Revision of Heterosavia, Stat. Nov., with Notes on Gonatogyne and Savia (Phyllanthaceae)

Revision of Heterosavia, Stat. Nov., with Notes on Gonatogyne and Savia (Phyllanthaceae)Author(s): Petra HoffmannSource: Brittonia, Vol. 60, No. 2 (Jul. 15, 2008), pp. 136-166Published by: Springer on behalf of the New York Botanical Garden PressStable URL: http://www.jstor.org/stable/30218783 .

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Revision of Heterosavia, stat. nov., with notes on Gonatogyne and Savia (Phyllanthaceae)

PETRA HOFFMANN

Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK e-mail: [email protected]

Abstract. Heterosavia (Phyllanthaceae) is segregated from Savia (tribe Bridelieae), recognized at generic rank, and placed in tribe Phyllantheae. Floral, fruit, leaf anatomical, leaf venation, and pollen characters of the neotropical taxa previously united as Savia including Gonatogyne are discussed and illustrated. Keys to the three genera and to the species of Heterosavia are presented. Four species (all new combinations), Heterosavia bahamensis, H. erythroxyloides, H. laurifolia, and H. maculata, are recognized. The new combinations Heterosavia laurifolia var. intermedia and H. maculata var. clementis are proposed. The names Heterosavia, H. erythroxyloides, H. laurifolia, Savia clementis, S. clusiifolia, S. clusiifolia var. fallax, and S. longipes are lectotypified. Distribution maps and conservation assessments (IUCN ratings) of Heterosavia species and varieties are provided.

Key Words: Gonatogyne, Heterosavia, morphology, Phyllanthaceae, Savia, taxonomy.

The circumscription of the genus Savia Willd. has been controversial for the past 150 years. It was consistently considered the largest genus of tribe Wielandieae Baill. ex Hurus. but subjected to repeated changes in composition (see below under Taxonomic history). Because of its alleged disjunct distribution and the relative rarity of many of the species concerned, little has been published on this group apart from regional floristic treat- ments (e.g., Alain, 1953, 1986; Leandri, 1958; Adams, 1972; Radcliffe-Smith, 1987) since its treatment in Pflanzenreich (Pax & Hoffmann, 1922). More recently, a number of systematic papers (Hoffmnann, 1994, 1998; Hoffmann & McPherson, 1998, 2007) were concerned mainly with the paleotropical members of this group.

Results of molecular phylogenetic analyses (Wurdack et al., 2004; Kathriarachchi et al., 2005) showed that Savia in its most commonly accepted, narrow circumscription (sensu Webster, 1994b, and Radcliffe-Smith, 2001) is triphyletic. All African species of Savia sensu Webster (1994b) are now united with Blotia Leandri and Petalodiscus Baill. in

Wielandia Baill., amounting to a total of 13 species (Hoffmann et al., 2006; Hoffmann & McPherson, 2007). Wielandia s.1. (clade F4b in Kathriarachchi et al., 2005) is sister to the southeast Asian Chorisandrachne Airy Shaw + Dicoelia Benth., and the three genera together constitute subtribe Wielandiinae of tribe Wielandieae (Hoffmann et al., 2006). They differ from all neotropical species of Savia sensu Webster and Brazilean Gonatogyne Klotzsch ex Miill. Arg. by being monoecious (versus dioecious), having exalbumimous seeds (versus seeds with copious endosperm), and possessing cyclocytic stomata with 3-6 subsidiary cells (versus paracytic).

Savia is now restricted to S. dictyocarpa Miill. Arg. and S. sessilifora (Sw.) Willd. (S. section Savia). Gonatogyne, considered a section of Savia by Pax and Hoffmann (1922) but segregated by later authors (Webster, 1994b; Radcliffe-Smith, 2001), was found to be the sister of Savia section Savia in the molecular analyses (Wurdack et al., 2004; Kathriarachchi et al., 2005). Savia + Gonatogyne and the sister clade consisting of Croizatia Steyerm.,

Brittonia, 60(2), 2008, pp. 136-166. ISSUED: 15 July 2008 x 2008, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.



2008] HOFFMANN: REVISION OF HETEROSAVIA 137

Discocarpus Klotzsch, and Tacarcuna Huft form the entirely neotropical subtribe Saviinae of tribe Bridelieae (sensu Hoffmann et al., 2006; clade F2b in Kathriarachchi et al., 2005). A revision of Savia and Gonatogyne will be published separately (Hoffmann and Cordeiro, unpubl. data).

Caribbean Savia section Heterosavia Urb. was shown to be strongly divergent from Savia section Savia (tribe Bridelieae) in the molecular studies and is instead sister to Flueggea Willd. (clade Flb in Kathriarachchi et al., 2005) in tribe Phyllantheae sensu Hoffmann et al. (2006). Savia section Heterosavia is therefore here elevated to generic rank as Heterosavia (Urb.) Petra Hoffm. The placement of sections Savia and Heterosavia in the same genus was mainly due to a general similarity both in vegetative characters as well as in the shared characters of the small, axillary-glomerulate flowers and the typically euphorbiaceous explosive fruits. Even after detailed analysis, the distinguishing macromorphological characters (see Key to the genera below) are subtle. However, pollen morphology (as discussed below) supports the molecular findings. Heterosavia is here taxonomically revised and its floral, fruit, leaf anatomical, leaf venation, and pollen characters as well as biogeography are discussed in conjunction with those of Savia and Gonatogyne. Heterosavia consists of four very similar species. Identifi- cation is problematic unless pistillate material is available (plants are dioecious). The taxa are vegetatively variable and seem at times to intergrade. It appears that ecological speciali- zation is almost more meaningful than morphological characters.

Taxonomic history

The genus Savia was first described by Willdenow (1806) for the single species S. sessiliflora from Hispaniola. Half a century later, Baillon (1858, 1861) added five new species from Madagascar. He divided the genus in section Eusavia Baill. (only Savia sessiliflora) and the two Madagascan sections Charidia Baill. and Petalodiscus Baill. In addition to these three sections, Miiller (1866) included in Savia also the monotypic genus Wielandia from the Seychelles as a fourth section. He placed the three species that

had been described in the meantime by Grisebach (1860, 1865) from Cuba in section Eusavia with Savia sessilif/ora. Bentham (1878) and Bentham and Hooker (1880) excluded Wielandia from Savia but tentative- ly included Miuller's (1873) new monotypic genus Gonatogyne from Brazil. Urban (1902), without reference to previous infrageneric classifications, considered Savia as a purely neotropical genus and described the new section Heterosavia and juxtaposed it with section Eusavia containing Savia sessiliflora. The former comprised Grisebach's species mentioned above, to which two species described by Britton (1904, 1916) and six of Urban (1924, 1930) were added in the following decades.

In the first edition of"Nattirliche Pflanzen- familien", Pax (1890) segregated from Savia all Madagascan species as genus Petalodis- cus, and restricted Savia to include only the neotropical species, including Gonatogyne. In "Pflanzenreich" (Pax & Hoffmann, 1922) and in the second edition of "Nattirliche Pflanzenfamilien" (Pax & Hoffinann, 1931), however, this decision was reversed and Savia was recognized in the widest sense in its history. Three sections were distinguished: 1) section Petalodiscus with free, petaloid disc lobes, 2) section Maschalanthus Pax with an annular disc, and 3) section Gonato- gyne (Miull. Arg.) Pax & K. Hoffm. with a disc consisting of numerous glands. Gonatogyne was later accepted by the same authors as a separate genus (Pax & Hoffmann, 1933). The largest section, Maschalanthus, comprised previous authors' sections Eusavia, Heterosavia, and Charidia as well as the two distinct species Pseudophyllanthus ovalis (E. Mey. ex Sond.) Voronts. & Petra Hoffm. and Phyllanthopsis phyllanthoides Voronts. & Petra Hoffm. in tribe Poranthereae; see Croizat, 1943: 11, Webster, 1967: 328; Hoffman, 1994; Vorontsova et al., 2005; Vorontsova & Hoffman, 2008. This unfortunate amalgamation of American and Madagascan taxa in one section has confused the taxonomy of Savia to the present day. Nomenclatoral issues with the section name Maschalanthus, as pointed out by Wheeler (1939), further complicate the matter. Maschalanthus Nutt. (typified by Phyllanthus polygonoides Nutt. ex Spreng.) is a synonym



138 BRITTONIA [VOL. 60

of Phyllanthus L. section Paraphyllanthus Miull. Arg. (Webster, 1970: 55). The correct author citation is therefore Savia section Maschalanthus Pax, not section Maschalan- thus (Nutt.) Pax as given, e.g., in Radcliffe- Smith (2001: 7).

The correct assignment of Savia s.1. taxa to genera or even subfamilies was also sometimes problematic. Savia sessiliflora was described several times in Phyllanthus (Richard, 1850; Lundell, 1942), and Savia dictyocarpa in Securinega Comm. ex A. Juss. (Kuhlmann, 1935). For Savia dictyocarpa, Smith and Downs (1964) also described the monotypic genus Kleinodendron L. B. Sm. & Downs, which they classified in the uniovu- late subfamily Acalyphoideae (now retained within Euphorbiaceae s.str.). A comparative wood anatomical study (Stern, 1967) failed to correct the misplacement. Webster (1982) eventually pointed out that Kleinodendron is conspecific with Savia dictyocarpa.

In the last pre-molecular classifications of Euphorbiaceae s.l., Webster (1994b) and Radcliffe-Smith (2001) recognized Blotia, Gonatogyne, Petalodiscus, and Savia at the generic level. Both unfortunately continued to place African and Madagascan species in Savia, an error that is corrected here. For a more detailed discussion of the old world taxa placed in Savia by various authors, see Hoffmann et al. (2006) and Hoffmann and McPherson (2007).

Morphology FLOWER MORPHOLOGY

The flowers of Gonatogyne, Heterosavia, and Savia are unisexual, usually petaliferous (with a biseriate perianth) and pentamerous with a trilocular gynoecium. The petals of staminate and pistillate flowers are alike in Gonatogyne whereas petals of pistillate flowers are thicker and longer relative to the sepals in Heterosavia and Savia than those of the staminate flowers. This sexual dimor- phism is strongest in Savia. Staminate flowers of Savia sessilifora have a mixture of normally developed spathulate petals and petals that are reduced to subulate structures that hardly extend beyond the disc (Michaelis, 1924: Fig. 1); often petals are absent. Some-

times all three states can be observed in the same flower, containing only some of the petals, some of which can be reduced. Staminate flowers of S. dictyocarpa lack petals (Fig. 1D) altogether, which was the likely reason for its incorrect generic affilia- tion in the past. Miuller (1874) had no staminate flowers when he first described this species correctly in Savia but the lack of staminate petals later led Kuhlmann (1935) to describe it again in Securinega, and Smith and Downs (1964) to describe it as a new genus Kleinodendron. Webster (1982: 536) wrote of the flower: "The perianth parts show a tendency towards spiral arrangement. ... In the staminate flower the perianth is usually 5- 7-merous and could be interpreted either as apetalous with 5-7 sepals, or as having 5 sepals with 0-2 petals in the inner 'whorl'...". When considering both species of Savia and examining the clawed spathulate petals careful- ly, the second interpretation becomes much more likely. The presence of well-developed petals in the pistillate flowers (Fig. 1G, H) and in both sexes of the sister genus Gonatogyne also supports this interpretation. The stronger degree of petal reduction in staminate relative to pistillate flowers ofHeterosavia and Savia is rare in Phyllanthaceae and Euphorbiaceae. Staminate petals are more frequently larger than pistillate ones (e.g., in Andrachne and Leptopus Decne., see Hoffmann, 1994: Fig. 1A-D, or in Croton L.). A case similar to Heterosavia and Savia is Paranecepsia Radcl.-Sm. (Euphorbiaceae) which has apetalous staminate flowers but well-developed petals in the recently discovered pistillate flowers (Vorontsova & Hoffmann, 2007).

All three genera possess a well-developed extrastaminal floral nectar disc in both sexes. In Heterosavia and Savia the disc is annular and more or less strongly altemipetalously crenate (Fig. 1A-E; Hoffmann, 1994: 5, Fig. 1F). The disc of Gonatogyne (Fig. 1F-H) consists of numerous (10-20) semi-globose to truncate, basally fused glands that are slightly variable in shape and size. The peculiar disc morphology was the main argu- ment for the recognition of Gonatogyne as a monotypic genus (Miiller, 1873: 13; Pax & Hoffmann, 1933; Webster, 1994b). The disc of Gonatogyne is markedly different from that of




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B. Pistillate flower, Alain 749 (HAJB). C. H. erythroxyloides, pistillate flower, Luna 525 (NY). D-E. Savia dictyocarpa. D. Staminate flower, part of perianth removed, Kuhlmann IBSP 55632 (SP). E. Pistillate flower, note the staminode and the different-sized petals, part of perianth removed, Kuhlmann 3687 (SP). F-H. Gonatogyne brasiliensis. F. Floral disc of a staminate flower, Hoehne IBSP 29617 (SP). G. Staminate flower, part of perianth removed, Hoehne 32203 (A). H. Pistillate flower, part of perianth removed, note the two staminodes, Hoehne 27387 (A). Scale bar= 1 mm.




its sister genus Savia but not dissimilar to that of species of closely related (see introduction) Discocarpus (see Hayden & Hayden, 1996: Figs. 10A, 11D).

Staminodes are present in all three genera discussed in this paper. They are alternipetalous, subulate structures without distal thick- enings. Staminodes are relatively common in Euphorbiaceae s.1. (Pax, 1924: 130; Pax & Hoffmann, 1931: 13). They were also found in Discocarpus (Klotzsch, 1841; Baillon, 1858) but were not mentioned by Hayden and Hayden (1996). The illustration in Klotzsch (1841: 201, Fig. 9Cc) clearly shows the staminodes bearing anthers with two thecae. Intriguingly, their position was reported as altemisepalous by Baillon (1858: 585, Fig. 22.1) and Miiller (1866: 223). The gynoecium is spheroidal to pyriform with 3 (rarely 2 or 4) locules. Each locule contains two pendulous anatropous ovules topped with an obturator. The ovary indumentum is diag- nostically significant at the species level. The styles in Heterosavia and Savia (Fig. 1B, E) are 2-fid whereas Gonatogyne has irregularly bent or twisted 4-fid styles (Fig. 1H) reminiscent of those of Pentabrachion Miull. Arg. (also in tribe Bridelieae; Pax & Hoffmann, 1922: 189, Fig. 15C, 1933; and pers. observ.). The pistillode is usually 3-fid in all three genera, but sometimes its branches are 2-fid again in Gonatogyne.

POLLEN MORPHOLOGY

Pollen of all species of Gonatogyne, Heterosavia, and Savia was examined for this study, and key measurements are given in Table I. In accordance with other characters, the individual species of both Heterosavia and Savia are palynologically uniform. Webster (1982: 537) described the pollen of Savia dictyocarpa as: "... subglobose, 3-colporate, angulaperturate, reticulate-semitectate, with diameters of 25-30 tm; they are very similar to the pollen grains of Savia sessilifora ... " However, there are significant differences between the macromorphologically similar Heterosavia and Savia. Heterosavia species (Fig. 2) have tectate-perforate pollen with an average polar/equatorial ratio of 1.3 and colpi that extend on average to 91% of the length of the entire pollen grain. In contrast, the

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pollen grains of Savia and Gonatogyne are microreticulate, more spherical (average pole/equator ratio= 1.15), and have shorter colpi relative to the pollen length (79% and 75%, respectively). Pollen of Heterosavia erythroxyloides (Griseb.) Petra Hoffm. and Savia sessilflora was comparatively examined by Punt (1962). He classified each in a different major type ("configuration"), within which Heterosavia erythroxyloides was grouped with other members of tribe Phyllantheae in the "Phyllanthus pentaphyllus subtype" whereas Savia sessiliflora was placed in the "Savia type" with the Madagascan species of Wielandia previously regarded as Savia, some members of tribe Bridelieae where Savia also belongs, as well as some other taxa unrelated within Phyllanthaceae. There are no previous data published on Gonatogyne pollen.

FRUIT AND SEED MORPHOLOGY

The fruits of Gonatogyne, Heterosavia, and Savia are explosively dehiscent schizocarps typical for most Phyllanthaceae, Picrodendraceae, and Euphorbiaceae. They are subglobose to ellipsoid, more or less 3-lobed (rarely 2- or 4-

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I.CBE D FIG. 2. Pollen of Heterosavia maculata var. clementis.

A. Equatorial view, mesocolpium. B. Equatorial view, colpus. C. Colpus, detail. D. Mesocolpium, detail. Clemente 4983 (NY). Scale bar=2 tm.

lobed), and brownish when mature. The mature fruits dehisce septicidally, loculicidally, and septifragously into three mericarps ("cocci") and a persistent columella. This autochorous dispersal expels the seeds several meters. It can be observed, accompanied by a loud noise, in drying herbarium material. Systematically significant differences are found in the fruiting pedicel (articulated or not), in the shape of the columella, in the shape of the dehiscence line splitting the septa, and in the persistence of the perianth. The shape of the columella depends on the dehiscence lines and on the shape of the seeds. In Gonatogyne (Fig. 3F) and Savia (Fig. 3D), the distal parts of the columella are narrow and parallel-sided in lateral view, with Gonatogyne having a far more pronounced basal thickening than Savia. Heterosavia (Fig. 3B), on the other hand, has a columella that is widely winged distally. The septa of Heterosavia and Savia split in a single, long and clean line that runs parallel to the pericarp (Fig. 3A, C), whereas those of Gonatogyne split irregularly and often in several places (Fig. 3E). The perianth falls off before fruit maturity in Gonatogyne (Fig. 3D) and Savia (Fig. 3F) but persists after fruit dehiscence in Heterosavia (Fig. 3B). Gonatogyne and Savia also share articulate pedicels (Fig. 3D, F). The described fruit characters are also diagnostic at generic level in other Phyllanthaceae genera.

The seeds are ecarunculate, more or less isodiametric, smooth, and dry (Fig. 4A, C, E). Stuppy (1996) only examined Savia sessiliflora among the taxa discussed here. He classified this species in his Phyllanthus group, Andrachne subgroup, together with Actephila Blume, Andrachne, Leptopus (now all tribe Poranthereae), and Phyllanthus welwitschianus Miull. Arg. (tribe Phyllantheae). The anatropous ovules go through unequal growth processes to form a more or less campylotropous seed whereby hilum and micropyle stay directly adjacent. The shape of the mature seed depends primarily on whether one or two seeds develop per locule. In Gonatogyne and Heterosavia both ovules usually ripen into seeds with a resulting triquetrous shape resembling segments of an orange (Fig. 4A, E) but in Savia one of the two ovules is usually aborted, resulting in a rounded, ellipsoid to globose seed (Fig. 4C).




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FIG. 3. Fruits. A, B. Heterosavia. A. H. bahamensis, mericarp with two seeds, Correll 47479 (NY). B. H maculata var. clementis, columella, note the persistent perianth, Clemente et al. 4036 (HAC). C, D. Savia sessiliflora. C. Mericarp. D. Columella, note the articulate pedicel, PFC (Bisse et al.) 38141 (HAJB). E, F. Gonatogyne brasiliensis. E. Mericarp, Hoehne 27387 (A). F. Columella, note the articulate pedicel, Sellow s.n. (G). Scale bar=5 mm.




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All three genera have similar embryos with copious endosperm and thin, flat, straight cotyledons (Fig. 4B, D, F, G). This is the predominant embryo type in Phyllanthaceae and the other Euphorbiaceae s.1. families. The cotyledons are oriented either parallel to the pericarp or to the septa depending on whether one (Savia) or two (Heterosavia and Gonatogyne) seeds develop per locule. Because the cotyledons rotate 900 in the exceptional cases when both ovules come to maturity in Savia, this orientation is not taxonomically determined and appears to de- pend entirely on the spatial conditions in the developing seed. The same in reverse happens when one of the ovules in Gonatogyne and Heterosavia aborts. The genera also differ with respect to the relative length of their radicles. It is shortest and broadest in Savia (5-6 times shorter than the cotyledons and about as long as wide; Fig. 4D), whereas radicles of Gonatogyne (Fig. 4F, G) and Heterosavia (Fig. 4B) are only 3-4 times shorter than the cotyledons and 3-4 times longer than wide.

LEAF MORPHOLOGY

Leaves in Gonatogyne, Heterosavia, and Savia are uniformly petiolate, simple, symmetrical, entire, penninerved, eglandular, and glabrous or with a simple indumentum. A clear-cut difference is that Heterosavia has adaxially channeled to 2-winged petioles whereas those of Gonatogyne and Savia are terete. A study of foliar morphology (Levin, 1986a, b, c) showed that, despite the relative uniformity of leaf characters in Phyllanthaceae, systematically meaningful results can be gained from their analysis. Apart from Levin's (1986a, b, c) comprehensive study, previous publications on leaf anatomy of Euphorbiaceae s.1. included only Savia sessiliflora out of the three genera discussed here (Rothdauscher, 1896: 135; Dehay, 1935).

Leaf venation. Leaf venation of the three genera was examined for this study using leaves from herbarium material that had been cleared over several weeks to months in 10% NaOH, and were briefly cleared further in 10% NaOC1. They were then stained in an

alcoholic solution of safranine and mounted in Canada balsam. Leaves from several specimens of each taxon were examined. The photo- graphs were taken directly from the slides onto photographic paper (without film). For terminology and categories used here see Hickey (1979). Taxon-specific variation was found in overall venation pattern, marginal venation, areolation, course and relative thickness of the individual orders of venation, and outer loops. Characters such as number and density of secondary veins, divergence angles, and presence and course of intersecondaries were, in contract, found to be as variable within as between taxa.

Overall venation in Heterosavia (Fig. 5) is brochidodromous as in the vast majority of Phyllanthaceae. There are clear differences in leaf venation between its macromorphologically very similar species. Venation yields the most divergent characters for distinguishing the two ecologically vicariant species Heterosavia bahamensis (Britton) Petra Hoffn. and H. maculata (Urb.) Petra Hoffinm. While the two rows of secondary loops in Heterosavia maculata (Fig. 5E) are always fused into a thick continuous fimbrial vein, the ultimate marginal venation of H. bahamensis (Fig. 5B) consists of free ending veinlets that are only partially straightened into a thin fimbrial vein. Heterosavia maculata also tends to have areolation with incompletely closed meshes compared with H. bahamensis that has (more highly ordered) imperfectly developed areoles and generally thinner and denser veins throughout. Heterosavia maculata shows stronger xeromorphic traits in its venation than H. bahamensis although both species are adapted to xerophytic habitats (serpentine and calcareous coastal scrub, respectively; see also under Biogeography and ecology). This also manifests itself in the thicker leaves and the peculiar shape of the outer stomatal cavity (see below and Fig. 10OB) in Heterosavia maculata. The venation of Heterosavia erythroxyloides (Fig. 5F) and H. laurifolia (Griseb.) Petra Hoffm. stands between the species mentioned above, with the incompletely closed meshes of H. erythroxyloides again probably linked to strong xeromorphic adaptation. A correlation of a reduced level of organization in the leaf




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FIG. 5. Leaf venation of Heterosavia, from cleared leaf material. A-C. H. bahamensis, A, B Coker 160 (NY), C Britton 1859 (NY). D, E. H. maculata var. maculata, PFC (Alvarez et al.) 57030 (B). F. H. erythroxyloides, PFC (Bisse et al.) 36062 (B). Scale bar= 1 cm (A, C, D, F) and 1 mm (B, E).

venation and xeromorphism is typical of many dicots (Hickey, 1971).

The brochidodromous leaves of Savia (Fig. 6A-D) often have well-developed areoles besides the imperfect ones, resulting in a generally more highly ordered higher venation than in Heterosavia. The marginal venation is a combination of fimbrial vein,

loops, and free ending veinlets (Fig. 6B). The two species of Savia show only slight quantitative differences. The venation of Gonatogyne (Fig. 6E, F) shows a tendency to dissolution of the basal secondary vein loops and continually curved secondary veins (compared to the sharp curves in the distal parts of the secondaries in the other two




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genera). Levin (1986a: 31) suggested that this "weakly brochidodromous" pattern may be a link between brochidodromous and eucamptodromous patterns. The teriaries form a loose, irregular network framing extraordi- narily fine (not continuously sclerified), incompletely closed higher-order venation. A similar secondary venation pattern was found in Discocarpus and Pentabrachion (Levin, 1986a), which are now also placed in tribe Bridelieae (sensu Hoffmann et al., 2006). Gonatogyne (Fig. 6F) has ultimate marginal veins that are free ending, exmedially branched, and distally conspicuously scleri- fled, interspersed with dissolving loops of 3rd and 4th order veins. This condition is reminiscent of Wielandia platyrachis (Baill.) Petra Hoffm. & McPherson and W elegans Baill., and was there termed "exmedial" ultimate marginal venation (Hoffmann & McPherson, 2007). It finds its strongest ex- pression in Wielandia bojeriana (Baill.) Petra Hoffm. & McPherson and W ranavalonae (Leandri) Petra Hoffm. & McPherson where the thick exmedial ultimate marginal veins run parallel to each other so as to resemble a comb.

Leaf anatomy. The leaf anatomical study of all taxa of Gonatogyne, Heterosavia, and Savia proved systematically relevant especially regarding the configuration of the stomatal ledges, the course of the anticlinal walls, the sclerification of higher vein orders, and the configuration of the petiolar vascular bundles. Sections were obtained by softening petioles and leaf fragments from herbarium material for ca. 7 days in equal parts of glycerin, 96% ethanol, and distilled water (Strassburger's mixture), and sectioning at 10-25 tm with a freezing microtome. Petiolar cross sections were made at three points for each taxon: at the base, in the middle, and distally just below the base of the leaf blade. To examine the epidermis, leaf fragments were macerated in Schulze's mixture (KCO103+ 60% HNO3) and 25% ammonia, and cleaned mechanically under a dissecting microscope. All were stained with aqueous solutions of safranine or toluidine blue, and mounted in glycerin jelly. Measurements were taken from slides prepared from two or three different

samples per species. Leaf surfaces were sput- tered with gold and photographed (150-4000x) using SEMs. Terminology follows Barthlott and Ehler (1977) for general epidermal and cuticle characters, and Wilkinson (1979) and van Cotthem (1971) for stomata. The differ- entiation between subsidiary cells and ordinary epidermis cells was at times problematic. The two types can be morphologically indistin- guishable (Fig. 8), but clearly behave differently when stained with toluidine blue, so that they cannot easily be placed in either category according to Wilkinson (1979: 97-98). All cells directly adjacent to guard cells are here referred to as subsidiary cells.

The leaf blades of Gonatogyne (Fig. 7D), Heterosavia (Fig. 7A, B), and Savia (Fig. 7C) are typical of most dicots: bifacial, hyposto- matic, with a one-layered epidermis, sclerenchymatic and/or collenchymatic vascular sheaths, and lacking a hypodermis. Epicutic- ular waxes are rare and if present, irregularly distributed and not constant for a given taxon. Trichomes on lamina and petiole are un- branched, uniseriate, and 1- to 5-celled (Fig. 8B). Savia has S-undulating anticlinal walls whereas those of Gonatogyne and Heterosavia are straight. Key measurements are given in Table II. Adaxial periclinal walls are thicker and straighter than their abaxial counterparts, with xeromorphic Heterosavia, particularly H. maculata, having markedly thicker periclinal walls than the other, mesomorphic species. The stomatal guard cells of Gonatogyne and Heterosavia are larger than those of Savia. All three genera have paracytic stomata (Fig. 8) that are predominantly laterocyclic, but also parallelocytic, brachy- paracytic or intermediate (see also Levin, 1986a: 38, 45, 70). A small proportion of stomata in each sample had more than two subsidiary cells, which is likely to be due to subsequent cell divisions as the anticlinal walls between them are thinner than those between other cells (see also Rothdauscher, 1896: 136). Rarely all three or four subsidiary cells were of equal thickness. Stomata sharing a subsidiary cell were observed regularly in all species. "Water stomata", which were recorded by Levin (1986a: 39, 55) only for some species of Andrachne, were observed in almost all species examined in this study.




FIG. 7. Leaf anatomy, light microscopy. A-D. Leaf blade cross sections. A. Heterosavia maculata var. clementis, note the fimbrial vein, PFC (Bisse et al.) 44142 (B). B. H. bahamensis, PFC (Areces et al.) 34173 (B). C. Savia sessiliflora, PFC (Areces et al.) 33128 (B). D. Gonatogyne brasiliensis, note the sunken stomata, Hoehne 27387 (SP). E-H. Petiole cross sections. E. Heterosavia bahamensis, PFC (Areces et al) 34173 (B). F. Savia sessilifora, Duke 7195 (MO). G, H. Gonatogyne brasiliensis, Rosa & Pires 3982 (SP). G. Basal part. H. Distal part. Scale bar=0.1 mm (A-D) and 0.5 mm (E-H).

They are considerably larger than normal stomata and are surrounded by four to six subsidiary cells. Differences between genera were also found in cross sections (Fig. 10) and scanning micrographs (Fig. 9) of the abaxial leaf surfaces. Inner and outer stomatal ledges are developed in Gonatogyne (Fig. 10C) and Heterosavia (Fig. 10D) whereas in Savia (Fig. 10A) only the outer stomatal ledges are present. Stomata of Heterosavia maculata (Figs. 9B, C and O10B) are a special case of the first type with the outer ledges incorporated in the massive cuticle. Transpiration is further minimized by having the stomata sunken into the epidermis in Gonatogyne (Figs. 9E and O10C) and Heterosavia maculata (Figs. 9B and O10B).

Sclerenchymatic sheaths and caps surrounding the vascular bundles are more stongly developed in xeromorphic Heterosa- via (Figs. 5E and 7A; except mesomorphic H. laurifolia) than in other taxa in this study. Gonatogyne presents the opposite end of this scale. Here the higher order venation is, except for the ultimate margin, only partially accompanied by sclerenchyma (Figs. 6F and 7D). While the supporting tissue surrounding the vascular bundles of the secondary and higher order veins in all three genera is exclusively sclerenchymatic, there is along the midvein also subepidermal collenchyma. This collenchymatic layer is responsible for

the prominent relief of the midvein in both live and dried leaves. In the petioles there are two types of collenchyma, the subepidermal fibers with large inner diameters also found in the lamina, and a clearly defined layer of perivascular collenchyma consisting of thick-walled fibers with very small inner diameters (Fig. 7E-H). Petiolar sclerenchyma is gradually replaced by collenchyma in Gonatogyne whereas Heterosavia and Savia often have substantial abaxial sclerenchymatic caps. The configuration of the petiolar vascularization in Euphorbiaceae s.l. is diverse, as shown by Dehay (1935), who only included Savia sessiliflora among the species studied here. Examining all other species of the three genera, petiolar vascular bundles of Heterosavia (Fig. 7E) and Savia (Fig. 7F) are found to have a U-shaped cross section throughout their length. In Gonatogyne (Fig. 7G, H) cross sections of the petiolar vascular bundle change from basally shallowly U-shaped with inrolled sides to cylindrical from the middle to the distal end of the petiole.

Biogeography and Ecology

Savia s.1. was frequently cited as an exam- ple of a disjunct African-American distribution (Pax, 1924: 174; Pax & Hoffmann, 1931: 29; Leandri, 1937: 95; Webster, 1967: 325,




FIG. 8. Abaxial leaf epidermis, light microscopy. A. Heterosavia maculata var. clementis, PFC (Bisse et al.) 44142 (B). B. Savia dictyocarpa, note the trichomes, Kuhlmann 3202 (SP). C. Gonatogyne brasiliensis, Hoehne 27387 (SP). Scale bar=50 .m.

1994a: 21; Howard, 1973: 35; Borhidi, 1985: 23, 1991: 262). Molecular phylogenetic analyses (Wurdack et al., 2004; Kathriarachchi et al., 2005) have disproved this theory and shown that Savia s.1. consists of several more or less unrelated clades within Phyllanthaceae, each with a coherent distribution. As with other Phyllanthaceae, there are no safely identified

fossils to allow inferences about their historical distributions. The biogeography and ecology of Savia and Gonatogyne will be discussed in a forthcoming publication (Hoffmann and Cordeiro, unpubl. data).

Heterosavia is restricted to the Caribbean, with the center of diversity in Cuba where all four species occur. Cuba is the largest and oldest of the Greater Antilles islands, and that with the most diverse flora and the highest percentage of endemics (Howard, 1973: 21, 31; Borhidi, 1991: 216). There are few recent collections of and hardly any ecological data on the Cuban endemic Heterosavia laurifolia (Fig. 14). The three remaining species, although morphologically very similar, are ecologically specialized. Heterosavia maculata (Fig. 15), also endemic in Cuba, occurs throughout the island but is limited to serpentine substrate (see discussion below). This is a typical distribution pattern of many Cuban endemics (Borhidi, 1991: 249). The morphologically most similar species Heterosavia bahamensis (Fig. 12) occurs only in calcareous coastal scrub. Apart from being found almost along the entire Cuban coast, it also occurs along the coasts of neighboring islands such as Jamaica, the Florida keys, the Bahamas, Caicos, Cayman, and Swan Islands. Flueggea acidoton (L.) G.L. Webster in Phyllanthaceae (Webster, 1984: 300) and Picrodendron baccatum (L.) Krug & Urb. ex Urb. in Picrodendraceae (Borhidi, 1991: 227) have a similar distribution and ecology. The morphologically more clearly defined fourth species, Heterosavia erythroxyloides (Fig. 13), occurs in the moun- tains of Cuba and Hispaniola, but in contrast to Heterosavia maculata is not restricted to the geologically oldest Cuban mountain ranges and more commonly found in the younger Sierra Maestra. It seems to prefer calcareous substrates and avoids serpentine.

Although Heterosavia maculata is reported as one of the 7% of Cuban endemics that occur exclusively on serpentine (Borhidi, 1991), the study of nickel concentrations in leaves of this species (see below) suggests that it occasionally occurs on soils of lower nickel availability than is usual for serpentine. Serpentine ecosystems are characterized by high concentrations of heavy metals (including nickel) on the one hand, and a lack of




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certain trace elements and a general shortage of nutrients on the other hand, resulting in slow metabolism. The typical xeromorphic aspect of the serpentine vegetation is better described as pseudoxeromorphic or peinomor- phic (= starvation shaped) (Borhidi, 1991). In a study of nickel accumulation in plants on serpentine habitats in Cuba leaf samples from 34 different specimens of Heterosavia and Savia were analysed (R. D. Reeves, pers. comm.; Reeves et al., 1996, 1999). Geologi- cally indifferent Savia sessilflora showed very low Ni content regardless of the substrate, and the calciphile Heterosavia bahamensis also had expectedly low Ni contents. The five examined samples of Heterosavia maculata var. clementis (Alain) Petra Hoffm. had low- to-intermediate Ni-values (40-850 gg/g). Twenty of the 25 examined samples of Heterosavia maculata var. maculata were Ni hyperaccumulators (1200-4900 gg/g). The fact that five samples of this taxon showed a low-to-intermediate Ni content means that var. maculata might not be a strict serpentine endemic, however, sometimes the two varieties are morphologically intergrading. Heterosavia maculata var. clementis has a somewhat distinct ecological profile in apparently favor- ing wetter habitats, and its large, reddish leaves are atypical of the normally small-leafed, grayish green serpentine vegetation (Howard, 1973: 9). Label data from herbarium sheets, the only source of ecological information for this species, is vague but records laterite rather than serpentine. It is possible that var. maculata occurs in habitats where serpentine is the direct substrate, whereas var. clementis grows on serpentine-derived laterite soils. This and the comparably higher precipitation of the collect- ing localities of var. clementis in northeastern Oriente (Fig. 15) could explain the morphological differences between the varieties. Borhidi (1991: 253) listed several infraspecific taxa with large-leafed forms restricted to the Sierra de Moa and Sierra de Baracoa while their small- leafed relatives are found elsewhere, especially in the Sierra de Nipe and Sierra Cristal.

All species of Heterosavia are evergreen (West & Arnold, 1956: 112 for H. bahamensis) whereas Savia is mainly deciduous. Hetersavia flowers during the rainy season in April to June, and fruits mostly between May and August (rarely to January).




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FIG. 9. Abaxial leaf surface, scanning electron microscopy. A. Heterosavia laurifolia, Wright 1430 (GOET). B, C. H. maculata var. clementis, Clemente 4982 (NY). D. Savia dictyocarpa, Glaziou 5986 (P). E, F. Gonatogyne brasiliensis, Hoehne 32203 (A). Scale bar= 10 xm (A, C, D, F) and 50 tm (B, E).

Key to the neotropical genera previously included in Savia s.l.

1. Leaves abaxially pilose throughout; sepals 4-5 mm long, twice as long as wide; filaments fused for 3/4 of their length; styles twice 2-fid; septa of cocci tearing irregularly; disc consisting of 10-20 basally fused glands; petiolar vascular bundles in cross section U-shaped at the base, cylindrical from the middle to lamina ............... Gonatogyne

1. Leaves glabrous or pilose only basally and along the midvein; sepals up to 2.5 mm long, about as long as wide; filaments free or fused for up to half of their length; styles 2-fid; septa of cocci with only one neat dehiscence line; disc 5-crenate, irregularly undulate or entire; petiolar vascular bundles in cross section U-shaped throughout. 2. Petiole 2-winged to channelled adaxially, glabrous; leaves evergreen; leaf blade elliptic to obovate, acuminate

to retuse apically, gradually narrowed into the petiole; perianth persistent in fruit; pedicels not articulated; columella distally distinctly 3-winged, 2-3 mm wide, 1.5 to 2 times as long as wide; seeds mostly 2 per locule, triangular in cross section with an acute micropylar pole; cotyledons 3 to 4 times as long as radicle; radicle 3 to 4 times as long as wide; anticlinal walls of leaf epidermis straight ....................... Heterosavia

2. Petiole terete, pubescent; leaves deciduous; leaf blade ovate, always acute to acuminate apically, abruptly narrowed into the petiole; perianth caducous in fruit; pedicels articulated; columella distally narrowly 3- angular, ca. 1 mm wide, 2 to 3 times as long as wide; seeds mostly 1 per locule, elliptic in cross section, rounded on all sides; cotyledons 5 to 6 times as long as radicle; radicle about as long as wide; anticlinal walls of leaf epidermis undulating ..................... ............

..................... Savia




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Taxonomic treatment

This revision is based predominantly on herbarium material. Measurements of floral parts are taken from softened flowers, all other data from dry material, if not stated differently.

Heterosavia (Urb.) Petra Hoffm., stat. nov. Savia sect. Heterosavia Urb., Symb. antill. 3: 284. 1902. Lectotype, here designated: Heterosavia erythroxyloides (Griseb.) Petra Hoffm. (= Savia erythroxyloides Griseb.)

Dioecious trees or shrubs with sparse to moderately developed simple indumentum; evergreen. Young twigs terete, with lenticels. Stipules persistent, deltoid to ovate. Petiole 2- winged to channelled adaxially, glabrous; petiolar vascular bundles in cross section U- shaped with steep sides throughout. Leaves evergreen, alternate, petiolate, simple; leaf blades symmetrical, entire, penninerved, obovate to elliptic or oblong, retuse to acuminate apically, acute to rounded basally, gradually narrowed into the petiole, glabrous, coriaceous; anticlinal walls of leaf epidermis straight; stomata paracytic, very rarely with 3 to 4 subsidiary cells. Flowers axillary, glomerulate, 2-5 x2-5 mm, 5-merous (rarely 4- or 6-merous) except for the gynoecium.

Pedicel not articulated. Staminate flowers 10 to 25 per inflorescence, subsessile. Pistillate flowers solitary, rarely in twos or threes, with terete pedicels 1-7 mm long. Sepals imbri- cate, quincuncial in pentamerous flowers, with hyaline margins. Petals of the staminate flowers shorter than the sepals, always well- developed, those of the pistillate flowers as long as or hardly shorter than the sepals. Disc extrastaminal, annular, alternipetalously crenate to entire, narrow, thick. Stamens alternipetalous; filaments free or basally fused for up to half of their length, subulate; anthers introrse, dehiscing longitudinally, sometimes pilose. Staminodes sometimes present, alternipetalous, subulate. Pistillode 3-fid to the base or nearly so, the branches erect. Ovary 3-, rarely 4-locular; styles 3, rarely 4, 2-fid to about half their length; stigmas acute to obtuse. Fruits explosive schizocarps, solitary, rarely in twos, subglobose, 3-, rarely 4-lobed, glabrous or pilose; dehiscence septicidal, locu- licidal, and septifragous; septa of cocci with only one neat dehiscence line running parallel to the pericarp; columella 1.5-2 times as long as wide, the basal third 3-angular in cross section, the distal part obconical, widely 3-winged, the wings alternating with the basal angles; perianth persistent in fruit. Seeds 2 per locule (sometimes of unequal size), ecarunculate, smooth, +




marbled, dull to shiny, x triangular with one acute angle in cross section, very rarely only 1 seed per locule, then seed + triangular without acute angle in cross section; endosperm copious in mature seeds; cotyledons thin, straight, 3 to 4

times as long as radicle; radicle 3 to 4 times as long as wide.

Four similar species in the Caribbean, from Florida to Swan Islands (Honduras).

Key to the species of Heterosavia

1. Ovary and fruit pubescent. 2. Pedicel of pistillate flowers 1-2 mm long; fruiting pedicel 2-5 mm long; ovary and fruit always densely pubescent;

leaf blade usually shorter than 5 cm, drying mostly gray-green adaxially and reddish brown abaxially; Cuba and Hispaniola 2. H erythroxyloides

2. Pedicel of pistillate flowers 3-7 mm long; firuiting pedicel 8-15(-22) mm long; ovary and fruit moderately pilose; leaf blade usually longer than 5 cm, drying gray-green on both surfaces; endemic to Cuba 3. H laurifolia

1. Ovary and fruit glabrous. 3. Pedicel of pistillate flowers 3-7 mm long; fruiting pedicel 8-15(-22) mm long; leaf blades elliptic, usually

acuminate apically, drying gray-green, chartaceous to thinly coriaceous; endemic to Cuba 3. H laurifolia 3. Pedicel of pistillate flowers 1-4 mm long; fruiting pedicel 2-6 mm long; leaf blades obovate, elliptic, oblong,

orbicular or obcordate, obtuse, rounded or retuse, rarely acute to short-acuminate apically, drying yellowish green, green, olive or red-brown, usually strongly coriaceous.

4. Leaf veins thin and densely set, leaf margin bordered by free ending veins and outer loops, rarely for short distances straightened into a thin fimbrial vein, always several rows of outer vein loops present; leaf blades often elliptic and only thinly coriaceous; on calcareous substrate; coastal; from Florida to Swan Islands (H onduras) . 1. H . baham ensis

4. Leaf veins thick and distantly spaced, leaf margin bordered by a thick continuous fimbrial vein, usually only one row of outer vein loops present; leaf blades often obovate and strongly coriaceous; on serpentine and more rarely on igneous rock; inland; endemic to Cuba 4. H maculata

Identification: Pistillate (flowering or fruiting) specimens of Heterosavia can be determined with greater certainty than staminate or sterile material that can sometimes only be named with a good knowledge of the genus. If in doubt, refer to locality and substrate.

Typification: Ekman specimens from Stock- holm (S) here recognized as holotypes of Urban taxa have the species name and/or the word "typus" in Urban's handwriting on the label. All other types from S are designated as lectotypes. This applies also for specimens with an additional label in Urban's hand with a new determination (as a species newly described by Urban), as these additional labels were probably sent to S by Urban and mounted on the sheets there. In these cases it is assumed Urban described the taxa using material that was destroyed in Berlin (B).

1. Heterosavia bahamensis (Britton) Petra Hoffm., comb. nov. Savia bahamensis Britton, Torreya 4: 104. 1904. TYPE: Bahamas, New Providence, West Street Road, near Nassau, thickets, 7-8 Apr 1904, N. L. Britton 84 (holotype: NY). (Fig. 11)

Savia clusiifolia Griseb. non auct., Nachr. K6nigl. Ges. Wiss. Georg-Augusts-Univ. 6: 164. 1865; Cat. pl.

Cub.: 15. 1866; Hoffmann, Taxon 43: 465., 1994, nom. rej. prop.; Greuter et al., Int. Code Bot. Nomen.: 401., 1994; nom. rej. Savia clusiifolia Griseb. var. genuina Miill. Arg. in DC., Prodr. 15 (2): 230. 1866, nom. inval. according to Art. 24.3. (Greuter et al., 1994). TYPE: Cuba, circa Punta Brava prope Matanzas, Jun 1849, F I. X Rugel 321 (lectotype, here designated: GOET; isolectotypes: BM, K). The specimen cited by Miuller (1866: 231) under this name (Wright 1431) is Heterosavia laurifolia var. intermedia (Miull. Arg.) Petra Hoffm.

Savia perlucens Britton, Bull. Torrey Bot. Club 43: 464. 1916. TYPE: Cuba, Isla de Pifios, Caleta Grande, 9 March 1916, N. L. Britton, P Wilson & H Leon 15330 (holotype: NY).

Savia apiculata Urb., Repert. Spec. Nov. Regni Veg. 28: 209. 1930. TYPE: Cuba, Prov. Oriente, Santiago de Cuba, at Cabo Cruz south of the town, limestone hills, 26 Jun 1924, E. L. Ekman 19079 (holotype: S; isotypes: G, K, NY, S, US). See note on typification following species key.

Illustrations: Small, Man. s. e. fl.: 777. 1933 (fl, fr); West and Arnold, Native trees Florida: 112. 1956 (fl, fr, habit); Correll and Correll, Fl. Bahama Archip.: 843, Figs. 350 a-n. 1982 (fl, fr, habit); Levin, Ann. Missouri Bot. Gard. 73: 45. 1986 (leaf venation); Scurlock, Native trees & shrubs Florida keys: 145. 1987 (photo: fr, habit). Note that Small (1933) and West and Arnold (1956) illustrate densely pilose fruits which




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FIG. 11. Heterosavia bahamensis. A. Habit, pistillate plant, Brace 345 (K). B. Habit, staminate plant, Rugel 618 (K). C. Internodes with columellae, Ekman 19079 (K). D. Pistillate flower, Britton & Brace 670 (K). E. Staminate flower, perianth partially removed, Rugel 618 (K). F. Mericarp. G. Seed, lateral view, Ekman 19079 (K). H. Details of leaf venation, Britton & Brace 670 (K). Scale bar=2 cm (A, B), 4 mm (C), 2 mm (D, F, G), 1 mm (E), and 6 mm (H). Drawn by Hazel Wilks.




are very rare (two of 241 specimens examined); fruits of Heterosavia bahamensis are normally entirely glabrous.

Tree or shrub, (0.5-)2-4(-8) m tall (label of Proctor 36833: 15 m tall, see note below); twigs + pubescent when young, glabrescent, sometimes some intemodes pilose and others glabrous. Stipules deltoid, acute, 2-3(-4) x 1- 2.5 mm, usually pubescent abaxially when young, glabrescent. Petiole 2-5(-8) x ca. 1 mm, glabrous, rarely with scattered hairs. Leaf blade obovate or elliptic, sometimes oblong, rarely orbicular to transverse elliptic, apically acute, obtuse, rounded, truncate or retuse, sometimes apiculate or short-acuminate, sometimes with a terminal or subapical glandular dark mucro ca. 0.1 mm long, basally acute, obtuse or rounded, rarely narrowly cuneate, (1-)2-4.5(-8)x (0.7-)1.2-2.5(-4.5) cm, not to hardly revolute, glabrous, rarely with scattered hairs abaxially, coriaceous to thinly chartaceous, discolorous or concolorous, yellowish green to olive-green adaxially, usually shiny, often lighter and duller abaxially, midvein moderately prominent to flat adaxially, strongly prominent abaxially, higher venation prominent on both surfaces or only abaxially. Staminate flowers (6-)10-25 per inflorescence, 3-4 x 2-3 mm; bracts deltoid to ovate, acute, 0.5-1.5 x ca. 1 mm, pubescent abaxially; buds sessile, ellipsoid to globose. Sepals 5, ovate or oblong, obtuse to rounded, 1.5-2 x 1-2 mm, pilose to pubescent abaxially, sometimes fimbriate. Petals 5, spathulate to obdeltoid, erose to lacerate, usually emarginate, 1-1.5 x 1-1.5 mm, shorter than the sepals. Disc + distinctly crenate or lobed, the lobes altemipetalous, to 1 mm long. Stamens 5; 2-4 mm long; filaments free or basally fused for ca. 0.5 mm, 1.5-3 mm long; anthers ovoid to ellipsoid, 0.7-1.2 mm long. Pistillode 1-2 x ca. 0.5 mm, as long as or (if filaments are fused) slightly longer than the sepals, sparsely pilose to pubescent. Pistillate Jflowers solitary, 3-4x3-4 mm; bracts as in staminate flowers; buds subsessile or short- pedicellate, globose to ellipsoid. Pedicel 1-4 x

0.7-1.5 mm, glabrous to pubescent. Sepals 5, + orbicular, ovate or deltoid, obtuse to rounded, 1-2 x 1-2 mm, pubescent abaxially, sometimes fimbriate. Petals 5, spathulate to orbicular or oblong, entire to slightly erose, 1-2 x 1-

1.5 mm, about as long as the sepals, glabrous or sparsely pilose abaxially, sometimes + fimbriate. Disc x distinctly crenate, to 0.5 mm thick, the lobes altemipetalous, 0.5-1 mm long. Staminodes rarely present (Ekman 19079, US), 0.6-1.2 mm long. Ovary globose or slightly pyriform, ca. 2 mm long, glabrous (Correll 42738 and 50287-A: ovary pubescent); styles sometimes basally fused, straight to recurved, spreading, 0.5-1.2 mm long, glabrous. Fruits 4-5(-6) x 6-7(-9) mm, glabrous (Correll 42738, NY: pubescent), reticulate; fruiting pedicel 2-6 x 1-1.5 mm, glabrous to pubescent; columella 4-5 mm long, to 3 mm wide; exocarp 0.1-0.2 mm thick; endocarp 0.3- 0.6 mm thick. Seeds 3-5.5 x2-3.5 mm, brown to reddish.

Distribution and habitat. Caribbean: Florida Keys (United States), Bahamas, Caicos Islands, Cuba, Cayman Islands, Jamaica, Swan Islands (Honduras); Fig. 12. Coastal dry scrub (dog- tooth limestone, manigua costera, monte seco), in hammocks of Florida keys, rarely in pine forests, always on limestone, at 1-50 m, rarely to 400 m altitude (to 750 m for morphologically aberrant Jamaican population, see note below). Heterosavia bahamensis is a signature species of a coastal plant association in Cuba (Linociero-Savietum bahamensis; Borhidi, 1991: 560).

Conservation status. Heterosavia bahamensis has the widest distribution of any taxon in the genus. It appears to be common in favorable conditions (241 collections were examined for this study) and is therefore here assigned the conservation rating Least Con- cern (LC), following the criteria of the International Union for the Conservation of Nature and Natural Resources (IUCN, 2001).

Vernacular names. Bahamas: Bastard Cocoa Plum (Brace 1970); Maiden-bush (Britton & Millspaugh, 1920: 218); Maiden Bush (Correll & Correll, 1982: 842); Wild Cherry (Byrne 365). Cayman Islands: Wild Coco Plum (Proctor, 1984: 520; Kings LC 64). Cuba: Hicaquillo hembra/macho (Clemente 7054, 7055); Hoja redonda (Urban, 1930: 210; Alain, 1953: 42; Ekman 7722, 8703,




84 800 760 I 72x

680

24o

o o

V20

ge-- 200

I . . . . . I I I 0 100 200 300 Kilometres

16x 84

180x 176

172x

68'

FIG. 12. Distribution of Heterosavia bahamensis.

9183); Icaquillo macho (Alain, 1953: 42); Siguaraya (Ekman 7722, 9183). United States (Florida): Bahama maidenbush (Little, 1953: 400; West & Arnold, 1956: 112); Maidenbush (Little, 1978: 241); Maiden-bush (Small, 1933: 777); Maiden Bush (Long & Lakela, 1971: 547; Scurlock, 1987: 145).

Selected specimens examined HONDURAS. SWAN ISLANDS: Coral Rock, 2 Apr 1913, Nelson 108 (GH).

BAHAMAS. ABACO: Above ocean at Schooxner Bay, windblown coastal coppice on hillside, 27 May 1979, Correll 50687 (MO, NY). ACKLIN'S ISLAND: 4434 (NY); Boathouse cove, 12 Dec 1905-6 Jan 1906, Brace 4468 (NY, US). Andros: Deep Creek, 27 Jun 1890 (A, GH: Fresh Creek, 5 Jun), Northrop & Northrop 610 (A, GH, K, NY), paratype. CAT ISLAND: E of Pidgeon Cay, limestone, thin red soil, tall thicket (4-5 m), 20 May 1968, Byrne 489 (A, MO, NY). CROOKED ISLAND: Vauxhall, 9-23 Jan 1906, Brace 4718 (NY). ELEU- THERA: Cape Eleuthera, coastal coppice, 18 May 1975, Correll & Hill 45304 (NY). EXUMA: W of Moss Town, coppice-covered high rocky hills, 14 Dec 1976, Correll 47916 (MO, NY). GRAND BAHAMA: West End, 16 Apr- 8 May 1905, Brace 3504 (K, NY, US). INAGUA: Mathew town to Lower Savannah, scrub, 4 Nov 1904, Nash &

Taylor 1439 (K, NY). LONG CAY: Foursyth Road, 7-17 Dec 1905, Brace 4221 (NY, US). MAYAGUANA: 10 miles W of Abraham's Bay, 5-6 Dec 1907, Wilson 7442 (GH, K, NY). NEW PROVIDENCE: Low Coppet, bastard crab wood, 24 Jun 1903, Coker 157 (NY), paratype. SAN SALVADOR: E of Granny Lake, E side of Island, open thicket, 14 Jun 1978, R.R. Smith 4906 (NY).

CAICOS ISLANDS (U.K.) NORTH CAICOS: Along road 3-4 miles SE of Bottle Creek, coppice along road, 11 Jun 1974, Gillis 12250 (A, B, MO). PINE CAY: NE side, in coppice on coastal ridge, 28 Aug 1974, Correll 43253 (NY).

CUBA. Without precise locality, 1860-64, Wright 1926 (BM, GH, G-DC [only microfiche seen], K, MO), syntype of Savia clusiifolia (often also other species mounted on sheets). CAMAGUlEY: Mun. Nuevitas, al sur de Playa Santa Lucia, monte seco, 7 May 1976, PFC (Areces et al.) 31218 (B, HAJB, JE); Peninsula Pastelillo, caliza, manigua costera, 27 Apr 1984, PFC (Bisse et al.) 54134 (B, HAJB, JE); Silla de Cayo, Cayo Romano, 9- 11 Oct 1909, Shafer 2519 (BM, GH, NY, US). HABANA: E of Tarari, coastal thickets, 14 Oct 1921, Ekman 13322 (G, K, NY); 10 Apr 1917, Lion 7155 (GH, HAC, NY); Isla de la Juventud, Camino de Cayo Piedras a punta del Este, caliza, monte seco, 5 Apr 1980, PFC (Bisse et al.) 41927 (B, HAJB, JE). LAS VILLAS, Casilda, coastal thicket, 16 Mar 1901, Britton & Wilson 5602 (NY); Along Camino de la Costa E of Castillo de Jagua,




limestone outcrop, 1-20 Jul 1950, Howard et al. 242 (A). MATANZAS: Varadero, near Cardenas, limestone, beach, 8 Feb 1916, Britton & Wilson 14044 (NY); Mun. Playa Larga, peninsula de Zapata, El Sibalo, monte seco, 20 Apr 1977, PFC (Bisse et al.) 34511 (B, HAJB, JE); Ad rupes supra Punta Brava, Rugel 618 (GOET, K, NY). ORIENTE: Ciudamar, near Santiago, manigua, Nov 1946, Clemente 5138 (GH, HAC, NY); Granma, Mun. Pil6n, ensefiada de la Mora, monte seco, 28 May 1988, PFC (Alvarez et al.) 65461 (B, JE); Guantinamo, Mun. Maisi, camino entre La Tinta y Jauco por el barranco, caliza, bosque de galeria y vegetaci6n de farrallones, 2 Jun 1982, PFC (Giinther) 47426 (B, HAJB, JE). PINAR DEL RIo: Insul. in Cabafias Bay, solo calcar, 25 May 1920, Ekman 10932 (MICH); Mun. Sandino, peninsula de Guanahacabibes, cerca de Cabo Corrientes, monte seco, Mar 1967, PFC (Bisse & Rojas) 3054 (HAJB, JE); Carabela Grande, Cabo San Antonio, 9 Apr 1924, Roig 3202 (HAC, NY).

CAYMAN ISLANDS (U.K.). CAYMAN BRAC: Cen- tral plateau S of Stake Bay village, ca. 60 R, dry rocky woodland, 2 May 1956, Proctor 15326 (BM). GRAND CAYMAN: Near Sand Bluff, East End, limestone karst, behind sandy beach ridge, 24 May 1967, Brunt 2003 (BM, NY). LITTLE CAYMAN: Inland from Jackson's Bay, rocky woodland, 5 Aug 1975, Proctor 35108 (BM, MO, US).

JAMAICA. CLARENDON: Along road 1-1.5 miles SSW of Inverness, toward Breadnut Gully, 200-300 ft, dry rocky woodland, 1 Oct 1974, Proctor 34191 (GH). COCKPIT COUNTRY (aberrant specimens): 5.3 miles by road N of Quick step, limestone, ca. 1250 ft, forested region, same locality, 5 May 1977, Correll 48638 (NY), 5 May 1977, Proctor 36833 (GH, MO); Ramgoat Cave district, ca. 1500 ft, dry rocky hillside, 4 Jul 1955, Howard & Proctor 14390 (A, BM, US). TRELAWNY: Island View Hill, Wilson Valley district, 1.5 miles N of WARSOP, limestone, 2000-2200 ft, wooded hilltop, 10 Apr 1961, Proctor 22184 (GH, NY, US).

U.S. VIRGIN ISLANDS. ST. THOMAS. Albion Mts., rough limestone formation, 2 Nov 1915, Harris 12196 (BM, GH, K, MO, NY, US).

U.S.A. (FLORIDA, MONROE CO.) BIG PINE KEY:

E side of Big Pine Key, hammock, 17 May 1922, Small 10501 (A?, GH, S, US). LITTLE TORCH KEY: hammock, 2 Jul 1956, Brizicky & Stern 346 (A, US). MIDDLE TORCH KEY: in hammock borders, 20 Mar 1982, Brumbach 9768 (GH, MO). No NAME KEY: hammock, near Clusia population, 22 Nov 1972, Gillis 11513 (A). RAMROD KEY: hammock, 15 Mar 1952, Caldwell 8782 (A, MO, UC-n.v.).

Heterosavia bahamensis is variable in leaf size and shape. Some specimens from Cock- pit country, Jamaica, are particularly distinct: Correll 48638, Proctor 36833 with to 13 cm long, acuminate leaves and 15 m tall; Howard & Proctor 14390 with to 8 cm long oblong to orbicular, rounded to retuse leaves, and Proctor 22183 and 22184 (presumably the same entity but with flowers and lacking leaves). These specimens were collected inland in a montane formation to 750 m altitude which is an atypical habitat for

Heterosavia bahamensis, but nevertheless dry limestone forest (Howard, 1973: 11-12) which is the species' exclusive substrate. This population could be taxonomically distinct but more material is needed. The typical form of Heterosavia bahamensis is found on the southeast coast of Jamaica.

The holotype and only specimen of Savia perlucens is a near-sterile specimen (with very young staminate buds) with a slightly aberrant leaf shape. However, its leaves resemble some specimens of Heterosavia bahamensis from Isla de Pifios (PFC (Bisse et al.) 45578 and 45662; Britton et al. 15265). Further arguments in favor of uniting the two species are the habitat as well as leaf venation, texture and color.

2. Heterosavia erythroxyloides (Griseb.) Petra Hoffm., comb. nov. Savia erythroxyloides Griseb., Mem. Amer. Acad. Arts, n.s., 8: 157. 1860. TYPE: Cuba, Prov. Oriente, prope Monte Verde, C. Wright 1433 (lectotype, here designated: GOET; isolectotype: GOET). Grisebach (1860) noted that the syntype Wright 1433 (here designated as lectotype) is pistillate, whereas the other original syntype Wright 1434 is staminate. This is not the case in all duplicates of these two collection numbers; some have other localities such as: K: La Palma, San Cristobal, NY: Cajalbana, NY: Vifiales, summit of mountain, the dates range from 1859 to 1864, and often there are also branches of Heterosavia maculata mounted on the sheets. Therefore, only the other GOET sheet is designated as an isolectotype, even though nine further possible duplicates from other herbaria were examined.

Savia erythroxyloides Griseb. var. parvifolia Urb., Repert. Spec. Nov. Regni Veg. 28: 209. 1930. TYPE: Cuba, Santa Clara, on top of Pico Potrerillo, ca. 1000 m, 30 Mar 1924 (fl

c3_), E. L. Ekman

18955 (holotype: S; isotype: S). See note following species key.

Tree or shrub, (1-)2-4(-8) m tall; twigs pilose to pubescent when young, glabrescent. Stipules deltoid, acute, (1-)2x1-1.5 mm, +pubescent abaxially. Petiole 2-5 x ca. 1 mm, glabrous. Leaf blades elliptic to




obovate, apically obtuse to rounded, sometimes short-acuminate to acute, rarely slightly retuse, sometimes with a terminal or subapical glandular dark mucro ca. 0.1 mm long, basally acute, (1-)2-5(-9) x 1-3(-5) cm, glabrous, rarely with scattered hairs along midvein abaxially, coriaceous to thinly chartaceous, mostly gray-green adaxially and reddish brown abaxially, midvein + prominent on both surfaces, higher venation not or hardly prominent adaxially, weakly prominent abaxially. Stami- nate flowers 6-20 per inflorescence, ca. 4 x 2- 3 mm; bracts deltoid, acute, 1-1.5 mm long, to 1 mm wide, pubescent adaxially; buds sessile, ovoid to globose. Sepals 5, ovate to deltoid, rarely obovate or lanceolate, acute to obtuse, 1.5-2 x 1-1.5 mm, pubescent abaxially. Petals 5, spathulate to rhombic, irregularly crenate, ca. 1 mm long, to 1 mm wide, shorter than the sepals. Disc irregularly lobed, the lobes ca. 0.5 mm long, or disc lacerate to the base. Stamens 5; 3 mm long; filaments free or basally fused for 1 mm, ca. 2.5 mm long; anthers ovoid, 1 mm long. Pistillode 1-1.5x 0.5(-1) mm, as long as or slightly longer than the sepals, pilose. Pistillate flowers solitary or in twos, 3-5 x ca. 4 mm; bracts and buds as in staminate flowers. Pedicel 1-2 x ca. 1 mm, pubescent. Sepals 5, rarely 6, ovate to deltoid, obtuse to acute, 1.5-2x 1-2 mm, pubescent abaxially. Petals 5, orbicular, often irregularly crenate, 1-1.5 x ca. 1 mm, shorter than the

sepals. Disc crenate, to 1 mm thick, the lobes altemipetalous, to 1 mm long. Ovary globose, ca. 2 mm long, densely pubescent; styles straight to recurved, spreading, flat, 2 mm long, to 0.5 mm wide, glabrous or sparsely pilose. Fruits 5-7 x 6-8 mm, densely pubescent, green maturing to reddish brown; fruiting pedicel 2- 5 x 1 mm, pubescent; columella 3-5 x2-3 mm; exocarp 0.1-0.2 mm thick; endocarp ca. 0.5 mm thick. Seeds 3-5 x 2-3 mm, brown.

Distribution and habitat. Cuba and His- paniola; Fig. 13. In forests (monte nublado, pluviosilva de montafia, bosque siempreverde mesofilo, monte semicaducifolio, pinares, mogotes, monte seco) at (300-)700 m to 1500 m altitude. Borhidi (1991: 418, 557) listed the species as characteristic of the limestone woods in eastern Cuba (Tabebuio- Coccothrinacetalia). Heterosavia erythroxyloides prefers humid evergreen habitats, generally at higher altitudes than the other species. A preference for calcareous substrate is likely, but cannot be confirmed due to the scarce geological data on the herbarium labels. None of the specimens are recorded from serpentine.

Conservation status. Eighty four specimens were examined. As well as occurring in scattered localities throughout Hispaniola, Heterosavia erythroxyloides has been collect-

84 I 80, ') 76 72x 24-

2O"

* * 2,0o20

I . . . . . I I I 0 100 200 300 Kilometres

16 I 84x

800 76x

72o 680

FIG. 13. Distribution of Heterosavia erythroxyloides.




ed in recent decades in eastern, central, and western Cuba. Because of its extensive distribution and lack of information on specific threats at least in Cuba, there seems no justification of according H. erythroxyloides a IUCN (2001) threat category and it is rated Least Concern (LC).

Selected specimens examined. CUBA. LAS VILLAS: Las Vegas de Matagua, Buenos Aires, above San Blas, 1900-2000 ft, 7 Apr 1928, Jack 5945 (A, HAC, NY, US); Sancti Spiritus, Mun. Trinidad, Sierra del Escam- bray, Lomas de los Manantiales, 2 km S de Topes de Collantes, 700 m, 3 Aug 1972, PFC (Bisse) 23275 (HAJB, JE); Loma del Burro, 13 Jul 1974, PFC (Areces & Berazain) 25016 (HAJB). ORIENTE: Ensenada de Mora, 330 m, shaded bank, 26-29 Mar 1912, Britton et al. 12996 (GH, K, NY, US); Mun. Yateras, Monte Cristi, altiplano, 700 m, May 1968, PFC (Bisse & Kohler) 8887 (HAJB, JE); Holguin, Mayari Abajo, Sierra de Nipe, mogotes de caliza cerca del terraplen de Miranda a los pinares de Mayari, 400 m, 3 Nov 1977, PFC (Bisse et al.) 36062 (B, HAJB, JE). PINAR DEL RIo: Vifiales, Sierra de Vifiales, Loma de la Bandera, jurassic limestone rocks, 9 Mar (or May?) 1924, Ekman 18665 (NY); Loma del Sabicii, Rangel, Rosario Mts., limestone cliff, 600 m (GH: 550 m), on exposed rocks, 27 Apr 1933, Lion & Roig 15975 (GH, HAC, NY); Mun. San Crist6bal, Sierra del Rosario, Santa Cruz de los Pifios, loma al oeste del camino de Cinco Pesos, caliza, 400 m, Dec 1974, PFC (Bisse & Klotz) 26531 (HAJB).

HAITI. DEPARTEMENT DE L'OUEST: Arrondissement Port-au-Prince, Massif de la Selle, Mome de l'Hospital, near Port-au-Prince, 1000 m, in a little patch of original vegetation, 21 Oct 1924, H 2228 (A, US); Arondissement Thomazeau, Massif du Mattheux, Grands Bois, road Cornillon to Thomazeau, limestone, ca. 750 m, 19 Mar 1926, H 5779 (G, GH, K, NY, US). DEPARTEMENT DE L'ARTBONITE: Arrondissement Marmelade, Vicinity of Marmelade, rocky lime summit, ca. 800 m, 20 Dec 1925, Leonard 8302 (GH, NY).

DOMINICAN REPUBLIC. AZUA PROV.: Cordillera Central, Valle de San Juan, at Seboruco, hillsides, ca. 600 m, 8 Sep 1929, Ekman 13529 (B, US). EL SEIBO PROV.: Los Haitises Frente a la Caseta de Guardaparques en Monte Bonito (de Trepada Alta), Parque Nacional Los Haitises, calcareo, 200-250 m, cima de un mogote, bosque latifoliado y humedo, arboles altos, rocoso, 190 00'N, 69030'W, 6 Aug 1985, Zanoni et al. 35455 (NY). PEDERNALES PROV.: Baoruco Mts., between Boucan Calice and Hojo de Pelempito, 800 m, on a rocky knoll, 13 Feb 1969, Alain 13875 (NY, US). PUERTO PLATA PROv.: Cordillera de Yaroa, ridge facing the Yaroa valley, limestone, 800-850 m, on top of the ridge, on exposed rocks, 11 May 1968, Alain 11215 (GH, NY, US). SANIANA PROV.: Cordillera Central, Los Haitises, on an Island near La Llanada, 29 Jun 1930, Ekman 15501 (B, MO).

3. Heterosavia laurifolia (Griseb.) Petra Hoffmn., comb. nov. Savia laurifolia Griseb.,

Nachr. Kinigl. Ges. Wiss. Georg-Augusts- Univ. 6: 164. 1865. TYPE: Cuba, Prov. Oriente, Monte Verde, C. Wright 1430 "13" (lectotype, here designated: GOET; isolectotypes: BM, G-DC, only microfiche seen, GH-n.v., GOET, K, MO, NY, US).

Savia clusiifolia auct. non Griseb., Miiller in DC., Prodr. 15(2): 230. 1866.

Tree or shrub, 2.4-3(-10?) m tall; twigs glabrous, only very young twigs pilose to pubescent. Stipules deltoid, acute, 1-2.5x 2 mm, + pubescent abaxially, glabrescent. Petiole (2-)3-6(-9) x 1-1.5 mm, glabrous. Leaf blades elliptic to oblong, rarely obovate, apically acuminate, acute to obtuse, more rarely rounded, sometimes with a terminal or subapical glandular dark mucro ca. 0.1 mm long, basally acute, rarely obtuse or rounded, (3-)5-10(-15) x(2-)2.5-4.5(-6) cm, glabrous, coriaceous to nearly chartaceous, usually gray-green, midvein prominent on both surfaces, higher venation slightly prominent on both surfaces. Staminate flowers 10-20 per inflorescence, ca. 5 x 3-4 mm; bracts deltoid, acute, 1-1.5 x 0.5-1 mm, pubescent adaxially; buds sessile, ovoid to globose. Sepals 5, rarely 4, ovate to deltoid, more rarely obovate or lanceolate, acute to obtuse, ca. 2x 1.5 mm, pilose abaxially. Petals 5, spathulate, irregularly crenate or lacerate, 1- 1.5x0.5-1.5 mm, shorter than the sepals. Disc crenate, the lobes altemrnipetalous, to 1 mm long. Stamens 5; 4-5 mm long; filaments basally fused for ca. 1 mm, 3- 4 mm long; anthers ovoid, 1-1.2 mm long. Pistillode ca. 1 x 0.5 mm, about as long as the sepals, glabrous or pilose. Pistillate flowers solitary, 3-4 x ca. 4 mm; bracts as in staminate flowers; buds subsessile, ovoid. Pedicel 3-7 x 1-2 mm, glabrous or sparsely pilose. Sepals 5, deltoid to ovate, acute to obtuse, 1.5-2 x 1- 2 mm, glabrous to sparsely pilose abaxially. Petals 5, rhombic to spathulate, irregularly crenate, 1.5-2 x ca. 1 mm, as long as the sepals. Disc crenate, to 1 mm thick, the lobes alternipetalous, to 1 mm long. Ovary globose, ca. 2 mm long, glabrous to pubescent; styles sometimes basally fused, recurved, spreading, flat, 1.5 xca. 0.6 mm. Fruits 7-8x9-10 mm, glabrous or reddish brown pilose, reticulate;




fruiting pedicel 8-15(-22) x ca. 1 mm, glabrous or pilose as the fruit; columella 5-7 x 2-4 mm; exocarp 0.1-0.2 mm thick; endocarp 0.5- 0.7 mm thick. Seeds 4-7 x 2-5 mm, brown.

Habitat. In forests at 200-550 m altitude. As in the previous species, a preference for limestone is likely but the data on the herbarium labels are insufficient to be sure.

Key to the varieties of Heterosavia laurifolia

1. Leaf blades elliptic to oblong, rarely obovate; fruits glabrous or reddish brown pilose; fruiting pedicels 8-15(-22) mm long, glabrous or pilose as the fruit H. laurifolia var. laurifolia

1. Leaf blades obovate, rarely elliptic; fruits glabrous; fruiting pedicels 5-7(-10) mm long, glabrous H laurifolia var. intermedia

3a. Heterosavia laurifolia var. laurifolia.

Savia clusiifolia Griseb. var. fallax Miull. Arg. in DC., Prodr. 15(2): 231. 1866. TYPE: Cuba, Prov. Oriente, Monte Verde, C. Wright 1430 (lectotype, here designated: G). This sheet bears the name of the new variety in Miiller's handwriting. This synonym is heterotypic although the number of the type collection is identical with that of Heterosavia laurifolia var. laurifolia. It is widely known that Wright's collections are heterogeneous and that plants with the same collection number in different herbaria are likely to be from different collections. However, all duplicates of Wright 1430 examined in this study are Heterosavia laurifolia var. laurifolia.

Savia clusiifolia Griseb. var. membranacea Miill.Arg. in DC., Prodr. 15(2): 231. 1866. Savia membranacea (Mill. Arg.) Urb., Repert. Spec. Nov. Regni Veg. 28: 211. 1930. TYPE: Cuba, Prov. Oriente, without precise locality, 1860-64 (fr), C. Wright 1924 (holotype: G- DC, only microfiche seen; isotypes: BM, GH-n.v., GOET, K, MO). The type of Savia clusiifolia var. membranacea is a paratype of S. laurifolia.

Savia impressa Urb., Repert. Spec. Nov. Regni Veg. 28: 211. 1930. TYPE: Cuba, Cienfuegos, Lomas de Sigua- nxa, at Rio Navarro, wooded mountainsides, 5 Jun 1922 (fr), E. L. Ekman 13927 (holotype: S; isotypes: NY, S). See note on typification following species key.

Savia longipes Urb., Repert. Spec. Nov. Regni Veg. 28: 211. 1930. TYPE: Cuba, Prov. Oriente, Bayate, Majagual (San German Sabonoso, Galb~s), 15 Jul 1916 (fr), E. L. Ekman 7377 (lectotype, here designated: S; isolectotypes: G, K, NY, US).

Leaf blades elliptic to oblong, rarely obovate, apically acuminate, acute to obtuse, more rarely rounded, basally acute, rarely obtuse or rounded, (3-)5-10(-15) x (2-)2.5- 4.5(-6) cm, usually gray-green. Fruits glabrous or reddish brown pilose; fruiting pedicels 8-15(-22) mm long, glabrous or pilose as the fruit.

Distribution. Endemic to Cuba; Fig. 14.

Conservation status. 34 specimens were examined, of which 10 are pistillate and can be distinguished with certainty from Hetero- savia erythroxyloides. The distribution of this variety, with an extent of occurrence of nearly 30,000 km2 and spanning almost the entire W-E extent of Cuba, precludes ranking it as threatened. Despite active collection programs in Cuba in recent decades, however, no specimens collected after 1956 were found during this study. Moreover, none of the herbarium labels indicate that Heterosavia laurifolia var. laurifolia grows in disturbed habitats. For these reasons, it is here assigned the IUCN (2001) category Near Threatened (NT).

Vernacular names Came de doncella, Vigueta colorada (Ekman 7377, 9556; Urban, 1930: 212).

Selected specimens examined. CUBA. LAS VILLAS: Mun. Cumanayagua, San Blas, La Sierra, 600-800 ft, 4 Mar 1929, Jack 6972 (A, HAC, K, NY, US); Sierra de Gavilanes, Banao, Sancti Spiritus Mts., 1-2 Aug 1916, Lion et al. 6614 (HAC, NY); Sancti Spiritus, Mun. Banao, Lomas de Banao, 27 Mar 1920, Luna 454 (HAC, NY); May 1920, Luna 525 (HAC, NY). MATANZAS: Pan de Matanzas, W of Matanzas bay, a limestone mountain, in forest, 30 May 1923, Ekman 16457 (MICH); Penin- sula Zapata, along the railroad from Central Australia to Bahia de Cochinos, limestone, low forest, 30 Jan 1924, Ekman 18330 (G, NY, US). ORIENTE: Santiago de Cuba, Mun. Mella, Bayate inter Arr. Libano et Rio Bayate, 16 Feb 1915, Ekman 4645 (US); Bayate, Cayo del Rey, at Arroyo Jimbambay, forest, 23 Feb 1915, Ekman 4707 (G, K); Holguin, Mun. Mayari, Sierra de Nipe, at the tunnel of Rio Piloto beneath the mountain, in steep limestone cliffs, 17 May 1915, Ekman 5783 (NY); Mun. Moa, Cupey, in forest, 4 Aug 1915, Ekman 6322 (G, K, NY, US); Mun. Mella, Sierra de Nipe, in the cation (gulch) of




Rio Jimbambay (Cayo del Rey), limestone, 26 Apr 1919, Ekman 9556 (G, HAC, K, NY); Guantinamo, Mun. Yateras, Monte Verde, 10 Jul 1919, Hioram & Mamel 2537 (HAC, NY); Holguin, Mun. Cueto, Alto Cedro, at Arroyo Blanco, dark soil, bosques humedos, 27 Jun 1931, Roig 5675 = Curbels 283 (HAC, NY). PINAR DEL RIo: Rangel, Sierra del Rosario, 500 m, woods, Jan 1953, Alain 2719 (GH, US); Top of Pico Tey, Alain 2735 (GH, HAC).

3b. Heterosavia laurifolia var. intermedia (Miill. Arg.) Petra Hoffm., comb. nov. Savia clusiifolia Griseb. var. intermedia Mill. Arg. in DC., Prodr. 15(2): 231. 1866. Type: Cuba, without precise locality, 1860-64, C. Wright 2009 (holotype: G-DC, only microfiche seen; isotypes: BM, GH, GOET, K, MO). The type of Savia clusiifolia var. intermedia is a paratype of S. laurifolia.

Leaf blades obovate, more rarely elliptic, apically acuminate, acute, obtuse, or rounded, basally acute, more rarely obtuse, 3-8 x 2-4 cm, green to gray-green. Fruits glabrous; fruiting pedicels 5-7(-10) mm long, glabrous.

Distribution. Cuba, Oriente province; Fig. 14.

Conservation status. Only five specimens of Heterosavia laurifolia var. intermedia have been located and examined. Because of the scarce label data and the somewhat question- able taxonomic status of this variety it is best

accorded the IUCN (2001) status of Data Deficient for the time being.

Additional specimens examined. CUBA. ORIENTE: Prope Monte Verde, Jan-Jul 1859, Wright 1431 (G, G-DC [only microfiche seen], GH, GOET, K, MO, NY); Palenquito, Yateras, GH: 400 m, HAC: ca. 550 m, woods, 20 Jul 1953, Alain 3105 (GH, HAC); Sierra de Nipe, at Loma Mensura, 725 m, in manascales, 14 Oct 1919, Ekman 9893 (MICH); Holguin, Mun. Mayari, Pinares de Mayari, mogotes y barranco del rio Piloto en su curso medio, caliza, 400 m, monte semicaducifolio, 1 Jun 1983, PFC (Bisse et al.) 50629 (HAJB, JE).

Heterosavia laurifolia var. intermedia dif- fers from the type variety in its mainly obovate leaves and the shorter fruiting pedicels. It stands morphologically between H. laurifolia var. laurifolia and H. maculata var. maculata, being slightly more similar to the former taxon.

4. Heterosavia maculata (Urb.) Petra Hoffm., comb. nov. Savia maculata Urb., Symb. antill. 9: 181. 1924. Type: Cuba, Pinar del Rio, Bahia Honda, south of town, towards the Sierra de los Organos, in cuabales, 8 Mar 1920, E. L. Ekman 10417 (holotype: S; isotypes: G, NY, S). See note on typification following species key.

Shrub or tree, 2-9 m tall; only very young twigs pubescent, soon glabrescent, sometimes individual intemrodes pilose and others glabrous. Stipules deltoid to ovate, acute, acu-

84xI 80o 762

o24

*o

o*o

20x

2;-:

I I I I

0 100 200 300 Kilometres

841 801 761

FIG. 14. Distribution of Heterosavia laurifolia var. laurifolia (solid circles) and var. intermedia (open circles).




minate or obtuse, (1-)2-3 x 1-2 mm, usually pubescent abaxially when young, glabrescent. Petiole (1-)3-5(-10)x 1-2 mm, glabrous, rarely with a few hairs. Leaf blades obovate, more rarely oblong to nearly elliptic, apically obtuse, rounded, truncate or retuse, more rarely very short-acuminate, basally acute or narrowly cuneate, rarely obtuse, (1.2-)2-9(- 11.5) x (0.8-)1.5-4(-5.5) cm, not to strongly revolute at the margin, glabrous, rarely with a few hairs abaxially, coriaceous, + concolorous, olive-green or reddish brown, midvein much less prominent adaxially than abaxially, higher venation on both surfaces or only abaxially prominent. Staminate flowers (4-) 10-20 per inflorescence, ca. 4 x 3-5 mm; bracts deltoid to ovate, acute, 1-1.5 x 0.5- 1 mm, pubescent abaxially; buds sessile, ellipsoid to globose. Sepals 5, ovate, oblong, deltoid, more rarely obovate, acute to rounded, 1.5-2.5 x 1-1.5 mm, sometimes fimbriate, pilose to pubescent abaxially. Petals 5, spathulate to + orbicular, irregularly crenate or lacerate, 1-1.5 x 0.5-1.5 mm, shorter than the sepals. Disc crenate, the lobes altemrnipetalous, ca. 0.5 mm long. Stamens 5; 3-5 mm long; filaments free or fused basally for ca. 1 mm, 2-4 mm long; anthers ovoid to

ellipsoid, 1-1.5 mm long. Pistillode 1-2x 0.5-0.8 mm, as long as or (if filaments fused) slightly longer than the sepals, glabrous or with a few hairs. Pistillate flowers solitary, more rarely in twos or threes, 2.5-4x3- 4 mm; bracts as in staminate flowers; buds sessile or short-pedicellate, globose to ellipsoid. Pedicel 1-2x 1-1.5 mm, glabrous to pubescent. Sepals 5, rarely 6, + orbicular, ovate or deltoid, acute to rounded, 1.5-2.5 x

1-2 mm, sometimes fimbriate, pilose to pubescent abaxially. Petals 5, rarely 6, orbicular to oblong, rhombic or spathulate, slightly erose, sometimes sparsely fimbriate, 1.5-2x 1-2 mm, about as long as the sepals. Disc entire to crenate, ca. 0.5 mm thick, the lobes alternipetalous, ca. 0.8 mm long. Ovary globose, ca. 2 mm long, glabrous (Ekman 19102, NY: 1 flower with sparsely pilose ovary); styles sometimes basally fused, recurved to revolute, spreading, 0.5-1 mm long, glabrous. Fruits 5-7 x 6-8 mm, glabrous (Wilson 9392, NY: 1 fruit with a few hairs at base of styles), reticulate; fruiting pedicel 2-5x 1-1.5 mm, glabrous to pubescent; columella 4-6 mm long, to 3 mm wide; exocarp 0.1-0.3(-0.5) mm thick; endocarp 0.4-0.8 mm thick. Seeds 4-6 x 2-4 mm, brown to black.

Key to the varieties of Heterosavia maculata

1. Young shoots 1-1.5 mm wide; leaf blades 2-5(-8)x 1.5-2.5(-3.4) cm, obovate, revolute or not, moderately coriaceous, olive-green or reddish brown abaxially, venation prominent adaxially; stipules to 1.5 mm wide; petiole (1-)3-5 x 1-1.5 mm; throughout Cuba. H maculata var. maculata

1. Young shoots 2-4 mm wide; leaf blades 5-10(-16) x 2-5(-6) cm, obovate to oblong or slightly elliptic, strongly revolute, strongly coriaceous, reddish brown abaxially, venation not or hardly prominent adaxially; stipules 1.5- 2 mm wide; petiole 3-10x 1-2 mm; Cuba, Oriente province only H maculata var. clementis

4a. Heterosavia maculata var. maculata.

Savia cuneifolia Urb., Repert. Spec. Nov. Regni Veg. 28: 210. 1930. TYPE: Cuba, Prov. Oriente, Sierra de Nipe, at Rio Piedra, 200-300 m, charrascales, 29-39 Nov 1924, E. L. Ekman 19102 (holotype: S; isotypes: G, NY, S). See note on typification following species key.

Savia clusiifolia auct. non Griseb., Alain in Leon & Alain, Fl. Cuba 3: 42. 1953.

Young shoots 1-1.5 mm wide. Stipules deltoid, acute to acuminate, (1-)2-2.5x 1- 1.5 mm. Petiole (1-)3-5x 1-1.5 mm. Leaf blades obovate, apically rounded, truncate or retuse, more rarely short-acuminate, basally

acute or narrowly cuneate, rarely obtuse, (1.2-)2-5(-8)x (0.8-)1.5-2.5(-3.4) cm, sometimes revolute, moderately coriaceous, olive-green or reddish brown on both surfaces, or olive-green adaxially and reddish brown abaxially, venation prominent adaxially. Staminate flowers: sepals 1.5-2 x ca. 1 mm, stamens 3-4 mm long. Pistillate buds with pedicel ca. 1 x0.5-0.7 mm. Pistillate pedicel ca. 2 x 1-1.5 mm. Fruiting pedicel 2- 5x 1-1.5 mm. Seeds 4-6 x 2-4 mm.

Distribution and habitat. Endemic to serpentine habitats in Cuba; Fig. 15. Semi-arid




84o 800

760x 24o

2OPO

"% .x~

220

I I I I

0 100 200 300 Kilometres

84x 80I 76,I

FIG. 15. Distribution of Heterosavia maculata var. maculata (solid circles) and var. clementis (open circles); half- filled circles indicate areas where both varieties have been collected.

montane rain forest on serpentine (Borhidi, 1991: 396), montes secos, montes siempre- verdes, pinares, bosques de galeria, pluviosil- vas de montafia, montes nublados, also scrub formations (charrascos, cuabales, matorrales xeromorfos subespinosos), often along streams; on serpentine, very rarely on igneous rock, at 60 to 1000 m altitude.

Conservation status. This variety is widely distributed in Cuba and has been much collected in recent decades (161 specimens examined), including several times in degraded rain forest. It is therefore assumed that it thrives even in disturbed habitats and is regarded as Least Concern (LC), by the criteria of the IUCN (2001).

Vernacular names. Plateado, Hicaquillo (Curbels/Herb. Roig 6193); Sapotillo (Ekman 5004).

Selected specimens examined. CUBA. CAMAGiEY: Mun. Camaguey, orillas del rio Yucatan, Jun 1967, PFC (Bisse & Rojas) 2901 (HAJB, JE); Ciego de Avila, Mun. Florencia, entre San Felipe y Guadalupe, suelo fersiali- tico pardo-rojizo sobre serpentina, charrascos, matorral xeromorfo subespinoso, 28 Apr 1987, PFC (Arias et al.) 61905 (B, JE). HABANA: San Miguel, in collibus solo eruptivo, 20 May 1920, Ekman 10923 (G, K, NY, US); Campo Florido, Loma Coca, in cuabales, 30 Apr 1924, Ekman 19007 (G, K, MICH, NY, US). LAS VILLAS:

10 km S of Santa Clara, serpentine, ravines, 1-20 Jul 1950, Howard et al. 126 (A, K, MICH, NY); Mun. Jatibonico, Arroyo Blanco, cuabales cerca de San Felipe, cuabales, 18 Nov 1975, PFC (Alvarez et al.) 28891 (B, HAJB, JE). MATANZAS: Canasi, falda sur de las Lomas de Galindo, serpentina, 200-300 m, Jul 1970, PFC (Bisse & Lippold) 17527 (HAJB, JE); Mun. Cirdenas, Lomas de Camarioca al sur de Cantel, suelo serpenti- noso, 19 Jul 1977, PFC (Bisse et al.) 34994 (B, HAJB, JE). ORIENTE: Bayate, prope Rio Piedra, on eruptives belonging strictly to Sierra de Nipe, thickets, 5 Jun 1915, 5912 (K, S), paratype of Savia cuneifolia; Mun. Baracoa, subida a la Mina Iberia, 300-700 m, pluviosilva, Mar 1968, PFC (Bisse & Kohler) 6142 (B, HAJB, JE); Moa, Las orillas del rio Jiguani, cerca del segundo asserio de la Melba, Apr 1968, PFC (Bisse & Kohler) 6752 (B, HAJB, JE); Holguin, Mun. Mayari, Sierra Cristal, valle del rio Levisa desde Mandinga hacia arriba, Apr 1968, PFC (Bisse & Kihler) 7416 (B, HAJB, JE). PINAR DEL RIo: La Palma, Loma Cajalbana, orillas de un arroyo, falda NE de la Cajalbana, serpentine barrens, 15 Jul 1950, Alain & Clemente 1485 (DAV-n.v., GH, HAC, HAJB, NY, US); Zambumbia hill, Rangel, Rosario Mts., serpentine soil, 510 m, 26 Aug 1926, L~on 12774 (GH, HAC, NY).

Heterosavia maculata var. maculata is variable in leaf shape and size. It is morphologically very similar to H. bahamensis but restricted to serpentine and never found on limestone.

4b. Heterosavia maculata var. clementis (Alain) Petra Hoffmin., stat. nov. Savia clementis Alain, Contr. Ocas. Mus. Hist.




Nat. Col. "De La Salle" 11: 2. 1952. TYPE: Cuba, Prov. Oriente, Moa, camino cerca de la Mina Delta, este de Punta Gorda, 21 Jul 1944, H. Clemente, H. Alain & H. Crisogono 4036 (lectotype, here designated: HAC; isolectotypes: DAV-n.v., HAC, NY).

Illustration: Alain, Contr. Ocas. Mus. Hist. Nat. Col. "De La Salle" 11: 2. 1952 (fr, habit).

Young shoots 2-4 mm wide. Stipules deltoid to ovate, acute to obtuse, 2-3 x 1.5- 2 mm. Petiole 3-10x 1-2 mm. Leaf blades obovate to oblong or slightly elliptic, apically rounded, obtuse, + retuse or short-acuminate, basally acute to cuneate, (2.5-)5-10(-16)x (1.5-)2-5(-6) cm, strongly revolute, strongly coriaceous, olive-green to reddish brown adaxially, reddish brown abaxially, venation not or hardly prominent adaxially. Staminate flowers: sepals 2-2.5xca. 1.5 mm; stamens 4-5 mm long. Pistillate buds sessile. Pistillate pedicel 1-2 x ca. 1 mm. Fruiting pedicel 2-3 x 1.5 mm. Seeds 5-6 x ca. 4 mm.

Distribution and habitat. Cuba, Oriente province, Sierra de Baracoa and Sierra de Moa; Fig. 15. Montane rainforest and pine forest (pluviosilva de montafia, pinares) and scrub formations (charrascos, cuabales, manascales) on serpentine, rarely on igneous rock, at 100 to 800 m altitude.

Conservation status. Thirty seven specimens of Heterosavia maculata var. clementis were examined. The extent of occurrence is only about 5000 km2 which would warrant a threat category, but most collections were made relatively recently, including two from degraded rain forest. The variety is therefore here assigned the IUCN (2001) category Near Threatened (NT).

Selected specimens examined. CUBA. ORIENTE: Sierra de Nipe ad Rio Piloto, ca. 350 m, in manascales, 14 Mar 1915, Ekman 5004 (S, US), paratype of Savia cuneifolia; Baracoa, al sur de la Loma del Yunque, 300- 400 m, pluviosilva, Apr 1970, 17076 (HAJB, JE); Holguin, Mun. Moa, alrededores del aserrio "La Melba", 30 Apr 1980, PFC (Bisse et al) 42421 (B, HAJB, JE); Holguin, Mun. Frank Pais, falda norte de la Sierra Cristal, subida a Palenque en la zona de Brazo Grande, suelo ferralitico, amarillento, lixiviado, pluviosilva des- truida, 7 Apr 1987, PFC (Bdissler et al.) 60500 (B, JE).

Alain (1952) gave only a single qualitative differential character for Savia clementis in the protologue: ovate, rounded stipules. Such stipules are sometimes also found in all other species of Heterosavia and provide no con- vincing basis for the recognition of a separate species. All other morphological characters intergrade with Heterosavia maculata var. maculata. The plants have, however, a stiff, thickly coriaceous habit not found elsewhere in the genus, large, often oblong, dark reddish leaves and shoots and a contiguous distribution area in Oriente province, and are therefore here recognized at varietal rank within H. maculata.

Acknowledgments

I gratefully acknowledge the following herbaria for granting me access to their collections: A, B, BM, CHAPA, G, GH, GOET, HAC, HAJB, HBG, JE, M, MEXU, MICH, MO, NY, P, S, US, and XAL. I thank Prof. E. K6hler (BHU) for his invaluable support, J. Ueckert (formerly BHU) for assistance for her friendship and support with photography and lab work, Dr I. Cordeiro (SP) for additional material of Gonatogyne and Savia, Prof. W. Greuter (B) and Dr. R. K. Brummitt (K) for help with nomenclatoral queries, Prof. R. D. Reeves (Massey Univer- sity, New Zealand), and Prof. A. M. J. Baker (University of Melbourne, Australia) for sharing and discussing their Ni content data with me, Prof. H. H. Hilger, Dr. S. Rogaschewski, Dr. W. Wenzel, Ms Domes, Ms Griiber, Ms Ritter, and Ms Weise (all Berlin) for access to and assistance with the SEM, H. Wilks (K) for drawing Fig. 11, M. Vorontsova (K) for help with IUCN ratings, and J. Stone (K) for preparing the figures for publication. G. Levin (ILLS) and an anonymous reviewer provided valuable comments on the manuscript. Field- work in Cuba was conducted in collaboration with the Jardin Botanico Nacional, La Habana.

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Revision of Heterosavia, Stat. Nov., with Notes on Gonatogyne and Savia (Phyllanthaceae)

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