Phytologia(February 1993) 74(2):61-103. TAXONOMIC OVERVIEW OF STEMODIA (SCROPHULARIACEAE) FOR NORTH AMERICA AND THE WEST INDIES Billie L. Turner & Clark C. Cowan Department of Botany, University of Texas, Austin, Texas 78713 U.S.A. ABSTRACT A tazononiic study of the species of Stemodia occurring in North America and the West Indies is rendered. Seventeen species are rec- ognized as native withinthis region. These include Stemodia angviata, S. bartsioides, S. chiapensis, 5. coahiiilensis, S. dttTantifolia, S. frati- cosa, S. jorullenais, 5. lanatcL, S. macTantha^ S. maritinm, S. multifida, S. palmeri, S. peduncidaris, S. pusilla, S. schottii, S. tenuifolia, and S. verticillata. Except for Stemodia coahuilenais and 5. multifida, which are currently under revisionby David Keil as part ofhis concept ofthe genus Leucoapora, a key to species, descriptions, distributional maps, and completesynonymyforeachoftheseis provided. One new species, Stemodia chiapensisB.L. Turner,isproposed. Thegenus Leucoapora istreatedaspartofStemodia(s.l.)but Schistophragmaisretained. This has necessitated the following new combinations: Schistophragma polystachya (Brandegee) B.L. Turner and Stemodia coahuilensis (Henr.) B.L. Turner. KEYWORDS: Scrophulariaceae, Leucospora,Schistophragma^Ste- modia 5<emo(ftabelongs tothefamilyScrophulariaceae,subfamilyAntirrhinoideae, tribe Gratioleae, whereit has been variously accepted,either as arather large, highly variable genus Stem,odia (s.l.) or as a less diverse, somewhat smaller genus Stemodia (s.s.). Most early workers more or less took the former stance (e.^., Bentham 1886; Wettstein 1891) but Minod (1918), in his revisionary study of the New World taxa, drastically reoriented taxonomic concepts or attitudes on the group by recognizing a monotypic Stemodiacra (typified by Stemodia maritim,a L.), an older name for Stemodia. Stemodia, however, was subsequently conserved over Stem,odiacra. Minod also segregated out of his 61 PHYTOLOGIA 62 volume 74(2):61-103 February 1993 Stemodiacomplex, four newly erected monotypic genera Chodaphyton, based ( upon Stemodia ericifolia [Kuntze] Hassler; Lindemia, based upon Stemodia verticillata (Miller) Hassler; Valeria, based upon Stemodia trifoliata [Link] Reichenbach [= Stemodia pratensis (Aublet) C. Cowan of the present treat- ment]; and Verena, based upon 'Stemodia hassleriana Chodat). He retained most of the remaining New World taxa of Stemodia (s.l.) in his concept of Stemodia. So treated, he recognized 31 species of Stemodia as occurring in the New World, thirteen of these occurring in the Americas north of South America. Most subsequent workers (e.^., Barroso 1952; Dawson 1974; D'Arcy 1979; Seymour 1976; Standley & Williams 1973) have continued to treat Stemodia in the broad sense, keeping in it most of the monotypic segregates proposed by Minod. We also subscribe to the wider view, with the exception that the Cuban species, Stemodia radicans Griseb., which was retained by Minod in Stemodia, is accepted as belonging to the genus Cheilophyllum., as first proposed by Pennell (1935). In addition, we include in Stemodia the genus Leucospora, which most recent workers have treated as distinct. Since Keil (in prep.) is currentlyrevising Leucospora(including Schistophragma) we have not treated in detail, nor mapped, species of the latter. We have also separated from Stem,odia three recently described closely related species, 5. reliquiarum D'Arcy, S. mutisii Fernandez-Alonso, and 5. costaricensis B.L. Turner, positioning these in a newly erected genus Darcya (Turner & Cowan 1993). Insummary,thepresenttreatmentrecognizesseventeenspeciesofStem,odia (s.l.) as native in the New World north of South America. (A single record of 5. pratensis from Trinidad is believed to be a recent introduction from the mainland of South America). Approximately nineteen species are recognized for South America, but these wiU be treated in a subsequent account (Turner & Cowan 1993). In addition, some ten to fifteen species are native to the Old World (including those in the recently submerged genus Morgania R. Br. (cf. Barker 1981), which we do not intend to treat. CHROMOSOME NUMBERS Chromosome counts for two New World species of Stem,odia have been reported, these being 2n = 44 for 5. multifida (Keil, pers. comm.) and 2n = 22 for S. verticillata (Fedorov 1974; reported as 5. parvifloraW.T. Alton). An additional Old Word species, Stemodia viscosa Roxb. has been reported as 2n = 28 (Subramanian k Pondmudi 1987) and 2n = 42 (Fedorov 1975). Ifthese numbers prove correct, the genus (s.l.) is likely to prove dibasic or perhaps multibasic, with numbers ofx= 11 and i= 14 (ifnot x = 7). Clearly, additional counts in the genus are sorely needed. Turner «fe Cowan; Stemodiain North America and West Indies 63 GENERIC CONSIDERATIONS As alreadynoted, Stemodiahas been variouslytreated, usuallyin thebroad sense, but Minod (1918) retained but a single species in the now conserved genus Stemodia (typified by 5. maritima), which he called Stem,odiacra P. Br. (1756). So treated, the next available generic name, Phaelypea P. Br. (1756) would have to be applied to the 40 or more species that make up Stemodia (s.l.). Regardless, ifStemodia were treated as monotypic, numerous new combinations under the obscure name Phaelypea (which is sufficiently obscure so as not to have been listed in Index Kewensis) would have to be made. Indeed, Stemodia(s.l.) appears to beamonophyleticassemblagelargely composed of species having a syndrome of characters, namely: 1) opposite subpinnate or pinnately veinedleaves; 2) flowers axillaryor in terminal spikes; 3) calyx with sepals ± alike and separate to the base; 4) corollas mostly lavender to purple, zygomorphic with well developed tubes; 5) anther thecae glabrous, divergent, with swollen connectives, or the thecae shortly stalked; 6) styles at anthesis 2-4 times as long as the stigmatic area; 7) stigmatic area enlarged and usually recurved, bifid; 8) capsule mostly loculicidally 4 valvate, ovoid to orbicvdar, 1.0-1.5 times as long as wide; 9) seeds pedicellate, ellipsoid to broadly obpyramidal, deeply 8 sulcate or not, the ridges parallel, never spirzd, the surfaces variously ornate. Most of the species of Stemodia (s.l.) will contain seven or more of these characters. As conceived here, the closest related taxon to Stemodia (s.l.) is the genus Schistophragma which is composed of three species, all of these native to Mexico and distinguished by elongate capsules with septicidal de- hiscence and seeds with helicalgrooves, characters which are not found in the 50 or more species ofStemodia (s.l.). Beyond Schistophragma, the genus most closely related to Stemodia (s.l.) might be Bacopa, but this is mostly specula- tive. Clearly, Stemodia (s.s.) and its cohorts aire in need of detailed character analysis, especially those derived from comparative DNA studies, before dras- tic generic redispositions are made, ifany. It is our personal opinion, however, that Stemodia(s.l.) is a monophyleticassemblage whose internal restructuring might best be shown through nomenclatural constructions at the infrageneric level. CHEMICAL INVESTIGATIONS Two New World species of Stem.odia, S. durantifolia (L.) Swartz and S. maritima, have reportedly been used in herbal medicine by local peoples, at leastinHaiti. Becauseofthis, attemptshavebeenmadetoelucidateanyactive ingredients in these species. Stemodane triterpenes and yet other compounds 64 P HYTO LO GIA volume 74(2):61-103 February 1993 were isolated but none of these appeared to possess especially significant an- tiviral or cytotoxicity attributes (c/. Weniger et al. 1982, Hufford et al. 1992, and references therein). TAXONOMY Stemodia L. (s.l.) 1759, nom. conserv. Cordium Sloajie 1707. Erinus Miller 1731. Stemodiacra P. Br. 1756. Phaelypea P. Br. 1756. Matourea Aublet 1775. Adenosma R. Br. 1810, not Adenosma Nees. Morgania R. Br. 1810. LeucosporaNutt. 1834. Chodaphyton Minod 1918. Lendneria Minod 1918. Valeria Minod 1918. Verena Minod 1918. m Annual or perennial herbs, shrublets or small scrambling shrubs to 3 high. Leaves oppositeor less often verticillate,simpleto bipinnatelydissected, mostlysubpinnately veined. Flowersaxillary,arranged (l-)2-4 to anode,often densely clustered along the upper stems forming well defined, usually inter- rupted, spikes. Sepals 5, ± adike, separate to the bsise or nearly so. Corollas mostly tubular, white to blue or violet, rarely somewhat yellowish, zygomor- phic, lobes usually shorter than the tube, variously pubescent without and within, rarely glabrous, the inner surface near the throat of the tube usually bestowed with elongate hairs with spatulate apices. Anther bearing stamens usuadly 4, the anther thecae glabrous, not closely adjacent or parallel, usu- ally separated by a swollen or enlarged connective,less often the thecae borne upon well developed stalks. Stylar shaft 2-10 times as long as the enlarged minutely bilobed stigmatic region, the latter usually reflexed, less often erect, or somewhat incurved, rarely markedlybilobed. Capsules ovoid, mostly some- what longer than wide, glabrous, 4 valvate with usually loculicidal dehiscence. Seeds numerous, ellipsoid to broadly obpyramidal, deeply longitudinally 6-8 sulcate or not, usually to some extent stipitate and variously ornate. Base chromosome numbers, i = 11 and 14 (from only 3 species). Type species, Stemodia maritima L. . Turner &: Cowan: Stemodia in North America, and West Indies 65 KEY TO NORTH AMERICAN AND WEST INDIAN SPECIES OF STEMODIA, S.L. (with the exclusion of Leucospora, which has two species both with dissected leaves and both ofwhich belong to Stemodiain the senior author's opinion; cf. comments under CONTROVERSIAL AND EXCLUDED NAMES at the end of this paper) 1. Leaves clearly petiolate, the blades tapering upon the petioles to the very base; calyx w/o bracts at the base; seeds deeply 6-8 sulcate •••(2) 1. Leaves sessile; calyx with 1 or 2 basal bracteoles; seeds not clearly sulcate. •••(11) 2. SufFruticose brittle stemmed herbs, shrublets or shrubs, mostly 0.5- m 2.0 high; leaves drying black; Belize, El Salvador, Honduras. . 5. fruticosa 2. Annualorperennialherbs (rarelysufFruticoseat base),mostly0.1-1.5 m high; leaves drying green or greenish; widespread. ..........(3) m 3. Perennial herbs mostly 0.3-1.5 high (4) 3. Annual herbs mostly 0.1-0.3(-0.4) m high (7) mm 4. Pedicels (both flower and fruiting) 1-6 long (5) mm mm 4. Pedicels 10-40 long; petioles 8-20 long; widespread. ••• (6) mm mm 5. Petioles 1-6 long; stems hirsute, the hairs 1 long or more; Jalisco, Michoacan, Mexico. S. tenuifolia mm mm 5. Petioles 8-20 long; stems puberulent, the hairs 0.4 long or less; Chiapas. 5. chiapensis 6. Leaves markedlypunctate beneath with amber colored glands; vesti- mm ture of stem mostly 1 high or less, and largely devoid ofglan- dular hairs; eastern Mexico (Hidalgo) to Chiapas and southwards to Panama. 5. peduncularis 6. Leaves moderately to inconspicuously punctate beneath with yellow mm or clear glands; vestiture of stem mostly 1-2 high and usually beneath this copiously covered with much shorter glandular hairs; western Mexico (Nayarit) to Chiapas and southwards to Honduras. . S. macrantha 7. Leaves markedly glandular punctate beneath; West Indies, Central Amer- ica (including Chiapas) and northwestern South America. . S. angulata 66 P HYT L GIA volume 74(2):61-103 February 1993 7. Leaves epunctate beneath or nearly so; Mexico (8) 8. Stemsstifflyerect,not muchbranched from thebase; petiolesmostly mm 3-6 long; Jalisco to Chiapas 5. joruUensis 8. Stems variously ascending to decumbent, much branched from the mm base; petioles mostly 6-20 long (9) mm mm 9. Corollas 2-3 long; pedicels mostly 1-2 long 5. verticillata mm mm 9. Corollas 6-16 long; pedicels mostly 3-30 long (10) mm mm 10. Corollas mostly 6-10 long; pedicels mostly 3-8 long; peti- mm oles mostly 3-9 long; coastal regions from Colima to Sinaloa, also in Cape Region of Baja California. 5. pxisilla mm mm 10. Corollas mostly 12-16 long; pedicels mostly 20-30 long; mm petioles mostly 9-18 long; inland regions of short tree forests, southern Sonora, northern Sinaloa and closelyadjacent Chihuahua. 5. palmeri 11. Stems and foliage densely white lanate 5. tomentoaa 11. Stems and foliage otherwise (12) 12. Annualtaprooted herbs 5-25 cmhigh; stems ± uniformlypubescent mm with short glandular hairs 0.2 longor less; northeastern Mexico and closely adjacent U.S.A 5. schottii 12. Perennial herbs mostly 20-100 cm high, frequently forming rhi- zomatous colonies; stems variously pubescent with longer hairs; widespread but not overlapping the range of 5. schottii (13) mm 13. Corollas mostly 11-15 long; locatlized semiaquatic inland species of southcentral and western Mexico (Durango to Michoacan and Mexico). . • 5. bartsioides mm 13. Corollas mostly 2-8 long; widespread weedy species (14) mm 14. Corollas 2.5-4.8 long, the tubes glabrous or nearly so; stems sufFruticose; anther thecae widely separated by pedicels up to 0.5 mm long 5. Tnaritima mm 14. CoroUais 5-8 long, the tubes clearly pubescent; stems herba- ceous; anther thecae close, merely separated by a globose connec- tive S. durantifolia Turner «& Cowan; Stemodiain North America and West Indies 67 STEMODIA ANGULATA Oerst. StemodiaangulataOerst.,Vidensk.Meddel. Dansk. Naturhist. Foren. Kjoben- harn 1853:22. 1854. Stemodiacra angulata (Oerst.) Kuntze, Rev. Gen. Pi 2:466. 1891. TYPE: COSTARICA. Cartago: "prope Cartago", w/o date, Oersted P^72(LECTOTYPE: C! [selected here]; Photolectotypes: F!,GH!; Isolectotype: K!). Oersted in his protoiogue also mentioned a Fendler collection which belongs to this taxon. Stemodia ageratifolia C. Wright in Sauville, Fl. Cubana 99. 1873. Ste- m,odia angulata Oerst. subsp. ageratifolia (C. Wright) Minod, Bull. Soc. Bot. Geneve, ser. II 10:191. 1918. Lindemia ageratifolia (C. Wright) Pennell, Proc. Acad. Nat. Sci. Phila. 75:13. 1923. TYPE: CUBA. Pinar del Rio: along margin ofarroyos, Luiz Lazo and Ar- royo Hondo, 1860-1864, Wright 299S{EOLOTYPE: GH!; Isotypes: G!,GH!,MO!,NY,US!). StemodiajorullenaisH.B.K. subsp. reptanaMinqd, Bull. Soc. Bot. Gen- eve, ser. II 10:190. 118. TYPE: NICARAGUA. Rivas: De de Omatepec, rues du village de Mayagulpa, 40 m, Oct 1869, P. Levy 154 (HOLOTYPE: G-BOIS!; Isotypes: C!,G!). Annual or short lived perennial (?) herbs mostly 5-30 cm high. Stems erect at first, those at the lower nodes often procumbent, moderately crinkly mm pilose with multiseptate hairs 1-2 long. Midstem leaves mostly 1.0-2.5 cm long, 0.7-1.5 cm wide; petioles 5-10 mm long; blades ovate to subdeltoid, subpinnatelynerved,gradingintothe petioles, sparsely pilose,glandular punc- tate beneath, the margins crenulodentate. Flowers axillary, arranged 1-3 at a node, the peduncles ebracteate, mostly 1-2 cm long, pubescent likethe stems. mm Sepals mostly 4-5 long, pilose, one ofthese somewhat larger and broader. mm Corollas mostly 7-9 long, white or pinkish, the tubes glabrous or nearly mm mm so, the lobes 1-2 long, sparsely pubescent. Anther thecae ca. 0.6 long, glabrous, separated by a small globose connective. Capsule ovoid, 4-5 mm mm high, (3-)4 valvate, the apices recurved. Seeds ca. 0.5 long, stipitate, longitudinally sulcate with ca. 6-8 ribs. DISTRIBUTION (Figures 1 and 2): Mexico (Chiapas), Cuba, Central America and northwestern South America, 100-1000 m; flowering all seasons. PenneU (1923), treated Stem.odia ageratifolia as a good species, believing this to be endemic to Cuba. Minod (1918) treated this as a subspecies of S. jomllensis. For additional discussion see the latter. REPRESENTATIVE SPECIMENS: MEXICO. Chiapas: Mpio. Cacahoatan, 1-2 km S ofCacahoatan, 90 m, 24 Nov 1980, Breedlove 47760 (CAS). 68 P HYT LO GIA volume 74(2):61-103 February 1993 Figure 1. American distribution of Stemodia angulata. Turner <fe Cowan: Stemodiain North Americaand West Indies 69 <^ Figure 2. Central American distribution of Stemodia angulata. 70 P HYT LO GIA volume 74(2):61-103 February 1993 WEST INDIES: CUBA. Pinar del Rio: Ekman 10638 (K). CENTRAL AMERICA: BELIZE, pine ridge near Manatee Lagoon, 3 Jan 1903, Peck 263(GH). COSTA RICA. Alajuela: 3 km EofAlajuela, 1000 m, 29 Dec 1974, Wilbur 19048 (DUKE). EL SALVADOR. Ahuachapan: vicinity of Ahuachapan, 800-1000 m, 9-27 Jan 1922, Standley 20013 (GH). GUATEMALA. Santa Ana: vicinity of Metepan, 370 m, 29 Jan-1 Feb 1947, Standley s.n. {¥). HONDURAS. Olancho: vicinity ofJuticalpa, 380-480 m, 5-16 Mar 1949, Standley 17490 {¥). NICARAGUA. Chontales: w/o date, Tate 223 (K). Managua: Sierras de Managua, 800 m, 6-14 Jan 1941, Grant 1002(GE). PANAMA. Darien: Rio Morti, ca. 6 mi upstream from Morti Abajo, ca. 100 m, 14 Feb 1967, Duke 10160 (DUKE,MO). SOUTH AMERICA: COLOMBIA. Bolivar: Frasquillo, on Rio Sinu, 20- 100 m, 5-6 Mar 1918, Pennell 4192 (C,K). El Valle: Cisneros, 300-500 m, 5 May 1939, Killip 35615 (F,PH,US). ECUADOR. Esmeraldas: Playa de Oro, Jul-Aug 1924, Thomas L30 (K). Los Rios: 14 km SE of Quevedo, 75 m, 22 Feb 1972, MacBryde 1122 (MO). PERU. Amazonaw: Labanda, Huampami, Rio Cenepa, Chacra, 600-700 ft, 3 Aug 1974, Ancuaah 712 (F,MO,TEX). STEMODIA BARTSIOIDES Benth. Stemodia bartsioides Benth., Bot. Reg. t. 1470. 1831. Stemodiacra bart- sioides (Benth.) Kuntze, Rev. Gen. PI. 2:466. 1891. TYPE: MEXICO. Michoacan(?): "Plain ofPopetongo", 1830, Graham 275(HOLOTYPE: K!; Photoholotype: NY!). Stiffly erect perennial herbs 15-40 cm high. Stems essentially unbranched except at the very baise, densely glandular pubescent, often arising from stout rhizomes. Midstemleaves mostly 2.0-3.5 cm long, 4-9 mm wide, sessile,lance- olate, widest near the middle, 2 or 3 at node, gradually reduced upwards, weakly subpinnately nervate, glandular pubescent, the surfaces also glandular punctate, the margins serrulate. Flowers mostly axillary along the upper half mm of the stems, arranged 2-4 at a node, the pedicels 1-8 long, glandular mm pubescent. Sepals 5-6 long, glandular pubescent, essentially alike, below these 1 or 2 bracts about as long or somewhat longer than the sepals. Corollas mm mostly 11-15 long, light to violet blue, sparsely pubescent, the lobes 3-6 mm mm long. Anther thecae purple, ca. 0.6 long, on minute stalks separated mm by a swollen connective. Capsules 3-4 long, broadly ovoid, 4 valvate, the