Notes on calycophyllous Rubiaceae. Part I. Morphological comparisons of the genera Chimarrhis, Bathysa, and Calycophyllum, with new combinations and a n e w species, Chimarrhis gentryana PIERO G~ DELPRETE Delprete, R G. (Department of Botany, University of Texas, Austin, TX 78713, USA). Notes on calycophyllous Rubiaceae. Part I. Morphological comparisons of the genera Chimarrhis, Bathysa, and Calycophyllum, with new combinations and a new species, Chimarrhis gentryana. Brittonia 48: 35-44. 1996.--A comparison of some genera of the Condamineeae (Rubiaceae) with a few taxa of closely related tribes (Rondeletieae, Calycophylleae, and Cinchoneae) revealed that some species of Chimarrhis (Condamineeae s. 1.), Bathysa (Rondeletieae), and CalycophyHum (Calycophylleae) are often misassigned to genera. The taxonomic significance of calycophylls is discussed; the generic boundaries of Chimarrhis, Bathysa, and Calycophyllum are reevaluated; and their similarities and differences are discussed. As a result, a new calycophyllous species of Chimarrhis from the Amazon, C. gentryana, is described, two of its species are transferred to Bathysa (B. Bathysoides, B. perija~nsis), and one species of Bathysa (B. difformis) is reduced to synonymy under Chimarrhis (C. turbinata). Key words: Calycophyllum, Chimarrhis, Bathysa, Semaphyllanthe, Calycophylleae, Condamineeae, Cinchoneae, Rondeletieae, Rubiaceae, calycophylls, semaphylls, pterophylls. The presence of enlarged calyx lobes for the purpose of attracting pollinators or seed dispersers has been reported in many families o f flowering plants (Plumbaginaceae, Dipterocarpaceae, etc.). Several genera o f the Rubiaceae have enlarged calyx lobes that are variously colored, which are here specifically called " c a l y c o p h y l l s " ("calyxborne semaphylls" in Leppik, 1977; "petMoid calyx-lobes" in Robbrecht, 1988). The often-used term " s e m a p h y l l " was established by Leppik (1956) and refers to "all colored leaves o f plants, like petals, sepals, bracts, ligulate flowers, etc., which serve to attract pollinators," and therefore not specific to calyx lobes. The function of expanded calyx lobes may be attraction of pollinators and/or seed disperses. In most rubiaceous genera, calycophylls are brightly colored during anthesis (red, pink, yellow, white to greenish white) and lose their pigmentation during seed dispersal (e.g., Po- gonopus, Pinkneya, Mussaenda, Warszewiczia, Calycophyllum, Capirona); in other genera the calyx lobes expand only after anthesis, when the fruit starts ripening (Jacksiopsis, Nematostylis, and some species o f Alberta). The latter group suggests that the function o f the enlarged calyx lobes (pterophylls) is for seed dispersal and therefore is not functionally h o m o l o g o u s to the calycophylls of the former group. Robbrecht (1988) published a list o f Rubiaceae genera with "petaloid calyx-lobes," and additional o b s e r v a t i o n s on c a l y c o p h y l l o u s A f r i c a n Rubiaceae can be found in Puff et al. (1984). As part o f a revisionary study o f the tribe Condamineeae s. 1. (Delprete, in prep.), the genera o f this tribe have been compared Brittonia, 48(1), 1996, pp. 35~14. 9 1996, by The New York Botanical Garden, Bronx, NY 10458-5126 ISSUED: 21 March 1996 36 BRITTONIA with some genera of the closely related Rondetelieae, Calycophylleae, and Cinchoneae. In each o f these tribes calycophytls are variably present in some genera. For this reason, the presence of calycophylls does not seem to be important in inferring the phylogeny of these tribes at the generic level or for the circumscription of certain genera. Nonetheless, calycophylls are found m m n l y in the genera with capsular fruits o f the subfamily Cichonoideae (Robbrecht, 1988). Because of the foregoing, some of the genera under study are redescribed below so as Io include the species with calycophytls, with fi~e necessary new c o m b i nations to a c c o m m o d a t e these species. The three genera here described have capsules with septicidal dehiscence. In the following descriptions ! used the term "disk-loculicidal dehiscence" to refer to the secondary dehiscence restricted to the disk that occurs in older capsules of s o m e species. The disk variabty exceeds the calyx, and its shape is reported from observation of mature capgules. Several herbarium specimens collected in the western A m a z o n basin and identified as Calycophytlum obovamm (Ducke) D u c k e were found to belong to Chimarrhis gentryana Delprete, described below for the first time. CHIMARRHIS Jacquin Chimarrhis Jacquin, Selec. Stirp. Amer. Hist~ 61. 1763. TYPE: Chimarrhis cymosa Jacquin. Pseudochiraarrhis Ducke, Archiv. Jard. l~ot. Rio Janeiro 4: 177. pL 23. 1925. TypE: Pseudochirnarthis turbinata (DC,) Ducke (=Chimarrhis turbinata DC,), Not Chimarrhis Baillon, Hist. pL 7: 493. I880. (incl. Sickingia Willdenow [=Simira Aublet] and Sprucea Bentham [= Simira Aublet]). Trees, often canopy trees; bole Irregular, with or without buttresses; buttresses sometimes v e r y large, especially in the A m a z o nian species; bark smooth to deeply fissured; w o o d yellow and v e r y hard. Stipules interpetiolar, free or connate at base, persistent or caducous. Blades elliptic to ovate to obovate, usually acuminate at apex; d o m a tia absent, or tuft o f hairs [raft-pit domatia [VOL. 48 in C. jarnaicensis (Urb.) Steyerm.]. Inflorescences axillary on terminal nodes, cory m b o s e and with opposite to subopposite lateral branches terminating in cymules. Flowers protogynous, sessile to pedicellate, v e r y fragrant; hypanthium obconical to turbinate. Calyx extremely reduced, truncate or with barely distinguishable lobes, persistent; calyx l o b e s 4 or 5(6); one lobe expanded into a white to greenish white calycophyll in a few species (C. brevipes and C. gentryana), in some of the flowers. Corolla funnelform and deeply lobed, with reflexed l o b e s , white, g r e e n i s h w h i t e or cream-white; corolla tube funne/form to ciatiform, with a ring o f pubescent hairs inside on the tube or on the base of the lobes; corolla lobes 4 or 5(6); aestivation narrowly imbricate, superficially resembling valvate aestivation. Stamens exserted; anthers button-shaped to elliphc, dorsifixed, dehiscing by lateral slits; filaments attached on the upper part of the tube~ slender and basally flattened, with a tuff of pilose to villous hairs at base. Style exserted; style branches protruding a b o v e corolla before anthesis, short, rounded to ovate, reflexed at maturity. Ovary 2-celled, ptacentation pettate on the septurn, turbinate to obovate. Capsules globose, oblong or obovoid, dehiscing septicidally; disk-loculicidal dehiscence present in old capsules o f some species. Seeds suborbicular, margin undulate, and with a narrow concentric wing. Pollen tricolporate; exine densely reticulate. Chimarrhis is a genus of approximately 13 species o f trees distributed f r o m Central America, the Caribbean, to tropical South America, being especially c o m m o n in the A m a z o n basir~ ( B o o m & C a m p o s , 1991; pers. obs.). Chimarrhis has been recently treated as belonging to the C o n d a m i n e e a e s. 1. (Hooker, 1873; Schumann, 1889, 1891; Robbrecht, 1988) or to the Rondeletieae (Verdcourt, 1958). With its yellow hardwood, Chimarrhis is a very important lumber tree in the A m a z o n Basin, where it can reach canopy level with dimensions o f up to 45 m tall and up to 2 m diameter. The m o s t comprehensive key to its South American and West Indian species was published by Steyermark (1965), a c c o m p a n i e d by brief notes on their distribtttion. 1996] DELPRETE: RUBIACEAE Adolpho D u c k e ' s collections were v e r y critical in m y studies on the phenology and distribution of Chimarrhis. He often collected one tree in the flowering stage and returned to the same individual several months later to collect its fruiting material, mounting both collections on the same herbarium sheet. Chimarrhis is often confused with other genera of the Cinchonoideae because of its minute flowers and v e r y small capsules. Three characters that readily separate Chimarrhis f r o m other genera are its axillary inflorescences on terminal nodes, peltate placentation on the septum, and seeds with central hilum and small fringed concentric wing. These characters also m a k e it difficult to position Chimarrhis into a tribe, so although its tribal position remains uncertain, it is temporarily maintained in the Condamineeae. CHIMARRHIS TURBINATA D e . , Prodr. 4: 404. 1830. Pseudochimarrhis turbinata (DC.) Ducke, Archi. Jard. Bot. Rfo Janeiro 3: 255. Chimarrhis duckei Rizzini (nomen. superfl.), Rev. Bras. Biol. 7: 277. 1947. TYPE: F R E N C H G U I A N A . Ile de Cayenne, Patris s.n. (LECTOTYPE, designated here: G). Elaeagia brasiliensis Standley in Gleason & Smith, Bull. Torrey Bot. Club 60: 395. 1933. TYPE: BRAZIL. Patti: Tapajos River Region, Boa vista, Fordlandia, 6 Sep 1931 [fr], Krukoff 1018 (rlOLOTYPE: NY; ISOTYPE: K). Bathysa difformis Benoist, Syn. Nov., BulL Mus. Hist. Nat. Paris 26: 185. 1920. Pseudochimarrhis difformis (Benoist) Benoist, Arch. Bot. 5, M6m. 1: 265. 1933. TYPE: FRENCH GUIANA. Vicinity of Saint-Jean-du-Maroni, 8 May 1914 [fl], Benoist 1190 (LECTOTYPE,designated here: P). In the original description of Bathysa difformis, Benoist noted that the inflorescences o f this taxon are axillary, a character that excludes this species f r o m Bathysa. A few years later, he transferred Bathysa difformis to Pseudochimarrhis Ducke. Unfortunately, Benoist did not note its relationships to C. turbinata, with which I consider it synonymous. Common name. French Guiana: SamaatiPalioudou (Paramaka, collector unknown 7608), Bois Chapelle (French, collector unknown 7608). 37 Chimarrhis gentryana Delprete, sp. nov. (Fig. 1) TYPE. PERU. Loreto: Maynas, Mishana, Rio Nanay, halfway between Iquitos and Santa Marfa de Nanay, ca. 150 m, 73~ 3~ upland forest mostly on white sand, 1 M a r 1979 [fl], Gentry & Aronson 25307 (HOLOTYPE: MO). Arbor cortice sulcatis inftorescentia axillari, Chirnarrhi brevipe similis, a qua presentiam differt capsulis ferrugineis aureo-pubescentibus (vs. albo-pubescentibus) anguste obconicis (vs. late obovoideis). Trees up to 15-25(30) m tall, with small butresses; bark deeply fissured, fibrous; w o o d yellow and extremely hard: Young leafy branchlets 4 - 8 m m thick, semi-succulent, terete to subterete, glabrous; older branches smooth, grayish pale brown. No lenticels seen. Stipules connate at base and not contorted in bud, often connected to petioles, deltoid and not acuminate, glabrous outside, microscopically ( 4 0 • ciliolate, glabrous with a basal triangular area o f colleters inside, 8.5-9 m m long, 6 . 5 - 1 0 m m wide at base, subcaducous, reddish green, leaving a scar encircling the stem and connected to the petiole scar, 1 - 2 m m wide. Blades (7)12-23 c m long, (4)8-13 c m wide, l e n g t h : w i d t h 1.5-1.7:1, pandurate, obtuse to rounded at base, concave at basal area, obtuse at apex, a c u m e n absent; dark green a b o v e and pale green below, semicoriaceous; drying reddish brown, semileathery to leathery; glabrous a b o v e and below. P r i m a r y and secondary veins glabrous and prominent below, secondary veins 7 10 each side; tertiary veins subparallel throughout. Petioles 6 - 1 2 m m long, 2 - 3 m m thick, terete to adaxially flattened or narrowly concave, glabrous. D o m a t i a absent. Inflorescences 2 per node, c o r y m b o s e and densely branched distally, with opposite to subopposite decussate branches terminating into cymules, forming a continuous flowering plane; 9.5-15 c m long, 7 . 5 11 c m wide, lateral branches 1-2 pairs, basal portion of axis not branched, 4.5-7.5 c m long. Rachis basally subterete and distally decussately compressed, rachis and branches white or golden minute-puberulent; flowers on distal branches in condensed c y m u les. Distal bracts absent or extremely re- 38 BRITTONIA [VOL. 48 D L3 A FIG. 1. Chimarrhis gentryana. A. Habit of inflorescence with mature leaf. B. Flower at anthesis. C. Detail of flower. D. Style. E. Anther. F. Mature capsule. ( A - E drawn from Huashikat 2136, MO; F drawn from Vdsquez & Arevalo 8969, TEX). 1996] DELPRETE: RUBIACEAE duced, deltoid to linear, up to 4 m m long and up to 1 m m wide; bracteoles subtending the flowers absent or reduced to microscopic scales. Flowers v e r y fragrant, sessile to short-pedicellate, pedicel 0-1.5 m m long, puberulent; hypanthium narrowly obconica], 1-1.5 m m long, ca. 1 m m wide, puberulent. Calyx truncate to short-lobed, 0 . 4 0.8 m m long, 1.3-2 m m wide, golden puberulent, pale green; calyx lobes barely distinguishable, 0-0.5 m m long, in s o m e flowers one lobe o f the calyx expanded into a calycophyll. Calycophylls ovate to elliptic; blades 3.2-5 c m long and 1.6-3 c m wide, with peltate venation, pale green to white; petioles 1.5-2 c m long. Corolla funnelform and deeply lobed, 3.5-4.5 m m long, white to greenish white; tube short cylindrical 1.5-2.5 m m long, 0.5-0.7 m m wide, glabrous outside and inside; corolla lobes 5, reflexed at anthesis, 2-2.5 m m long, 0.6-0.8 m m wide at base, narrowly oblong with acute apex, glabrous outside, with a basal area white-pilose and with dista] zone glabrous inside. Stamens 5, obviously exserted, equal length; filaments attached 1 - 2 rnm f r o m the base of the tube; anthers elliptic, 0.6--0.7 m m long, 0.3-0.5 m m wide, dorsifixed at medial zone, base rounded, apex acute to short mucronate; illaments 3.5-5 m m long, terete, basally flattened and connate to throat, sparsely whitepilose at base. Style exserted, 5 - 6 m m long, terete, densely strigose, adpressed-retrorse; style branches extruding as two lips above corolla before anthesis, oblate, 0.3-0.6 m m long, s t i g m a t i c s u r f a c e m i c r o s c o p i c a l l y (40• papfllose. Capsules narrowly obconical to turbinate, acute at base, rounded at apex, 3.5-4.5 m m long, 1.5-3 m m wide, capsule rust-brown, densely golden puberulent, without lenticels; disk white-pubescent, not exceeding the calyx; disk-loculicidal dehiscence present in old capsules. Seeds with central hilum, with fringed concentric wing, 8 - 1 2 wide including wing. N of Pinglo, Quebrada Caterpiza, 200 m, 19 Feb 1980 (fl), Huashikat 2136 (MO). Loreto: Prov. Maynas, Mishana, Rfo Nanay, 75~ 3~ 130 m, 25 Jul 1984 (fr), Vdsquez et aL 5405 (NY); road Nauta-Iquitos, Nauta, 73~ 4~ 200 m, 28 Mar 1987 (fr), Vdsquez & Arevalo 8969 (F, TEX). BRAZIL. Amazonas: Mun. S~o Paolo de Oliven~a, basin of creek Belem, Oct-Dec 1936 (fr), K r u k o f f 8 7 8 2 (MO). The late A1 Gentry was a m o n g the m o s t significant botanists of the twentieth century. His dedication to fieldwork was extraordinary and his field k n o w l e d g e was phenomenal. It is with great honor and nostalgia that I dedicate this species to this dear colleague and never-forgotten friend. Chimarrhis gentryana occurs in the western A m a z o n Basin at relatively low elevations ( 1 5 0 - 3 9 0 m) and represents one of the m i d - c a n o p y trees o f that area. Common name. P e r r : T a y u m u k u n t (Huambisa, Huashikat 2136). Chimarrhis gentryana is m o s t similar to C. brevipes Steyermark of the Venezuelan A m a z o n (known only f r o m its type in fruiting stage); the latter differing f r o m the previous in having capsules broadly obovoid (vs. narrowly obconical), 4 . 5 - 6 m m long and 2.5-3.5 m m wide (vs. 3.5-4.5 m m long and 1.5-3 m m wide), rust-brown (vs. beigebrown), and sparsely white-puberulent (vs. densely golden-puberulent). One isotype (US) o f C. brevipes has one calycophyll attached to an aborted flower. Future collections o f flowering material o f C. brevipes will better clarify the relationship between these two species, which are similar in having broadly pandurate glabrous leaves with 5 - 9 secondary veins on each side. Chimarrhis gentryana has been often c o n f u s e d with Calycophyllum obovatum (Ducke) D u c k e probably because o f its white calycophylls, minute white flowers with exserted button-shaped anthers, obovate leaves with short stout petioles, and intrapetiolar triangular stipules. CALYCOPHYLLUM Specimens examined: ECUADOR. P a s t a z a : UNOCAL petroleum exploration well site, Mazaramu, 1~ 76~ 390 m, 28 Apr 1990 (fr), Beck et al. 1055 (QCNE); Kapawf, Rfo Pastaza, 76~ 2~ 'S, 235 In, 14-20 Jul 1988 (fr), Lewis et al. 13543 (QCNE). PERU. Amazonas: Valley of Rfo Sagrado, 65 krn 39 De Candolle Calycophyllum De Candolle, Prodr. 4: 367. 1830. TYPE: Calycophyllum candidissimum (Vahl) De Candolle Eukylista Bentham, Hook. J. Bot. 5: 230. 1853. TYPE: Eukylista spruceana Bentham (= Calyco- 40 BR1TTONIA phyllum spruceanum J. D. Hooker ex K. Schu- [VOL. 48 A m e r i c a for construction timber. D u c k e (1937) produced a key to the k n o w n species Semaphyllanthe L. Andersson, Ann. Missouri Bot. Gard. 82: 421. 1995. TYPE:Semaphyllantheobov- and Steyermark (1964) published a k e y to ata (Ducke) L. Andersson (=Calycophyllum the species o f the C. obovatum-acreanum obovatum (Ducke) Ducke. complex. Additional recent notes on Calycophyllum can be found in Taylor (1992) Trees (rarely shrubs), sometimes canopy and Andersson (1994, 1995). trees, a few species with small buttresses; As described above, Calycophyllum is a bark often exfoliating in long stripes; w o o d genus of approximately 10 species of trees white and very hard. Stipules interpetiolar, that occur in tropical Central and South free or connate at base, caducous. Blades America. Calycophyllum has been treated elliptic, ovate to obovate, obtuse to acumi- as belonging to the tribe Cinchoneae (Schunate at apex. Inflorescences terminal, cor- mann, 1889, 1891; Verdcourt, 1958; Roby m b i f o r m to p a n i c u l a t e , w i t h lateral brecht, 1988). In their attempt to circumbranches terminating in cymules. Flowers scribe the tribe Cinchoneae, Andersson & protandrous, often with two small m e m b r a - Persson (1991) r e m o v e d Calycophyllum, nous bracts including (or not including) the along with Alseis, Schizocalyx, and Wittflowers. Calyx truncate to reduced to min- mackanthus, placing these in the newly esute teeth, persistent; calyx lobes 5 or 6(8); t a b l i s h e d C a l y c o p h y l l e a e . A c c o r d i n g to in a few species, one lobe expanded into a their analysis, the Alseis-Calycophyllum c r e a m to yellowish white calycophyll, in clade was defined by six characters: c a m s o m e o f the flowers. Corolla funnelform to panulate corolla, distinctly exserted stacampanuliform and deeply lobed; corolla mens, filaments equalling anthers, buttontube narrowly funnelform; corolla lobes shaped anthers, exserted style, and long, (4)5-6(8), aestivation contorted to narrowly simple style branches. All of these characimbricate. Stamens alternate to the petals ters are found in Chimarrhis, which also and exserted; anthers button-shaped, dorsi- has very similar pollen grains (Aiello, fixed, dehiscing by lateral slits; filaments at- 1979; Andersson, 1993; pers. obs.) and diftached on the upper part of the tube, very fers mainly f r o m the previous group in havslender, glabrous throughout or with a tuft ing peltate placentation, seeds with narrow o f pilose to villous hairs at base. Style ex- concentric wing (vs. strongly bipolar), and serted; styie branches opening after c o m - axillary-subterminal inflorescences (vs. terplete maturity of the anthers, long, reflexed minal). at maturity. Ovary 2-celled, placentation In his recent synopsis o f the Cinchoneae imbricate, narrowly obovate. Capsules ob- complex based on cladistic analysis, Anovoid to narrowly obovoid, slightly later- dersson (1995) r e m o v e d six species f r o m ally c o m p r e s s e d , d e h i s c i n g septicidaliy; Calycophyllum to the newly established Sedisk-loculicidal dehiscence present in old maphyllanthe (with C. obovatum as type capsules of some species. Seeds elliptic, s p e c i e s o f the n e w g e n u s ) . A n d e r s s o n with bipolar wing. Pollen tricolporate; ex- (1995) reported that "'Semaphyltanthe difine densely reticulate. fers f r o m Calycophyllum in having free (vs. Calycophyllum is easily recognized by its c a l y p t r a t e ) stipules, p l a n e (vs. s a c c a t e ) reddish brown bark exfoliating in long ver- bracteoles, a distinct calyx (vs. reduced to tical stripes and exposing the inner deep- minute teeth), contorted (vs. imbricate) cogreen layer. Its trunks are usually twisted, rolla aestivation, filaments inserted in the even though quite rectilinear. O f all the spe- center of the corolla tube or lower (vs. at cies examined I have never o b s e r v e d indi- top), and filaments bristly hirsute toward viduals with buttresses, even a m o n g older base (vs. glabrous throughout)." In m y obtrees. The w o o d o f Calycophyllum is white servations o f Ducke 266 (NY; the voucher (vs. yellow in Chimarrhis) and of good o f C. obovatum studied by Andersson) and c o m m e r c i a l value. S o m e species [C. can- Ducke H.J.B.R. 22837 (R; type of C. obovdidissimum (Vahl.) DC. and C. spruceanum atum), C. obovatum has stipules that v a r y (Benth.) Hook.] are cultivated in tropical f r o m free to connate, calices with a variable rll ~rllCl). 1996] DELPRETE: RUBIACEAE basal tube, these often reduced to minute lobes, and filaments glabrous to basally pubescent. I studied C. candidissimum f r o m fresh material (Delprete & Apreza 6358) and from herbarium specimens, and I observed that the corolla aestivation is variably imbricate or contorted, filaments are variably glabrous or pubescent, and the filament insertion is variably medial or distal to the corolla tube. I also o b s e r v e d that the calices are minutely cupular or reduced to barely distinguishable teeth, and the filaments are glabrous to basally pubescent in C. spruceanum (Smith 4043, Hatschbach 29506). As a result o f the above observations, I conclude that Semaphyllanthe and Calycophyllum are synonymous. Calycophyllum obovatum (Ducke) D u c k e is described below because it has been often confused with Chimarrhis gentryana. (Ducke) OBOVATUM Ducke, Trop. Woods 49: 2-3. 1937. Warszewiczia obovata Ducke, Notizbt. Bot. Gart. Berlin-Dahlem 11: 475. 1932. Semaphyllanthe obovata (Ducke) L. Andersson, Ann. Missouri Bot. Gard. 82: 421. 1995. TYPE: Brazil. U p p e r Rio Negro region, vicinities o f Camanfios, in sil- CALYCOPHYLLUM vis humilioribus leviter paludosis ubi abundat palmae Leopoldina piassaba Wallace, 20 N o v 1929 (fl), Ducke Herb.Jard.Bot.R{o 22837 (HOLOTYPE: B, destroyed; Isorvfe: R). Trees up to 20 m tall and 22 c m dbh; bark smooth, reddish, peeling o f f in vertical stripes; w o o d extremely hard, yellowish white. Stipules connate at base (rarely free) and often connected to petioles, deltoid and not acuminate, glabrous outside, readily caducous, reddish green, leaving a scar encircling the stem and connected to the petiole scar, 1-2 m m wide. Blades 14-20 cm long, 6-11 c m wide, length : width 1.5-1.7:1, obovate, acute at base, obtuse at apex; dark green above and pale green below, semicoriaceous; drying reddish brown, semileathery; glabrous above and below. Primary, secondary, and tertiary veins glab r o u s and p r o m i n e n t b e l o w , s e c o n d a r y veins 7 - 1 0 each side; tertiary veins parallel throughout. Petioles 2 5 - 3 5 m m long, 3 - 4 41 m m thick, terete to adaxially flattened or narrowly concave, glabrous. D o m a t i a absent. Inflorescences terminal and solitary, c o r y m b o s e and densely branched distally, with opposite branches terminating into cymules, (7)14-20 c m long; lateral branches 6 - 1 8 c m long with terminal cymules 4 . 5 - 6 c m wide. Rachis basally subterete and distally decussatoly compressed, rachis and branches white or golden minute-puberulent; flowers on distal branches in condensed cymules. Distal bracts absent to foliose, deltoid to obovate (when foliose); bracteoles subtending the flowers absent or reduced to microscopic scales. Flowers fragrant, sessile to short-pedicellate, pedicel 0-1.5 m m long, puberulent; hypanthium narrowly obconical, 1-1.5 m m long, ca. 1 m m wide, puberulent. Calyx truncate to short-lobed, 0.3-0.6 m m long, 1-1.7 m m wide, golden puberulent, pale green; calyx lobes barely distinguishable, 0-0.5 m m long, in some flowers one lobe of the calyx expanded into a calycophylL Calycophylls obovate, blades 3.5-5 c m long and 1.4-3 c m wide, with peltate venation, white to c r e a m white; petioles 1-2 c m long. Corolla funnelform and deeply lobed, 3.5~4.5 m m long, white to greenish white; tube short cylindrical, 1.5-2.5 m m long, 0.4-0.6 m m wide, pubescent outside and glabrous inside; corolla lobes 5, rounded and with a fringed margin, 0.6-0.7 m m wide, sparsely puberulent outside, glabrous inside. Stamens 5, obviously exserted, equal length; filaments attached 1.5-2 m m f r o m the base o f the tube; anthers elliptic, 5 m m long and 3 m m wide; dorsifixed, base rounded, apex acute to short mucronate; filaments terete, basally flattened and connate to throat, sparsely white pilose at base (sometimes glabrous throughout). Style exsertod, ca. 4 m m long, terete, glabrous; style branches opening after complete release of pollen, ca. 1 m m long, stigmatic surface microscopically ( 4 0 • papillose. Capsules narrowly obovoid, acute at base, narrowly rounded at apex, 8 - 1 5 m m long, 3.5-5 m m wide, capsule b l a c k , s p a r s e l y m i n u t e - p u b e r u l e n t , without lenticels; disk black and glabrous, not exceeding the calyx; disk-loculicidal dehiscence present in old capsules. Seeds strongly bipolar, with acute apices, 8 - 1 3 42 BRITTONIA m m long and 0 . 9 - 2 m m wide including wing. Specimens examined: VENEZUELA. Amazonas: San Carlos de Rfo Negro, ca. 20 km S of confluence of R/o Negro and Brazo Casiquiare, l~ 67~ 119 m, 4.3 km NNE of Solano rd., 29 Jan 1980 (fr), Clark & Maquirino 7310 (NY), 9 Aug 1980 (fr), Clark & Maquirino 7656 (NY); Dpto. Atabapo, Rfo Cuchaken, 3~ 67~ 1000 m, Oct 1989, Velazco 767 (NY); Dpto. Atabapo, E slope of Cerro Cucurito, river bank of Carlo Yagua, 3~ 66~ 120 m, 28 Jun 1979 (fr), Huber 3883 (NY). BRAZIL. Amazonas: Rfo Negro, Taraqu~i, 5 Mar 1959 (fl), G. S. Rodrigues 190 (NY); Rio Curicuriary, Cajti waterfalls, 29 Feb 1936 (fl), Ducke 266, Herb.Jard.Bot.Rfo 29034 (NY), Yale School of Forestry 32642 (NY---2 sheets). Calycophytlum obovatum is a small tree that is usually 8 - 1 2 m tall and 10-15 c m dbh, with one specimen reported to be 20 In tall and 22 c m dbh (Clark & Maquirino 7656). As in other species of Calycophyllure, the bark of this species exfoliates in long vertical strips, exposing an inner deepgreen layer. Calycophyllum obovatum is reported f r o m very low elevations ( 1 0 0 - 1 5 0 m), usually in s w a m p y areas of southern Venezuelan A m a z o n and Brazilian Catingas. Common names. Venezuela: G u a y a b o (Velazco 767), guayab6n (Clark & Maquirino 7310, 7656). Brazil: Pfio mulato da catinga (Ducke, 1937). B A T H Y S A Presley Bathysa Presley, Bot. B e m e r k 84. 1844. (Vellozo) PresTYPE: Bathysa sr ley. Schoenleinia Klotzsch in Haine, Arzneigew. 14, t 15. 1846. TYPE: Schoenleinia cuspidata (Saint-Hihire) Klotzsch. Voigtia Klotzsch, 1. c. TYPE: Voigtia australis (SaintHilaire) Klotzsch. Trees, rarely shrubs; bole irregular, with or without buttresses; buttresses sometimes v e r y large, especially in the A m a z o n i a n species; bark fissured, light brown to grayish; w o o d yellow to reddish and v e r y hard. Stipules interpetiolar, free or connate at base, persistent or caducous. Blades elliptic to ovate to obovate, usually acuminate at apex; domatia absent, or tuft of hairs. Inflorescences terminal and solitary, c o r y m b o s e and with opposite to subopposite lateral [VOL. 48 branches terminating in cymules. Flowers protandrous, sessile to pedicellate, flagrant; hypanthium obconical to turbinate. Calyx t r u n c a t e or w i t h b a r e l y d i s t i n g u i s h a b l e lobes, persistent; calyx lobes 4 or 5; one lobe expanded into a c r e a m to yellowish white calycophyll in a few species, in s o m e of the flowers. Corolla funnelform to subrotate, white, greenish white, or creamwhite; corolla tube funnelform to ciatiform; with a ring of pubescent hairs inside on the tube or on the base o f the lobes; corolla lobes 4 or 5, rounded; aestivation imbricate. Stamens central to the adjacent petals or alternate to the petals, exserted; anthers elliptic, dorsifixed, dehiscing by lateral slits; illaments attached on the upper part o f the tube, slender and basally flattened, with a tuft of pilose to villous hairs at base. Style exserted; style branches opening after c o m plete maturity o f the anthers, long, narrowly o b l o n g , r e f l e x e d at m a t u r i t y . Ovary 2celled, seeds horizontally attached on the septum (not peltate), obconical to oblong, with m a n y ovules in each locule. Capsules globose, oblong or obovoid, dehiscing septicidally; disk-loculicidal dehiscence present in old capsules of s o m e species. Seeds with lateral hilum, uncinulate, flattened at polar ends, rarely with small wing. Pollen tricolporate, exine densely reticulate. As described above, Bathysa is a genus of approximately 14 species of trees (rarely shrubs) mainly f r o m South A m e r i c a (especially the A m a z o n basin), with one species f r o m Central America. Bathysa has been treated as belonging to the tribe Rondeletieae (Schumann, 1889, 1891; Verdcourt, 1958; Robbrecht, 1988). Bathysa is another important lumber tree in the A m a zon Basin, with hardwood that varies f r o m yellow to reddish, and reaching up to 40 m height. A key to the Brazilian k n o w n species was published by Schumann (1889), but a m o d e r n monographic treatment is still needed. Because Bathysa has been often confused with Chimarrhis, the following combinations are necessary. Bathysa bathysoides (Steyermark) Delprete, comb. nov. Chimarrhis bathysoides Steyermark, Mem. New 19967 DELPRETE: RUBIACEAE York Bot. Gard. 12(3): 181, fig. 29. 1965o TYPE: V E N E Z U E L A . Amazonas: Cerro Sipapo (Parfique), along trail near Base Camp, 25 Jan 1949 (tt), Maquire & Politi 28626 (HOLOTYPE: NY; 1SOTYPES: B, GH, U S - - 2 sheets). Specimens examined: COLOMBIA. M e t a : Sierra de la Macarena, carlo Ciervo, 500 m, 4 Jan 1950 (fl), Philipson, Idrobo & Jaramillo 2018 ( U S - - 2 sheets). Vaup~s: ARo Vanp6s, vicinity o f Miraflores, 300 m, 29 Jan 1944 (fl), Gutierrez & Schultes 704 (NY). V E N E Z U E L A . A m a z o n a s : Sierra Parima, vicinity of Simarawochi, Rio Matacuni, 3~ 64~ 6-7 km from Brazilian border, 800 m, 18 Apr-23 May 1973 (fr), Steyermark 107036 (E G, MO, NY); Dpto. Atures, W side o f Valley o f Rio Coro-Coro, N N W o f Yutaje, 5~ 66~ 500-1000 m, 26 Feb 1987 (fr), Liesner & Hoist 21414 (NY); foothills o f S Guiana Highlands, Carlo Iguana, 5~ 65~ 200 m, 23 Jan 1976 (fl), Lister & Colchester 2113 ( K ~ 2 sheets); Rio Orinoco, confluence with Rio Ugueto, 2~ 63~ 3 6 0 4 2 0 m, 4 Jan 1952 (fl), Cruxent 139, 140, 141, 163, 164 (NY). Bolivar: Dtto. Cedefio, Serrania de los Pijiguaos, 6~ 66~ 500-600 m, 29 Jan 1989 (fl), Cuello 641, 691 (NY); Dtto. Cedefio, 20 km E o f Tfiriba, 6-11 Dec 1970 (fl), MarcanoBerti 2578 (SP). BRAZIL. A m a z o n a s : Upper Rio Negro basin, along Rio Castanho, tributary of Rio Paduiri, 100-140 m, 16-24 Feb 1946 (fr), Cardona 1379 (NY). BOLIVIA. L a Paz: Prov. Larecaja, Tuiri, near Mapiri, left bank of Rio Mapiri, 500-750 m, 12-30 Sep 1939 (fr), Krukoff 10746 (NY). ~n the original description of this species, Steyermark (t965) did not report its solitary terminal inflorescence and its well-developed persistent cupular calyx, both characters typical of Bathysa. In Chimarrhis the inflorescences are always axillary and its calyx extremely reduced. Its finely and densely parallel tertiary venation make this species easily distinguishable among the other species of Bathysa. B a t h y s a p e r i j a g n s i s (Steyermark) Delprete, comb. nov. Chimarrhis perija~nsis Steyermark, Acta Bot. Venez. 8: 248. 1973. TYPE: V E N E Z U E L A . Zulia: Sierra Perijfi, forest near Rio Omira-Kun~ (Tnmuriasa), SW of Pishikako, 1440-1460 m, 28 Mar 1972 (fl), Steyermark & Dunsterville 105658 (HOLOTVr'E: VEN-n.v.). Specimens examined: V E N E Z U E L A . Zulia: Sierra Perijfi, forest near Rio Omira-Kunfi (Tumuriasa), SW of Pishikako, 1440-1460 m, 22-23 Mar 1972 (st), Steyermark & Dunsterville 105555 (NY, US), SW of Pishikako and Irfa, 1470-1560 m, 25 Mar 1972 (fr), Steyermark & Dunsterville 105655 (E US). The two main characters that differenti- 43 ate this species from Bathysa bathysoides are the denser spreading pubescence o f the calyx and rachis and the corolla length. Future collections might well prove that these two taxa should be reduced to varietal rank. Bathysa perijaensis is k n o w n only from Sierra Perijgt, close to the C o l o m b i a n border, at elevations much higher ( 1 4 0 0 - 1 6 0 0 m) than B. bathysoides, w h i c h is mainly o f A m a z o n i a n distribution. Common names. Venezuela: Tubabanoche (Steyermark & Dunsterville I05655), Rubabanoche (Steyermark & Dunsterville 105555). BATHYSA P~TrIERI (Standley) Steyerrnark, Mere. N e w York Bot. Gard. 12(6): 185. 1965. Chimarrhis pittieri Standley, Field Mus. Bot. 8: 53. 1930. TYPE: V E N E Z U E L A . Dto. Federal: Caruao, 7 N o v 1925 (fr), Pittier 11954 (HOLOTYPE: US, frag--E US--photos at F, G; ISOTYPES: G, NY). Representative specimens examined: VENEZUELA. Distrito F e d e r a l : Cerro Nalguat~i, N slopes, vicinity of Quebrada Basenilla, 900-1000 m, 27 Jun 1964 (fl), Manara 44 (E NY); Cerro Naiguat~i, N slopes, Quebrada Frontina, 900-1100 m, 2 Nov 1963 (fr), Steyermark 91843 (NY, US); Cerro Nalguatfi, between La Chocha and Las Delicias, 800 m, 31 Mar 1972 (fr), Morillo & Manara 2138 (E M O ~ 2 sheets); Carayaca, Dec 1965 (fr), Aristeguieta 5454 (E MO, N Y ~ 2 sheets). M i r a n d a : Caracas, E1 Cafetal, 10 Jul 1966 (fl), Aristeguieta 6170 (US); Alto Prado, Quebrada Manzanares, creek o f Quebrada Baruta, 10~ 66~ 1300 m, 8 Oct 198X (fr), Steyermark et al. 123593 (MO); Cerro Baciller, Quebrada Corozal, S o f Santa Cruz, 10~ 65~ 20-26 Mar 1978 (fr), Steyermark & Davidse 116496 (MO, NY); Dto. Pa6z, Fila La Tigra, Quebrada San Juan, 10~ 65~ 500 m, 2-7 Sep 1977 (fr), Gonzdlez & Ortega 1347 (NY); rd. Las Caracas-Cabo Codera-Higuerote, 17 Jun 1947 (fr), Aristeguieta 4847 (NY). Steyermark (1965) transferred this species from Chimarrhis to Bathysa, a decision with which I am in c o m p l e t e agreement. This Venezuelan endemic is usually a tree 5 - 1 2 m tall and is easily recognized by its membranous leaves and w e l l - d e v e l o p e d cupular calyx. Common name. Venezuela: Lengua de vaca (Pittier 11954). Acknowledgments I am grateful to the directors and staff o f the f o l l o w i n g institutions for loans o f ma- 44 BRITTONIA terial and for help in field work: AAU, B, BR, CAS, CAY, CHOCO, COL, CR, DAV, D S , F, G , G B , G H , I N P A , J B G P , K , L L , M , M O , N Y , R Q C A , Q C N E , R , S, T E X , U B , U C , U S , W. [ t h a n k B i l l i e T u r n e r , A l a n Prather, and Guy Nesom for reviewing the manuscript and the latter for reviewing the Latin diagnosis. I am particularly grateful to Charlotte Taylor and Brian Boom for helpful comments on the final revision of t h e m a n u s c r i p t . T h e i l l u s t r a t i o n is p r o v i d e d by the author. Literature Cited Aiello, A. 1979. A reexamination of Portlandia (Rubiaceae) and associated taxa. J. Arnold Arbor. 60: 38-123. Andersson, L. 1993. Pollen characteristics of the tribe Calycophylleae, Cinchoneae, and Hillieae (Rubiaceae). Nordic J. But. 13: 405-417. . 1994. Calycophylleae. Pages 82-85. In: R. Harling & L. Andersson (eds.). Flora of Ecuador 50: 82-85. . 1995. Tribes and genera of the Cinchoneae complex. Ann. Missouri But. Gard. 82: 409-427. - & C. Persson. 1991. Circumscription of the tribe Cinchoneae (Rubiaceae)--a cladistic approach. PL Syst. Evol. 178: 65-94. Boom, B. M. & M. T. V. do Amaral Campus. 1991. 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Die Natiirlichen Pflanzenfamilien 4(4, 5). Standley, P. C. 1929. Rubiaceae. Studies of american plants, Publ. Field. Mus., But. 4: 264-298, 324345. Steyermark, J. A. 1964. In: B. Maguire and collaborators. Botany of the Guayana Highlands. Mem. New York But. Gard. 10: 186-278. 91965. In: B. Maguire and collaborators, Botany of the Guayana Highlands. Mere. New York But. Gard. 12: 230-436. Taylor, C. M. 1992. Notes on the Rubiaceae of Peril. Novon 2: 438-442. Verdcourt, B. 1958. Remarks on the classification of the Rubiaceae. Bull. Jard. But. Brnxelles 28: 209281.