Critical revision of the \u003ci\u003eDeppea\u003c/i\u003e complex (Rubiaceae, Hamelieae)

$Page 1: Critical revision of the \u003ci\u003eDeppea\u003c/i\u003e complex (Rubiaceae, Hamelieae)$
Acta Botanica Hungarica 46 (1–2), pp. 77–89, 2004

CRITICAL REVISION OF THE DEPPEA COMPLEX(RUBIACEAE, HAMELIEAE)

A. BORHIDI1, 3, 4, J. DARÓK2, M. KOCSIS2, SZ. STRANCZINGER1 and F. KAPOSVÁRI2

1Institute of Ecology and Botany, Hungarian Academy of Sciences, Vácrátót, Hungary2Biological Institute, Botanical Department, University of Pécs, Hungary

3Research Group of “Biological Adaptation”Hungarian Academy of Sciences–University of Pécs, Hungary

4Laboratory of Phanerogams, Science Faculty of the UNAM, México

(Received 11 March, 2004)

A monographic treatment was published about the genus Deppea Cham. et Schltdl. includ-ing the genus Edithea Standl. and several new species modifying the original generic con-cept and description creating a polyphyletic complex. The authors discuss the broad ge-neric concept of the treatment revalidating the genus Edithea Standl. describing the Editheafloribunda subsp. leuconeura subsp. nova, with the inclusion of Deppea miahuatlanica Lorenceas Edithea miahuatlanica (Lorence) Borhidi comb. nova. The authors suggest the exclusion ofDeppea panamensis Dwyer maintaining the description and delimitation of the entomophi-lous Deppea in its original, classic sense, and the separation of the obviously different orni-thophilous Omiltemia (Deppea) scoti Kirkbr. and Deppea splendens Breedlove and Lorenceinto two monotypic genera: Bellizinca scoti and Csapodya splendens.

Key words: Bellizinca, complex, Csapodya, Deppea, Edithea, Hamelieae, México, polyphyletic

INTRODUCTION

In the last decades three critical taxonomic treatments has been publishedwithin the framework of the Flora Mesoamerica Project, related to the tribeHamelieae: Hamelia (Elias 1976), Omiltemia (Kirkbride 1984b) and Deppea (Lo-rence and Dwyer 1988). While the treatment of Hamelia seems to be a very de-tailed, thorough study based on a concise, correctly shaped generic concept,the last two treatments rise several problematic – partly not well documented,partly unilaterally evaluated – conclusions, which need of a re-examinationand re-discussion of the whole material.

THE DEPPEA COMPLEX

The genus Deppea was described by Chamisso and Schlechtendal in theLinnaea (5: 167. 1830) based on the species Deppea erythrorhiza Cham. et Schltdl.from the state Veracruz of México. To the end of the eighties the number of the

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detected and described species has elevated to more than 20, ranging betweenMexico and Panama, but the overwhelming majority is Mexican endemics.Even the monographers, Lorence and Dwyer added 5 new species (D. marti-nez-calderoni, D. purpurascens, D. hernandezii, D. oaxacana, D. guerrerensis), andrecently the first author of this study further two ones (D. sousae, D. longifolia)(Borhidi et al. 2003, 2004). All these species may be characterised by a well de-fined basic ground plan of the corolla shape and size and the structure and po-sition of the androeceum, having a rotate or short funnelform corolla withshort tube up to 1 cm long and longer extended corolla lobes, open throat, cy-lindrical filaments, inserted above the middle of the tube or near the throat(not near the base as it is stated by Lorence and Dwyer in the generic diag-nose!) and exerted anthers. The fruit is a capsule with strong several longitudi-nal and one circular bundles permitting only an apical-loculicidal dehiscence.

The authors of the monograph admit that “During the course of the revi-sion for Flora Mesoamericana it was found that about 25 names for Deppea ex-ist in the literature, several of which represent taxa clearly not belonging to thegenus” (1988: 389). Therefore the authors excluded 7 names. Instead of themincluded 4 or 5 other species having different ground model of flower with dif-ferent characters than listed above and cannot be maintained within the frameof Deppea. According to my view the Deppea sensu Lorence and Dwyer (1988)is a polyphyletic complex of species belonging to 3 or 4 different genera. TheDeppea complex of Lorence and Dwyer includes several alien taxa, like: Deppeapanamensis, D. schultzei, D. miahuatlanica (described later by Lorence 1997), D.scoti and D. splendens. The inclusion of the Brazilian Schenckia blumenaviensisinto Deppea is also questionable.

The main argument of Lorence and Dwyer for doing so is the similarstructure and dehiscence of the fruit. As they pointed out: “We believe thatmode of capsule dehiscence provides one of the most important characters forseparating the genera in this alliance. It is stressed that capsule dehiscence re-fers to its condition at the time of seed release, after which the capsule begins todeteriorate and may split along additional sutures” (1988: 391). Lorence andDwyer invented the “fruit-taxonomy” based fundamentally on fruit morphol-ogy and dehiscence, neglecting or underestimating the floral characters.

I cannot agree with this kind of taxonomy, because fruit is a dispersal or-gan which cannot be responsible for the genetic processes controlled basicallyby the morphology and position of the sexual organs (corolla, androeceum,pistil, ovarian disc, etc.) by selecting the pollinating agents being mostly differ-ent from the distributors. Fruits are much less diverse than flowers, thus can-not be correlated so closely with the extremely high genetic diversity. Themost sophisticated “dehiscentological dissertation” cannot explain the geneticand taxonomic relation between two taxa without considering floral charac-

78 BORHIDI, A., DARÓK, J., KOCSIS, M., STRANCZINGER, SZ. and KAPOSVÁRI, F.

Acta Bot. Hung. 46, 2004

ters. I consider very improbably that the Deppea species of small entomo-philous flowers could have the slightest genetic relation to the colourful big-flowered, hummingbird-pollinated Edithea, Deppea scoti and D. splendens.

Table 1 gives the main structural differences between the members of theDeppea complex.

Based on the characters listed above the following key is proposed for thetaxa of the Omiltemia and Deppea complexes:

1a Corolla rotate or short funnelform, tube very short, mostly shorter thanthe lobes Deppea

1b Corolla tubular or salverform, tube elongate, several times longer thanthe lobes 2

2a Calyx lobes linear or filiform, mostly deciduous, fruit elongate, cylind-rical, apically then septicidally dehiscent 3

CRITICAL REVISION OF THE DEPPEA COMPLEX 79


Table 1Main floral characters in the Deppea s. l.

Characters Deppea s. str. Edithea D. scoti D. splendensNumber of species 21 2 1 1Calyx lobes 0.2–7 mm 0.8–2 mm 8–18 mm 15–25 mm

green green green redCorolla shape rotate or short

funnelformsalverform tubular tubular to

salverformCorolla tube length 0.4–8 mm 10–35 mm 14–26 mm 50–55 mm

lobes length 2.5–14 mm 1.5–4.5 mm 9–15 mm 8–14 mmtube/lobes 0.1–1.2 8–15 1.8–2.5 4–7

Filaments shape cylindrical flattened cylindrical flattenedlength 0.5–6 mm 2.5–6 mm 15–22 mm 15–22 mm

Filaments insertion in or nearthe throat

below thethroat

in the baseof the tube

in the middleof the tube

Anthers exerted inserted inserted insertedStyle glabrous glabrous glabrous villousOvary disc ring dome-shaped bilobate 4-lobateOvary inside hairy hairy glabrous glabrousSeed oblong lenticular lenticular triangular to

moniliformInsertion subcentral central central lateralPerpendicular wall transversally

thickenedobtuse, pitted obtusely

undulatepitted

Horizontal wall obtuselypitted

densely pittedand lineolate

reticulate reticulate

2b Calyx lobes not linear or filiform, persistent; fruit globose or obovate,apically or apically then loculicidally dehiscent 4

3a Corolla tube asymmetrical, glabrous, filaments elongate Omiltemia

3b Corolla tube symmetrical, hirsute, anthers subsessile Pseudomiltemia

4a Calyx lobes triangular or lanceolate to 2 mm long, anthers subsessile inthe throat of the corolla tube Edithea

4b Calyx lobes foliaceous, 8–25 mm long, filaments 1–2 cm long 5

5a Inflorescence erect; calyx lobes green, very unequal in pairs, corolla tubularfilaments cylindrical, born at the base of the corolla Bellizinca

6b Inflorescence scandent, calyx lobes red, subequal, corolla salverform, fi-laments flattened, born above the middle of the corolla tube Csapodya

Bellizinca Borhidi, genus novum, hoc loco

Frutex vel arbor parva, 2–4 m alta, ramuli teretes vel compressi et 2-sulcati, pilisfulvis hirsuto-villosi vel glabrescentes, stipulae persistentes, deltoideo-acutae, apiceglandulosae, dorso hirsutae, margine collateris 4–8 digitalibus suffultae. Folia opposi-ta, petiolata, inaequalia, petiolis dense fulvo-hirsutis vel villosis, ovata vel ovato-elliptica, basi obtusa vel rotundata saepe valde obliqua, apice acuminata et falcata,chartacea, leviter discolora, viridia et utrinque hirsuto-villosa; nervis lateralibus bro-chidodromeis, venatione conspicua saepe rubella, margine tenui, ciliata. Inflorescen-tiae monochasiae, simplices, scorpioideae, terminales vel pseudo-axillares, pauciflorae,pedunculo hirsuto-villoso, flores subsessiles vel breviter pedicellati, hypanthium obco-nicum vel turbinatum sulcatum et hirsuto-villosum; calycis lobi 4, foliacei, forma mag-nitudineque valde inaequales, venosi in sicco saepe rubelli utrinque strigilloso-hirsuti,margine ciliati; corolla flava, tubulosa utrinque glabra vel extus sparse pilosa, in ala-bastro 1.5–2 cm, sub anthesi 2–4 cm longa, tubus corollae lobulis 4, ovatis, erectis etacuminatis in alabastro contortis duplo longior, stamina 4, inclusa, antherae longae,dorsifixae, apice obtusae, basi 2-lobatae; filamenta larga cilindrica, basi corollae in ani-llum incrassatum adnata; stylus glaber, stigma 2-lobatum, exsertum; discus glaber.Capsula turbinata, obovato-oblonga, leviter 2-sulcata et 6–8-costata, basi acuta, lobiscalycis persistentibus coronata, apice loculicide et posterior septicide dehiscens apertu-ra apical rhombea formans. Semina rhombea, testa rubro-brunnea, reticulata.

Genus endemicum mexicanum.Nomen genericum ex adiectivo italiano bello -a (i.e. pulcher, -a, -um) et ex

nomine folclorico hungarico Lysimachiae (e familia Gentianaceae) flore flavosimillima compositum.



Species typica: Bellizinca scoti (J. H. Kirkbr.) Borhidi comb. nov., hoc loco. – Bas.:Omiltemia scoti J. H. Kirkbr., Syst. Bot. 9: 411. (1984). Tipo: México: Puebla,above Teotitlán del Camino on road to Huautla de Jiménez, 2,000–3,250 m, 3 Aug.1961, C. E. Smith jr., F. A. Peterson and N. Tejeda 4137; holotipo: US n.v., iso-tipo: GH n.v. – Syn.: Deppea scoti (J. H. Kirkbr.) Lorence, Allertonia 4: 398. (1988).

Shrubs or treelets 2–4 m tall,twigs terete or compressed and 2-sul-cate hirtellous-villosulous with fulvoushairs or glabrate; stipules persistent,acutely deltoid, glandular at apex,externally densely hirtellous, themargin with 4–8 digitate colleters;Leaves opposite, petiolate, subequalor unequal at a node, petioles denselyfulvous hirtellous or villosulous, la-mina ovate or ovate-elliptic, slightlyfalcate, obtuse or rounded at thebase, the apex acuminate, secondaryveins 7–12 pairs, brochidodromous,venation visible to tertiary adaxiallyand to quaternary abaxially, costa andveins often reddish. Inflorescenceerect, terminal or pseudoaxillary, anunbranched scorpioid monochasium,2–10-flowered, pedunculate, the pe-duncle hirtellous-villosulous; hypan-thium obconical or turbinate, ribbed,the cup shallow, the lobes 4, folia-ceous, venose, strongly unequal, thelarger pair broadly ovate, apicallyacuminate, the smaller pair lanceo-late or narrowly elliptic, both sur-faces strigillose-hirtellous, each sinuswith a conical gland; corolla yellow,drying yellow, glabrous on both sur-faces, tubular or salverform at an-thesis, 2.4–4 cm long, tube about 2times as long as the 4 ovate lobes, sta-mens included, anthers narrowlyovoid, apically obtuse, basally bi-



1

2

3

Figs 1–3. Bellizinca scoti, seed, 1 = ×190, 2 =×700, 3 = ×3,000

lobed, the filaments cylindrical, 1.5–2.2 cm long, thickened and connate at thebase into a ring attached very near to the base of the tube. Style glabrous,stigma bilobed, exserted; the disc bilobate. Capsules turbinate, slightlybisulcate, weakly 6–8-costate, basally acute. Seeds rounded, 0.3–0.4 mm longand wide with central insertion, perpendicular walls obtusely undulate, hori-zontal ones densely and irregularly reticulate (Figs 1–3).

Monotypic Mexican genus.

Specimens studied: OAXACA. Municipio Teotitlan: 20 km NE of Teotitlan del caminoon road to Huautla de Jiménez, D. E. Breedlove et F. Almeda 59924 (MEXU). – 26.1 km W ofTeotitlan del camino, B. Bartholomew et al. 3177 (MEXU). – 3 km al E de El Puerto, carreteraa Huautla, P. Tenorio 5862 (MEXU). – Puerto de la Soledad, 30 km al NE de Teotitlan, P. Te-norio and C. Romero de T. 12622 (MEXU). – Teotitlan, Puerto de la Soledad, 30 km al NE dela carretera a Huautla, P. Tenorio 14391 (MEXU).

Csapodya Borhidi, genus novum, hoc loco

Frutex vel arbor parva usque ad 8 m alta, ramuli glabri, lenticellati, longitudina-liter striati; stipulae crassae, brunneae, deltoideo-subulatae, extus hirsutae, intus vi-llosae, collateris marginalibus 6–8 suffultae, deciduae. Folia ternata vel opposita sub-aequalia vel inaequalia, petiolata; lamina elliptica vel ovato-elliptica, basi cuneata velattenuata, apice acuta et acuminata saepe leviter falcata, membranacea vel subcharta-cea, sparse pilosa demum glabra, nervis lateralibus utroque latere 7–10, brochidodro-meis, subtus villosulis, nervo medio prominente, flavescente vel rubello. Inflorescentiaterminalis, pendula, dichasialis, corymbosa, usque ad 21 cm longa et 14 cm lata,16–25-flora, pedunculis 3.5–8.5 cm longis, rachidis glabris bracteis linear-subulatis,ciliatis, usque ad 8 mm longis suffulta; flores 1–3 cm longe pedicellati, hypanthiumturbinatum, glabrum, basi obtusum vel rotundatum, lobi calycis 4, aequales vel subae-quales, erecti, foliacei, venosi, acuti, in sicco rubelli, margine ciliati, 15–25 mm longi et3–7 mm lati, in sinubus glandula minutissima praediti. Corolla hippocrateriforme,flava vel orangeata, in alabastro 5–6 cm longa, utrinque glabra, in anthesi tubus 5–5.5cm longus, in 2/3 superiori 5–7 mm dilatatus, lobi 4, 0.8–1.4 cm longi et 3–4 mm lati,in alabastro contorti, sub anthesi rectangulariter expansi vel reflexi, acuti vel acumi-nati; stamina glabra, antherae lineares, 7–8 mm longae, apice obtusae, basi sagittatae,fauce insertae vel usque dimidium exsertae, filamenta 1.5–2 cm longa, aplanata, tubosupra medio affixa; stylus 5–6.5 cm longus, staminis ± aequilongus, basi dense villo-sus, stigma ovoideum. Ovarium 2-loculare ovulis numerosis, ovarii discus cónicus,4-lobatus; capsula subglobosa, 6–8 mm longa et 5–7 mm lata, 2-sulcata, leviter6-costata, dehiscentia apicali-loculicida supra anillum calycinum lobulis persistenti-bus, rubris; pedicellum fructifrum recurvatum. Semina angulosa vel moniliformis,testa brunnea, foveolato-reticulata.



Genus monotypicum Mexicanum.Nomen genericum in honorem Verae Csapody botanicae Hungaricae

studiosissimae, in plantis germinantibus expertissimae atque in illustrationeflorae Europae Mediae et Meridionali-Orientalis excellentissimae, atque in ho-norem István Csapody botanici Hungarici et ingenieri forestalis illustrissimiin protectione naturae laboriosissimi et in humanis splendidissimi dedicavi.

Typus generis: Csapodya splendens (Breedlove et Lorence) Borhidi, comb.nov., hoc loco. – Bas.: Deppea splendens Breedlove et Lorence, Phytologia 63: 43.fig. 1. (1987). Tipo: México: Chiapas, Mun. de Motozintla de Mendoza SW ofCerro Mozotal, 11 km NW of junction of road to Siltepec, alt. 2,100 m, 18 Sep1976, D. E. Breedlove 40258; holotipo: CAS n.v., isotipos: MEXU!, MO!.

Shrubs or small trees 5–8 m tall twigs glabrous, minutely lenticellate, lon-gitudinally wrinkled, nodes crowded, annular; stipules thick, brown, deltoid-subulate, deciduous, externally sparsely hirsutulous, internally villosulouswith 6–8 digitate marginal colleters. Leaves ternate or opposite subequal or



4

6

5

7

Figs 4–7. Csapodya splendens, seed, 4 = ×120, 5 = ×370, 6 = ×550, 7 = ×1,200

unequal, petiolate; lamina elliptic to ovate-elliptic, the apex acuminate, slightlyfalcate, the base narrowly cuneate to attenuate, the secondary veins 7–10 paresfestooned brochidodromous, veins and costa abaxially prominent, yellowishto purplish, the lamina membranaceous to thinly chartaceous, vein axilsslightly barbate abaxially. Inflorescence terminal, pendulous, cymose-corym-biform, 16–25-flowered, up to 21 cm long and 14 cm wide, pedunculate, theaxes glabrate subtended by linear-subulate bracteoles to 8 mm long. Flowerspedicellate, hypanthium glabrous, turbinate drying reddish, calyx cup 0.5 mmdeep, lobes 4, subequal, erect, foliaceous, venose, drying reddish, narrowlyovate-elliptic to lanceolate, 1.5–2.5 cm long, acute; corolla yellow to orange,salverform, glabrous on both surfaces, tube 5–5.5 cm long, flared to 5–7 mmwide in the distal 2/3, lobes 4, much shorter than the tube, contorted in bud,spreading at anthesis or recurved, ovate-deltoid, acute to acuminate; stamens4, glabrous, the filaments flattened, 1.5–2 cm long, affixed 1.5–2 cm below thefaux above the middle of the tube; anthers linear sagittate at the base, the apexobtuse, inserted in the throat; style 5–6.5 cm long, equalling or slightly exceed-ing the stamens, densely villous in the basal half, stigma ovoid; ovary 2-locu-lar, placentas bar-like, longitudinally inserted, ovules numerous per locule,ovary disc dome-shaped, 4-lobate; capsule subglobose, compressed perpen-dicular to the septum, 6–8 mm long and 5–7 mm wide, weekly 6-costate, withapical-loculicidal dehiscence above the calyx-ring, lobes persistent in fruit;fruiting pedicels recurved; seeds numerous, triangular to moniliform, later-ally inserted in the middle, perpendicular walls simple, pitted, horizontalwalls densely reticulate (Figs 4–7).

Monotypic Mexican genus.

Specimens studied: OAXACA. Municipio Motozintla de Mendoza: SW side of CerroMozotal, 11 km NW of the junction of the road to Motozintla, along the road to El Porvenirand Siltepec, D. E. Breedlove 25705 (MEXU), 25758 (MEXU), D. E. Breedlove and R. F. Thor-ne 31119 (MEXU), D. E. Breedlove and B. Bartholomew 55758 (MEXU).

For giving a reason to include this species into the genus Deppea, Lorenceand Dwyer wrote the following: “In spite of its extreme floral morphology,Deppea splendens corresponds with the genus Deppea in all essential characters,notwithstanding the usually whorled leaves a feature also occurring in D.hernandezii and occasionally in D. erythrorrhiza and D. hintonii. The pendulousinflorescence, large, red calyx lobes and long orange or yellow salverform co-rollas with partly included stamens obviously represent adaptations to hum-mingbird pollination, in contrast to the majority (if not all pers. remark) ofDeppea species with small erect or inclined flowers having yellow or white co-rollas with a short tube and exserted stamens characteristic of entomophily”. Itis obvious that Lorence and Dwyer saw aright the differences between this



species and the other Deppeas, but – in our view – they did not judge correctlywhat are the essential characters determining the genetic flow responsible fortaxonomic relations.

Deppea Cham. et Schltdl. 1830

Slender shrubs or subshrubs rarely treelets up to 7 m tall, sparsely ordensely pubescent, trichomes simple, septate, uniseriate; raphides present inall parts, cut wood rarely turning pink or reddish. Leaves opposite rarely inwhorles of 3, thin, petiolate, those of a pair often unequal; stipules small inter-petiolar, triangular, persistent, the margin and inner surface often with glan-dular colleters. Inflorescence axillary or terminal, cymose, mono- or dichasial,corymbiform or thyrsiform, or rarely 1-flowered. Flowers 4-merous, pedicel-late, hypanthium, turbinate obconic or cylindrical, the persistent calyx lobesminute or sometimes foliaceous, green, equal or strongly unequal, each sinususually with a small gland; corolla yellow or rarely purple or white, rotate orshortly funnelform, tube mostly shorter than the lobes, aestivation sinistror-sely contorted in bud, the lobes spreading at anthesis; stamens 4, the filamentscylindrical, inserted below the open throat or above the middle of the tube, an-thers exserted, dorsifixed at the base, narrowly ovoid, or narrowly ellipsoid,tetrasporangiate; ovary bilocular, placenta elongate, vertical, peltately affixedto the septum above the middle; ovules numerous; style slender glabrous,stigma entire or bilobed, disc entire or shallowly annular. Fruit a small drycapsule, obovate, obconic or cylindric, turbinate, often bisulcate, 8-costate, ca-lyx lobes persistent apical-loculicidally dehiscent, with longitudinal emergentvascular bundles united into a circular ring at the calyx, enclosing the scleren-chymatous endocarp; seeds numerous, oblong or oblong-angulate with sub-central insertion, perpendicular walls with transversal thickenings, horizontalwalls obtusely pitted.

Mexican and Central American genus with over 20 species, the majorityin Mexico.

Deppea longifolia Borhidi, spec. nov., hoc loco

Tipo: México: Oaxaca, Mpio: Comaltepec; SW slope of Cerro Relámpago just aboveRío Soyolapan, near Federal Electricity Commission Camp, down trail that descends fromHwy 175; 17°29’15”N, 96°24’05”W, 1,750–1,780 m, 3 Dec. 1993, B. Boyle, D. Massart, E. Mo-rales 2652; holotipo: MEXU, isotipo: MO.



Frutex usque ad 2 m alta, ramuli cylindracei, glabri, in sicco nigri; stipulaeoblongo-lanceolatae vel triangulares, acutae, ferrugineo-pubescentes; petioli valdeinaequales, unus altro 3-plo longiores, 0.7–3 cm longi; folia oblongo-lanceolata veloblongo-elliptica, 6–15 cm longa et 1.5–3.5 cm lata, utroque extremo longe angustata,apice acuta et apiculata, basi longissime cuneata, membranacea, supra pilis minutissi-mis albo-punctata subtus ad nervos pilosa, nervi laterales utroque latere 13–17, nervisintercalaribus usque ad 25 pares, quasi rectangulariter abeuntes, deinde valde arcuatiet ascendentes utrinque obsoleti; inflorescentiae axillares, cymoso-corymbosae, laxissi-me multiflorae, pedunculatae, pedunculis 4–5 cm longis, in sicco nigrescentibus, uni-lateraliter ferrugineo-pilosis, ramulis inflorescentiae valde patentibus, panicula ple-rumque longitudine latior, ramificationibus ebracteatis vel bracteis minutissimis etcaducis; Hypanthium obovatum, glabrum, 0.8 mm longum, lobi calycini 4, late ovati,rotundati, 0.2–0.3 mm longi et lati, glabri et margine ferrugineo-pulverulenti; corollabreviter infundibuliformis, 4.5–6 mm longa, in alabastro obtusa, tubus corollinus te-nuis, 2 mm longus et 1 mm latus, lobi 4, tubo 1.5–2-plo longiores, ovati, apice rotunda-ti, 3–4 mm longi et 2–3 mm lati, venulosi, glabri; stamina 4, fauce inserta, filamentacompressa, 0.5 mm longa, antherae lineares, exsertae, 2–2.5 mm longae, dorsifixaesubbasales, appendicibus rotundatis basalibus 0.2 mm longis suffultae; stylus 4–5 mmlongus, superne dilatatus et pulverulentus, stigma integrum, globosum. Fructus nonvisus.

Next species Deppea grandiflora, from that it differs by having stems andinflorescences drying blackish, elongate leaves with secondary nerves muchmore numerous and obsolete, and flowers half as large as in D. grandiflora.

Edithea Standl. 1933

Shrubs to 2.5 m tall, the twigs subterete, pubescent when young, glabrate;stipules deciduous, deltoid, acuminate, externally villosulous, the margin andapex with 2–3 pairs of digitate colleters. Leaves opposite, subequal, petiolate.Inflorescence terminal, dichasial, corymbiform, pedunculate, flowers numer-ous pedicellate, ebracteolate, 4-merous; hypanthium turbinate; calyx 4-parted,lobes erect, equal or subequal, acutely deltoid triangular or subulate; corollasalverform, apiculate in bud, the tube narrow, several times longer than thelobes, slightly widened distally, lobes erect or slightly outcurved, or rectangu-larly extended; stamens 4, inserted in the throat, the filaments very short, flat-tened, glabrous, attached below the throat, anthers subsessile linear or ellip-soid, apically acute, basally bilobed, dorsifixed at the middle, style included,stigma swollen, bilobate, the disc dome-shaped. Ovary 2-locular, ovules nu-merous in each cell, imbricate, placenta peltately attached to the septum. Cap-



sule turbinate obovate or obconic, acute at the base, strigillose, bisulcate,8-costate. Seeds rounded or elliptic-angulate with subcentral insertion, per-pendicular walls obtuse, pitted, horizontal walls densely pitted and lineolate(Figs 8–10).

Endemic genus of Mexico with 2 species in Guerrero and Oaxaca.

1a Inflorescence 30–180-flowered; corolla finely hirsutulo-sericeous externally,tube 4–6 times longer than the lobes; calyx lobes triangular-subulate, 1–2mm long E. floribunda

1aa Flowers orange or yellow; petioles subequal, lateral nerves 10–13 pairs,slender, rufous villosulous, calyx lobes linear-lanceolate, slightly unequal

subsp. floribunda

1ab Flowers red; petioles unequal, lateral nerves thick, 8–11 pairs, white, whitehairy or glabrous, calyx lobes triangular, equal subsp. leuconeura

1b Inflorescence 10–20-flowered; corolla glabrous externally, tube 8–10 timeslonger than the lobes; calyx lobes triangular, up to 1 mm long

E. miahuatlanica

Edithea floribunda Standl., Trop. Woods 34: 58. 1933

Tipo: México: Guerrero, Malinaltepec, Mar 1930, L. Schultze 363; holotipo: B destrui-do; fragmento: F n.v., foto: MEXU!). – Syn.: Omiltemia floribunda (Standl.) Kirkbr., Syst. Bot.9: 413. (1984). – Deppea schultzei Lorence, Allertonia 4: 422. (1988).

subsp. floribunda

Petioles of a pair at a node subequal, leaves densely rufous villosulous oncosta and secondary veins, these 10–13 pairs, calyx lobes linear-lanceolate,slightly unequal, corolla orange or yellow, fruit more elongate. Distr.: Mexico:Guerrero, endemic.

subsp. leuconeura Borhidi, subsp. nov., hoc loco

A typo petiolis inaequalibus, foliis ad nervum medium albo-hirsutis vel glabrisnervis lateralibus utroque latere 8–11 magis distantibus, albis et crassis valdeprominentibus, lobis calycinis triangularibus aequalibusque, corolla rubra, fructusnervis vascularibus crassioribus differt.



Petioles of a pair at a node unequal, leaves white hirsute or glabrate on thecosta and secondary veins these 8–11 pairs more distant, white, thick, stronglyprominent, calyx lobes equal, triangular, corolla red, fruit con thicker vascularbundles.

Distr.: Mexico: Oaxaca, endemic.

Holotypus: México, Distr. SantiagoJuxtlahuaca, Mpio. Santiago Juxtlahuaca.Loc. Poblado El Manzanal, a 15 km de ladesviación de Santa Rosa, carr. a San Mi-guel Cuevas, Coord.: 17°13’N, 98°03’W,alt.: 2,325 m, 18 Febr. 1995. J. I. Calzada19772 (MEXU).

Edithea miahuatlanica(Lorence) Borhidi, comb. nov.,

hoc loco

Bas.: Deppea miahuatlanica Lorence,Novon 7: 47. (1997). Tipo: México: Oaxaca:Distr. de Miahuatlán, Mpio. De San Jeróni-mo Coatlán, 17.9 km al SW de San JerónimoCoatlán, carretera Miahuatlán-Playa Larga,1,800 m, 16°12’N, 96°57’W, 13 Dec. 1987, R.Torres C. y A Campos 10847; holotipo:PTBG n.v., isotipo: MEXU!).

*

Acknowledgements – The authors ex-press their thanks to the director and cura-tors of the MEXU, ENCB and FCME for thepermission to study the materials of theircollections, to Nelly Diego-Pérez project-leader of the Proyecto de la Flora del EstadoGuerrero for organising and helping thework in the Taxonomic Laboratory of theFacultad de Ciencias de la UNAM. Thestudy has been supported by theCONACyT, the Hungarian Scientific Re-search Fund (OTKA T032036) and by theInstitute of Ecology and Botany of the Hun-garian Academy of Sciences.



8

10

9

Figs 8–10. Edithea floribunda, seed, 8 = ×140,9 = ×500, 10 = ×4,300

REFERENCES

Borhidi, A., Darók, J., Kocsis, M. and Stranczinger, Sz. (2004): Critical revision of theOmiltemia complex (Rubiaceae, Hamelieae). – Acta Bot. Hung. 46(1–2): 69–76.

Borhidi, A. and Barreto Valdés, A. (2002): Introducción a la taxonomía de la família Rubia-ceae en la flora de México. – Acta Bot. Hung. 44: 237–280.

Borhidi, A., Martinez Cordillo, M. and Cruz Darién, R. (2003): Deppea sousae sp. n. (Rubia-ceae) en la flora del estado Guerrero (México). – Acta Bot. Hung. 45: 31–35.

Breedlove, D. E. and Lorence, D. H. (1987): New species of Deppea (Rubiaceae) from Chia-pas, Mexico. – Phytologia 63(1): 43.

Kirkbride, J. H. jr. (1984a): Manipulus Rubiacearum III. Deppeeae, a new tribe of Rubioi-deae (Rubiaceae). – Brittonia 36: 317–320.

Kirkbride, J. H. jr. (1984b): Review of Omiltemia (Rubiaceae). – Syst. Bot. 9(4): 410–414.Lorence, D. H. (1997): New taxa, a new name, and a new combination in Rubiaceae from

Southern Mexico and Mesoamerica. – Novon 7: 46–58.Lorence, D. H. and Dwyer, J. D. (1988): A revision of Deppea (Rubiaceae) – Allertonia 4(7):

389–434.Rova, J. H. E., Delprete, P. G., Andersson, L. and Albert, V. A. (2002): A trnLF cpDNA se-

quence study of the Condamineeae-Rondeletieae-Sipaneeae complex with implica-tions of the phylogeny of the Rubiaceae. – Amer. J. Bot. 89(1): 145–159.

Standley, P. C. (1918): Omiltemia longipes Standl. – J. Wash. Acad. Sci. 8: 427.Standley, P. C. (1918–1934): Rubiaceae. – North American Flora 32: 1–300.Standley, P. C. (1933): Edithea, a new genus of Mexican shrubs of the family Rubiaceae. –

Trop. Woods 34: 1–3.Standley, P. C. (1940): Kohleria filisepala Standl. – Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 107.



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Critical revision of the \u003ci\u003eDeppea\u003c/i\u003e complex (Rubiaceae, Hamelieae)