G. GODLEY. 1968. 21: Methylcytisine/cytisine ratio: less than 0.5, 0; more Transoceanic dispersal in Sophora and than 0.5, 1. -- -other-genera. Natllie(Loi:ld.) 218: 495­ 22:-Anagirine-percentage:- below I%, I; over 1%, O. - 23: Exine ornamentation: heterobrochate, 0; homobro­ 496. chate, 1. TAYLOR, D. W. 1990. Paleobiogeographic 24: Germination: epigeal, I; hypogeal, O.