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Leaf architecture of the genus Didymaea Hook. f. (Rubiaceae)

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Abstract

We examined 532 specimens representing eight species of Didymaea (Rubiaceae) to describe their leaf architecture. Characters observed in the herbarium specimens and cleared leaves were leaf base, apex, and margin; trichome type and distribution; vein orders from primary to higher order veins, veinlets, and ultimate marginal vein. Six morphometric variables were analyzed using variance and principal component analysis to identify which characters are useful to distinguish species. Most of the species present brochidodromous venation pattern, except D. hispidula, which has eucamptodromous venation pattern; Didymaea floribunda is unique in having cylindrical and bicellular trichomes. Principal component analysis revealed that 78% of the total variation is mainly related to leaf maximum width. Variance analysis showed significant differences between D. microflosculosa and the other species. Architectural features were useful for differentiating all the species, and an identification key for the species of Didymaea based only on leaf characters is presented.

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Authors and Affiliations

  1. División de Posgrado, Instituto de Ecología A. C., km 2.5 Antigua Carretera a Coatepec no. 351, Veracruz, 91070, Xalapa, Mexico

    Jaime Pacheco-Trejo

  2. Instituto de Biología, Universidad Nacional Autónoma de México, Apartado Postal 70-233, 04510, México, Mexico

    Teresa Terrazas & Helga Ochoterena

  3. Departamento de Botánica, Instituto de Biología, Universidad Nacional Autónoma de México, Circuito exterior s/n, Ciudad Universitaria,, Apartado Postal 70-233, Coyoacán, 04510, México Distrito Federal, México

    Teresa Terrazas

Authors
  1. Jaime Pacheco-Trejo
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  2. Teresa Terrazas
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  3. Helga Ochoterena
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Correspondence to Teresa Terrazas.

Appendices

Appendix 1

Didymaea specimens examined. Acronyms following Holmgren et al. (1990). Those used to prepare cleared leaves are marked with an asterisk (*).

Didymaea alsinoides

MEXICO: Hidalgo, R. Hernández & D. Rodríguez 5436 (MEXU).

Tamaulipas, A. García & R. Bracho 829 (MEXU*). Veracruz, F. Ventura 17753 (MEXU*, XAL); J. Becerra 311 (XAL); 445 (XAL); 471 (XAL); 637 (XAL); 657 (XAL); W. Marquéz 46 (MEXU); W. Marquéz, J. Dorantes & A. Calles 46 (MEXU*); W. Marquéz, J. Dorantes & A. Choles 46 W (MEXU) F. Vázquez 2272 (XAL).

Didymaea crassifolia

MEXICO: Oaxaca, D. Lorence, A. García & C. Allen 4702 (MEXU*).

Didymaea floribunda

MEXICO: Chiapas, D. E. Breedlove 51360 (MEXU); 5970 (DS, F); E. Cabrera & H. de Cabrera 6006 (MEXU); B.Y. López & F. Ramos 204 (MEXU); A. Méndez & M. C. Martínez. 9837 (CAS, MO); C. Santíz Ruíz 178 (CAS, NY); 946 (CAS). Distrito Federal; G. J Goodman 3454 (F); E. Lyonnet 1390 (US); 1600 (US); 1640 (US); 3035 (US); E. Matuda 18617 (MEXU*); J. Rzedowski 27340 (F, MO, NY); A. A. Ventura 788 (MEXU); 1021 (MEXU, MO); 2033 (MEXU, NY); 2231 (XAL); 2344 (MEXU, MO); 2514 (XAL); 2584 (MEXU); 2619 (MEXU, MO, NY); 2770 (MEXU); 2835 (MEXU, NY); 2893 (MEXU, NY); 3135 (XAL); 3510 (MEXU, XAL); s. coll., s. n. (MEXU). Estado de México, B. Beauchamp s. n. (MO); J. I. Calzada 22065 (MEXU); G. B. Hinton 1725 (NY); 4071 (F); 4076 (US); 4205 (F); S. D. Koch 7791 (MEXU); D. Lorence & D. Tejero 4886 (MEXU*); E. Matuda 26467 (MEXU); 27747 (MEXU); 28952 (MEXU); G. C. Pringle 9824 (F, MEXU, MO, NY, US); J. Rzedowski 28192 (DS); 36250 (MEXU*, XAL); R. Torres & F. Ornelas 10099 (MEXU); G. Webster 21172 (GH, MEXU). Guerrero, N. Herrera 894 (MEXU); E. M. Martínez & G. Silva 5752 (MEXU); J. S. Miller & G. Campos 2842 (MEXU, MO). Hidalgo, C G. Pringle 13016 (F, MEXU, US). Jalisco, D. E. Breedlove & F. Almeda 45638 (CAS); R. Cuevas 1299 (MEXU); C. Díaz 3424 (MEXU*); H. H. Iltis, R. Gúzman, J. Doebley & A. Lasseigne 494 (MEXU); J. A. Machuca 6204 (MEXU); 6347 (XAL); R. McVaugh 22036 (NY). Michoacán, G. Arsène 5502 (MO, US); 5507 (MO, US); 8479 (NY); 9039 (F, MO, US); E. Carranza & E. García 4178 (MEXU, XAL); H. Díaz 1207 (MEXU); J. M. Escobedo 1035 (MEXU); 1195 (MEXU); E. García 3147 (MEXU); E. García & E. Pérez 2764 (F, MEXU, XAL); G. B. Hinton 17167 (MEXU, XAL); H. H. Iltis & R. Koeppen 482 (K); H. H. Iltis, R. Koeppen & F. Iltis 420 (GH, NY, US); M. J. Jasso 628 (MEXU); 914 (MEXU) 1770 (MEXU); J. N. Labat 1367 (MEXU); Wm. C. Leavenworth 338 (F, NY); E. Martínez, J. C. Soto, G. Silva & T. P. Ramamoorthy 4674 (MEXU); E. Mayo 319 (MEXU, XAL); 608 (MEXU); M. E. Motte 383 (MEXU); E. Pérez 522 (MEXU*); 722 (XAL); M. Pérez 208 (MEXU); J. Rzedowski 39756 (MEXU, XAL); 44894 (MEXU, XAL); S. Zamudio 5451 (MEXU*). Morelos, F. A. Barkley, G. L. Webster & C. M. Rowell 7376 (MEXU, MO); O. M. Clark 7309 (MO, NY); I. Díaz & R. Noriega 1055 (MEXU); D. P. Gregory & G. Eiten 66 (GH, MEXU*, MO); G. B. Hinton 17151 (NY, MO); 17167 (MEXU, XAL); C. G. Pringle 13017 (CAS, F, MEXU*, MO). Oaxaca, A. Campos 4750 (MEXU); O. F. Clarke 18512-12 (F); J. Conrad & R. Conrad 3175 (MO); C. Conzatti 2300 (F, MEXU); W. R. Ernst 2537 (US); H. H. Iltis, R. Koeppen & F. Iltis 1238 (MO); D. Lorence 4746 (MEXU*); C. G. Pringle 4716 (CAS, MEXU, MO); E. Ramírez 833 (MEXU); E. Ramírez & P. Ramírez 466 (MEXU); J. Reyes 2685 (MEXU); J. Rivera 2102 (MEXU). Puebla, C. A. Purpus 1693 (F, MO, NY, US). Veracruz, S. Avendaño & J. I. Calzada 421 (F, XAL, MEXU); M. Cházaro & F. Texon 3217 (XAL); M. Cházaro & R. Acosta 3711 (XAL); R. Hernández 1362 (F, MEXU, XAL); D. Lorence & T. P. Ramamoorthy 4294 (MEXU, MO); J. L. Martínez & G. Castillo 1247 (XAL); J. L. Martínez & F. Vázquez 259 (XAL); 316 (MEXU*, XAL); 619 (MEXU, XAL); E. Matuda 1317 (F); F. Müller 197 (GH); 1519 (NY); M. Rosas 1462 (GH, MEXU); B. L. Turner & F. Barrie 15301 (XAL). Sine loc. E. Burgeau 98 (K); L. Liebmann L. 232 (US); 260-261 (MO); C. A. Purpus 202 (MO, US); 1693 (F, MO, US); Sessé, Mociño, Castillo & Maldonado 535 (F); 536 (F); Schaffner 177 (K).

Didymaea hispidula

COSTA RICA: Prov. Cártago; G. Davidse, G. C. Herrera & R. H. Wagner 26065 (MEXU, MO); R. Torres 123 (US). Prov. Heredia, W. H. Hatheway 1326 (US). Prov. Punta Arenas, G. C. Herrera & R. H. Wagner 26160 (MO); G. Herrera & J. Cordero 3637 (F, MO). Prov. San José, A. M. Evans, D. B. Lellinger & F. Bowers 122 (US); G. Davidse & M. H. Grayum 28789 (MO); W. H. Hateheway 1326 (US); P. C. Standley 34139 (US); P. C. Standley & J. Valerio 43386 (US).

GUATEMALA: Dpto. Chimaltenango, B. Palm 123 (F); F. Skutch 187 (US). Dpto. Chiquimula, J. A. Steyermark 31459 (F). Dpto. Guatemala, L. M. Andrews 473 (NY). Dpto. Huehuetenango, A. R. Molina 21181 (F, NY); A. Van Hook 2 (F); L. O. Williams, A. Molina & T. P. Williams 22140 (F); 22383 (F). Dpto. Jalapa, P. C. Standley 76912 (F); J. A. Steyermark 32428 (F). Dpto. Jutiapa, J. A. Steyermark 31927 (F). Dpto. Quetzaltenango, P. C. Standley 84187 (F); M. Véliz 7306 (MEXU*, MO); L. O Williams, A. Molina & T. P. Williams 22798 (F, NY); 25492 (F). Dpto. Quiché, A. Molina, A. R. Molina & J. A. Molina 30281 (F, MO); A. Molina & A. R. Molina 25058 (F). Dpto. Sacatepequéz, P. C. Standley 65188 (F). HONDURAS: Dpto. F. Morazán, A. R. Molina 1493 (F).

MEXICO: Chiapas, D. E. Breedlove 22972 (MO); 23016 (DS); 27702 (DS); 27772 (DS); 41330 (DS); 53769 (CAS, MO); D. E. Breedlove & F. Almeda 47794 (DS); M. C. Carlson 2419 (MEXU*); R. M. Laughlin 2333 (DS).

PANAMA: Prov. Chiriquí, T. B. Croat 13726 (GH, MO); T. S. Cochrane, A. B. Cochrane & R. R. Kowal 6290 (MO); W. G. D´Arcy 9996 (MO); B. A. Stein 1300 (MEXU*); K. Sytsma & W. D. Stevens 2087 (MO, NY); K. Sytsma & W. D. Stevens 2167 (MO); R. E. Woodson, P. H. Allen & R. J. Seibert 826 (F, MO, NY). Prov. Boca del Toro, S. Knapp 1617 (MEXU*).

Didymaea linearis

MEXICO: Jalisco, F. Boutin & M. Kimnach 3127 (MEXU*); Oaxaca, D. Lorence, A. García & C. Allen 4745 (F, MEXU*, MO); D. Lorence & R. Torres 4005 (MEXU*); G. Webster, G. Breckon & S. Lynch 17485 (MEXU).

Didymaea mexicana

MEXICO: Distrito Federal, E. Lyonnet. 526 (CAS, MO, NY, US); 1650 (US); C. R. Orcutt 3793 (F, K, MO); A. Panti-Madero 122 (MEXU*); J. E. Rivera-H. & A. Espinosa 3299 (MEXU); N. J. Rose & R. Hay 5328 (F, US); L. G. Ruíz 11 (F); A. Ventura 1685 (MEXU); 2255 (MEXU*, XAL); 2373 (MEXU, NY); 2989 (MEXU, NY); 3436 (MEXU, XAL); 3478 (MEXU, XAL). Estado de México, J. H. Beaman 2075 (GH, US); G. B. Hinton 1304 (NY); 3690 (NY, MO); E. J. Lott 284 (MEXU); S. D. Koch &. J. Magaña 75248 (CAS, F, MO); E. Matuda 19423 (F); 32462 (MEXU); J. Rzedowski 24186 (F); R. Vega 142 (CAS, MO); 251 (MEXU); E. Ventura 1450 (XAL); L. Villers 5 (MEXU*); J. N. Weaver 730 (US). Guerrero, J. Rzedowski 23648 (MEXU*). Hidalgo, F. R. Barrie & T. Atkinson 917 (MEXU); T. S. Cochrane 8475 (MEXU); R. G. García 648 (MEXU); R. Hernández 414 (MEXU); M. Medina 560 (MEXU, NY); 693 (MEXU); C. G. Pringle s. n. (F, MO); J. N. Rose, J. H. Painter & J. S. Rose 8701 (US). Jalisco, J. F. Santana, M. Wetter, D. De Niz & J. Cevallos 5208 (MEXU). Michoacán, S. Arias 846 (MEXU); E. Carranza 3912 (MEXU, XAL); M. G. Cornejo & G. Ibarra 236 (MEXU); M. G. Cornejo, J. Martínez, G. Ibarra & M. A. Salinas 785 (MEXU); 842 (MEXU); M. J. Jasso 900 (MEXU, XAL); 1778 (MEXU). Morelos, F. A. Barkley, G. L. Webster & C. M. Rowell 7385 (MEXU); H. H. Iltis, R. Koeppen & F. Iltis 242 (MO); E. Matuda 25585 (MEXU); G. Quiram 32 (MEXU). Oaxaca, W. R. Anderson & C. Anderson 4851 (GH, NY); D. Lorence, A. García & D. Frame 4593 (F, MEXU, MO, XAL); D. Lorence, A. García, G. Martin & R. Cedillo 3645 (MEXU); E. W. Nelson 1067 (US); C. R. Orcutt s. n. (MO); S. S. Tillet 637-110 (DS, US). Puebla, E. J. Alexander & E. Hernández 2261 (MEXU, NY); E. K. Balls & W. B. Gourlay B4479 (K, NY, US); N. Fröderström & E. Hulten 1259 (F); C. L. Gilly & A. J. Sharp 18 (CAS); J. N. Rose & R. Hay 6253 (NY, US); C. E. Smith, F. A. Peterson & N. Tejeda 3862 (F, US). Querétaro, H. Díaz-Barriga 6915 (MEXU). Tlaxcala, L. Bello 1779 (MEXU); Y. Ramos s. n. (F); A. Ruíz 308 (MEXU). Veracruz, E. K. Balls 4784 (K, US); C. P. Cowan, L. Dorr & L. Barnett 3900 (MEXU, NY); H. Galeotti 4400 (K); L. Liebmann 255 (MEXU, US); M. Martínez 8492 (MEXU); J. L. Martínez, R. Acevedo, G. Gómez 1112 (XAL); A. J. Sharp 45586 (MEXU); F. Ventura 18851 (MEXU, XAL); 3871 (MEXU, XAL). Sine loc. M. Martínez 8492 (MEXU); Sessé, Mociño, Castillo, Maldonado 534 (F); Sumichrast 472 (F).

Didymaea microflosculosa

EL SALVADOR: Dpto. Ahuachapan, M. Renderos & R. Villacorta 557 (MO); R. Villacorta & M. Renderos 2724 (MO). Dpto. Santa Ana, R. A. Carballo & J. Monterrosa 165 (MO); R. A. Carballo 483 (MO); J. L. Linares & C. A. Martínez 1429 (MO); J. Monterrosa & R. Carballo 254 (MO); R. Villacorta & M. Calderón 361 (MO); 550 (F, MO).

GUATEMALA, Dpto. Amatitlán, W. A. Kellerman 6660 (F). Dpto. Guatemala, P. C. Standley 58484 (F); 80546 (F). Dpto. Chimaltenango, F. Skutch 380 (F, NY, US); s. coll. 1330 (F). Dpto. El Progreso, J. A. Steyermark 43499 (F). Dpto. Quetzaltenango, J. A. Steyermark 33978 (F, US). Dpto. Quiché, Heyde 2892 (K, US). Dpto. Sololá, J. A. Steyermark 46874 (F, US); J. A. Steyermark 47230 (F).

HONDURAS: Dpto. F. Morazán, S. F. Glassman 1623 (F); 1958 (F); S. Lagos-Witte, C. Torres & P. R. House 32 (MO); J. L. Linares & F. J. Hubbard 3419 (MEXU*); I. Martínez 79 (MO, NY); A. Molina 204 (F, GH, MO); 1029 (F, GH, US); 1256 (F); 8596 (F); 10132 (F, US); 11052 (F); A. Molina & A. R. Molina 27946 (F); A. Molina, L. O. Williams, W. C. Burger & B. Wallenta 17019 (F, NY, US); P. C. Standley 13250 (F); 14053 (F); 20690 (F); P. C. Standley & L. O. Williams 677 (F); N. Vargas & F. A. Barkley 45055 (MO); L. O. Williams & A. Molina 11246 (F, NY); 13720 (F, GH); 13734 (F, MO, US); L. O. Williams, A. Molina & E. D. Merril 15624 (F). Dpto. El Paraíso, A. R. Molina 27487 (F); J.Valerio 1766 (F).

MEXICO: Chiapas, M. Chazaro, P. Hernández & G. Rodríguez 5354 (XAL); E. Martínez 20777 (MEXU*); E. Martínez & A. Reyes 20336 (MEXU).

Didymaea microphylla

GUATEMALA: Dpto. Chimaltenango, J. H. Beaman 4037 (GH, US). Dpto. Quetzaltenango, J. A. Steyermark 34188 (NY); 34888 (F); M. Véliz, N. Gallardo & M. Vásquez 9596 (MEXU). Dpto. Sacatepequéz, P. C. Standley 65250 (F); M. Véliz, N. Gallardo & M. Vásquez 8364 (MEXU); 8491 (MEXU*). Dpto. San Marcos, E. Martínez & M. Ramírez 20829 (MEXU*); 20842 (MEXU); M. Véliz, R. Luarca & M. Vásquez 10565 (MEXU).

MEXICO: Chiapas, D. E. Breedlove 40333 (MO).

Appendix 2

Leaf architecture comparison among Didymaea and potentially related genera within Rubioideae (number of species recognized/studied) RUB Rubieae, ANTH Anthospermeae, PAE Paederieae, THE Theligoneae.

Table 5
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Pacheco-Trejo, J., Terrazas, T. & Ochoterena, H. Leaf architecture of the genus Didymaea Hook. f. (Rubiaceae). Plant Syst Evol 281, 137–149 (2009). https://doi.org/10.1007/s00606-009-0195-3

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