taxonID	type	description	language	source
56153F74FFF75974FF7FFE6C05E985C3.taxon	description	Trees to 8 m tall, mostly epipetric. Older (woody) stems subquadrate to terete, irregularly fissured, sometimes ± scurfy, often lenticellate, lacking trichomes; younger (herbaceous) stems terete (when fresh), becoming flattened on drying, sparsely lenticellate, irregularly fissured, lacking trichomes. Leaves seasonally deciduous, clustered at shoot apex, petiolate, petioles to 90 mm long, sometimes tinged with pink or red, blades subsucculent, narrowly ovate to ovate to elliptic, 85 – 157 mm long, 31 – 76 mm wide, 1.8 – 2.7 × longer than wide, acute to acuminate at apex, acute to attenuate at base, surfaces glabrous (or rarely with very few scattered flexuose eglandular trichomes mostly on veins on abaxial surface, the trichomes especially evident on immature leaves), midvein sometimes pinkish to reddish purple (especially adaxially), margin entire. Inflorescence a terminal sessile or pedunculate thyrse to 38 cm long (including peduncle, if present, and excluding corollas), peduncle (if present) pubescent with erect to flexuose glandular trichomes 0.05 – 0.7 mm long (glandular pubescent), rachis glandular pubescent and often with scattered erect to flexuose eglandular trichomes to 0.7 mm long as well; dichasia opposite or alternate, pedunculate, 1 – many-flowered, to 120 mm long (excluding corollas), dichasial peduncles 15 – 60 mm long, pubescent like rachises. Bracts caducous (rarely present), lanceolate to lance-ovate, 6.5 – 7. 5 mm long, 2.5 – 2.8 mm wide, abaxially pubescent like rachis (except sometimes with more eglandular trichomes). Bracteoles and secondary bracteoles caducous, lance-subulate to linear to ovate-elliptic, 2 – 7 mm long, 0.9 – 2 mm wide, abaxially pubescent like rachis or with more eglandular trichomes. Flowers pedicellate, pedicels 25 – 50 mm long, glandular pubescent. Calyx 10 – 16 mm long, lobes fused at base for 1.5 – 3 mm, subhomomorphic to subheteromorphic, membranaceous, lanceolate to ovate, 7. 5 – 12 mm long, 3 – 7 mm wide, subequal or with posterior lobe either slightly larger or smaller than lateral lobes, acute at apex, abaxially pubescent with erect to flexuose glandular and eglandular trichomes 0.05 – 0.3 mm long, posterior lobe subplanar to subconduplicate. Corollas greenish, 35 – 65 mm long, externally pubescent with erect to flexuose glandular and eglandular trichomes 0.05 – 0.3 mm long, tube 25 – 30 mm long, narrow proximal portion 7 – 12 mm long, 7 – 11 mm in diameter near midpoint, throat 15 – 25 mm long, 26 – 30 mm in diameter at mouth, lobes spreading to recoiled, ovate to subcircular, 15 – 24 mm long, 16 – 24 mm wide, rounded and emarginate at apex. Stamens 4, 65 – 80 mm long, filaments distally glabrous, pubescent with eglandular trichomes near base, thecae 8 – 11 mm long; staminode 0. Style 75 – 77 mm long, glabrous distally, pubescent near base with eglandular trichomes, stigma subequally 2 - lobed and ± funnelform, lobes elliptic to subspheric, 2 – 3 mm long, 1.5 – 3 mm wide. Capsules 32 – 42 mm long, 6 – 9. 5 mm in diameter, pubescent with erect to flexuose eglandular and subsessile to stipitate glandular trichomes less than 0.05 – 0.4 mm long, stipe 2 – 5. 2 mm long. Seeds 16 – 24 per capsule, 5. 5 – 7 mm long, 4. 5 – 6 mm wide, surfaces smooth. PHENOLOGY. — Flowering: March; fruiting: March. Field observations on Daniel & Steinmann 11913 reveal that no flowers were open between 16: 00 – 17: 30 on the first day of observations, but several very large floral buds were present. On the following day, all of the large buds were found to be fully open at 08: 30, and no visitors were noted over the course of an hour.	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFF75974FF7FFE6C05E985C3.taxon	distribution	DISTRIBUTION AND HABITAT. — West-central Mexico (Michoacán; Fig. 8); plants occur on exposed limestone, often on steep slopes of arroyos, in tropical subdeciduous forests at elevations from 200 to 590 meters. ILLUSTRATIONS. — Figure 2 A – E; Richardson (1972: 66, fig. 2). CONSERVATION. — Except for L. rzedowskianum, which is known from a single site, L. brevicalyx has the smallest known EOO (0.27 km 2) among its congeners. At the site of Daniel & Steinmann 11913, about 100 plants were seen growing, mostly epipetrically, in an area of ca. 5000 m 2 on exposed limestone slopes in forest. No protected land appears to be within the species EOO. Evident threats to this species appear to be large-scale mining in the vicinity of Daniel & Steinmann 11913 (ca. 1.5 km distant) and deforestation for agriculture (including cattle) that is evident within the small EOO and in adjacent regions. Given the size of the EOO and apparent major threat (i. e., deforestation for agriculture), a single location is evident. A preliminary conservation assessment of Critically Endangered (CR; B 1, a, b; IUCN 2017) is proposed for L. brevicalyx.	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFF75974FF7FFE6C05E985C3.taxon	discussion	DISCUSSION. — Leaves are absent on the field specimens examined. Information on young stems and leaves provided above is from cultivated plants (Daniel & Steinmann 11913 cv). Leaves of these plants may not have attained the maximum size encountered in natural settings. The inflorescence, which appears before the leaves, arises from the apex of the previous season’s woody growth and bears a stalked (pedunculate) thyrse or a sessile rachis subtended by axillary dichasia (sessile thyrse). A stem forms with the flush of new clustered leaves later in the season. Most calyces have the three lobes that are characteristic of the genus. Five lobes with three prominent lobes and two reduced lobes are evident on Hinton et al. 15843; the reduced lobes are 3 to 6 mm long and 0.8 to 2 mm wide. Five-lobed calyces in inflorescences are likely teratological and suggest suppression rather than fusion of lobes for typical plants with three-lobed calyces. ADDITIONAL SPECIMENS EXAMINED. — MEXICO. Michoacán: Mpio. Aquila, KM 6. 7 carretera Aquila – Coalcomán, 18 ° 36 ’ 38 ” N, 103 ° 28 ’ 32 ” W, P. Carillo-Reyes & V. Steinmann 5484 (CAS, RSA); between Coalcomán and Aquila, 5.6 km NE of Aquila, 18 ° 36 ’ 32.54 ” N, 103 ° 28 ’ 31.38 ” W, T. Daniel & V. Steinmann 11913 (CAS, COLO, MEXU, MO, NY, US), cultivat- ed plants of this collection grown from seed in San Francisco, California, 11913 cv (CAS); Aquila [ca. 18 ° 35 ’ 49.39 ” N, 103 ° 30 ’ 14.64 ” W], G. Hinton et al. 15843 (RSA, US).	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFF65949FF7FFAB604C982F9.taxon	description	Shrubs to trees to 3.5 m tall, often epipetric. Older (woody) stems irregularly striate, ± lenticellate, lacking trichomes; younger (herbaceous) stems subterete to subquadrate, glabrous. Leaves seasonally deciduous, ± clustered just proximal to inflorescence, petiolate (or distal leaves subsessile), petioles to 70 mm long, blades subsucculent, ovate to elliptic (to obovate-elliptic), 70 – 285 mm long, 30 – 95 mm wide, 1.7 – 3.7 × longer than wide, acuminate to subfalcate at apex, rounded to acute (distal leaves) to attenuate (proximal leaves) at base, surfaces glabrous, margin entire and sometimes with inconspicuous marginal swellings / encrustations. Inflorescence a terminal (raceme to) racemose thyrse to thyrse to 580 mm long (including peduncle and excluding corollas), peduncle to 240 mm long, glabrous, rachis glabrous; dichasia expanded or sometimes modified by very short expansion between pairs of succeeding flowers with the congested dichasial axis becoming a ± linear racemelike lateral short-shoot to 10 mm long, opposite, sessile to pedunculate, (1 –) many-flowered, to 70 mm long (excluding corollas), dichasial peduncles (if present) to 24 mm long, glabrous. Bracts caducous, subfoliose, ovate to lanceolate, 18 – 52 mm long, 3. 5 – 18 mm wide, reduced in size toward apex, abaxially glabrous. Bracteoles caducous, triangular, 2 – 7 mm long, 1.5 – 2.5 mm wide, abaxially glabrous. Flowers pedicellate, pedicels 30 – 43 mm long, glabrous. Calyx 18 – 26 (– 29 in fruit) mm long, lobes fused at base for 1 – 1.5 mm, subheteromorphic, subsucculent, linear to lance-linear, 15 – 25 mm long, 3 – 5 mm wide, (subrounded) to acute at apex, abaxially glandular-punctate (punctations to 0.1 mm in diameter, sometimes sparse) but lacking elongate trichomes, posterior lobe subplanar to subconduplicate, 15 – 23 mm long, equal to or short- er than lateral lobes, 3 – 5 mm wide, lateral lobes 17 – 25 mm long, 3. 2 – 5 mm wide, all lobes (subrounded to) acute at apex. Corolla light green or greenish yellow with lobes usually becoming slightly maroon-tinged and darker maroon at distal tips with age, 35 – 40 mm long, externally glabrous, tube 21 – 23 mm long, narrow proximal portion 5 – 8 mm long, 6 – 11 mm in diameter near midpoint, throat 14 – 17 mm long, 19 – 23 mm in diameter at mouth, lobes recurved to recoiled, oblong to ovate-elliptic, 12 – 14. 5 mm long, 8.5 – 10. 5 mm wide, rounded and bifid at apex. Stamens 4, 55 – 77 mm long, distally glabrous, pubescent near base with eglandular trichomes, thecae yellowish green, 6.5 – 9. 5 mm long; staminode 1, ± triangular, 0.1 – 1.5 mm long. Style 60 – 67 mm long, glabrous, stigma unequally 2 - lobed and ± obliquely funnelform, 1.5 – 2 mm long, lobes ± elliptic, 1 – 1.5 mm long, 0.6 – 1 mm wide. Capsule (20 –) 23 – 26 mm long, 4 – 6 mm in diameter, glabrous, stipe 1.5 – 2 mm long. Seeds up to 16 per capsule, 4 – 4.5 mm long, 3 – 4 mm wide, surfaces with subconic to low rounded protrusions or becoming smooth. PHENOLOGY. — Flowering: January – March; fruiting: February – March, June. In the two populations of this species observed for ca. one hour each, floral buds were present on the plants at both sites during daylight hours (morning for Daniel 8294 and afternoon for Daniel & Butterwick 5905), and mature fallen flowers were present on the ground at one site (Daniel 8294). Likewise, Brewer et al. 7176 notes that the corollas fall in the morning.	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFF65949FF7FFAB604C982F9.taxon	distribution	DISTRIBUTION AND HABITAT. — East-central Belize (Belize, Cayo; Fig. 10); plants occur on steep limestone slopes in evergreen seasonal forests (tropical evergreen seasonal broadleaf lowland forest fide Meerman and Sabido 2001) at elevations from 10 to 120 m. ILLUSTRATIONS. — Figure 12; Leonard (1936: 197, fig. 1). CONSERVATION. — Daniel (1997) noted that the population at the type locality had been destroyed by quarrying activities, but that other populations had been located on several of the small, isolated, and steep limestone hills in the southern portion of the coastal plain in Belize District. Louteridium chartaceum has since (i. e., in 2005) been located in Cayo District, as well. Plants occur in at least three protected areas, and the species’ EOO (189 km 2) includes a portion of another one. Ecologist and botanist Steven Brewer (in litt., 14 May 2018) indicates that the species “ is common though not abundant on ridge-tops and exposed limestone along the north side of the Maya Mountains in Cayo and Belize districts. ” Brewer (in litt., 22 May 2018) also notes that in this region, potential threats to all of the limestone forests include land conversion via agriculture (both in and around protected areas), encroachment or settlement in protected areas, and fire (either natural or “ escaping ” flames from regular burning used to clear nearby agricultural fields or bush). In the same communication, Brewer also observed that, “ L. chartaceum (though I doubt the same is true for L. donnell-smithii) is surprisingly resilient from the effects of even very hot fires. Natural light fires are occasional on those limestone hills and so it seems that most species there are tolerant or resilient in the face of light fires. ” Scant change in vegetation cover within the EOO is evident from the historical imagery (1969 to 2016) seen via Google Earth Pro (2018). If the three protected areas in which the species occurs are treated either as a single or multiple locations, and the unprotected areas are treated as another location under threat (i. e., conversion of nearby, and partly within the EOO, land to agriculture as an inferred or projected threat), a preliminary conservation assessment of Endangered (EN; B 1, a, b) can be proposed for L. chartaceum.	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFF65949FF7FFAB604C982F9.taxon	discussion	DISCUSSION. — Plants grown in San Francisco from cuttings of Louteridium chartaceum from the type locality (Daniel & Butterwick 5905 cv) produced floral buds that attained 35 mm in length but abscised prior to opening. Both thyrses and racemose thyrses are present on Brewer et al. 7176. Bracteoles of a pair are sometimes fused basally for about 1 mm, at least along one side. ADDITIONAL SPECIMENS EXAMINED. — BELIZE. Belize: Runaway Creek Nature Reserve, E of Coastal Road, 17 ° 18 ’ 44 ” N, 088 ° 27 ’ 23 ” W, S. Brewer et al. 7176 (MO-image seen); along Coastal Hwy. near milepost 18, ca. 5 km W of Gales Point toward La Democracia, ca. 17 ° 11 ’ N, 088 ° 22 ’ W, T. Daniel 8294 (BR, CAS, MEXU, MICH, MO, US); Gracie Rock Hill near Rockville Quarry between Western Hwy. and Sibun River, ca. 30 km SW of Belize City [ca. 17 ° 23 ’ 13.67 ” N, 088 ° 27 ’ 1.91 ” W], T. Daniel & M. Butterwick 5905 (C, CAS, K, MICH, MO, NY), cultivated plants of this collection grown from cuttings in San Francisco, California, 5905 cv (CAS); Gracie Rock, 1.5 – 4 mi S of Mile 22 on Western Hwy., J. Dwyer 10959 (LL, MEXU, MO), R. Liesner & J. Dwyer 1485 (BM, DUKE, MO, NY, TEX). Cayo: Manatee Forest Reserve, Banks of Indian Creek, at confluence of Yaha and Indian Creek, 1 km W from Daylight Cave, 17 ° 12 ’ 54.2 ” N, 088 ° 33 ’ 19.4 ” W, H. Baden & D. Harris 3 (BM).	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFCB594FFF7FFBBA055082EF.taxon	description	Perennial herbs to shrubs to 2 m tall, terrestrial. Older (woody) stems subquadrate, lenticellate, lacking trichomes; younger (herbaceous) stems subquadrate to flattened, glabrous. Leaves not all seasonally deciduous, ± evenly distributed along stems, petiolate, petioles to 85 mm long, blades subsucculent, (ovate-elliptic to) elliptic to obovate, 125 – 335 mm long, 43 – 153 mm wide, 2.2 – 3.2 × longer than wide, acute to acuminate to acuminate-falcate at apex, attenuate at base, surfaces glabrous, margin entire to sinuate. Inflorescence a terminal pedunculate (racemose thyrse to) thyrse to 335 mm long (including peduncle and excluding corollas), peduncle to 230 mm long, glabrous, rachis glabrous; dichasia often modified via sympodial expansion and appearing like lateral branches, opposite or alternate, (subsessile to) pedunculate, 1 – many-flowered, to 117 mm long (excluding corollas), dichasial peduncles 1 – 15 mm long, glabrous. Bracts persistent, lance-ovate to ovate to elliptic (or subfoliose proximally), 6 – 70 mm long, 2 – 24 mm wide, abaxially glabrous or glandular punctate (but lacking elongate trichomes), sometimes with lenticel-like encrustations as well. Bracteoles persistent, ovate to elliptic, 3. 5 – 11 mm long, 2 – 6. 5 mm wide, abaxial surface like that of bracts, those of a pair sometimes basally fused for ca. 1 mm (at least along one side). Flowers pedicellate, pedicels 6 – 58 mm long, glabrous or sometimes covered with crystal-like punctations or globules or pubescent with stipitate glandular trichomes (sometimes sparse) 0.1 – 0.2 mm long. Calyx pale green to white, 23 – 65 mm long, lobes subheteromorphic to heteromorphic, membranaceous, acute to subacuminate at apex, abaxially glabrous or pubescent (at least near base) with sparse glandular trichomes like those of pedicels and conspicuously or inconspicuously glandular-punctate, punctations <0.1 mm in diameter, posterior lobe planar or ± conduplicate, oblong-lanceolate to ovate to elliptic, 23 – 65 mm long, longer than lateral lobes, 10 – 36 mm wide, lateral lobes lance-ovate to ovate to ovate-cordate to elliptic, 21 – 52 mm long, 7 – 27 mm wide. Corolla greenish white to pale yellow, 40 – 60 mm long, externally pubescent with erect to flexuose glandular (0.1 – 0.5 mm long) and eglandular (0.1 – 1 mm long) trichomes, trichomes sometimes very sparse, tube 25 – 46 mm long, narrow proximal portion 3 – 17 mm long, 4. 5 – 10 mm in diameter near midpoint, throat 22 – 43 mm long, 17 – 23 mm in diameter at mouth, lobes spreading to recoiled, broadly triangular to broadly ovate to subcircular, 8 – 16 mm long, 6 – 12 mm wide, round- ed to acute and ± emarginate at apex. Stamens 2, 52 – 57 mm long, filaments glabrous distally, not seen proximally, thecae 9 – 15 mm long; staminodes (if present) not seen. Style 50 – 55 mm long, glabrous distally, not seen proximally, stigma subequally to unequally 2 - lobed, lobes linear-oblong, 3 – 5. 2 mm long, 1.2 – 1.5 mm wide. Capsule 27 – 35 mm long, 6 – 6.5 mm in diameter, glabrous, stipe 7 – 12 mm long. Seeds up to 16 per capsule, 4 – 6. 2 mm long, 4 – 6 mm wide, surfaces smooth to wrinkled. PHENOLOGY. — Flowering: throughout the year; fruiting: June – March. Folsom et al. 5464 notes that corollas open in the evening; González 7918 notes that flowers were closed at 17: 00 but opened by 19: 00.	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFCB594FFF7FFBBA055082EF.taxon	distribution	DISTRIBUTION AND HABITAT. — Costa Rica (Cartago, Guanacaste, Heredia, Limón, Puntarenas, and San José) and Panama (Bocas del Torro, Chiriquí, Colón, and Guna Yala), primarily (or exclusively?) on the Caribbean versant (Fig. 9); plants occur in and along streams in lowland to montane wet forests and cloud forests at elevations from 15 to 1700 m. ILLUSTRATIONS. — Figure 4 A, B; Durkee (1978: 232, fig. 20); Durkee (1985: 14, fig. 12). CONSERVATION. — Louteridium costaricense has the largest EOO (61,058 km 2) among congeners. Its EOO includes at least 23 protected areas (16 in Costa Rica and seven in Panama), and the species has been collected in five of them. Although there are undoubtedly threats to the species in various portions of its extensive geographic range, considering its overall distribution and relative abundance, the species is here assessed as Least Concern (LC; IUCN 2017).	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFCB594FFF7FFBBA055082EF.taxon	discussion	DISCUSSION. — Corollas of Louteridium costaricense are more tubular and generally less open than in congeners. No significant differences were encountered between plants from Costa Rica and those from Panama. Considerable variation is evident in the shape and width of corolla lobes from oblong-lanceolate to lance-ovate and 7 to 12 mm wide (e. g., Tonduz 8123) to ovate-cordate and 21 to 32 mm wide (e. g., Herrera & Gamboa 8581). ADDITIONAL SPECIMENS EXAMINED. — COSTA RICA. Cartago: Cantón de Turrialba, Valle del Reventazón, Grano de Oro, El Sies, Moravia de Chirripó, 09 ° 49 ’ 50 ” N, 083 ° 26 ’ 40 ” W, P. Campos & A. Campos 213 (MO); “ Limon, Siquirres, Las Brisas de Pacuarito, camino a Cerro Tigre, ” 09 ° 57 ’ 00 ” N, 083 ° 25 ’ 50 ” W [= Cartago Province], G. Herrera 8840 (K); “ Limon, Siquirres, Pacuarito, Las Brisas, ” 09 ° 56 ’ 10 ” N, 083 ° 25 ’ 20 ” W [= Cartago Province], G. Herrera et al. 8646 (K). Guanacaste: Cordillera de Tilarán, 1 km N de Las Nubes de Río Chiquito, Zona Monteverde, 10 ° 22 ’ N, 084 ° 51 ’ W, W. Huber & W. Zuchowski 8681 (MO); “ Puntarenas, ” Cordillera de Tilarán, Monteverde, San Gerardo Biological Station, Sendero Congo, 1 km W of Station, 10 ° 22 ’ N, 084 ° 48 ’ W [= Guanacaste Province], D. Penneys 454 (CAS). Heredia: Finca La Selva, OTS Field Station on Río Puerto Viejo just E of its jct. with Río Sarapiquí [10 ° 25 ’ 53 ” N, 084 ° 00 ’ 13 ” W], B. Hammel & J. Trainer 13226 (CAS, DUKE, MO). Limón: Cantón Siquirres, Finca de E. Berlin, Pocora, cuenca del Río Destierro, [ca. 10 ° 10 ’ 5.11 ” N, 083 ° 35 ’ 41.15 ” W], J. González 7918 (LSCRimage seen), J. González & J. Chaves 8728 (LSCR-image seen), J. González et al. 8157 (LSCRimage seen); Cantón Siquirres, La Alegría, Alto Botella, [ca. 10 ° 05 ’ 45.64 ” N, 083 ° 36 ’ 58.36 ” W], J. González et al. 7967 (LSCR-image seen), 9618 (LSCR-image seen); Cantón Siquirres, Florida, Alto Botella, [10 ° 05 ’ 19.97 ” N, 083 ° 33 ’ 26.45 ” W], J. González et al. 9353 (LSCR-image seen); Cantón Siquirres, Pocora, Cuenca del Río Destierro, Cordillera de Talamanca, Cantón de Matina, 200 m aguas abaja de la confluencia de Quebrada Cañabral con Río Barbilla, 10 ° 00 ’ 10 ” N, 083 ° 25 ’ 30 ” W, G. Herrera 2286 (DUKE, MO); Almirante, Cerro Chiqui, 09 ° 43 ’ 40 ” N, 083 ° 18 ’ 30 ” W, G. Herrera & W. Gamboa E. 8581 (F, K, MEXU, MO, NY, US); 10 km SW of Siquirres on road to Turrialba, E of ridge top, J. MacDougal 1110 (DUKE); Cantón de Limón, Cordillera de Talamanca, Cerro Muchilla, Fila Matama, entrando por pueblo El Progresso, 09 ° 47 ’ 40 ” N, 083 ° 06 ’ 30 ” W, R. Robles & A. Chacón 2765 (MO). Puntarenas: vicinity of Finca Las Cruces, SE of San Vito on road to Via Neilly, [ca. 08 ° 47 ’ 14.82 ” N, 082 ° 57 ’ 25.04 ” W], R. Weaver et al. 1777 (DUKE). San José: from La Montura to Los Chorritos, L. Gomez et al. 20929 (MO). PANAMA. Bocas del Torro: Fish Creek Mts., vicinity of Chiriqui Lagoon [ca. 08 ° 58 ’ 59.68 ” N, 082 ° 13 ’ 48.70 ” W], H. von Wedel 2283 (MO). Chiriquí: Fortuna Dam site [ca. 08 ° 44 ’ 46.90 ” N, 082 ° 14 ’ 38.50 ” W], J. Folsom & R. Dressler 5464 (MO); La Fortuna hydroelectric project, ridge behind camp, [08 ° 41 ’ 25.09 ” N, 082 ° 13 ’ 47.82 ” W], B. Hammel 2228 (MEXU, MO). Colón: along Río Guanche, 3 – 7 km above bridge [ca. 09 ° 29 ’ N, 079 ° 38 ’ W], B. Hammel et al. 4916 (MO); ca. 2 – 3 mi up Río Guanche [ca. 09 ° 30 ’ 30 ” N, 079 ° 39 ’ 30 ” W], H. Kennedy & R. Foster 2180 (MO). Guna Yala (San Blas): Nusagandi, El Llano – Cartí Road, along creek on Atlantic slope [ca. 09 ° 18 ’ 18.56 ” N, 078 ° 59 ’ 20.14 ” W], G. de Nevers & B. González 3656 (DUKE, MEXU, MO); El Llano – Cartí Road, KM 19. 1 [ca. 09 ° 20 ’ 51.93 ” N, 079 ° 01 ’ 54.94 ” W], G. de Nevers et al. 6195 (CAS, DUKE, MEXU, MO); trail to Cerro Obu (Habu of maps) from Río Urgandi (Río Sidra) [ca. 09 ° 23 ’ 59.51 ” N, 078 ° 49 ’ 40.88 ” W], G. de Nevers et al. 8007 (CAS, MO); El Llano – Cartí hwy., ca. 23 km N of El Llano, R. Dressler 4310 (DUKE, ENCB, F, MO); Nusagandi, El Llano – Cartí road, 09 ° 19 ’ N, 078 ° 55 ’ W, H. van der Werff 7019 (CAS, MO).	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFCD594DFF7FFBAA053785AF.taxon	description	Shrubs to trees to 12 m tall, sometimes epipetric. Older (woody) stems quadrate, lenticellate, irregularly striate-sulcate, lacking trichomes; younger (herbaceous) stems subquadrate-sulcate, sparsely lenticellate, irregularly fissured, evenly pubescent with erect to flexuose simple and dendritic (sparse) eglandular trichomes <0.1 – 0.5 mm long. Leaves seasonally deciduous, ± clustered at apex of old growth or at apex of an otherwise leafless shoot of new growth, petiolate, petioles to 65 mm long, blades subsucculent, ovate to elliptic to broadly elliptic, 76 – 190 mm long, 40 – 122 mm wide, 1.4 – 2.5 × longer than wide, apiculate to acuminate at apex, rounded to acute to attenuate at base, adaxial surface pubescent with simple and dendritic eglandular trichomes, trichomes soon becoming ± restricted to proximal portion or to midvein, abaxial surface pubescent (especially along veins) with dendritic trichomes to 0.5 mm long, midvein often pinkish or reddish, margin entire (sometimes undulate and appearing subcrenate). Inflorescence a terminal subsessile to pedunculate raceme to 220 mm long, peduncle to 50 mm long, pubescent like young stems, rachis pubescent like young stems; dichasia opposite or alternate, sessile, 1 - flowered, to 47 mm long (excluding corollas). Bracts caducous, lanceolate to lance-ovate, 11 – 16 mm long, 2. 5 – 6 mm wide, abaxially pubescent with simple and dendritic trichomes 0.05 – 0.2 mm long. Bracteoles caducous, lanceolate to lance-elliptic, 7 – 9 mm long, 2 – 3 mm wide, abaxially pubescent like bracts. Flowers pedicellate, pedicels 21 – 46 mm long, pubescent like rachis or with the trichomes to 1 mm long. Calyx 17 – 32 mm long, lobes fused at base for 1 – 1.5 mm, subhomomorphic to subheteromorphic, membranaceous, subelliptic to ovate-elliptic to subrhombic-obovate, rounded to acute at apex, abaxially pubescent with mostly dendritic trichomes 0.1 – 0.5 mm long, posterior lobe planar, 17 – 32 mm long, 10 – 19 mm wide, usually slightly larger and sometimes more conspicuously venose than lateral lobes, major veins often maroon, lateral lobes 20 – 31 mm long, 8 – 18 mm wide. Corolla light green or greenish yellow, sometimes with maroon on limb (especially at base of lobes) and distal portion of throat, externally glabrous (inconspicuously glandular punctate but lacking elongate trichomes), 50 – 62 mm long, tube 35 – 37 mm long, narrow proximal portion 11 – 15 mm long, 6 – 10. 5 mm diameter near midpoint, throat 20 – 24 mm long, 25 – 35 mm diameter at mouth, lobes recurved to recoiled, broadly ovate to subtriangular, 13 – 20 mm long, 10 – 21 mm wide, entire at apex. Stamens 4, 60 – 80 mm long, filaments glabrous distally, glabrous or pubescent with eglandular trichomes near base, thecae 8 – 10. 5 mm long; staminode 1, rodlike, 0. 6 – 40 mm long. Style 70 – 101 mm long, distally glabrous, pubescent with eglandular and glandular trichomes near base, stigma equally 2 - lobed, lobes broadly elliptic to broadly ovate-triangular, 1 – 2 mm long, 1 – 1.4 mm wide. Capsule 21 – 28 mm long, 6.5 – 9. 5 mm in diameter, densely pubescent with erect glandular trichomes 0.05 – 0.5 mm long and with an overstory (sometimes sparse) of erect to flexuose (sometimes dendritic) eglandular trichomes to 1.4 mm long, stipe 2.5 – 3.5 mm long. Seeds up to 16 per capsule, 5. 2 – 7 mm long, 5 – 6. 4 mm wide, surfaces smooth. PHENOLOGY. — Flowering: February – March; fruiting: March – April. Based on field observations and cultivated plants (Daniel et al. 11894 cv), flowering occurs on leafless (or nearly leafless) plants during the dry season. Near the end of the dry season (e. g., late March – April) when flowering is waning and fruits are mature, a new flush of vegetative growth appears from axils of clustered leaf scars at the base of the inflorescence, which eventually falls away. New internodal stem elongation (e. g., the young stems of the description above) takes place from the axil of a leaf scar on the old growth and terminates in a cluster of new leaves and / or between at least one of the pairs of leaves in the cluster and the remaining cluster. Field observations of Daniel et al. 11894 over three days (24 – 26 February 2012) revealed: day 1 between 08: 00 – 09: 00 (light) — corollas mostly fallen, only a few from previous night still attached to tree; day 2 between 19: 00 – 20: 00 (dark) — all corollas open, ca. 100 seen, bats active around plants but none seen visiting flowers, no floral odor detected and no nectar visible in saccate tube of undissected corollas, stigma extended ca. 1 cm beyond anthers on fresh flowers, stigma of 7 flowers examined for pollen (all pollinated); day 3 between 17: 30 – 18: 30 (light) — many corollas open and many others still in bud, corollas open fully (including recurving of corolla lobes) in 15 to 20 seconds, open flowers actively visited by Cinnamon Hummingbird (Amazilia rutila, species det. by Jeff Chemnick), birds probe for nectar once or twice at same flower before moving on, nectar is located behind a barrier (seen in dissected flowers) at the base of the tube, nectar not visible in the saccate throat, birds appear to contact anthers with head or back and presumably contact the stigma on some visits as well, birds visit between 5 – 10 flowers on a single plant or on multiple plants before moving away from an area or resting on a branch, 2 flowers were observed to open and when subsequently visited by a hummingbird (one flower visited once, other flower visited twice) they were checked for pollination (no pollen observed on stigma of either), small bees or flies also observed visiting flowers but they only contact anthers, buds continue opening until full darkness (at 18: 30); day 3 between 18: 30 – 20: 30 (dark) — hummingbirds no longer active, bats very active around plants but none observed visiting flowers. On a single flower of Daniel et al. 11894 cv grown in San Francisco, corolla lobes began to separate at 15: 50 and the corolla was fully open (lobes spreading 90 ° with respect to the throat or reflexed) with the stamens and style fully exserted by 16: 15. By 18: 00, all corolla lobes were at least partially recoiled. At 23: 00, 6.5 μl of nectar was encountered in the nectar chamber, and the stigma was dusted with pollen from the anthers. At 07: 15 the next day, 72.1 μl of nectar was recovered from the nectar chamber. The corolla abscised and fell from the persisting flower at 07: 45. Thus, the corolla of the pollinated flower persisted for ca. 16 hours.	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFCD594DFF7FFBAA053785AF.taxon	distribution	DISTRIBUTION AND HABITAT. — Southern Mexico (Oaxaca; Fig. 8) in the Sierra Madre del Sur and the Isthmus of Tehuantepec; plants occur on limestone (often karstic) slopes of streams in tropical deciduous forests and tropical subdeciduous forests at elevations from 600 to 750 m. ILLUSTRATIONS. — Figure 2 F – L; Daniel (2017: 143, fig. 8). CONSERVATION. — A discussion and preliminary conservation assessment of Endangered (En) was proposed for this species (B 1, a, b; IUCN 2017) by Daniel (2017).	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFCD594DFF7FFBAA053785AF.taxon	discussion	DISCUSSION. — Louteridium dendropilosum is unique among congeners by its dendritic trichomes, which are present on both vegetative and reproductive organs. Trichomes of other species may consist of one or more cells and be either glandular or eglandular, but they are not branched. Bracts and bracteoles are caducous prior to maturation of the flowers associated with them; thus, they are rarely encountered and are not present on any of the wild-collected specimens. The data on their shape, size, and pubescence noted above were taken just prior to their dehiscence from the young inflorescence of a cultivated plant (Daniel et al. 11894 cv). ADDITIONAL SPECIMENS EXAMINED. — MEXICO. Oaxaca: Distr. Pochutla, Mpio. San Miguel del Puerto, Arroyo Arena, ca. 100 m downstream from jct. Río Laja, ca. 3 km SE of Rancho Dioon toward Xadani, 15 ° 58 ’ 51.33 ” N, 096 ° 05 ’ 53.91 ” W, T. Daniel, et al. 11894 (CAS, MEXU), 12187 (CAS, MEXU), cultivated plants of 11894 grown from seeds in San Francisco, California, 11894 cv (CAS); Mpio. El Barrio, 9 km N [sic] de El Barrio, Cerro Palmasola, antena microondas [ca. 16 ° 44 ’ 32.52 ” N, 095 ° 05 ’ 36.04 ” W], R. Fernández N. 4189 (IEB, NY); Distr. Pochutla, Mpio. San Miguel del Puerto, Arroyo Arena, 15 ° 58 ’ 39.7 ” N, 096 ° 05 ’ 54.9 ” W, J. Pascual 1396 (MEXU, SERO, TEX); Distr. Pochutla, Mpio. San Miguel del Puerto, 300 m de la terracería sobre la vereda rumbo Río la Laja, 15 ° 58 ’ 49.9 ” N, 096 ° 06 ’ 6.9 ” W, A. Saynes V. et al. 3831 (MEXU, SERO); Distr. Juchitán, Mpio. El Barrio, parte alta del Cerro Palmasola, junto a la antena de microondas [ca. 16 ° 44 ’ 32.52 ” N, 095 ° 5 ’ 36.04 ” W], S. Zamudio R. 6352 (CAS, IEB).	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFCF5947FF7FFB6B07378451.taxon	description	Shrubs or trees to 12 m tall, terrestrial or epipetric. Older (woody) stems subquadrate to quadrate, irregularly striate-fissured, pubescent or glabrate; younger (herbaceous) stems quadrate to quadrate-sulcate, densely pubescent with erect to flexuose eglandular trichomes to 2 mm long (distal 1 or several internodes sometimes with glandular trichomes as well). Leaves not all seasonally deciduous, evenly distributed on young shoots, petiolate, petioles to 190 mm long, blades membranaceous, ovate to broadly ovate to cordate, 140 – 465 mm long, 85 – 316 mm wide, 1 – 2 × longer than wide, acuminate at apex, acute to rounded to cordate at base, surfaces densely pubescent with cauline type trichomes, margin crenate-dentate with rounded (to acute) teeth up to 2 mm long. Inflorescence a terminal pedunculate raceme to thyrse to 450 mm long (including peduncles and excluding corollas), peduncle to 250 mm long, pubescent with flexuose glandular and eglandular trichomes to 2.8 mm long (viscid), rachis viscid; dichasia opposite, sessile or pedunculate, (1 –) 3 – 5 (– many) - flowered, to 130 (– 185) mm long (excluding corollas), dichasial peduncles to 5 (to 35 or to 90 at proximal nodes) mm long, viscid. Bracts foliose to subfoliose and often persistent proximally, reduced in size and caducous (usually only scars present) distally, proximalmost pair sessile to subsessile, lanceolate to cordate, similar to leaves except smaller (e. g., 45 × 32 mm), distal pairs ovate to lanceolate, 10 – 22 mm long, 2. 5 – 9 mm wide, abaxially pubescent like leaves (i. e., trichomes mostly eglandular). Bracteoles and secondary bracteoles usually present on young inflorescences but becoming deciduous as inflorescence matures, linear to lanceolate, 3. 5 – 13 mm long, 1 – 3. 4 mm wide, abaxially pubescent like bracts or viscid. Flowers pedicellate, pedicels to 105 mm long, viscid with trichomes mostly glandular. Calyx 22 – 35 (– 40 in fruit) mm long, lobes fused at base for 1 – 3 mm, heteromorphic, membranaceous, abaxially viscid where exposed in bud, posterior lobe conduplicate, ovate, 21 – 37 mm long, equaling to longer than lateral lobes, 8 – 15 mm wide, acuminate to subfalcate at apex, lateral lobes lunate, 17 – 33 (– 37) mm long, 6 – 10. 5 mm wide, acute to subacuminate at apex. Corolla cream to green-yellow with maroon veins to pale maroon, 41 – 57 mm long, externally viscid or pubescent (sometimes sparsely so) with erect to flexuose glandular trichomes 0.05 – 0.3 mm long, tube 25 – 34 mm long, narrow proximal portion 8 – 14 mm long, 5. 7 – 9 mm in diameter near midpoint, throat 15 – 24 mm long, 25 – 38 mm in diameter at mouth, lobes recoiled, broadly ovate to subtriangular, 16 – 20 mm long, 13 – 15 mm wide, acute or emarginate at apex. Stamens 2, 65 – 85 mm long, filaments glabrous distally, pubescent proximally with eglandular trichomes, thecae 10 – 15 mm long; staminodes 2, 0.5 mm long. Style 76 – 85 mm long, glabrous (or with a few glandular trichomes near base), stigma subequally to unequally 2 - lobed and often ± funnelform, lobes broadly elliptic to obovate to 3 - pronged, 1 – 2. 2 mm long, 0.7 – 2.1 mm wide. Capsule 21 – 31 mm long, 5 – 8 mm in diameter, pubescent with erect glandular trichomes 0.1 – 0.8 mm long, stipe 1.5 – 2.5 mm long. Seeds 12 – 16 per capsule, 4.5 – 5.5 mm long, 4 – 5 mm wide, surfaces minutely papillose in longitudinal rows. PHENOLOGY. — Flowering: November – June; fruiting: December – June. Brewer et al. 5748 notes corollas opening at night, just after dark. Davidse & Brant 32089 and Gregory 612 note corollas opening at dusk or in the evening and withering by sunrise. Daniel (2010) noted floral color forms with different flowering times (see below). Fredy Archila (in litt. January 2012) observed both bats and moths visiting flowers of L. donnell-smithii in and around Cobán, Guatemala over a period of years.	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFCF5947FF7FFB6B07378451.taxon	distribution	DISTRIBUTION AND HABITAT. — Southern Mexico (Chiapas), Belize, Guatemala, Honduras (Fig. 10); plants occur, usually on limestone (often karstic), on slopes and along streams in lowland moist to wet forests, mesophytic montane forests, montane rain forests, and cloud forests at elevations from 1 to 1600 (to 2100) meters. ILLUSTRATIONS. — Figures 13, 4 C, D; Lindau (1895: 294, fig. 118); Gibson (1974: 407, fig. 90). LOCAL NAMES. — “ Carreton ” (Lentz et al. 2428), “ matacucuyuchi ” (Miranda 7163); “ oxox če ” (Williams & Wilson 40437 and Wilson 40943, Quecchí name); “ palpalte ” (Ochoa 80); “ tabaco silvestre ” (Contreras 5510); “ vejiga ” (Standley 91236); “ ulul k’uum ” (Ventur 177). USES. — Williams & Wilson 40437 notes that the young growth is eaten; Ochoa 80 indicates that “ esta planta es comida por el jabali. ” CONSERVATION. — Louteridium donnell-smithii is the most commonly collected species of the genus, and it is also infrequently cultivated in botanical gardens. Its EOO (32,499 km 2) encompasses all or parts of at least 40 protected areas (10 in Belize, 19 in Guatemala, 9 in Honduras, two in Mexico). Collections of the species have been made in 13 of these areas. The species has been abundantly collected in most portions of its EOO, and in some regions it was sometimes noted on collection labels or observed (TFD) to be common. Fredy Archila of Cobán, Guatemala indicates (in litt. Jan 2012) that in the vicinity of Cobán there were hundreds of trees of L. donnell-smithii in the 1980 s. Although most of these are no longer extant, some still remain as street trees in that city. Based on these data, this species is accorded a preliminary conservation assessment of Least Concern (LC).	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFCF5947FF7FFB6B07378451.taxon	discussion	DISCUSSION. — In the protologue, Watson (1888) provided the collection information cited above. The holotype has been cited by various authors (e. g., Richardson 1972; Daniel 1995) as being at US. Indeed, a specimen there (accession: 1320518 with image number: 00136468) was annotated by D. Gibson and T. Daniel as the holotype. This somewhat fragmentary specimen, originally in H. Rusby’s herbarium, bears von Türckheim’s number 856 (as indicated in the protologue), but notes a collection date of February 1886. The date of collection of the specimen from Pansamalá indicated in the protologue is May 1887. The specimen at GH, where Watson conduct- ed his studies, bears the date noted in the protologue and an annotation written by Watson stating, “ Louteridium Donnell-Smithii, Watson-n. gen. ” Therefore the specimen at GH indicated above conforms to the information in the protologue. The earlier annotations by Gibson and Daniel of the specimen at US are considered to be erroneous, and the specimen at GH noted above is treated as the holotype. This flowering and fruiting specimen lacks leaves, which are described in the protologue. However, drawings of the leaves are on another sheet at GH bearing the same barcode number. If the drawings are original material, the holotype would be considered as mounted on two sheets, and if they were made subsequently, the sheet with an inflorescence, flowers, and fruits would be considered as the holotype. Other collections that bear von Türckheim’s number 856, but with a date of February 1886, are located at the following herbaria: K, NY, P, PH, US (although one of three specimens at US has both “ 1887 ” and February 1886 on it). Whether these represent isotypes with an incorrect date (i. e., actually collected in May 1887) or the same collection number was used for gatherings of this species during different visits to Pansamalá on various dates remains unknown. Daniel (2010) noted variation in the timing of anthesis, color of corollas, and diurnal visitation to dark-colored flowers by hummingbirds in this species. There is a possible correlation with cream to green-yellow corollas at lower elevations and chestnut to maroon corollas at higher elevations. Based on specimens of L. donnell-smithii at F, which are mostly from Guatemala, plants with cream to green-yellow corollas occur from 75 to 630 m, whereas those with chestnut to maroon corollas occur at 1000 to 2100 m. Figures 4 C and D show these extremes of corolla coloration in Guatemala. Other flowers show various degrees of maroon coloration on cream-yellowish corollas. Based on a larger sample of herbarium specimens that include data on corolla color and elevation, it would appear that both the cream to green-yellow and chestnut to maroon color forms occur in Chiapas, Belize, and Guatemala, but that there is no clear difference in elevational ranges of these forms in Chiapas and Belize. However, it is somewhat difficult to correlate corolla color and elevation from herbarium specimens because of variation in the terminology of colors used by collectors. Additional studies that address the different (or at least partially so) flowering times of the different color forms and their primary pollinators, as well as any correlation of floral color with elevation, are desirable. The inflorescence of L. donnell-smithii is commonly a somewhat contracted thyrse at the proximal nodes (with dichasial peduncles usually to 5 mm long) and a raceme (with sessile dichasia) distally. Several collections (e. g., Armour & Chable 6047, Hawkins 981, Vargas et al. 1183) have exceptionally elongate peduncles (to 90 mm long) with expanded dichasia (e. g., secondary peduncles to 55 mm long). These latter dichasia are usually only at the proximal 1 to 3 nodes. ADDITIONAL SPECIMENS EXAMINED. — MEXICO. Chiapas: Las Nubes, F. Billiet 8889 (BR); Mpio. Ocosingo, W end of Laguna Ocotal Grande, [ca. 16 ° 50 ’ 26.15 ” N, 091 ° 28 ’ 5.95 ” W], D. Breedlove 15709 (DS, F, LL, MICH, NY, US); Mpio. Las Margaritas, western side of Laguna Miramar E of San Quintín, [ca. 16 ° 24 ’ 24.45 ” N, 091 ° 18 ’ 6.60 ” W], D. Breedlove 33282 (DS, ENCB, F, LL, MEXU, MICH); Mpio. La Independencia, valley of Santa Elena along rd. to Ixcán, 30 km E of paved rd. at Montebello, [ca. 16 ° 15 ’ 57.16 ” N, 091 ° 49 ’ 20.79 ” W], D. Breedlove 41991 (DS, MO); Mpio. La Trinitaria, 10 km ENE of Dos Lagos above Sta. Elena, D. Breedlove 49721 (CAS, ENCB), D. Breedlove & F. Almeda 57578 (CAS, ENCB, NY); Mpio. Ocosingo, near lake at Naja, [ca. 16 ° 59 ’ 4.48 ” N, 091 ° 35 ’ 11.70 ” W], D. Breedlove 49962 (CAS, MEXU); Mpio. Ocosingo, 70 km SW of Palenque on rd. to Ocosingo along the Jol Uk’um, [ca. 17 ° 10 ’ 33.18 ” N, 092 ° 06 ’ 54.71 ” W], D. Breedlove 50875 (CAS, ENCB), D. Breedlove & F. Almeda 48259 (CAS); Mpio. La Trinitaria, KM 21 – 23 along road from Lago Tzizcao toward Santa Elena, [ca. 16 ° 06 ’ 20.94 ” N, 091 ° 34 ’ 48.32 ” W], D. Breedlove & M. Bourell 68070 (BR, CAS, MEXU, US), D. Breedlove & T. Daniel 71197 (CAS); Mpio. Ocosingo, 1 km S de Naja, camino a Monte Líbano, [ca. 16 ° 58 ’ 25.71 ” N, 091 ° 35 ’ 15.33 ” W], E. Martínez 17416 (MEXU, MO); Mpio. Ocosingo, Naha, 15 km N de Monte Líbano, camino a Chancala, E. Martínez S. 18025 (MEXU); Mpio. Ocosingo, Cañón del Colorado, E. Martinez S. & C. Ramos 26235 (K, MEXU, XAL); Mpio. Las Margaritas, 1 km S de Flor del Río, E. Martínez S. et al. 24768 (B); Mpio. La Trinitaria, KM 18, Col. Cuauhtémoc, [16 ° 05 ’ 16.55 ” N, 092 ° 02 ’ 57.11 ” W], A. Méndez G. [= A. Ton] 8026 (MEXU); Mpio. Trinitaria, Col. Cuauhtémoc, KM 20 a Montaña Alta, A. Méndez G. [= A. Ton] 8893 (IEB, MEXU, MO, RSA, XAL); Monte Líbano, 60 km E de Ocosingo, [ca. 16 ° 54 ’ 7.71 ” N, 091 ° 37 ’ 10.27 ” W], F. Miranda 7163 (MEXU); Mpio. Las Margaritas [= Mpio. Palenque?], NW del ejido José Castillo T., [ca. 17 ° 24 ’ 59.51 ” N, 091 ° 50 ’ 10.35 ” W], M. Ochoa 80 (MEXU); Mpio. La Independencia, Col. Fco. Madero, ca. de la carretera, [ca. 16 ° 17 ’ 23.49 ” N, 091 ° 54 ’ 38.13 ” W], T. Ramamoorthy et al. 1930 (CAS, MEXU); near jct. of Río Perlas and Río Jataté at San Quintín and near Laguna Miramar, [ca. 16 ° 23 ’ 27.68 ” N, 091 ° 20 ’ 21.96 ” W], E. Sohns 1684 (MICH, NY, UC, US); Km. 36 – 38 al E Tziscao o a 7 – 9 km al E de Amparo Agua Tinta, sobre el camino Montebello a Bonampak, [ca. 16 ° 07 ’ 19.90 ” N, 091 ° 26 ’ 36.06 ” W], O. Téllez & J. Villaseñor 6642 (CAS, ENCB, MEXU); Mpio. Ocosingo, 3 km W de Plan de Ayutla, [ca. 16 ° 47 ’ 59.90 ” N, 091 ° 18 ’ 58.16 ” W], F. Vázquez B. & S. Avandano R. 1650 (ENCB, MEXU, XAL); Mpio. La Trinitaria, ca. 56 km E of La Trinitaria, 9.4 km E of Lago Tziscao, 16 ° 06 ’ N, 091 ° 35 ’ W, T. Wendt et al. 2409 (CAS, MEXU, MO, TEX). BELIZE. Cayo: Maya Mts., central Bladen Nature Reserve, 16 ° 29 ’ 41 ” N, 088 ° 54 ’ 43 ” W, S. Brew- er et al. 5748 (MO); 2.5 mi past Guacamallo Bridge, road to Millionario [16 ° 50 ’ 05 ” N 089 ° 02 ’ 12 ” W], J. Dwyer & R. Liesner 12313 (BM, DUKE, MEXU, MO, NY); S of San Luis and E of Camp Six [16 ° 54 ’ 35 ” N 089 ° 00 ’ 22 ” W], J. Dwyer et al. 360 (MEXU, MO); vicinity of Grano de Oro lumber camp, 6 mi S of Millionario [16 ° 40 ’ 22 ” N 089 ° 01 ’ 35 ” W], A. Gentry 7791 (MO); Central Maya Mts., Macal river, vicinity of Guacamollo Bridge, 16 ° 55 ’ N, 088 ° 55 ’ W, R. Glaholt 11 (MO); Las Cuevas Research Station to Monkey Tail River, 16 ° 44 ’ N, 088 ° 59 ’ W, D. Harris 7888 (BM, MO); between San Antonio and Santa Helena, 17 ° 06.290 ’ N, 089 ° 03.457 ’ W, D. Lentz et al. 2428 (NY); Ceibo Chico, drill site, 16 ° 31 ’ 21 ” N, 089 ° 07 ’ 04 ” W, M. Peña et al. 947 (MO); Grano de Oro Camp, M. Peña-Chocarro et al. 1435 (BM, MO); Chiquibul Forest Reserve, 2.2 mi SE of Holtec Camp, G. Proctor 30106 (BM, LL); 3 km S de New Maria camp, T. Ramamoorthy et al. 2965 (MEXU); between Las Cuevas and Río Ceibo Chico, 16 ° 43 ’ 02 ” N, 088 ° 59 ’ 02 ” W, R. Rees et al. 2 (BM, MEXU, MO); Puente Natural, C. Whitefoord 9125 (BM, MO); Chiquibul Forest Reserve, Natural Arch, C. Whitefoord 10044 (BM, US); Chiquibul Forest Reserve, Monkey Tail Branch, C. Whitefoord 10051 (BM); Chiquibul Forest Reserve, road from Caracol junction to New Maria Camp, C. Whitefoord 10136 (BM, US). Toledo: river trail from Blue Creek village to Blue Creek’s source (ca. 1 km N of village), ca. 26 km NW of Punta Gorda, [ca. 16 ° 12 ’ 13.03 ” N, 089 ° 02 ’ 49.12 ” W], T. Daniel & M. Butterwick 5891 (CAS); Maya Mts. foothills, Solomon Camp, vicinity of jct. Richardson Creek and Bladen Branch, 16 ° 32 – 33 ’ N, 088 ° 45 – 46 ’ W, G. Davidse & A. Brant 32089 (CAS, MEXU, MO, US), Maya Mts., canyon along Bladen Branch, from Richardson Creek to Quebrada de Oro, 16 ° 31 – 33 ’ N, 088 ° 46 – 49 ’ W, G. Davidse & A. Brant 32300 (CAS, MO); southern Maya Mts., Bladen Nature Reserve, upper Bladen Branch, 16 ° 29 ’ 31 ” N, 088 ° 54 ’ 37 ” W, G. Davidse & M. Meadows 35742 (CAS, MO); southern Maya Mts., Bladen Nature Reserve, just S of Upper Bladen Branch and 1.3 km (air) SE of “ AC Camp, ” 16 ° 28 ’ 17 ” N, 088 ° 55 ’ 04 ” W, G. Davidse et al. 35881 (MO); Mira-Mar Hill, Edwards Road beyond Columbia [ca. 16 ° 19 ’ 06 ” N 089 ° 06 ’ 57 ” W], P. Gentle 6355 (F, K, LL, MO, UC, US); Camp 1, 3 – 4 km SE of Union Camp, 16 ° 22 ’ 58 ” N, 089 ° 07 ’ 10 ” W, T. Hawkins 1387 (CAS); NE side of Blue Creek, upstream from village of Blue Creek, 16 ° 14 ’ N, 089 ° 00 ’ W, S. Hill 20270 (MO, NLU, NY, US); SW Maya Mts., Columbia River Forest Reserve, Union Camp, 16 ° 23 ’ N, 089 ° 09 ’ W, B. Holst 4045 (CAS, MO, MEXU); without locale, Peck 780 (K, GH, NY); Columbia Forest Reserve, vicinity of forest camp, ca. 6 mi S of Cabro, in upper Rio Grande drainage area [ca. 16 ° 19 ’ N 089 ° 03 ’ W], G. Proctor 36148 (MO); Fern Hill [ca. 5 miles from Punta Gorda fide Lowden 1970: 850], W. Schipp 1110 (BM, F, G, K, MICH, MO, NY, UC). GUATEMALA. Alta Verapaz: antes de Campur, carretera a Cahabón, [ca. 15 ° 35 ’ 19.06 ” N, 090 ° 02 ’ 46.43 ” W], J. Castillo Mont 1197 (NY); near Finca Sepacuite, O. Cook & R. Griggs 205 (US); planted along street in Cobán, 15 ° 28.517 ’ N, 090 ° 22.349 ’ W, 1230 m, T. Daniel et al. 11335 (CAS, COLO, NY); Sierra de Chamá, along hwy. from Cobán to Chisec, N of Cobán, off of hwy. ca. 1.5 km N of Río Sachichá along rural road, 15 ° 36.019 ’ N, 090 ° 23.333 ’ W, T. Daniel & M. Véliz 11337 (BIGU, CAS, MO, US); Sierra de Chamá, ca. 2 km above Aldea Sehubub (= 7 km from Chiquixji; = 35 km NE of Cobán), 15 ° 35.928 ’ N, 090 ° 17.559 ’ W to 15 ° 36.842 ’ N, 090 ° 17.955 ’ W, T. Daniel et al. 11356 (BR, CAS, F, K, MO, NY); Chucaneb, J. Donnell-Smith 1620 (US); Pansamalá, J. Donnell-Smith 1621 (G, K, US); Sepacuite, G. Goll 155 (US); Sebol, [ca. 15 ° 47 ’ 7.25 ” N, 089 ° 56 ’ 14.34 ” W], C. Lundell 18310 (CAS, F, K, LL, MO, PH, US); road to Finca Trece Aguas, ca. 1.1 mi N of church in Senahú, [ca. 15 ° 25 ’ 01.41 ” N, 089 ° 49 ’ 14.17 ” W], J. Luteyn & F. Almeda 3516 (DUKE, F); trail between Sepacuité and Secanquim, [ca. 15 ° 26 ’ 41.71 ” N, 089 ° 44 ’ 45.44 ” W], W. Maxon & R. Hay 3282 (K, NY, US); near Senahú, W. Maxon & R. Hay 3296 (NY, US); NO de Tactic, 6 km a Estor, A. Molina R. & A. R. Molina 12303 (F); between Sepacuité and Secanquim, H. Pittier 320 (NY, US); Tactic, [ca. 15 ° 19 ’ 44.02 ” N, 090 ° 21 ’ 6.69 ” W], P. Standley 71094 (F); E of Tactic on road to Tamahú, [ca. 15 ° 18 ’ 47.45 ” N, 090 ° 18 ’ 26.68 ” W], P. Standley 71249 (F), 71342 (F); along Río Frio, S of Santa Cruz, P. Standley 90192 (F); between Tatic and divide, on road to Tamahú, P. Standley 90593 (F, US); vicinity of Cobán, [ca. 15 ° 27 ’ 53.66 ” N, 090 ° 21 ’ 2.74 ” W], P. Standley 91236 (F); between Cobán and Chimoté, near Rubeltein, J. Steyermark 44171 (US); Pansamalá, H. von Tuerckheim 856 (NY, US); Cobán, H. von Tuerckheim II- 2033 (NY, US); Cubilquitz, [ca. 15 ° 39 ’ 56.02 ” N, 090 ° 25 ’ 38.84 ” W], H. von Tuerckheim 7936 (K, MO, NY, US); Senahu, Finca Aguas, [ca. 15 ° 24 ’ 53.59 ” N, 089 ° 48 ’ 39.16 ” W], M. Véliz 95.4359 (BIGU, MEXU, MO); near San Juan Chamelco, ca. 15 ° 26 ’ N, 090 ° 16 ’ W, L. Williams & M. Wilson 40437 (F, MEXU); near “ La Presa ” ca. 6 – 8 km SE of Cobán, [ca. 15 ° 28 ’ N, 090 ° 19 ’ W], L. Williams et al. 40323 (F); 1 – 8 km NW of Cobán, [ca. 15 ° 28 ’ 53.47 ” N, 090 ° 22 ’ 55.17 ” W], L. Williams et al. 42031 (BM, F, MO, NY); vicinity of San Juan Chamelco, L. Williams et al. 43189 (F, US); Finca Tres Aguas, C. Wilson 180 (F); vicinity of San Juan Chamelco, [ca. 15 ° 25 ’ 52.07 ” N, 090 ° 20 ’ 10.64 ” W], M. Wilson 40943 (F). Baja Verapaz: bei Parula [Purulhá], [ca. 15 ° 13 ’ 52.37 ” N, 090 ° 13 ’ 27.20 ” W], F. Lehmann 1430 (BM); along Río Frio near San Julían [possibly in Alta Verapaz], L. Williams et al. 43563 (F, MICH). El Progresso: near Finca Caieta, [ca. 14 ° 59 ’ 00.03 ” N, 089 ° 53 ’ 59.98 ” W], J. Steyermark 43768 (F, MICH, US). Huehuetenango: Barillas, Aldea San José Maxbal, trail to Laguna Maxbal, 15 ° 58 ’ N, 091 ° 18 ’ W, M. Christenhusz et al. 5488 (MO); San José Nueva Frontera – Yolnajab, camino hacia Laguna Yolnajab, Nenton, 16 ° 03 ’ 30 ” N, 091 ° 32 ’ 47 ” W, J. Morales & M. García 4377 (MO); Cerro Chiblac, between Finca San Rafael and Ixcán, Sierra de los Cuchumatanes [ca. 15 ° 46 ’ 25 ” N, 091 ° 13 ’ 58 ” W], J. Steyermark 49180 (F, US). Izabal: 25 km SSW of Puerto Barrios, [ca. 15 ° 34 ’ 21.26 ” N, 088 ° 37 ’ 0.05 ” W], D. Gregory 612 (CAS, F, US); Modesto Méndez, [ca. 15 ° 53 ’ 29.49 ” N, 089 ° 13 ’ 46.15 ” W], W. Harmon & J. Fuentes 2089 (MO); Biotopo Chocón Machacas, Livingston, 15 ° 45 ’ 56 ” N, 088 ° 55 ’ 55 ” W, J. Morales 1959 (MO); Río Dulce, 2 – 4 mi W of Livingston, [ca, 15 ° 48 ’ 12.69 ” N, 088 ° 46 ’ 37.09 ” W], J. Steyermark 39562 (F); Chocón River, S. Watson 292 (US). Petén: Dolores, S of village, [ca. 16 ° 30 ’ 51.17 ” N, 089 ° 24 ’ 56.87 ” W], E. Contreras 2097 (LL, MO, US); N of Lake Macanché, foot of Cerro Rocoso at Mancanché [ca. 16 ° 58 ’ 04 ” N 089 ° 38 ’ 20 ” W], E. Contreras 5510 (BM, F, K, MO, NY, WIS, US); La Cumbre, W of KM 145, [ca. 16 ° 04 ’ 51.97 ” N, 089 ° 21 ’ 3.37 ” W], E. Contreras 6646 (F, LL, MO, PH, US); Tikal District, ruins, Naranjo, O. Cook & R. Martin 79 (US); along Río Dulce – Flores highway near KM 327, ca. 10 km NW of San Pedro Cadenas, 15 ° 58 ’ 08.6 ” N, 089 ° 16 ’ 42.7 ” W, T. Daniel & M. Véliz 11255 (BIGU, CAS, MICH, US); Strasse von Flores zur Río Dulce-Brücke, Strecke zwischen Poptún und San Luis, 12 km S von Poptún, H. Kurz & R. Frisch 147 (W), 148 (W); Sayab, Fallabón – Yaxha road, [ca. 17 ° 04 ’ 01.49 ” N, 089 ° 22 ’ 56.17 ” W], C. Lundell 2081 (F, MICH); Arroyo Dolores, ca. 200 m de Dolores, R. Ortíz 1569 (F, MO, NY, US); San Luis, [ca. 16 ° 11 ’ 37.30 ” N, 089 ° 25 ’ 48.59 ” W], P. Ventura 177 (F). Zacapa: Volcán de Monos, [ca. 15 ° 08 ’ 58.95 ” N, 089 ° 30 ’ 18.66 ” W], J. Steyermark 42400 (F, US). Department undetermined: eastern portions of Vera Paz and Chiquimula, S. Watson s. n. (K). HONDURAS. Comayagua: Villa de Taulabé, Quebrada la Caliche, [ca. 14 ° 39 ’ 56.69 ” N, 087 ° 59 ’ 26.20 ” W], C. Alduvín 101 (MO); Siguatepeque, [ca. 14 ° 36 ’ 10.86 ” N, 087 ° 47 ’ 54.39 ” W], J. Edwards P- 589 (F); Quebrada el Caliche, SE de la Villa Taulabé, [ca. 14 ° 41 ’ 02.38 ” N, 087 ° 57 ’ 02.73 ” W], D. Ruiz 138 (NY, TEFH); near Taulabé, P. Standley et al. 7001 (F); Pito Solo, Lago Yojoa, [ca. 14 ° 47 ’ 39.06 ” N, 087 ° 58 ’ 38.47 ” W], J. Valerio R. 2940 (F); near Pito Solo, L. Williams & A. Molina R. 12335 (F); along Lake Yojoa, near Pito Solo, L. Williams & A. Molina R. 17730 (F). Copán: Montaña Los Zapotes, W of Los Zapotes, 18 km N of Copán Ruinas on road to Agua Caliente and 2 – 3 km W of Los Zapotes school, [ca. 14 ° 59 ’ 13.83 ” N, 089 ° 09 ’ 04.57 ” W], T. Hawkins 981 (CAS, MO); Montaña Espiritu Santo, 15 ° 05 ’ N, 088 ° 55 ’ W, T. Hawkins & D. Mejía 185 (EAP, MO). Cortés: entre Pito Solo y Agua Azul, Lago Yojoa, [ca. 14 ° 51 ’ 07.26 ” N, 087 ° 57 ’ 14.09 ” W], A. Molina R. 10619 (EAP, F); camino entre Tapiquilares y Las Crucitas, 20 km S de San Antonio de Cortés, al pie de la montaña La Nieve, [ca. 14 ° 59 ’ 22.86 ” N, 088 ° 02 ’ 19.81 ” W], C. Nelson et al. 8128 (MO). Santa Bárbara: eastern slopes of Cerro Santa Bárbara, [ca. 14 ° 53 ’ 14.92 ” N, 088 ° 07 ’ 01.01 ” W], P. Allen et al. 6047 (CAS, EAP, F, GH, US); Cerro Sta. Bárbara, R. Armour & A. Chable 6047 (BM, F); Punta Gorda, Lago Yojoa, 14 ° 53 ’ N, 088 ° 00 ’ W, R. Evans 1001 (CAS); Punta Gorda, Lago Yojoa, 14 ° 53 ’ N, 088 ° 00 ’ W, R. Liesner 26754 (MO).	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFC55944FF7FFAC204328308.taxon	description	Trees to 10 m tall. Older (woody) stems subquadrate, irregularly fissured-striate; younger (herbaceous) stems stubby, quadrate, glabrous. Leaves deciduous, clustered at apex of old growth, petiolate, petioles to 73 mm long, blades membranaceous, ovate to elliptic, 90 – 220 mm long, 35 – 100 mm wide, 2 – 2.7 × longer than wide, acute to acuminate at apex, rounded to attenuate and asymmetric at base, surfaces glabrous, margin entire. Inflorescence of axillary and / or terminal subsessile to pedunculate thyrses (main axis with lateral pedunculate dichasia appearing as branches) to 250 mm long, peduncle to 50 mm long, pubescent with erect to flexuose eglandular trichomes 0.05 – 0.2 (– 1) mm long, rachis pubescent with erect to flexuose eglandular trichomes 0.1 – 0.4 (– 1) mm long; dichasia opposite, pedunculate, 1 – 3 (or more) - flowered, to 63 mm long (excluding corollas), dichasial peduncles 3 – 20 mm long, pubescent like rachis. Bracts caducous, lance-elliptic to elliptic, 12 – 17 mm long, 3.5 – 5 mm wide, abaxially pubescent with erect to flexuose eglandular trichomes 0.05 – 0.2 mm long. Bracteoles and secondary bracteoles usually present on young inflorescences but becoming deciduous as inflorescence matures, lance-elliptic to elliptic, 7 – 14 mm long, 2 – 3. 5 mm wide, abaxially pubescent like bracts. Flowers pedicellate, pedicels 10 – 50 mm long, pubescent like rachis. Calyx 17 – 28 mm long, lobes subheteromorphic, membranaceous, abaxially pubescent with erect (to ± antrorse) eglandular trichomes 1 – 2 (– 2.3) mm long, posterior lobe planar to ± conduplicate, ovate-elliptic, 17 – 25 mm long, 11 – 15 mm wide, usually slightly larger than lateral lobes, acuminate at apex, lateral lobes ovate-elliptic to elliptic to obovate, 15. 5 – 22 mm long, 9 – 15 mm wide, rounded to acute at apex, veins of lobes prominent (ca. 9 major veins). Corolla “ pale green ” to greenish yellow, 50 – 55 mm long, externally glabrous, tube 29 – 33 mm long, narrow proximal portion 9 – 13 mm long, 9 – 16 mm in diameter near midpoint, throat 15 – 20 mm long, 30 – 35 mm in diameter at mouth, lobes spreading to recurved, “ narrowly ovate to subcircular ” (fide Miranda and McVaugh 1962), 18 – 24 mm long, 17 – 27 mm wide, round- ed to acute at apex. Stamens 4, 72 – 90 mm long, filaments glabrous distally, pubescent proximally with eglandular trichomes, thecae 10 – 12 mm long; staminode (if present) not seen. Style 50 – 55 mm long, glabrous, stigma lobes 1.5 mm long, shape and width not determined. Capsule 25 – 30 mm long, 7 – 10 mm in diameter, pubescent throughout with erect to flexuose to antrorse eglandular trichomes 0.05 – 0.2 mm long, stipe 1.5 – 2 mm long. Seeds up to 16 per capsule, 7 – 8 mm long, 5 – 6 mm wide, surfaces minutely papillose. PHENOLOGY. — Flowering: December – February; fruiting: February.	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFC55944FF7FFAC204328308.taxon	distribution	DISTRIBUTION AND HABITAT. — West-central Mexico (Jalisco, Michoacán; Fig. 8); plants occur on limestone slopes in “ high forest dominated by Brosimum ” in regions of tropical subdeciduous and deciduous forests at elevations from 500 to 1400 meters. ILLUSTRATIONS. — Figure 14; Miranda and McVaugh (1962: 183, fig. 2). CONSERVATION. — Louteridium koelzii is currently known from a very limited region of western Mexico (EOO = 5.3 km 2; AOO = 12 km 2). The species is not known to occur in a protected area, and it has not been recollected since 1975 (i. e., McVaugh 26180; although see discussion below about an undetermined plant from a nearby region of Jalisco). Based on current knowledge, there appear to be two subpopulations about 80 km apart, one in Jalisco and one in Michoacán. Comparing historic landsat images (2006 to 2017) via Google Earth Pro (2018) in the region of the known collections from Jalisco, it is evident that deforestation for agriculture has reduced the natural vegetation there substantially (ca. ≥ 30 %), thus creating a threat to this species, at least in that region. No specific threat has been identified for the subpopulation in Michoacán. Thus, there are two locations for the species, and a threat is evident for one of them. Based on these observations, a preliminary conservation assessment of Endangered (EN; B 1, a, b) is proposed for L. koelzii.	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFC55944FF7FFAC204328308.taxon	discussion	DISCUSSION. — An undetermined specimen of Louteridium from western Mexico (Jalisco: Mpio. Jilotlán de los Dolores, Cerro de San Antonio, 15 km N de Tazumbos, matorral subtropical en suelos con pedregocidad de toba volcánica, 900 m, 6 Feb 1982, F. Santana Michel 1081, IBUG) does not seem to pertain to either species known from nearby regions (i. e., L. brevicalyx or L. koelzii) or other species of the genus. The plant, which lacks mature flowers, is geographically closest and morphologically most similar to L. koelzii. Like that species, it occurs in an apparently dry habitat and has seasonally deciduous foliage. Both of these characteristics suggest that the undetermined plant has four stamens. Other characteristics in common with L. koelzii include erect to flexuose eglandular trichomes on the rachis, peduncles, and pedicels; ovate-elliptic calyx lobes, and externally glabrous corollas. It differs from specimens of L. koelzii studied herein by having pubescent young stems, shorter calyces (14 – 15 mm long, although the flowers are not fully mature), and longer (35 – 40 mm) capsules that are pubescent only at the apex. Disposition of this plant awaits further studies and / or collections. ADDITIONAL SPECIMENS EXAMINED. — MEXICO. Jalisco: ca. 12 – 13 km SW of Pihuamo, R. McVaugh 26180 (MICH); 8 mi SW of Pihuamo, R. McVaugh & W. Koelz 1797 (MEXU, MICH, US). Michoacán: 15 – 16 km SE of Aserradero Dos Aguas, W of Aguililla [ca. 18 ° 45 ’ 51.11 ” N, 102 ° 49 ’ 22.26 ” W], R. McVaugh 24740 (MICH).	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFC6595FFF7FFD8F06FE8767.taxon	description	Louteridium conzattii Standl., Contr. U. S. Natl. Herb. 23: 1338. 1926. TYPE. — MEXICO. Oaxaca: Distr. Tuxtepec, de Jalapa a La Raya, margen izquierda del Río Grande, [ca. 18 ° 01 ’ 16.00 ” N, 096 ° 34 ’ 25.98 ” W], 450 m, 3 - XI- 1919 (buds), C. Conzatti 3788 (holotype: US- 1014038!; isotypes: MEXU! US- 1014039!). Shrubs to trees to 12 m tall, terrestrial. Older (woody) stems subquadrate, irregularly striate-fissured, glabrous or glabrate; younger (herbaceous) stems quadrate to quadrate-sulcate, internodes glabrous or with sparse flexuose to antrorse eglandular trichomes to 0.5 mm long, nodes glabrous or with some flexuose eglandular trichomes to 0.5 mm long. Leaves not all seasonally deciduous, evenly distributed on new growth, petiolate, petioles to 220 mm long, blades membranaceous, broadly ovate to ovate to ovate-elliptic to elliptic to broadly elliptic, 40 – 360 mm long, 21 – 170 mm wide, 1.3 – 2.9 × longer than wide, acuminate to acute-apiculate to caudate at apex, acute to round- ed to subcordate at base, adaxial surface glabrous, abaxial surface glabrous or pubescent along major veins with trichomes like those at nodes, margin ± entire to crenate. Inflorescence a terminal pedunculate (racemose thyrse to) thyrse to 390 mm long (including peduncle and excluding corollas), peduncle to 190 mm long, glabrous or pubescent like young stems, rachis glabrous or pubescent with flexuose to flexuose-antrorse eglandular trichomes 0.2 – 0.8 (– 1) mm long (villous), trichomes sometimes restricted to nodes; dichasia modified by sympodial expansion into contract- ed to ± expanded unbranched or branched short-shoots to 25 (– 46) mm long, opposite, subsessile to pedunculate, 3 – many-flowered, to 120 mm long (excluding corollas), dichasial peduncles 1 – 22 mm long, glabrous or villous. Bracts often caducous, sessile to petiolate, ovate to lance-elliptic to linear, 20 – 48 mm long, 2 – 16 mm wide (or the proximalmost pair sometimes foliose and larger), abaxially glabrous or pubescent with antrorsely appressed eglandular trichomes. Bracteoles and secondary bracteoles usually persistent, lance-ovate to lanceolate to linear-lanceolate, 10 – 25 mm long, 1 – 8 mm wide, abaxially glabrous or pubescent with a few antrorsely appressed eglandular trichomes to 0.5 mm long along midvein or pubescent with latter type trichomes over entire surface. Flowers pedicellate, pedicels to 85 mm long, glabrous or villous. Calyx 17 – 39 mm long, lobes appearing distinct or fused at base for 1 – 2 mm, heteromorphic, ± membranaceous, abaxially appearing glabrous although covered with minute sessile glands (glandular punctate; punctations <0.05 mm in diameter) and posterior lobe sometimes sparsely villous along keel, posterior lobe conduplicate, ovate, 16 – 36 mm long, ± equaling to slightly longer than lateral lobes, 9 – 14 mm wide, acute to attenuate to subfalcate at apex, lateral lobes lunate, 16 – 32 mm long, 6 – 12 mm wide, rounded to acute to subacuminate at apex. Corolla usually greenish and maroon (also described as white with purplish lobes and pale rose in plants from San Felipe Usila, Oaxaca; see discussion below), 38 – 54 mm long, externally puberulent with sessile to subsessile glands 0.05 – 0.1 mm long and often with glandular and eglandular trichomes to 0.2 mm long as well, tube (19 –) 30 – 35 (– 42) mm long, narrow proximal portion 8 – 14 mm long, 5 – 10 mm in diameter near midpoint, throat 15 – 23 mm long, 24 – 33 mm in diameter at mouth, lobes recoiled, subtriangular to broadly ovate to elliptic, 10 – 16 mm long, 8 – 14 mm wide, rounded at apex. Stamens 2, 60 – 77 mm long, filaments glabrous distally, pubescent proximally with eglandular trichomes, thecae 9 – 13 mm long; staminodes 2, 1 – 3 mm long. Style 72 – 85 mm long, glabrous (or sometimes with sessile to subsessile glands to 0.05 mm long near base), stigma subequally to unequally 2 - lobed, lobes linear-elliptic to elliptic to obovate-elliptic, 1 – 3. 8 mm long, 0.6 – 1.5 mm wide. Capsule 16 – 24 mm long, 4 – 8 mm in diameter, sparsely puberulent with sessile to subsessile glands to 0.05 mm long, stipe 1 – 2 mm long. Seeds up to 16 per capsule, 4 – 5. 5 mm long, 3.5 – 5 mm wide, surfaces smooth or covered with subconic papillae. PHENOLOGY. — Flowering: September – May; fruiting January – May. Based on greenhouse observations of Breedlove & Daniel 70879 gh, corollas open after 17: 00 and all fall by 11: 00 the next morning.	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFC6595FFF7FFD8F06FE8767.taxon	distribution	DISTRIBUTION AND HABITAT. — Southern Mexico (Chiapas, Oaxaca, Puebla, Tabasco, Veracruz) and Guatemala (Alta Verapaz, Petén) (Fig. 11); plants occur on slopes and along watercourses on limestone (often karstic) in primary or secondary lowland rain forests (selva alta perennifolia), montane rain forests, cloud forests, and evergreen seasonal forests (including selva mediana subperennifolia) at elevations from 50 to 1200 m. The distribution of this species appears to be entirely on the versants of the Gulf of Mexico and the Caribbean Sea. Daniel et al. (2012) report- ed this species from Guatemala. Plants from Guatemala previously attributed to L. mexicanum pertain to L. purpusii (see discussion in Daniel 1995). LOCAL NAMES. — “ K’o’och batz’ ” (Paniagua 539); “ oj-oj ” (Kunkel 161); “ palo aguañoso ” (Ramos & Martínez S. 620); “ sabal tzununte ” (Palacios E. 9631); “ Uc’o’och chuba’atsir wis ” (Levy T. & Durán F. 398). USES. — Leaves applied against the body for fever (Wendt et al. 3614); a drink made from the boiled leaves is used for “ dolor orinar ” (Paniagua 539). ILLUSTRATIONS. — Figures 4 E, 15. CONSERVATION. — Louteridium mexicanum is the most commonly collected species in Mexico, and has an EOO of 35,531 km 2. At least nine protected areas are located entirely or partially within its EOO, and the species has been collected in two of them. Frequency data from herbarium collection labels varies from scarce to infrequent to common to abundant. Although unidentified threats are undoubtedly present in portions of its geographic distribution, a preliminary conservation assessment of Least Concern (LC) is proposed for this species.	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFC6595FFF7FFD8F06FE8767.taxon	discussion	DISCUSSION. — Daniel (1995) discussed the probable state from which the type was collected; this would appear to be Veracruz rather than Chiapas. Linden collected in both states (Ossenbach 2009), but no “ Zacualpan ” has been located in Chiapas (D. Breedlove, personal communication in 1994). Zacualpan (or Zacuapan or Zacuapam [ca. 19 ° 12 ’ 46.44 ” N, 096 ° 51 ’ 8.73 ” W], see Sousa S. 1969; not the town of Zacualpan at 20 ° 26 ’ 1.71 ” N, 098 ° 20 ’ 57.69 ” W in northwestern Veracruz), Carl Sartorius’ large hacienda located between Jalapa and Córdoba in west-central Veracruz, with its headquarters at El Mirador, is known to have been visited by several early Mexican collectors, including Linden (Ossenbach 2009). Because it is ca. 42 km north of the currently known distributional range of L. mexicanum (Fig. 11), and because of the ambiguity of the locality based on Linden’s indication of Chiapas on the collection labels, Zacuapam (shown in Fig. 11 with a question mark) is not included in the calculation of the EOO for the species. Based on living flowers of Breedlove & Daniel 70879 cv, the coloration of corollas of this species can be more fully described as follows: the narrow proximal portion of the tube (covered by the calyx) is green; the throat is green with maroon veins and tinged with maroon between the major veins; the lips (with recoiled lobes) are mostly maroon. Daniel (1995) noted considerable variation in pubescence, leaf shape, bract persistence, and bract shape among specimens from Chiapas, Mexico. One of the two collections (Breedlove 24803, Palacios E. 385) that he indicated showed some intermediacy with L. donnell-smithii in pubescence was sampled in the phylogenetic analysis. Palacios E. 385 was strongly supported as nested within the clade of other sampled plants of L. mexicanum (Fig. 1). Thus, both plants are here included within L. mexicanum. Somewhat atypical plants from the vicinity of San Felipe Usila, Oaxaca (e. g., Ibarra M. et al. 3758, 3765; Calzada et al. 16579) are distinctive by their long calyces (30 – 39 mm) with the posterior lobe conspicuously attenuate-falcate apically, corollas pale rose to white with purplish lobes and pubescent with glandular and eglandular trichomes to 0.2 mm long, and rachis (i. e., Calzada et al. 16579) often somewhat scurfy to minutely puberulent with sessile to subsessile glandular trichomes less than 0.05 mm long. In Calzada et al. 16579, the calyx and pedicels are like more typical specimens of L. mexicanum, and the rachis often has longer eglandular trichomes to 0.1 mm as well. Although similar calyces and trichomes on corollas are sometimes seen elsewhere among specimens of L. mexicanum, the corolla color of these collections appears to be unusual compared to other Mexican representatives of the species. However, corollas of Foerther et al. 10940 from Guatemala are described as brownish violet. In pubescence of the rachis, these plants somewhat resemble L. purpusii. On the whole, they appear more like L. mexicanum, in which species they are treated here. ADDITIONAL SPECIMENS EXAMINED. — MEXICO. Chiapas: Mpio. Ocosingo, 4.4 km SO de San Javier, 16 ° 48 ’ 49 ” N, 091 ° 04 ’ 04 ” W, G. Aguilar M. et al. 4713 (MEXU); Mpio. Solosuchiapa, 2 – 4 km below Ixhuatán toward Pichucalco, [ca. 17 ° 18 ’ 47.23 ” N, 093 ° 00 ’ 11.57 ” W], D. Breedlove 24177 (DS, ENCB, MO, NY), 29799 (DS, LL, MEXU, MICH, MO, RSA), D. Breedlove & F. Almeda 56805 (CAS, ENCB, LL, NY); Mpio. Berriozábal, 13 km N of Berriozábal near Pozo Turripache and Finca El Suspiro, [ca. 16 ° 52 ’ 25.01 ” N, 093 ° 19 ’ 8.64 ” W], D. Breedlove 24803 (DS, ENCB, MO), D. Breedlove & T. Daniel 70879 (CAS), cultivated plants of this collection grown from cuttings in San Francisco, California, 70879 cv (CAS), D. Breedlove & R. Thorne 30786 (DS, ENCB, MO), D. Breedlove et al. 66158 (CAS); Mpio. Palenque, near Agua Azul, D. Breedlove 56091 (CAS, LL, MEXU); Mpio. Ocosingo, near ruins of Bonampak at jct. road from Lacanjá to Echeverría and road from Chancalá, [ca. 16 ° 48 ’ 04.57 ” N, 091 ° 06 ’ 21.97 ” W], D. Breedlove & F. Almeda 58020 (CAS, LL, MEXU, MO); Mpio. San Fernando, 20 km NW of San Fernando just beyond Colonia Cuahtémoc, [ca. 16 ° 58 ’ 42.74 ” N, 093 ° 20 ’ 28.46 ” W], D. Breedlove & M. Bourell 68269 (CAS); Mpio. Ocosingo, 70 km SW of Palenque toward Ocosingo along the Jol Uk’um, [ca. 17 ° 10 ’ 48.02 ” N, 092 ° 06 ’ 39.86 ” W], D. Breedlove & B. Keller 49526 (CAS, MEXU, MO); Mpio. Ocozocoautla de Espinosa, 26 – 28 km N of Ocozocoautla toward Mal Paso, [ca. 16 ° 56 ’ 22.87 ” N, 093 ° 27 ’ 55.46 ” W], D. Breedlove & A. Smith 22469 (DS, MEXU, MICH, MO, RSA); Mpio. Ocosingo, 4 km E del crucero San Javier, 16 ° 48 ’ 44 ” N, 091 ° 03 ’ 13 ” W, J. Calónico S. et al. 25022 (ENCB, MEXU); Mpio. Berriozábal, ca. 12 km NW of Berriozábal, ca. 16 ° 51 ’ N, 093 ° 19 ’ W, T. Daniel 8375 (CAS, MEXU); Mpio. Ishuatán, carretera 195, 45 km N [S] de Pichucalco, [ca. 17 ° 18 ’ 26.14 ” N, 093 ° 00 ’ 27.05 ” W], S. Gliessman 77 - 7 (MEXU); Mpio. Pichucalco, 20 km Teapa – Pichucalco, [ca. 17 ° 32 ’ 45.83 ” N, 093 ° 04 ’ 44.80 ” W], M. Ishiki et al. 2035 (CAS); Mpio. Ocosingo, Lacandona de Lacanhá – Chansayab, 130 km SE de Palenque, por la carretera fronteriza hasta el crucero San Javier, después 8 km hacia el oeste, 16 ° 44 ’ N, 091 ° 05 ’ W, S. Levy T. & A. Durán F. 398 (MEXU); Mpio. Berriozábal, 12 km N de Berriozábal, E. Martínez S. & M. Soto 24216 (MEXU); Mpio. Ishuatán, 9 km N de Tapilula, [ca. 17 ° 17 ’ 16.71 ” N, 093 ° 00 ’ 38.70 ” W], E. Martínez S. et al. 3198 (CAS, ENCB, MEXU, MO, WIS); Mpio. Yajalón, Yajalón – Petalsinco, carretera de terraceria KM 5, A. Méndez T. [= A. Ton] 7134 (IEB, MEXU, MO, RSA, XAL); Mpio. Ocozocoautla, 9 km NW de Emilio Rabasa, 500 m W de El Aguajito, 16 ° 57 ’ 32 ” N, 093 ° 40 ’ 31 ” W, S. Ochoa G. 3860 (XAL); Mpio. San Fernando, cerros adelante de Col. Cuauhtémoc, [ca. 16 ° 56 ’ 39.34 ” N, 093 ° 13 ’ 33.83 ” W], E. Palacios E. 385 (CAS); Mpio. Ocozocoautla, Reserva El Ocote, al O mas adelante de la caseta, E. Palacios E. 9631 (CAS); Mpio. San Fernando, 1.4 km N de Cuauhtémoc, 16 ° 56 ’ 32 ” N, 093 ° 13 ’ 30 ” W, R. Palestina 1747 (XAL); Mpio. Ocosingo, Lacanjá – Chansayab, Vereda de Pancho López Kin, B. Paniagua 539 (MEXU); Mpio. Berriozábal, 13 km N de Berriozábal, O. Téllez V. et al. 7591 (MEXU); Mpio. Yajalón, Yajalón – Petalzinco, [ca. 17 ° 12 ’ 13.88 ” N, 092 ° 23 ’ 17.85 ” W], A. Ton 7134 (XAL); Mpio. Tila, Finca Morelia, [ca. 17 ° 17 ’ 13.71 ” N, 092 ° 26 ’ 08.79 ” W], A. Ton 7427 (MEXU). Oaxaca: Mpio. San Pedro Ixcatlán, Cerro Camarón, al W de Cerro Quemado, J. Calzada 10293 (XAL); Mpio. Chiltepec, Ejido de Rancho Faisán, vereda para La Rejoya, [ca. 17 ° 56 ’ 57.43 ” N, 096 ° 09 ’ 47.56 ” W], J. Calzada 14456 (CAS, MEXU); Mpio. San Miguel Soyaltepec, Cerro Tepezcuintle, J. Calzada & S. Anta 17515 (MEXU); Mpio. San Felipe Usila, 2 km del poblado Cerro Verde, carretera al campamento Arroyo Tambor, J. Calzada et al. 16579 (MEXU); Distr. Tuxtepec, Presa Temazcal, camino Temazcal – Vertedor a 6 km, L. Cortes & R. Torres C. 66 (ENCB, MEXU); Mpio. Soyaltepec, 2 km de la Hidroelectrica al Vertedor de la Presa Temascal, L. Cortes & R. Torres C. 617 (MEXU); Distr. Tuxtepec, Mpio. Acatlán, Cerro Buenos Aires, lado oeste de la Presa Temazcal (Miguel Alemán), L. Cortes et al. 165 (ARIZ, CAS, MEXU); Uxpanapa Region, E of Sarabia toward Uxpanapa, 4.2 mi NE of Río Corte, 17 ° 09 ’ N, 095 ° 15 ’ W, T. Croat & D. Hannon 63176 (CAS, DUKE, MEXU, MO, TEX); Mpio. San Miguel Soyaltepec, Cerro Tepezcuintle, 18 ° 09 ’ 34.1 ” N, 096 ° 20 ’ 50.5 ” W, C. Cruz E. & G. Juárez G. 2599 (MEXU, SERO); Mpio. Santa María Chimalapa, Matías Romero, Colonia Cuauhtémoc, [ca. 17 ° 06 ’ 06.64 ” N], A. Delgado S. 942 (ENCB, MEXU, US); Mpio. San Felipe Usila, Cerro Verde, [ca. 17 ° 53 ’ 13.44 ” N, 096 ° 30 ’ 05.71 ” W], A. Hanan A. & R. de Santiago 854 (MEXU, SERO); Mpio. Santa María Chimalapa, ca. 10 km O de Sta. María por la vereda a Chicosaja, 16 ° 53 ’ N, 094 ° 46 ’ W, H. Hernández G. 1054 (CAS, ENCB, MEXU, MO, XAL); Mpio. San Felipe Usila, Cerro Verde, 8 km NNE of San Felipe Usila, 17 ° 57 ’ N, 096 ° 30 ’ W, G. Ibarra M. et al. 3758 (BM, MEXU, MO, SERO); Mpio. San Felipe Usila, Nueva Santa Flora, 17 ° 55 ’ N, 096 ° 26 ’ W, G. Ibarra M. et al. 3765 (BM, MEXU, MO, SERO); Distr. Tuxtepec, Mpio. Ayotzintepec, 3.7 km NE de Ayotzintepec, Cerro Cueva de los Duendes, 17 ° 42 ’ 26 ” N, 096 ° 07 ’ 59 ” W, G. Juárez G. 3771 (SERO); Mpio. Santa María Tlalixtac, Río Cóndor, brecha entre Santa María Tlalixtac y Chiquihuitlán de Benito Juárez, 17 ° 57 ’ 14.5 ” N, 096 ° 43 ’ 4.5 ” W, G. Juárez G. & C. Cruz E. 937 (SERO, MEXU); Mpio. San Miguel Soyaltepec, Cerro Tepezcuintle, Papaloapam, 18 ° 08 ’ 46.6 ” N, 096 ° 21 ’ 37.3 ” W, G. Juárez G. & A. Martínez F. 788 (SERO, MEXU); Mpio. Matías Romero, Río Coatzacoalcos at Cuauhtémoc, 18 km ESE of Palomares, 17 ° 06 ’ N, 094 ° 53 ’ W, M. Nee 29710 (F, NY, XAL); Distr. Tuxtepec, “ Cerro de la Cruz ” near Temascal, E arm of Presa Alemán, 18 ° 13 ’ N, 096 ° 24 ’ W, D. Neill 5399 (CAS, MEXU, MO); Mpio. Santa María Jacatepec, predio La Joya del Obispo, [ca. 17 ° 51 ’ 33.80 ” N, 096 ° 11 ’ 53.88 ” W], C. Ramos & E. Martínez S. 620 (MEXU); Santa María Chilchotla, Cuahtémoc, ca. 4 km NE of Santa María Chilchotla, near Clemencia and Sta. Rosa, 18 ° 14 ’ N, 096 ° 50 ’ W, S. Solheim & S. Reisfield 1363 (MEXU, NY, XAL, WIS); Mpio. San Pedro Ixcatlán, Sierra Mazateca, Cerro Camarón, 1 km E of village of Cerro Quemado, 18 ° 09 ’ N, 096 ° 35 ’ W, S. Solheim & S. Reisfield 1499 (MEXU, XAL, WIS); Distr. Tuxtepec, Presa Miguel Alemán, Cerro de la Cruz, [ca. 18 ° 12 ’ 20.03 ” N, 096 ° 25 ’ 35.09 ” W], R. Torres C. et al. 2133 (MO); Distr. Tuxtepec, Mpio. Santa María Jacatepec, camino a Cosolapa San Antonio, Ejido de San Felipe Tilpa, 13.3 km SO de La Reforma, 17 ° 51 ’ N, 096 ° 03 ’ W, R. Torres C. & L. Cortes A. 11480 (CAS, MEXU); Mpio. Santa María Jacatepec, bajada del predio El Aguila a San Agustín, entrada por La Reforma, 28 km SW de Tuxtepec, carr. a Matías Romero, 17 ° 50 ’ N, 96 ° 06 ’ W, R. Torres C. & E. Martínez S. 11093 (IEB, MEXU, MO, TEX); Mpio. Matías Romero, 7.2 km O de Esmeralda, 17 ° 08 ’ N, 094 ° 50 ’ W, T. Wendt et al. 3614 (ARIZ, CAS, ENCB, LSU, MEXU, MO, NY, TEX, XAL). Puebla: Mpio. San Sebastián Tlacotepec, 18 ° 26 ’ 12 ” N, 096 ° 50 ’ 31.8 ” W, L. Caamaño O. 6602 (HUAP-image seen). Tabasco: Mpio. Teapa, 0.34 km E de la Univ. Aut. Chapingo, 17 ° 31 ’ 31 ” N, 092 ° 55 ’ 33 ” W, J. Calónico S. et al. 21459 (BM, MEXU); Mpio. Teapa, Cerro del Madrigal, 3 km E de Teapa, camino a Talcotalpa, 17 ° 35 ’ 42 ” N, 092 ° 55 ’ 26 ” W, R. Fernández N. 2214 (ENCB, RSA); Mpio. Teapa, Sierra El Madrigal, Centro Regional Tropical Puyacatengo de la Univ. Autónoma de Chapingo, 17 ° 31 ’ 02 ” – 17 ° 32 ’ 30 ” N, 092 ° 54 ’ 10 ” – 092 ° 56 ’ W, A. Hanan A. et al. 326 (MEXU); Mpio. Tacotalpa, 3.2 km después de cruzar la panga del Río Oxiolotán, yendo de Tapijulapa a Oxoiotán, 17 ° 26 ’ 30 ” N, 092 ° 45 ’ 30 ” W, S. Hernández & A. Espejo 139 (ENCB, IEB, MEXU, MO, XAL); Mpio. Macuspana, Parque Nac. Agua Blanca, KM 64 carretera Villahermosa – Escárcega, 17 ° 38 ’ N, 092 ° 30 ’ W, M. López P. et al. 3 (ENCB, XAL); Mpio. Teapa, KM 6 de la entrada al ejido La Unión hacia Ixtapangajoya, [ca. 17 ° 33 ’ 34 ” N, 092 ° 57 ’ 39 ” W], M. Magaña & A. Suárez F. 724 (ENCB, MEXU, XAL); Mpio. Macuspana, Centro Recreativo Agua Blanca, [ca. 17 ° 41 ’ 15 ” N, 092 ° 27 ’ 42 ” W], M. Magaña et al. 1198 (ENCB, MEXU, XAL); Mpio. Macuspana, Parque Nac. Agua Blanca, KM 64 carretera Villahermosa – Escárcega, 17 ° 38 ’ N, 092 ° 30 ’ W, L. Martínez G. 12 (ENCB, MEXU); Mpio. Teapa, Cerro Madrigal, 500 m E de Puyacatengo, Univ. Aut. Chapingo, 17 ° 31 ’ 34 ” N, 092 ° 55 ’ 17 ” W, E. Martinez S. et al. 34736 (BM, MEXU); Mpio. Teapa, 1 km NE de Puyacatengo, 17 ° 31 ’ 39 ” N, 092 ° 55 ’ 22 ” W, E. Martínez S. et al. 34899 (BM, BR, CAS, FLAS, MEXU, PH); Mpio. Teapa, El Azufre, 16 km E de Teapa, 17 ° 35 ’ N, 092 ° 50 ’ W, P. Tenorio L. et al. 5577 (ARIZ, MEXU, RSA, TEX); Mpio. Macuspana, Agua Blanca, F. Ventura A. 20941 (CAS, ENCB, IEB, MEXU, NY); Mpio. Teapa, El Madrigal, 17 ° 35 ’ 07 ” N, 092 ° 56 ’ 21 ” W, F. Ventura A. 20989 (ENCB, MEXU, NY, XAL); Mpio. Macuspana, Agua Blanca, S. Zamudio 183 (XAL). Veracruz: Mpio. Cuichapa, carretera de terracería entre Cuichapa y Omealca [ca. 18 ° 45 ’ 51.96 ” N, 096 ° 49 ’ 53.57 ” W], J. Calzada & A. Vovides 2396 (ENCB, F, MEXU, MO, XAL); Mpio. Jesús Carranza, KM 6 del camino Cedillo – Río Alegre, 17 ° 10 ’ N, 094 ° 40 ’ W, B. Dorantes 3917 (ENCB, MEXU, XAL); Mpio. Hidalgotitlán, kms 0 – 2 del camino Plan de Arroyos – Alvaro Obregón, 17 ° 15 ’ N, 094 ° 40 ’ W, J. Dorantes et al. 2801 (ENCB, F, NY, XAL); Mpio. Hidalgotitlán, 0 – 2 km del Campo Cedillo, rumbo a Río Alegre, 17 ° 19 ’ N, 094 ° 40 ’ W, J. Dorantes et al. 2936 (F, XAL); Mpio. Jesús Carranza, KM 10 del camino Cedillo – Fco. Villa, 17 ° 10 ’ N, 094 ° 40 ’ W, J. Dorantes et al. 3938 (ENCB, F, MEXU, MO, XAL); Mpio. Hidalgotitlán, camino Cedillo – La Laguna, J. Dorantes et al. 4138 (F, MEXU, XAL); Mpio. Tezonapa, 5 – 6 km SE de Motzorongo, 18 ° 40 ’ N, 096 ° 40 ’ W, R. Gobles G. 208 (XAL), 299 (XAL); Mpio. Hidalgotitlán, 3 km SW of Campamento La Laguna, 17 ° 16 ’ N, 094 ° 32 ’ W, M. Nee 29995 (F, MEXU, NY, XAL); Mpio. Amatlán de los Reyes, Cerro de Amatlán [ca. 18 ° 49 ’ 53.93 ” N, 096 ° 54 ’ 3.08 ” W], H. Oliva & V. Villar 624 (IBUG, XAL); Mpio. Coetzala, 1 km E de Coetzala, 18 ° 46 ’ 50 ” N, 096 ° 54 ’ 52 ” W, A. Rincón G. & C. Durán E. 1508 (MEXU, XAL); Mpio. Hidalgotitlán, Río Soloxuchil, entre Hnos. Cedillo y La Escuadra, 17 ° 17 ’ N, 094 ° 38 ’ W, M. Vásquez et al. 322 (F); Mpio. Hidalgotitlán, brecha Hnos. Cedillo – La Laguna, B. Vázquez 17 (F, MEXU, NY, XAL); Mpio. Jesús Carranza, 3 km E del Río Chalchijapa por la carretera Sarabia – Cedillo, M. Vázquez T. 1585 (XAL); Mpio. Jesús Carranza, 2 km N del Poblado 2, Ejido F. J. Mina, 17 ° 16 ’ N, 094 ° 40 ’ W, M. Vázquez T. et al. V- 2568 (CAS, MEXU, XAL); Chicomapa [ca. 18 ° 33 ’ 39, 096 ° 51 ’ 08 ” W], V. Vázquez T. 548 (ENCB, F, IEB, MEXU, XAL). GUATEMALA. Alta Verapaz: Mpio. Rubeltein, Montaña Sacranix, Wegstrecke zwischen der Finca Xalcata (= Saqmoc) und der Finca Sacté, S des Río Sachichaj (W der Strasse Cobán – Chisec), H. Foerther et al. 10940 (BM, W); Sacté, 15 ° 30 ’ N, 090 ° 27 ’ W, I. Kunkel 161 (BR).	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFDC5953FF7FFF6D02458028.taxon	description	Shrubs or trees to 3 m tall, epiphytic or epipetric. Older (woody) stems subquadrate to subterete, lenticellate, lacking trichomes; younger (herbaceous) stems quadrate-sulcate, glabrous. Leaves not all seasonally deciduous, evenly distributed along stems, petiolate, petioles to 20 mm long, blades coriaceous-subsucculent, oblanceolate to obovate, 50 – 200 mm long, 15 – 63 mm wide, 3.3 – 5 × longer than wide, acute-apiculate (to caudate) at apex, attenuate to long-attenuate at base, adaxial surface glabrous, abaxial surface with scattered flexuose to antrorse eglandular trichomes to 1 mm long (or sometimes nearly glabrous), margin entire. Inflorescence a terminal pedunculate thyrse to 27 cm long (including peduncle and excluding corollas), peduncle 3. 5 – 15 cm long, glabrous, rachis glabrous; dichasia modified by sympodial expansion and appearing like dichotomous branches with zig-zag nodes, alternate, pedunculate, 1 – many-flowered, to 100 mm long (excluding corollas), dichasial peduncles to 55 mm long, glabrous. Bracts caducous, lance-ovate to elliptic, 7 – 15 mm long, 4 – 8 mm wide, abaxially glabrous. Bracteoles and secondary bracteoles persistent, ovate to ovate-elliptic to broadly elliptic, 6 – 11 mm long, 5 – 7. 5 mm wide, abaxially glabrous (or inconspicuously glandular-punctate), those of a pair connate along one side for up to 1 / 2 their length and proximally subsaccate, sometimes conspicuously imbricate. Flowers pedicellate, pedicels to 30 mm long, glabrous. Calyx 24 – 39 (– 48 in fruit) mm long, lobes heteromorphic, apparently subsucculent, abaxially glabrous, posterior lobe conduplicate, oblong to subelliptic to oblanceolate, 22 – 45 mm long, longer than lateral lobes, 7. 5 – 14 mm wide, saccate or with a flap-like appendage to 2.5 mm long at base, rounded to acute at apex, lateral lobes lance-ovate to lance-linear to linear-oblong, 19 – 44 mm long, 5 – 8. 8 mm wide, acute at apex. Corolla pale greenish, 45 – 72 mm long, externally glabrous, tube 25 – 35 mm long, narrow proximal portion of tube 0.5 – 3 mm long, 6 – 8. 5 mm in diameter near midpoint, throat 24 – 33 mm long, 28 – 40 mm in diameter at mouth, lobes spreading to recurved, subtriangular to broadly ovate, 8 – 21 mm long, 10 – 17 mm wide, rounded and emarginate at apex. Stamens 2, 70 – 90 mm long, filaments glabrous distally, densely pubescent at base with eglandular trichomes, thecae 13 – 17 mm long; staminodes (if present) not seen. Style 74 – 90 mm long, glabrous, stigma unequally 2 - lobed, lobes ovate to linear-elliptic to linear-oblanceolate, 2. 5 – 6 mm long, 1.1 – 2. 8 mm wide. Capsule 27 – 40 mm long, 5.5 – 7. 2 mm in diameter, glabrous, stipe 5 – 13 mm long. Seeds up to 18 per capsule, 4.5 – 5 mm long, 4.3 – 4. 5 mm wide, surfaces smooth or with irregular ridges. PHENOLOGY. — Flowering: April – December; fruiting: June – December. Flowers of Daniel & Wendt 5804 were observed to be fully open after dark.	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFDC5953FF7FFF6D02458028.taxon	distribution	DISTRIBUTION AND HABITAT. — Southern Mexico (Chiapas, Veracruz; Fig. 8); plants occur on slopes on limestone (often karstic) in lowland rain forests, lower montane rain forests, and montane rain forests at elevations from 100 to 1500 meters. ILLUSTRATIONS. — Figures 4 G, H, 16; Miranda (1954: 141). CONSERVATION. — Louteridium parayi has an EOO of 2,210 km 2. Plants occur in at least one large protected area in Chiapas, and the EOO includes another small one in Verazcruz. Frequency information on collections varies from locally infrequent to locally frequent. Known occurrences reveal two subpopulations separated by ca. 110 km. Historical landsat imagery (1984 to 2017) via Google Earth Pro (2018) reveals that the habitat of the western subpopulation in the Uxpanapa region of Veracruz has undergone extensive conversion from forested land to local agriculture and pasturage. The same imagery does not clearly show a similar large-scale conversion of forest in the region of the eastern subpopulation in Chiapas. Thus, there are two locations, in one of which a threat is evident. Based on these data, a preliminary conservation assessment of Endangered (EN; B 1, a, b; IUCN 2017) is proposed.	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFDC5953FF7FFF6D02458028.taxon	discussion	DISCUSSION. — Although a single specimen was cited in the protologue, and Miranda (1954: 125) indicated that the types of plants described in his publication were deposited at MEXU, there are two specimens of Paray & Miranda 7834 in that herbarium. On one of them, “ tipo ” is written in the same handwriting as the rest of the label (i. e., by Miranda), and in his 1955 note on this specimen Miranda indicated that flowers and part of the inflorescence were lost from the specimen. In his note, he also refers to the isotype, which contains flowers and an inflorescence. Therefore, it seems clear that the sheet bearing “ tipo ” is the holotype, as indicated above. Louteridium parayi is perhaps the most morphologically distinctive species in the genus. It possesses several unusual or unique characteristics: posterior lobe of the calyx subsaccate or with a basal appendage to 2.5 mm, a highly modified inflorescence, fused and basally subsaccate bracteoles, narrow proximal portion of the corolla tube very short to nearly nonexistent, and leathery oblanceolate to obovate leaves. The modified inflorescence of this species appears to be a thyrse, but because the lateral and modified dichasia are alternate (i. e., only one per node of the rachis), the thyrse appears dichotomously branched. The major branches in the inflorescence are interpret- ed as dichasial lateral branches that expand through non-symmetrical sympodial growth into often elongate and somewhat zig-zag “ dichasial shoots. ” The inflorescence terminates in a dichasium. Although no staminodes were seen in several dissections of corollas, it is possible that one or more small appendagelike staminodes could not be discerned among the dense trichomes at the base of the filaments. Indeed, an aberrant bud with extra corolla lobes possessed three fully formed stamens. ADDITIONAL SPECIMENS EXAMINED. — MEXICO. Chiapas: Mpio. Berriozábal, 13 km N of Berriozábal near Pozo Turipache and Finca Suspiro, [ca. 16 ° 52 ’ 10.30 ” N, 093 ° 17 ’ 21.61 ” W], D. Breedlove 20260 (DS), 24816 (DS, F, MICH, MO, NY, RSA), 26336 (DS, ENCB), 67024 (BR, CAS, MEXU, MO, US), D. Breedlove & T. Daniel 70889 (CAS), D. Breedlove & R. Thorne 30789 (DS, ENCB, LL, MO); Mpio. Ocozocoautla de Espinosa, 30 km NW of Ocozocoautla, Cerro del Ocote, [ca. 16 ° 55 ’ 51.78 ” N, 093 ° 28 ’ 15.37 ” W], D. Breedlove 28970 (DS, ENCB); Mpio. Ocozocoautla, Reserva del Ocote, Cerro La Colmena, al NE del Rancho Corocito, J. Calzada et al. 9623 (F, MEXU, XAL); Mpio. Berriozábal, entre Col. Vista Hermosa y Rancho Flor de Corazón, [ca. 16 ° 54 ’ 25.98 ” N, 093 ° 26 ’ 56.10 ” W], E. Palacios E. 1727 (CAS, IBUG, MEXU); El Suspiro, cerca de Berriozábal, [ca. 16 ° 50 ’ 33.95 ” N, 093 ° 18 ’ 07.62 ” W], L. Paray 204 (ENCB); Mpio. Ocozocoautla de Espinosa, Cerro El Banadero, 7 km carr. Horizonte – El Ciprés, 16 ° 53 ’ N, 093 ° 32 ’ W, F. Vázquez B. 952 (MEXU, XAL). Veracruz. Mpio. Jésus Carranza, Zona de Uxpanapa, ca. 1 km S of Sarabia – Uxpanapa road toward pueblo of Río Alegre, [ca. 17 ° 12 ’ 07.46 ” N, 094 ° 41 ’ 34.34 ” W], T. Daniel & T. Wendt 5804 (CAS, DUKE, K, MEXU, MICH); Mpio. Minatitlán, Zona de Uxpanapa, 13.7 km E of La Laguna toward Uxpanapa, then 6.5 km N toward Belisario Domínguez, [ca. 17 ° 18 ’ 47.80 ” N, 094 ° 23 ’ 55.68 ” W], T. Daniel & T. Wendt 5805 (CAS); Mpio. Hidalgotitlán, Hnos. Cedillo – La Escuadra, por el Río Soloxuchil, 17 ° 16 ’ N, 094 ° 37 ’ W, B. Dorantes 957 (ENCB); Mpio. Hidalgotitlán, 0 – 2 km del camino Plan de Arroyios – Alvaro Obregón, 17 ° 15 ’ N, 094 ° 40 ’ W, B. Dorantes 2850 (F, XAL); Mpio. Hidalgotitlán, KM 3 camino a La Escuadra, 17 ° 15 ’ N, 094 ° 40 ’ W, B. Dorantes 3466 (ENCB, XAL); Mpio. Hidalgotitlán, KM 2 Plan de Arroyo – Río Alegre, 17 ° 15 ’ N, 094 ° 40 ’ W, J. Dorantes 3040 (ENCB, IBUG, IEB, MEXU, XAL); Mpio. Hidalgotitlán, Camino Campto. Hnos. Cedillo – Plan de Arroyo, J. Dorantes 3625 (XAL, WIS); Uspanapas, 14 km E of Campamento La Laguna (Poblado D. S.), 7.5 km N along small road through rubber plantation, [ca. 17 ° 18 ’ 48.43 ” N, 094 ° 23 ’ 56.21 ” W], B. Hammel & M. Merello 15573 (CAS, ENCB, MO); Mpio. Uxpanapa, a 13.8 km E de La Laguna por la terracería a Uxpanapa, luego 6.8 km N por el camino a Belisario Domínguez, 17 ° 18 ’ 48 ” N, 094 ° 23 ’ 56 ” W, M. Ishiki et al. 2230 (K, MEXU, US); Mpio. Hidalgotitlán, Hnos. Cedillo – La Escuadra por el Río Soloxuchil, 17 ° 16 ’ N, 094 ° 37 ’ W, M. Vázquez T. 957 (ENCB, UC, XAL); Mpio. Jésus Carranza, 2 km N del Poblado 2, Ejido F. J. Mina, 17 ° 16 ’ N, 094 ° 40 ’ W, M. Vázquez 2617 (CAS), M. Vázquez et al. V- 2566 (CAS, MEXU, MO, XAL); Mpio. Minatitlán, 6.6 km N de la terracería La Langua – Río Grande, sobre el camino nuevo a Ejido Belisario Domínguez, el cual sale de la terracería 14.7 km E de La Laguna, 17 ° 20 ’ N, 094 ° 22 ’ W, T. Wendt & A. Villalobos C. 2540 (CAS, ENCB, LSU, MEXU).	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFD15952FF7FFEEB059A8707.taxon	description	Shrubs or trees to 9 m tall, terrestrial. Older (woody) stems subquadrate-sulcate, ± scurfy and lenticellate, often puberulent like young stems; younger (herbaceous) stems quadrate to quadrate-sulcate and ± irregularly striate, glabrous or ± evenly puberulent with erect to retrorse to antrorse eglandular trichomes to 0.05 mm long. Leaves apparently not all seasonally deciduous, ± evenly distributed along young stems, petiolate, petioles to 40 mm long, blades membranaceous, ovate-elliptic to obovate-elliptic to obovate, 130 – 380 mm long, 40 – 115 mm wide, 2.1 – 4.3 × longer than wide, acuminate to acute-apiculate at apex, acute to attenuate at base, adaxial surface glabrous or puberulent like young stems, abaxial surface paler than abaxial surface, glabrous or pubescent with mostly antrorse eglandular trichomes 0.05 – 0.1 mm long along major veins, margin often undulate, entire to sinuate to crenate to crenate-dentate. Inflorescence a terminal pedunculate raceme to racemose thyrse to 63 cm long (including peduncles and excluding corollas), peduncle to 20 cm long, glabrous or pubescent like young stems, rachis puberulent with (retrorse to) erect to flexuose to antrorse eglandular trichomes to 0.05 mm long; dichasia modified by very short expansion between pairs of succeeding flowers with the congested dichasial axis becoming a ± fan-shaped to multi- and tortuously-branched racemelike lateral short-shoot to 15 mm long, opposite (and alternate?), sessile to subsessile, 3 – many-flowered, to 60 mm long (excluding corollas), dichasial peduncles (if present) to 2 mm long, pubescent like rachis. Bracts caducous, not seen. Bracteoles and secondary bracteoles caducous, lance-linear to linear-elliptic, 5 – 13 mm long, 1 – 2. 5 mm wide, abaxially pubescent like rachis. Flowers pedicellate, pedicels to 55 mm long, puberulent with erect to retrorse (to antrorse) eglandular trichomes to 0.05 mm long. Calyx 22 – 40 mm long, lobes distinct or fused at base up to 1 mm, heteromorphic, membranaceous, abaxially puberulent like pedicels and sometimes with a few sparse stipitate glands up to 0.2 mm long as well, also inconspicuously glandular-punctate, posterior lobe conduplicate, ovate to elliptic to obovate, 21 – 38 mm long, slightly shorter than to slightly longer than lateral lobes, 8. 5 – 15 mm wide, acuminate to falcate at apex, lateral lobes sublunate, 22 – 37 mm long, 6 – 9 mm wide, acute to subfalcate at apex. Corolla cream to yellow-greenish and tinged with maroon, 45 – 52 mm long, externally pubescent with stipitate glandular trichomes 0.05 – 0.3 mm long (sometimes with flexuose eglandular trichomes to 0.2 mm long on or near margins of lobes as well), tube 25 – 33 mm long, narrow proximal portion 3 – 5 mm long, 7 mm in diameter near midpoint, throat 22 – 30 mm long, 25 – 35 mm diameter at mouth, lobes spreading to recurved, triangular to ovate, 12 – 21 mm long, 10 – 13 mm wide, rounded at apex. Stamens 2, 65 – 75 mm long, filaments glabrous distally, pubescent proximally with eglandular trichomes, thecae 10 – 12 mm long; staminodes (if present) not seen. Style 85 – 92 mm long, glabrous distally (not seen proximally), stigma equally 2 - lobed, lobes elliptic, 1.4 – 2 mm long, width not determined. Capsule 22 – 30 mm long, 5 – 7. 5 mm in diameter, densely pubescent with stipitate glandular trichomes (0.05 –) 0.1 – 0.4 mm long, stipe 3 – 3.5 mm long. Seeds up to 12 per capsule, 4.5 – 5.2 mm long, 4.5 – 5 mm wide, surfaces smooth. PHENOLOGY. — Flowering and fruiting: August – March.	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFD15952FF7FFEEB059A8707.taxon	distribution	DISTRIBUTION AND HABITAT. — Southern Mexico (Chiapas), Guatemala (Fig. 10); plants occur in wooded ravines in montane rain forests on the Pacific versant at elevations from 600 to 2000 m. ILLUSTRATION. — Figure 4 F. LOCAL NAME. — “ palo de agua, ” (Quarles van Ufford 137). CONSERVATION. — Louteridium purpusii has an EOO of 944 km 2. Two prominent threats are evident throughout most or all of its known distribution, coffee agriculture and volcanic eruptions. Numerous active volcanos lie within and adjacent to the entire EOO of this species. Most of the occurrences of the species are within the prime growing elevations (i. e., 900 to 1800 m) for Coffea arabica L., and within two important coffee-growing regions (the Soconusco region of Chiapas, Mexico and the departments of San Marcos and Quetzaltenango in Guatemala). Mexico and Guatemala are two of the top 10 coffee-producing countries in the world (Anonymous 2003, 2018). Increasing demand for coffee worldwide (e. g., Lorenzetti 2016) has or could result in increased conversion of natural vegetation in the EOO of this species to agriculture. Thus, coffee production would appear to be the more immediate threat to L. purpusii. However, based on either threat, there would appear to be a single location for the species. A preliminary conservation assessment of Endangered (EN; B 1, a, b; IUCN 2017) is proposed for this species.	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFD15952FF7FFEEB059A8707.taxon	discussion	DISCUSSION. — This species resembles some plants of L. mexicanum (see discussion under that species), but differs by its more obovate leaves and pubescence. Additional morphological tendencies that distinguish these species include the more racemose (vs. thyrsoid) inflorescences of L. purpusii (with dichasial peduncle lengths of 0 to 2 (vs. 1 to 22) mm; the caducous bracteoles (vs. usually persistent in L. mexicanum); and the generally longer styles (85 to 92 vs. 72 to 85 mm long), capsules (22 to 30 vs. 16 to 24 mm long), and capsular stipes (3 to 3.5 vs. 1 to 2 mm long). The distributional ranges of these two species are not known to overlap and are restricted to different versants of Mexico and Guatemala. ADDITIONAL SPECIMENS EXAMINED. — MEXICO. Chiapas: Mpio. Unión Juárez, 17 km E of Cacahoatán [ca. 15 ° 1 ’ 56.19 ” N, 092 ° 5 ’ 27.65 ” W], D. Breedlove & A. Smith 31613 (DS, MEXU, MO); Finca Nubes [ca. 15 ° 03 ’ 43.30 ” N, 092 ° 08 ’ 39.20 ” W], F. Miranda 1723 (LL, US); from [Finca San Antonio] Chicharras, E. Nelson 3749 (US); Finca San Antonio [Chicharras; ca. 15 ° 7 ’ 26.12 ” N, 092 ° 14 ’ 45.30 ” W], L. Quarles van Ufford 137 (U). GUATEMALA. Quezaltenango: Sta. María Ikibál [not located, possibly Santa María de Jesús], C. Bernoulli & O. Cario 2254 (K); lower S slope of Volcán Santa María, 0.3 mi W of jct. Finca Pirineos road with hwy. 95 at KM 97 [ca. 14 ° 41 ’ 5.41 ” N, 091 ° 32 ’ 50.04 ” W], C. Broome 728 (CAS, DUKE, F, MICH, US); de la represa de Santa María al Volcán Pecul [ca. 14 ° 43 ’ 26.73 ” N, 091 ° 30 ’ 42.09 ” W], Equipo CECON 1063 (MO); San José Buena Vista, Costa Cuca, [ca. 14 ° 41 ’ 53.74 ” N, 091 ° 46 ’ 35.82 ” W], L. Rodríguez 444 (P); Palmar, [ca. 14 ° 39 ’ 12.22 ” N, 091 ° 35 ’ 25.38 ” W], A. Skutch 1451 (F, US); Finca Pireneos, below María de Jesús, [ca. 14 ° 41 ’ 38.02 ” N, 091 ° 32 ’ 30.77 ” W], P. Standley 68259 (F, US); along old road between Finca Pirineos and Patzulín, P. Standley 86714 (F); 86726 (F), 86749 (F), 86750 (F, US), 86763 (F); along Quebrada San Gerónimo, Finca Pirineos, lower S-facing slopes of Volcán Santa María, between Santa María de Jesús and Calahuaché, J. Steyermark 33435 (F); off Hwy. 95 at KM 197, old road to Finca Pirineos, ca. 0.5 mi from main hwy., D. Stone 3086 (DUKE); Colomba, Finca San Francisco Pie de la Cuesta, 14 ° 43 ’ 41.05 ” N, 091 ° 43 ’ 04.45 ” W, L. Velásquez & W. López 1792 (CAS). San Marcos: Finca El Porvenir along Río Chopal, S-facing slopes of Volcán Tajumulco [ca. 14 ° 57 ’ 36.58 ” N, 091 ° 56 ’ 40.73 ” W], J. Steyermark 37501 (F); El Tumbador, finca Nueva Granada, 14 ° 51 ’ 19.9 ” N, 091 ° 53 ’ 05.9 ” W, L. Velásquez et al. 658 (BIGU, CAS).	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFD05950FF7FF9F30441804F.taxon	description	[ca. 17 ° 17 ’ 45.05 ” N, 099 ° 28 ’ 17.54 ” W], 750 m, 19 - I- 1964 (flr), H. Kruse 1380 (holotype: ENCB- 002790 - shoots bearing flowers only-image seen; isotypes (shoots bearing flowers only): EAP!, ENCB- 002789!, FCME!, MEXU!, MICH!, MO!, US!). Shrubs to trees to 4 m tall. Older (woody) stems irregularly fissured-striate, lenticellate, lacking trichomes; younger (herbaceous) stems quadrate-sulcate, glabrous (or with the surface roughened but lacking trichomes). Leaves seasonally deciduous, clustered at branch apices (and appearing quadrate at some nodes), petiolate, petioles 20 – 90 mm long, blades apparently subsucculent, lanceolate to ovate to elliptic, 82 – 260 mm long, 35 – 110 mm wide, 2 – 2.6 × longer than wide, (rounded to) acuminate at apex, cuneate to subattenuate at base, abaxial surface minutely glandular-punctate (sometimes not evident) and sparsely pubescent (especially near base) with flexuose eglandular trichomes 0.2 – 0.7 mm long or trichomes absent on mature leaves, adaxial surface very sparsely pubescent (if at all) with similar trichomes, margin entire to subsinuate, sparsely ciliate with trichomes like those of abaxial surface. Inflorescence a terminal sessile or pedunculate thyrse to 303 mm long (including peduncle, if present, and excluding corollas), peduncle (if present) to 52 (or more?) mm long, glandular-puberulent with glandular trichomes to 0.1 mm long, rachis pubescent with erect glandular trichomes 0.05 – 0.2 mm long; dichasia opposite, pedunculate, mostly 1 – 3 - flowered, to 50 (– 70 in fruit) mm long (excluding corollas), dichasial peduncles 6 – 14 (– 29 in fruit) mm long, pubescent like rachis. Bracts caducous (not seen). Bracteoles caducous (not seen). Flowers pedicellate, pedicels 23 – 30 (– 48 in fruit) mm long, pubescent like rachis. Calyx 5 – 9 mm long, lobes fused for 1 – 3. 5 mm at base, subheteromorphic, apparently subsucculent, abaxially pubescent with erect to flexuose glandular trichomes 0.05 – 0.2 mm long, posterior lobe subconduplicate, broadly ovate to subtriangular, 3.5 – 4.5 mm long, 3 – 4. 3 mm wide, sometimes smaller than lateral lobes, subacute to acute at apex, lateral lobes ovate to subtriangular, 4 – 4.8 mm long, 3 – 4 mm wide, acute at apex. Corolla green and maroon-purple, 35 – 39 mm long, externally puberulent with erect glandular trichomes to 0.1 mm long, tube 17 – 30 mm long, narrow proximal portion 4. 5 – 9 mm long, 6 – 11 mm in diameter near midpoint, throat 11 – 20 mm long, 20 – 25 mm in diameter at mouth, lobes spreading to recurved, subtriangular to ovate, 10 – 13 mm long, 9 – 12 mm wide, rounded at apex. Stamens 4, 55 – 75 mm long, filaments glabrous (at least distally, proximal portion not seen), thecae 7. 5 – 8 mm long; staminode (if present) not seen. Style 80 – 82 mm long, glabrous, stigma 2 - lobed, lobes ± funnelform, 1.5 mm long, shape and width not determined. Capsule 38 – 47 mm long, 4.7 – 6 mm in diameter, pubescent throughout with erect glandular trichomes to 0.1 mm long, stipe 4 mm long. Seeds up to 24 per capsule, 4 – 4.5 mm long, 3 – 4 mm wide, surfaces minutely granulate to ± scurfy. PHENOLOGY. — Flowering: January; fruiting: January – March. Kruse 1380 noted that the flowers were “ pronto caedizas, ” possibly referring to their being soon deciduous during the day, and thus likely largely nocturnal.	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFD05950FF7FF9F30441804F.taxon	distribution	DISTRIBUTION AND HABITAT. — Western Mexico (central Guerrero; Fig. 8); plants occur on limestone slopes in tropical subdeciduous forests (selva mediana subcaducifolia) at elevations near 750 m. CONSERVATION. — Daniel (2017) discussed the rationale for and provided a preliminary conservation assessment of Data Deficient (DD) for L. rzedowskianum, which is known only from a single site. ILLUSTRATIONS. — Rzedowski (1973: 53, fig. 4).	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFD05950FF7FF9F30441804F.taxon	discussion	DISCUSSION. — Daniel (2017) discussed the necessity for validating this name, its typification, and the sole collection locality. ADDITIONAL SPECIMENS EXAMINED. — MEXICO. Guerrero: Mpio. Chilpancingo, Rincón de la Vía, cerca de Agua de Obispo, 9 - VIII- 1964 (vegetative) and 23 - III- 1963 (fruiting), H. Kruse 1380 (EAP, ENCB, FCME, MEXU, MICH, MO, US); Mpio. Chilpancingo, Rincón de la Via, 27 - I- 1970, H. Kruse 2701 [catalog # 4755] (MEXU); same locale, 14 - VII- 1970, H. Kruse 2701 - b [catalog # 5033] (B, FCME, MEXU).	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFD25957FF7FFECB04C98747.taxon	description	Perennial herbs to shrubs to 2.5 m tall, possibly epipetric (Richardson 1972: 63). Older (woody) stems prostrate (fide Richardson 1972: 70), subquadrate or becoming irregularly fissured, lacking trichomes; younger (herbaceous) stems quadrate-sulcate to quadrate-alate, glabrous. Leaves apparently not all seasonally deciduous, ± evenly dispersed along young stems, petiolate, petioles to 85 mm long, blades subsucculent, ovate to elliptic, 80 – 260 mm long, 22 – 115 mm wide, 1.9 – 4 × longer than wide, acuminate to subfalcate to caudate at apex, subattenuate to attenuate at base, surfaces glabrous, margin entire to subsinuate-crenate. Inflorescence a terminal pedunculate thyrse to 250 mm long (including peduncle and excluding corollas), peduncles to 84 mm long, evenly pubescent with erect to flexuose glandular trichomes 0.1 – 0.3 mm long, rachises similarly pubescent; dichasia sometimes modified by sympodial expansion and appearing as lateral branches (especially from proximal nodes of inflorescence), opposite, pedunculate, mostly 3 – many-flowered, up to 115 mm long (excluding corollas), dichasial peduncles 3 – 45 mm long, pubescent like rachis. Bracts caducous, linear-lanceolate, 9 – 19 mm long, 1 – 2. 3 mm wide, abaxially pubescent like rachis. Bracteoles and secondary bracteoles caducous, lanceolate to linear-lanceolate, 9 – 14 mm long, 2 – 3 mm wide, abaxially pubescent like rachis. Flowers pedicellate, pedicels to 46 mm long, pubescent like rachis. Calyx 32 – 47 mm long, lobes subheteromorphic, membranaceous in texture (i. e., neither succulent nor coriaceous), apically acuminate, abaxially pubescent like rachis, posterior lobe planar, lance-ovate to ovate, 33 – 45 mm long, 10 – 17 mm wide, slightly to conspicuously larger than other lobes, lateral lobes lance-ovate to ovate, 29 – 39 mm long, 8 – 14. 5 mm wide. Corolla whitish to greenish yellow, 55 – 60 mm long, externally puberulent with scattered glands to 0.1 mm long, tube ca. 25 – 30 mm long, narrow proximal portion 3 – 6 mm long, 3 – 9 mm in diameter, throat 20 – 25 mm long, 20 – 30 mm diameter at mouth, lobes spreading to recurved, triangular to ovate, 15 – 27 mm long, 5. 2 – 9 mm wide, tapered and ± acute at apex. Stamens 2, 50 – 63 mm long, filaments glabrous distally (not seen proximally), thecae 11 – 15 mm long; staminodes (if present) not seen. Style 48 – 73 mm long, glabrous distally, pubescent with glandular trichomes proximally, stigma unequally 2 - lobed, lobes often recoiled, ± oblong, 1.5 – 3.5 mm long, width not determined. Capsule 22 – 28 mm long, diameter not determined, pubescent with glandular trichomes 0.05 – 0.2 mm long, stipe 1 – 4 mm long. Seeds up to 20 per capsule, 3.5 – 5 mm long, 3 – 4 mm wide, surfaces smooth or with low subconic tubercles. PHENOLOGY. — Flowering: March – June, September, December; fruiting: March – April, September. Dressler (s. n.) notes that flowers are nocturnal. Richardson (1972) observed flowers of L. tamaulipense over 14 hours (from 18: 00 to 08: 00) in September of 1969. He noted: 1) that corollas began opening at sunset and falling in the early morning; 2) the lack of a floral scent; 3) that no nectar was found in the gibbous throat of corollas; 4) the nectary had an intensely sweet taste; and 5) hummingbirds visited flowers at dusk and in the early morning.	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFD25957FF7FFECB04C98747.taxon	distribution	DISTRIBUTION AND HABITAT. — Northeastern Mexico (Tamaulipas; Fig. 8); plants occur on rocky limestone slopes in “ tropical semi-evergreen and evergreen forest ” (Richardson 1972) at elevations from 340 to 560 (to 1400) meters. The vegetation at collection sites has been characterized on specimen labels as wet forest, cloud forest, and “ selva mediana subperennifolia. ” CONSERVATION. — Louteridium tamaulipense is a local endemic that is known in the vicinity of the Sierra de Guatemala (= Sierra de Cucharas) in the northern Sierra Madre Oriental of Tamaulipas, where its known distribution (EOO = 43 km 2) is confined to the Reserva de la Biosfera El Cielo. Valiente B. et al. 285 noted that the species was locally very abundant. Several threats have been noted for the biosphere reserve, including: unplanned agricultural and forestry exploitation still being undertaken, overgrazing by livestock, increases in population density in surrounding buffer zones, poorly regulated eco-tourism, and poaching (Anonymous 2007). Most (or all?) of the collections have been made in the buffer zones outside of the two core areas (in which most human travel and exploitation are prohibited) of the reserve. The buffer zones, including the EOO, include roads, villages, and tourist facilities. However, historical landsat images (2005 to 2016) via Google Earth Pro (2018) reveal relatively little change in vegetation cover in the EOO. Thus, based on current knowledge of the species, it is not known to qualify for a threatened category in spite of its very small EOO. Based on the information summarized above, the species is provisionally assessed as Near Threatened (NT).	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
56153F74FFD25957FF7FFECB04C98747.taxon	discussion	DISCUSSION. — Richardson’s (1972) observations on floral phenology and visitation are very similar to those noted above for L. dendropilosum in Oaxaca. Louteridium tamaulipense shows some similarities to L. costaricense, and can be distinguished from that species of southern Central America by characters in the following couplet:	en	Daniel, Thomas F., Tripp, Erin A. (2018): Louteridium (Acanthaceae: Acanthoideae: Ruellieae: Trichantherinae): Taxonomy, Phylogeny, Reproductive Biology, and Conservation. Proceedings of the California Academy of Sciences 65 (2): 41-106, DOI: 10.5281/zenodo.13155705
