Avicennia (Acanthaceae: Avicennioideae) in North America and Mesoamerica

Avicennia (Acanthaceae: Avicennioideae) in North America and Mesoamerica Thomas F. Daniel Department of Botany, California Academy of Sciences, 55 Music Concourse Drive, Golden Gate Park, San Francisco, CA 94118 U.S.A.; Email: [email protected] A taxonomic revision of Avicennia, an acanthaceous genus of mangroves, in North America and Mesoamerica recognizes three species, two native (A. bicolor and A. germinans) and one locally naturalized in southern California (A. marina subsp. australasica). Herbarium specimens from throughout the region were used to formulate descriptions of each species, obtain distributional and ecological data, and as a source for pollen in order to characterize and illustrate palynological features of each species. A distribution map, an illustration of the most common species (A. germinans), and photos showing diagnostic attributes of each species are provided. Una revisión taxonómica de Avicennia, un género de los manglares en la familia Acanthaceae, en Norteamérica y Mesoamérica reconoce tres especies, dos nativas (A. bicolor y A. germinans) y una naturalizada localmente en el sur de California (A. marina subsp. australasica). Los ejemplares de herbario de toda la región se utilizaron para formular descripciones de cada especie, obtener datos distributivos y ecológicos, y como fuente de polen con el fin de caracterizar e ilustrar caracteres palinológicos de cada especie. Además, se ofrecen un mapa de distribución, una ilustración de la especie más comune (A. germinans), y fotos que muestran los atributos diagnósticos de cada especie. Avicennia has been treated previously in several families, most prominent among them Verbenaceae and Avicenniaceae. Molecular phylogenetic analyses (Schwarzbach and McDade 2002; Borg et al. 2008; McDade et al. 2008) and studies of floral structure and development (Borg and Schönenberger 2011) reveal the genus to be monophyletic, nested among Acanthaceae s.l., and sister to either subfamily Acanthoideae or more likely to subfamily Thunbergioideae. The genus is here treated as subfamily Avicennioideae. Although its taxonomic relationships have been controversial, Avicennia is well established as a genus of mangrove species. Indeed, they are usually treated in an ecological sense among the so-called “true” mangroves (Tomlinson 1986), which taxa generally occur exclusively in mangrove communities and exhibit both morphological and physiological specializations to the highly saline substrate. It is likely that morphological adaptations to a highly specialized environment have obscured its familial affinities. Other species of Acanthaceae in different tribes of subfamily Acanthoideae are considered to be true mangroves (e.g., Acanthus ebracteatus Vahl and A. ilicifolius L. of Acantheae fide Wang et al. 2011; but treated as mangrove associates by Tomlinson 1986) or mangrove associates (Bravaisia berlandieriana (Nees) T.F. Daniel and B. integerrima (Spreng.) Standl. of Ruellieae). The economic importance of mangrove plants in general, and species of Avicennia in particular, was correctly pointed out by Moldenke (1960: 144) in citing and refuting another author’s statement that plants of Avicennia are of little economic importance; Moldenke stated, “…memPROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5, pp. 163–189, 4 figs. April 29, 2016 163 bers of this genus are of tremendous economic importance to man and his economy because of their constant battle with the sea and their great success in extending and eventually building up the surface of the land adjacent to the sea in subtropical and tropical climes.” Indeed, as discussed by numerous authors (e.g., Tomlinson 1986; Odum and McIvor 1990; Costanza et al. 1997; Chivian and Bernstein 2008; Polidoro et al. 2010; and Cavanaugh et al. 2014) important ecological services provided by mangroves include: serving as home and/or nurseries for numerous marine and terrestrial organisms (e.g., fish, mammals, birds, crustaceans, worms, and insects), protecting coastlines from storm surges (e.g., buffering wave action, preventing flooding and soil erosion, and decreasing saltwater invasion), waste treatment (e.g., nutrient recycling and pollution control), carbon sequestration, and recreation (e.g., eco-tourism and sport fishing). The value of ecosystem services provided by mangrove communities worldwide has been estimated to be at least US $1.6 billion annually (Polidoro et al. 2010). The ecological and economic importance of mangroves, among which several species of Avicennia are noteworthy by their abundance and widespread distribution, will only increase with both growth of human populations in coastal areas and sea-level changes associated with climate alterations due to global warming. Estimates for recent degradation and loss of mangrove vegetation worldwide is significant. Between 20 and 35 percent of mangrove habitats have been lost since about 1980, and the current rate of annual disappearance of this vegetation is estimated to be between one and eight percent (Polidoro et al. 2010). The loss of mangrove communities in Mexico has been estimated as from 15,000 sq. km. in the 1970s to ca. 5,000 sq. km. by the late 1990s (Spalding et al. 1997). Major anthropogenic threats to mangrove communities include coastal development (e.g., tourist/resident infrastructure, mariculture, agriculture, and conversion for harbor/industrial uses), pollution (e.g., sewage effluents, oil spills, and agricultural/industrial/urban runoff), fresh water diversions, and sea level changes due to global warming (e.g., Cintrón M. and Schaeffer N. 1992). For the North American and Neotropical regions, distribution models under future climate scenarios (e.g., Cerón S. et al. 2015) predict a pole-ward shift for mangroves (including Avicennia germinans), an overall contraction of species distributions, and a decline in species richness. This regional taxonomic study used data from more than 650 specimens from North America and Mesoamerica (southern Mexico and Central America) in 22 herbaria to formulate descriptions of species, reproductive phenological periods, distributions, habitats, and other pertinent information. In the few instances where only an image of a specimen was studied, this is noted by “-image!” Pollen from each species was examined with scanning electron microscopy and characterized using terminology of Walker and Doyle (1975). Local names noted herein were obtained from herbarium specimens studied. Many additional local names and uses for these species can be found among references in the literature cited. Avicennia L., Sp. Pl. 1: 110. 1753; Gen. Pl. ed. 5, 49. 1754. TYPE.— Avicennia officinalis L. Bontia L., Sp. Pl. 2: 638 (“938”). 1753. TYPE.— Bontia daphnoides L. Upata Adanson, Fam. 2: 201. 1763. ≡ Avicennia L. Sceura Forssk., Fl. Aegypt.-Arab. 37. 1775. TYPE.— Sceura marina Forssk. Halodendrum Thouars, Gen. Nov. Madagasc. 8. 1806. TYPE.— Halodendrum thouarsii Roem. & Schult. Hilairanthus Tiegh., J. Bot. (Morot) 12: 358. 1898. TYPE.— Not designated (two species cited). Shrubs or trees lacking cystoliths and with erect, aerial, and sometimes branched roots (pneumatophores) up to 4 dm tall under and beyond canopy (and infrequently also with aerial stilt roots), cystoliths absent. Leaves opposite, leathery, entire, margin flat to revolute. Inflorescence of axillary and/or terminal dense (sometimes subcapitate) pedunculate dichasiate spikes or panicles of 164 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 dichasiate spikes; dichasia opposite, 1-flowered, sessile. Bracts ± leathery, concavoconvex. Bracteoles concavoconvex. Flowers sessile, protandrous. Calyx 5-lobed, lobes ± free, concavoconvex, imbricate, equal to subequal in size. Corollas white to yellow to orange, tube expanded distally, shorter than limb, limb 2-labiate or actinomorphic, 4or 5-lobed, lobes erect to reflexed, oblong to obovate, contorted (left contort aestivation) in bud. Stamens 4, at least 2 exserted from mouth of corolla (in ours), oriented in pairs below upper lip of corolla or oriented symmetrically around corolla with each equally distant from the others; anthers 2-thecous, thecae of a pair ± equally inserted, parallel, ± equal in size, lacking basal appendages, each dehiscing toward lower lip (i.e., flower nototribic) or toward each other (i.e., flower pleurotribic) by a longitudinal slit; pollen suboblate to euprolate, 3-colporate, exine reticulate; staminodes 0. Style not evident or elongating with age of flower; stigma 2-lobed, lobes slightly unequal. Capsule podlike, leathery, ± ellipsoid to ovoid (often asymmetric), ± compressed, unilocular, retinacula absent, dehiscence not explosive, usually occurring at time of detachment or soon after being shed. Seed 1 (–2) per capsule, filling fruit and consisting mostly of ripe embryo with 2 conspicuous folded cotyledons and a pubescent radicle (cryptoviviparous). Moldenke (1960, 1973) and Duke (1991) listed additional generic synonyms. The generic name is derived from Ibn Sina (=Avicenna), 980–1037, a Persian physician, philosopher, and naturalist. At least eight (e.g., Duke 1991; Duke et al. 1998) species of Avicennia are recognized in maritime regions of the tropics and subtropics worldwide. The genus is monophyletic, and includes some of the most prominent and widely distributed “true” mangroves (cf. Tomlinson 1986). Plants occur along shorelines and in tidally influenced waterways in two mostly tropical regions: Atlantic/Caribbean/eastern Pacific (3 spp.) and Indo-western Pacific (5 spp.). Although elevations up to 150 meters have been noted on herbarium labels of American plants, occurrences at elevations much above sea level would appear to be exceptional (if accurately reported) or more likely erroneous. Leaves of Avicennia accumulate an external crystalline layer of excreted salt between rains. Three types of major roots are evident on plants: cable roots, anchoring roots, and pneumatophores. Cable roots extend up to several meters horizontally underground from the base of the trunk and give rise to downward growing anchoring roots and upward growing pneumatophores. At least the cable and anchoring roots also give rise to finer roots within the substrate. Pneumatophores (aerial, pencil-like roots that are evident at least at low tide and that facilitate the uptake of atmospheric oxygen) and cryptoviviparous seeds (in which the embryo swells and breaks through the seed coat, but not through the fruit wall, prior to the fruit falling from the plant) are characteristic of the genus. Macromorphological variation among several species of Avicennia is extensive, and often has been used in taxonomic circumscriptions. It has been shown that habitat conditions (e.g., variations in salinity and topographic position) and geography can correlate with morphology of the habit, leaves, and/or flowers (Sherrod and McMillan 1985; Duke 1991; Turner et al. 1995). Based on limited sampling (Fig. 1), pollen shape varies from suboblate to euprolate (i.e., with polar diameter:equatorial diameter [P:E] from 0.86 to 1.90). Much of this variation is evident within a single sample (i.e., Salywon 1188 for A. germinans) and probably represents harmomegathic variation. Key to the Native and Naturalized Species of Avicennia in North America and Mesoamerica 1a. Flower actinomorphic; corolla yellowish to orangish, upper lip entire or bifid with the division to 0.2 mm long, the four corolla lobes ovate-triangular to ovate-elliptic; stamens inserted in distal half of corolla tube near base of lobes, oriented equally distant from each other and DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 165 166 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 dehiscing toward the central gynoecium, 1.5–2 mm long, filaments 0.5–0.8 mm long; naturalized in southern California. . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. marina subsp. australasica 1b. Flower ± 2-labiate; corolla white to cream (often with yellow in throat), upper lip bifid to 2-parted with the division to 3 mm long, the 4–5 lobes oblong to obovate; stamens inserted at midpoint or in proximal half of corolla tube, oriented in pairs adjacent to upper lip of corolla and dehiscing toward lower lip, 2–5 mm long, filaments 1.5–4 mm long; native in southeastern U.S.A., Mexico, and Central America. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2a. Internodes of young stems glabrous; 1-flowered dichasia ± evenly distributed along rachis, internodes near midspike 3.5–7 (–9) mm long, rachis clearly visible; internal surface of corolla lobes glabrous; 2 stamens exserted from mouth of corolla and 2 stamens included in corolla tube (or only partly exserted from it); style not evident after corolla dehisces; capsule blackish when dry. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. bicolor 2b. Internodes of young stems usually scurfy with shiny whitish trichomeor scale-like projections to 0.05 mm long; 1-flowered dichasia congested at or toward apex of rachis (± headlike), internodes near midspike mostly 0.8–4 mm long, rachis not or barely or only partially visible; internal surface of corolla lobes densely pubescent (at least in distal half); all 4 stamens exserted from mouth of corolla; style usually conspicuous after (and often before) corolla dehisces; capsule usually grayish when dry. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. germinans 1. Avicennia bicolor Standl., J. Wash. Acad. Sci. 13: 354. 1923. TYPE.— PANAMA: Coclé: Aguadulce, outskirts of tidal belt, 5 XII 1911, H. Pittier 4968 (holotype: US!; isotypes: F-image!, K-image!, BM! NY! P! US!). Figures 1A–C, 2, 3D. Shrubs to 3 m tall or trees to 10 (–23) m tall. Young stems of reproductive shoots glabrous (trichomes sometimes persisting around leaf scars, but internodes glabrous). Leaves petiolate, blades ovate to elliptic to broadly elliptic (to obovate), 62–175 mm long, 31–86 mm wide, length:width = 1.2–2.6, (emarginate to) rounded to subacute at apex, rounded to acute to subattenuate at base, surfaces often conspicuously discolorous (abaxial surface lighter), punctate-pitted (sometimes inconspicuously so abaxially), adaxial surface lacking trichomes, abaxial surface covered with a dense scurfy layer that sometimes includes longer ± appressed eglandular trichomes. Inflorescence of axillary (from distalmost leaves) and terminal pedunculate panicles of elongate spikes, panicles ± open, to 115 mm long (including peduncle and excluding corollas) and to 120 mm wide, peduncles 25–40 mm long, glabrous or distally becoming pubescent like branches and rachises, panicle branches subtended by triangular-concave inflorescence bracts (sometimes caducous) 1–3 mm long, pubescent like rachises, fertile portion of spike 12–35 mm long, rachises clearly visible, internodes near midspike 3.5–7 (–9) mm long, densely pubescent with minute (˂ 0.05 mm long) DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 167 FIGURE 1 (left). Pollen of Avicennia spp. A–C. Avicennia bicolor (Sediles 461). A. Interapertural view. B. Apertural view. C. Polar view. D–G. Avicennia germinans (D, Palmer 484; E–G, Salywon 1188). D. Interapertural view. E. Apertural view. F. Polar view. G. Interapertural view. H–K. Avicennia marina subsp. australasica (Moran 28024). H. Interapertural view. I. Apertural view. J. Polar view. K. Apertural view. Scales = 5 µm. Pollen measurements (n = number of grains measured, P = polar diameter, and E = equatorial diameter). Many additional grains from each specimen were examined, but data summarized below were made on a subset of those oriented to permit accurate measurements and photographic documentation. Avicennia bicolor — Sediles 461 (CAS, n = 5): P = 19–22 μm, E = 19–24 μm, P:E = 0.86–0.92. Avicennia germinans — Daniel & Araque 9478 (CAS, n = 2): P = 23 μm, E = 24–25 μm, P:E = 0.92; de Nevers et al. 6554 (CAS, n = 2): P = 25 μm, E = 26–28 μm, P:E = 0.89; Palmer 484 (CAS, n = 1): P = 46 μm, E = 24 μm, P:E = 1.90; Salywon 1188 (CAS, n =4): P = 27–42 μm, E = 24–29 μm, P:E = 0.93–1.75. Avicennia marina subsp. australasica — Moran 28024 (CAS, n = 4): P = 28–41 μm, E = 24–27 μm, P:E = 1.04–1.70. glandular trichomes and with antrorse (to flexuose) eglandular trichomes to 0.1 mm long. Bracts triangular-ovate to subcircular, 1–2.3 mm long, abaxially pubescent like rachis. Bracteoles similar to bracts. Flowers mostly 8–10 per spike (but up to 22 per spike). Calyx 2–4 mm long, lobes broadly ovate to subcircular, abaxially pubescent like rachis. Corollas 4.5–7 mm long, internally white to cream, externally glabrous (proximal 2/3 of tube) and densely pubescent with appressed eglandular trichomes to 0.2 mm long (distal 1/3 of tube and limb), tube 2–2.5 mm long, limb ± 2-labiate with 4–5 lobes, upper lip apically 2-parted (with division up to 2 mm long) and/or wider than lobes of lower lip, all lobes oblong to obovate, glabrous internally, 3–4.5 mm long. Stamens 4, inserted at midpoint or in proximal half of corolla tube, 2 exserted from mouth of corolla tube and 2 included in or only partially exserted from corolla tube, oriented in pairs near upper lip of corolla with thecae opening toward lower lip, 2–2.5 mm long, filaments 1.5–2 mm long, anthers presented at 2 heights, thecae 0.3–0.6 mm long; pollen suboblate to oblate spheroidal, polar diameter (P) 19–22 µm, equatorial diameter (E) 19–24 µm, P:E = 0.86–0.92. Style not evident, stigma lobes 0.2–0.4 mm long. Fruit greenish yellow, black when dry, ovoid to ellipsoid, 15–29 mm long, 7–17 mm wide, ± sparsely pubescent with antrorsely appressed eglandular trichomes to 1 mm long, these sometimes more or only evident distally on mature fruits. PHENOLOGY.— Flowering: September–May; fruiting: February–August. DISTRIBUTION AND HABITATS.— Pacific coast of southern Mexico (Chiapas, ca. lat. 16°02ʹ26ʺN), Central America, to southern Panama (Los Santos and Darién, ca. lat. 7.98°N; Fig. 2); plants occur along and near shorelines in coastal mangrove swamps (mangals) and salt marshes at elevations at or near sea level. In addition to the provinces and departments of Central American nations from which specimens have been examined (noted below), this species potentially also occurs or occurred in the following Pacific coastal political units: Guatemala (Escuintla, Jutiapa, Retalhuleu, San Marcos, Santa Rosa, Suchitepequez), El Salvador (La Libertad, La Paz, La Unión, San Miguel, San Vicente, Sonsonate, Usulután), Nicaragua (Carazo, Managua), and Panama (Chiriquí, Veraguas). Gibson (1970) included Avicennia bicolor Standl. in her account of the Guatemalan taxa because of its expected occurrence in the country. Although it undoubtedly occurred or still occurs there, the species has yet to be collected in Guatemala. The southernmost occurrence of Avicennia bicolor has been attributed to the Pacific coast of Colombia (e.g., Sanders 1997; Duke 2010; Aymard 2015). The sole collection cited by Aymard (2015; Forero & Gentry 794 at COL) and another Colombian collection identified as this species (Gentry & Juncosa 41115 at COL), both appear to pertain to A. germinans based on images of these collections supplied by COL. Unless other collections that conform to A. bicolor have been made in Colombia, the southern extent of this species appears to be in Panama at ca. 7.98°N, on both sides of the Gulf of Panama (i.e., in the provinces of Los Santos [Dwyer 5079A] and Darién [Duke 5488]). LOCAL NAMES.— “Madre sal” (DeRiemer s.n.); “madresal prieto” (Santamaría D. & Romero B. 1a); “palo de sal” (Sediles 457); “palo de sal hoja ancha” (Sediles 455, 460). CONSERVATION STATUS.— The extent of occurrence (EOO) of Avicennia bicolor is 262,479 km2, although a major portion of that area consists of open ocean and inhospitable upland habitats. The species has been assessed as VU (vulnerable) by the IUCN (Duke 2010) based on a documented continuing decline in population under criterion A. DISCUSSION.— This species is readily recognized by the combination of its glabrous young stems, relatively remote dichasia in the inflorescence (Fig. 3C), bilaterally symmetric flowers with internally glabrous corolla lobes, and black fruits (when dry; Fig. 3D). The Mexican occurrence of this species has sometimes been overlooked (e.g., Breedlove 1986; Spalding et al. 1997) or minimized (e.g., treated as a synonym of A. germinans for practical pur168 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 poses; López P. and Ezcurra 2002). Throughout its distributional range it often grows with A. germinans (e.g., Pittier 4968, the type, grows with Pitier 4969, a collection of A. germinans from the same locale). Rabinowitz (1978) noted that where these species are sympatric there is apparently no zonation between them. Like those of A. germinans, flowers of A. bicolor have been noted to be fragrant (e.g., Knapp 1244). Corollas are usually described by collectors as white to cream; they are sometimes noted to have a yellow throat (like A. germinans; e.g., Borg and Schöenenberger 2011). Williams 78 from Panama shows exceptionally long and floriferous inflorescences with the fertile portion of spikes to 65 mm long and with up to 22 flowers (vs. usually 4–8) per spike. Tomlinson (1986) treated A. tonduzii as a synonym of A. bicolor, and others have followed his influential taxonomic account (e.g., Aymard 2015). In his key to species of Avicennia and description of A. bicolor, Tomlinson indicated that corollas of this species were conspicuously pubescent within (as they are on the type of A. tonduzii, but not on the type of A. bicolor). Distinctions, if any, between A. tonduzii and A. germinans are not readily apparent, and others have treated these names as synonymous. Additional information about A. tonduzii is provided below under A. germinans. ADDITIONAL SPECIMENS EXAMINED.— COSTA RICA. Guanacaste: Nandayura, Península de Nicoya, Playa Bejuco, 09°49ʹ56ʺN, 085°20ʹ34ʺW, A. Fernández 1762 (MO); Abangares, Cuenca del Abangares, San Buenaventura, 10°10ʹ29.8253ʺN, 085°09ʹ31.4524ʺW, L. González & A. Garita 3909 (MO); Tamarindo, Playa Tamarindo, 10°18ʹN, 085°51ʹW, W. Haber & W. Zuchowski 8961 (F, MO); Port Parker, J. Howell 10242 (CAS); Refugio Silv. Tamarindo, Estero Tamarindo, Santa DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 169 FIGURE 2. Map showing distribution of Avicennia spp.in North America and Mesoamerica. The generalized distribution of A. germinans is shown. The distribution of A. germinans in Cuba and other West Indian islands is not shown. 170 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 FIGURE 3. Avicennia spp. A. Flower of A. germinans in Florida (photo by Bob Peterson, cropped, creative commons license, <https://www.flickr.com/photos/pondapple/7270558948/in/photostream>). B Flower of A. marina subsp. australasica in Australia (photo by M. Fagg, cropped, source: Australian National Botanic Gardens at <http://www.anbg. gov.au/photo>). C. Inflorescence of A. biflora (Sediles 461, CAS). D. Fruits of A. bicolor (top; Barrera 8, CAS), A. germinans showing dehiscence (middle; Ferris 5396, DS), and A. marina subsp. australasica (bottom; Nickerson 6445a from New Zealand, CAS). Cruz, 10°19ʹ40ʺN, 85°49ʹ20ʺW, Q. Jiménez 859 (K, MO); P.N. Santa Rosa, Cantón de La Cruz, Península de Santa Elena, Estero Grande, 10°54ʹ59ʺN, 085°47ʹ03ʺW, J. Morales 4142 (F, MO); P.N. Santa Rosa, Playa Naranjo, 10°47ʹ53ʺN, 085°40ʹ44ʺW, J. Morales et al. 1262 (F); Puerto Jesús, R. Pohl & G. Davidse 10588A (F, MO, US); Cantón La Cruz, P.N. Santa Rosa, Peninsula de Santa Elena, Murciélago, 10°55ʹ20ʺN, 85°44ʹ15ʺW, F. Quesada 84 (K, MO); P.N. Santa Rosa, Playa Naranjo, N. Zamora et al. 1140 (F, MO). Puntarenas: Golfo de Nicoya Externo, Isla San Lucas, Punta Cañón, J. Morales & D. Santamaria 12387 (MO); Garabito, Cuenca del Jesús María, Garabito, alrededores de Playa Punta Loros, 09°51ʹ26.3900ʺN, 084°41ʹ29.8110ʺW, A. Rodríguez & V. Ramírez 6688 (MO). ELSALVADOR. Ahuachapán: without locale, Padilla 333 (US); Las Salinas, K. DeRiemer 1625 (US), 1626 (US); Las Chacaras, en La Barra de Santiago, K. DeRiemer s.n. (US). HONDURAS. Choluteca: Playas de Cedeño, F. Padilla 101 (BM).Valle: Puerto Soto, 12 km from El Tular, A. Molina R. 21457 (BM, NY, UC, US); Puerto Soto, 15 km WSW de San Lorenzo, C. Nelson 1323 (MO). MEXICO. Chiapas: Paderón, Tonala, E. Matuda 16353 (US); Mpio. Pijijiápan, Estero San José, 15°43ʹ39.50ʺN, 093°29ʹ50.20ʺW, S. Santamaría-Damián & E. Romero-Berny 1a (MEXU), 1b (MEXU). NICARAGUA. Chinandega: Mpio. El Viejo, Reserva Natural de Cosigüina, San Remigio entre El Congo y Bella Vista, 13°01ʹN, 097°35ʹW, I. Coronado G. & R. Rueda 3626 (MEXU, MO); Mpio. Puerto Morazán, de Morazán hasta 5 km en dirección Ttonalá, 12°49ʹN, 087°09ʹW, R. Rueda et al. 17322 (MEXU, MO). León: Isla del Venado, 3.3 km de Las Peñitas, Barrera 8 (CAS); Las Peñitas, Isla Juan Venado, 12°13ʹN, 086°53ʹW, I. Coronado G. & R. Rueda 4943 (MO); Las Peñitas, Isla Juan Venado, 12°17ʹN, 086°53ʹW, I. Coronado G. et al. 6979 (MO); Isla del Venado, 8.2 km de Las Peñitas, 60 m del Estero Las Peñitas, Sediles 445 (NY), 456 (NY); 3.3 km de Las Peñitas, Isla del Venado, Sediles 454 (MO), 459 (MO), 461 (CAS); 3 km de Las Peñitas, en la Isla del Venado, Sediles 455 (F), 460 (K); 8.1 km de Las Peñitas, en la Isla del Venado, 60 m del Estero Las Peñitas, Sediles 457 (P); 8.3 km del Las Peñitas, en la Isla del Venado, 80 m del Estero Las Peñitas, Sediles 458 (US). Rivas: “Marsella,” 11°16–17ʹN, 85°52–53ʹW, M. Araquistain 3828 (MO, P); San Juan del Sur, entre Las Playas de Marsella y Rivas, 11°17ʹN, 085°54ʹW, R. Rueda et al. 1435 (MO). PANAMA. Coclé: ca. 2 km del Puerto, camino entre el puerto de Aguadulce hasta el pueblo, M. Correa A. 4314 (MO); Isla del Pozo, salinas of Río Estero Salado, 08°11ʹN, 080°30ʹW, S. Knapp et al. 3401 (MEXU, MO); below Aguadulce, E. Tyson 7262 (FSU, MO). Darién: ca. 10 mi S of El Real on Río Pirre, J. Duke 5488 (MO). Herrera: Cienega El Mangle, NE of Paris, S. McDaniel 8020 (FSU, MO). Los Santos: Monagre Beach, J. Dwyer 5079A (MO). Panamá: Isla Casaya, J. Duke 10372 (MO); San José Island, ca. 55 mi SSE of Balboa, Playa Grande, I. Johnston 1259 (DUKE, MO, P, US); Porto Posada, R. Williams 78 (NY). Panamá Oeste: Punta Chame, W. D’Arcy 10241 (MO); Punta Chame, 10–15 mi from Pan American Hwy., 08°40ʹN, 079°45ʹW, S. Knapp 1244 (MEXU, MO). 2. Avicennia germinans (L.) L., Sp. Pl., ed. 3, 2: 891. 1764. Bontia germinans L., Syst. Nat., ed. 10, 2: 1122. 1759. TYPE.— “Habitat in Indiis” [JAMAICA], P. Browne s.n., LINN Herb. No. 813.2 (lectotype, designated by Stearn [1958: 35]: LINN-image!). Figures 1D–G, 2, 3A,D. Avicennia nitida Jacq., Enum. Syst. Pl. 25. 1760. Avicennia officinalis var. nitida (Jacq.) Kuntze, Revis. Gen. Pl. 2: 502. 1891. Hilairanthus nitidus (Jacq.) Tiegh, J. Bot. (Morot) 12: 358. 1898. TYPE.— See discussion. DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 171 ELSALVADOR. Ahuachapan: Las Chacaras, La Barra de Santiago, K. DeRiemer 1645 (US); San Francisco Menéndez, Garita Palmera, zanjón El Aguacate, 13°43ʹN, 090°04ʹW, D. Rodríguez & E. Escobar 1875 (MO); Santuario de las Aves, 13°42ʹN, 90°00ʹW, R. Villacorta y E. Montalvo 817 (K, MEXU). La Libertad: El Amatal, San Diego, 13°25ʹN, 089°14ʹW, J. González 316 (MO); Estero de San Diego, J. González & A. Pérez 193 (MEXU). La Paz: Estero de Jaltepeque, P. Allen 7278 (US); near mouth of Río Jiboa, F. Choussy 1593 (US); El Zapote, Costa del Sol, 13°21ʹN, 89°W, J. González & M. Hernández 326 (MEXU, MO). La Unión: rocky beach, La Union, A. Beetle 26263 (K, UC); ca. 1 km S of Barrancones, 13°26ʹ08ʺN, 087°47ʹ32ʺW, G. Davidse et al. 37355 (MEXU, MO); coast near La Union, V. Grant 716 (F); vicinity of La Unión, P. Standley 20786 (US). Sonsonate: S of Acajutla, P. Allen 6837 (F, US); Estero San Juan, K. DeRiemer 1617 (US). Usulután: Jiquilisco, El Tercio, 13°15ʹN, 088°31ʹW, R. Carballo & L. Cabrera 831 (LAGUimage!). GUATEMALA. Escuintla: Iztapa, Canjón Morón, M. Arrecis 106 (CAS, MEXU, MO); San José, J. Donnell Smith 2510 (K, NY, US); San José, W. Maxon & R. Hay 3659 (US); Had. Las Fianzas, G. Salas 367 (US); Puerto de San José, J. Véliz & M. Véliz 94.4076 (CAS, MEXU). Izabal: near Puerto Barrios, P. Standley 72167 (F, NY). Jutiapa: Las Lisas, Barra el Ahumado, M. Arrecis 74 (CAS, MEXU, MO), 121 (CAS), 122 (MEXU); Iztapa, Zanjón Morón, M. Arrecis 105 (CAS). Retalhuleu: Río Ocosito en límite Tilepa, Ocós, San Marcos y Manchón, M. Arrecis 47 (MO); Champerico, P. Standley 66563 (F), 87592 (F, NY). San Marcos: Mpio Ocós, almendrales, Tilapa, M. Arrecis 50 (MO), Ocós, J. Steyermark 37803 (F). Santa Rosa: Las Lisas, M. Lara s.n. (MO); Las Lisas, Camaronera Mayasal, 13.80723°N, 090.21703°W, J. López & R. Jiménez 120 (CAS); Parque de la Barra Hawaii, Aldea el Dormido, 13.84113°N, 090.34751°W, J. López et al. 100 (CAS). HONDURAS. Atlántida: ca. 5 km NE of Tela near Telatinza, ca. 15°48ʹN, 087°26ʹW, T. Daniel & J. Araque 9478 (CAS, NY). Choluteca: Ratón Island, A. Molina R. 22779 (US), 23287 (DS, F, NY, US); Cedeños Beach, A. Molina R. et al. 31978 (F); Punta Ratón, 70 km NW de Cd. Choluteca, C. Nelson et al. 3259 (MO). Colón: Río Guaimoreto, 4.5 km NE of Trujillo on road to Castilla, 15°57ʹ30ʺN, 085°54ʹ30ʺW, J. Saunders et al. 625 (F, MO, NY). Gracias a Dios: Puerto Lempira, Laguna de Caratasca, A. Díaz Z. 212 (MEXU); Puerto Lempira, Laguna de Caratasca, M. Espinal 142 (MO). Islas de la Bahía: Isla de Barbareta, C. Nelson & G. Cruz 8411 (US); Isla de Roatán, playa al E de Roatán, C. Nelson & E. Romero 4591 (MO). Valle: cerca de Isla Zacate Grande, D. Hazlett 916 (MO); San Lorenzo, A. Molina R. 8635 (F); Golfo de Fonseca, Puerto Soto, 12 km from El Tular, A. Molina R. 21454 (F, NY). MEXICO. Baja California: Bahía San Francisquito [28.40814°N, 110.57079°W], R. Moran 12625 (SD-not seen). Observed by I. Wiggins at Bahía de los Ángeles (see Turner et al. 1995). Baja California Sur: Playa Santispac, carr. transpeninsular, B. Arteaga et al. 130 (MEXU); Magdalena Bay, T. Brandegee s.n. (DS, UC); El Mogote, peninsula extending into La Paz Bay, 24°8– 11ʹN, 110°19–26ʹW, A. Carter 2721 (DS, K, UC); Isla Carmen, vicinity of Las Salinas, ca. 25°59ʹN, 111°07ʹW, A. Carter 5924 (UC); Isla Carmen, Puerto Balandra, ca. 26°00.5ʹN, 111°10.5ʹW, A. Carter & R. Ferris 3733 (CAS, UC); Bahía Concepcion, ca. 14 mi S of Mulegé, M. Dillon et al. 1954 (F); Bahía de La Paz, Chametla, 24°09ʹN, 110°06ʹW, R. Domínguez C. 443 (MEXU); Sierra de la Giganta, ca. 12–14 mi S of Mulegé, near Bahía Santispac, T. Elias et al. 10812 (F, NY); 16 mi S of Mulegé at Conception Bay, R. Ferris 8680 (DS); Mpio. La Paz, Punta Prieta cerca de Pichilingue, 14 km NE de La Paz, F. González M. et al. 8163 (MEXU); La Paz, I. Johnston 3045 (CAS); Coronados Island, I. Johnston 3758 (CAS, K, UC); Carmen Island, Puerto Balandra, I. Johnston 3821 (CAS, K, UC); San Evaristo Bay, I. Johnston 4089 (CAS, K, UC); Puerto Escondido, I. Johnston 4293 (CAS, K, UC); Magdalena Bay, H. Mason 1909 (CAS, DS, K); 178 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 Ballandra Bay, Carmen Island, 26°00.5ʹN, 111°10.5ʹW, R. Moran 3926 (DS, UC), 9168 (MEXU); El Mogote, peninsula in La Paz Bay, ca. 24°10ʹN, 110°20ʹW, R. Moran 7151 (CAS, DS, K); Concepcion Bay, F. Shreve 7099 (F); Isla San José, costa SW, 24°54ʹN, 110°38ʹW, M. Sousa P. 218 (MEXU); Isla del Carmen, lado W, Puerto Balandra, 26°01ʹN, 111°11ʹW, M. Sousa P. 242 (MEXU); Magdalena Bay near Medano Amarillo, J. Thomas 7940 (CAS, UC); Bahía Concepción, Playa Los Cocos, 26.87763°N, 111.97499°W, D. Valvov 2005088 (MEXU); Bahía de Concepción between Mulegé and head of bay, I. Wiggins 5455 (CAS, DS, F, UC); Estero Salinas, arm of Almejas Bay, S of Magdalena Bay, I. Wiggins 11487 (CAS, DS, UC); 1.8 mi E of La Paz toward Pichilinque Bay, I. Wiggins 14563 (CAS, DS); S shore of Bahía de La Paz, I. Wiggins 16174 (DS); Puerto Escondido, 15 mi S of Loreto, I. Wiggins 17526 (DS); N side of Bahía Astiones, W side of Isla San José, I. Wiggins 17672 (DS); NE part of Isla San Francisco [24°50ʹ32.36ʺN, 110°33ʹ59.94ʺW], I. Wiggins 17765 (DS); Bahía de la Concepción, beach at Punta Guadalupe, I. Wiggins & D. Wiggins 18017 (DS); N side of Santispaquis Cove, Bahía de Concepción, I. Wiggins & D. Wiggins 18239 (CAS, DS). Campeche: between Sabancuy and Cd. del Carmen, R. Burnham & R. Spicer 146 (MEXU); 30 km W de Hecelchakan, camino a Isla Jaina, E. Cabrera & H. de Cabrera 13345 (MEXU); 6 km NE de Champotón, carretera Cd. del Carmen, E. Cabrera et al. 8497 (MEXU); Isla de Jaina, ca. 54 km W de Hecelchacan, E. Cabrera C. et al. 11975 (MEXU); Mpio. Calkini, Isla Punta, 20°30ʹN, 90°W, C. Chan V. & J. Flores 420 (XAL); “Panga” de Zacatal, J. Chavelas P. & C. Zamora S. ES-4752 (MEXU); Mpio. Hecelchakan, 6 km antes de la costa, en la carretera de Pomuch, Isla de Jaina, 20°14ʹN, 090°24ʹW, E. Gongora 546 (UC, XAL); Mpio. Campeche, Palmas, Cd. de Campeche, 19°52ʹN, 090°30ʹW, C. Gutiérrez B. 5876 (XAL); Mpio. Cd. del Carmen, 5 km NE de Sabankuy, 19°03ʹ00ʺN, 091°08ʹ00ʺW, C. Gutiérrez B. 7390 (MEXU); Puerto Real, F. Menendez L. 467 (MEXU); S de Campeche, F. Miranda 7944 (MEXU); Mpio. Calkiní, Isla Arena, 27 km de Takuché, 20°37ʹ25ʺN, 090°25ʹW, M. Narváez 1365 (MEXU); Mpio. El Carmen, Punta Cochinitos, Laguna San Francisco, 18°26ʹN, 091°46ʹW, D. Ocaña N. & A. Novelo R. 158 (MEXU); carretera Champotón–Isla del Carmen, ca. 15 km de Champotón, T. Pennington & J. Sarukhán K. 9405 (K); carretera Champotón–Campeche, T. Pennington & J. Sarukhán K. 9623 (K); Mpio. Champotón, camino a El Zapote, 4 km desde el entronque con la carr. Campeche–Champotón, 19°20ʹN, 090°45ʹW, R. Rico G. 141 (XAL); 2 km N of Cd. del Carmen, J. Sauer 2440 (F); Champotón, W. Steere 1751 (CAS, MICH); Mpio. Tenabo, granja camaronera de Tenabo, entre KM 27 y el mar, 20°01ʹ30ʺN, 090°13ʹ06ʺW, P. Zamora C. et al. 5824 (XAL); Mpio. Cd. del Carmen, Isla del Centro de Cayo Arcas, SE parte, S. Zamudio 103 (MEXU, XAL). Colima: no collections seen, but species recorded from the state (see: http://www.projectsabroad.org/_downloads/uk/conservation-management-plan/mexico-conservation-managementplan-2014.pdf). Chiapas: Mpio. Tonalá, W side of Mar Muerto opposite Paredón, D. Breedlove 20771 (DS); Mpio. Tonalá, E shore of Mar Muerto, N of Paredón, D. Breedlove & R. Thorne 20806 (DS, NY); Mpio. Arriaga, balneario La Gloria, A. Espejo & S. Hernández 2904 (MEXU); Acapetahua, cerca al Embarcadero Las Garzas, 15°12ʹ38.7ʺN, 92°48ʹ39.1ʺW, H. Gómez D. 2293 (K); Las Garzas, Acapet, E. Matuda 2728 (K, NY); Paderon, Tonala, E. Matuda 16279 (US); Mpio. Acapetahua, 0.5 km antes de La Palma, 15°12ʹ16, 092°48ʹ37ʺW, S. Ochoa G. et al. 4535 (MEXU); Mpio. Tonala, Col. Miguel Hidalgo, afueras de Puerto Arista, 15°55ʹN, 93°50ʹW, V. Rico-Gray & I. Espejel 298 (F, MEXU); Mpio. Pijijiapan, Salina atras del Chocohuital, 15°30ʹN, 093°15ʹW, V. Rico-Gray & I. Espejel 305 (MEXU, XAL); Mpio. Tapachula, Estero de Puerto Madero, 14°45ʹN, 092°35ʹW, V. Rico-Gray & I. Espejel 346 (F, MEXU); Mpio. Puerto Madero, Puerto Madero, E. Ventura & E. López 91 (MEXU, XAL); Las Margaritas, Pijiiapan, G. Zavala P. & M. Illescas 19 (MEXU). Guerrero: Mpio. Copala, Laguna de Chautengo, J. Almazán 248 (FCME); Mpio. Zihuatanejo, Playa La Ropa, Bahía de Zihuatanejo, 17°40ʹN, 101°34ʹW, G. CastilDANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 179 lo G. 1137 (UC), 6273 (MEXU, XAL); Mpio. Jose Azueta, Barra del Potosí, 17°40ʹN, 101°34ʹW, G. Castillo C. & P. Zamora C. 6511 (XAL); Mpio. Zihuatanejo, entre la Punta Ixtapa y el Cerro El Rialito, 17°40ʹN, 101°39ʹW, G. Castillo C. et al. 6566 (MEXU); Mpio. Cuajinicuilapa, Punta Maldonado, N. Diego 2213 (FCME); Mpio. Acapulco de Juárez, El Arenal, Laguna de Tres Palos, N. Diego 4161 (FCME, MEXU); Mpio. Petatlán, Cerro Huamilule, (Morro de la Laguna Potosí), N. Diego & R. Oviedo 6636 (FCME); Mpio. Cruz Grande, W de Las Penas, R. Fonseca 1625 (MEXU); Mpio. Cruz Grande, Los Tamarindos, Laguna de Chautengo, R. Gutiérrez 4 (FCME); Mpio. Tecopan de Galeana, Laguna Nuxco, extremo SE, F. Lorea 5280 (XAL); Mpio. Atoyac de Alvárez, Arenal de Palos, Laguna de Mitla, L. Lozada P. 418 (FCME, XAL); Laguna del Potosí, F. Menendez L. 448 (CAS); Mpio. Petatlán, 4 km de Petatlán dirección Tecpan, A. Nuñez 605 (MEXU, XAL); Mpio. José Azueta, Cerro Huamilule, en Barra el Potosí, 17°31ʹ56ʺN, 101°27ʹ08ʺW, S. Peralta 426 (MEXU, FCME); Mpio. Tecpan de Galeana, Nuxco, laguna, 17°15ʹ07.6ʺN, 101°49ʹ18ʺW, S. Peralta 444 (FCME). Jalisco: Puerto Vallarta, R. Acevedo R. 1487 (NY, XAL); Mpio. La Huerta, Laguna de Corte, 19°19ʹ00ʺN, 104°56ʹ20ʺW, G. Castillo C. et al. 10745 (MEXU); Mpio. La Huerta, Playa Tenacatita, 19°17ʹ00ʺN, 104°51ʹ50ʺW, G. Castillo C. et al. 10848 (MEXU); Mpio. Tomatlán, Playa Chalacatepec, 19°38ʹ50ʺN, 105°12ʹ20ʺW, G. Castillo C. et al. 10967 (MEXU, XAL); Mpio. Tomatlán, Laguna Xola, 19°43ʹ10ʺN, 105°15ʹ20ʺW, G. Castillo C. et al. 10978 (XAL); Salina al N de Chamela, M. González G. 152 (CAS); Mpio. La Huerta, La Manzanilla, L. de Puga 15542 (XAL); Barra de Navidad, J. Rzedowski 14605 (DS). Michoacán: Mpio. Aquila, Estero de Maquili, B. Guerrero C. 676 (XAL); Las Salinas [vic. of delta of Río Balsas, fide McVaugh 1951], E. Langlassé 146 (K); Mpio. Coahuayana, Boca de Apiza, C. Soto N. et al. 7116 (MEXU). Nayarit: Mpio. Bahía de Banderas, Bahía de Banderas, 20°47ʹN, 105°15ʹW, G. Castillo C. 5818 (MEXU); Mpio. Bahía de Banderas, Laguna del Quelele, 20°44ʹN, 105°18ʹW, G. Castillo C. 6012 (MEXU); Mpio. Bahía de Banderas, Club de Golf Flamingos, cerca de Bucerías, M. Cházaro B. & R. Romero 8472 (XAL); SE of San Blas through Matanchen, SE toward Río San Cristobal, C. Davidson 7608 (CAS); Pochote, Santiago Ixcuintla, 21°55ʹ20.1ʺN, 105°30ʹ30.9ʺW, DIAAPROY S.A. de C.V. 47383 (MEXU); Tres Marías Islands, Magdalena Island, F. Elmore 1132 (F); vicinity of San Blas, R. Ferris 5396 (DS); Mpio. Santiago Ixcuintla, Mezcaltitán, J. González O. 5537 (DS); Tres Marías Islands, Isla María Magdalena, H. Mason 1793 (CAS, F, K, NY); Mexcaltitlán, Y. Mexia 1008 (CAS, UC); Mpio. Santiago Ixcuitla, Isla de Mexcaltitán, 21°50ʹ36ʺN, 105°24ʹ42ʺW, A. Miranda & G. Villegas 2053 (MEXU); Mpio. San Blas, ca. 2 mi E of San Blas on Hwy. 54, D. Norris & D. Taranto 13329B (CAS); Mexcaltitán, J. Ortega 5537 (K); Isla María Magdalena, O. Solís 9 (MEXU). Oaxaca: Mpio. Santa María Huatulco, Estero Cacaluta, 15°43ʹ20ʺN, 096°09ʹ40ʺW, G. Castillo C. et al. 9782 (MEXU, XAL); Chacahua Bay, F. Elmore D21a (DS), D22 (UC); Distr. Juchitán, Mpio. Chahuites, camino Chahuites–Las Salinas, A. Flores M. 1101 (CAS); Mpio. Chahuites, Rancheria Trejo, M. García B. 486 (XAL); Mpio. Huamelula, 4 km por la carr. Pochutla–Salina Cruz, 4 km después Huamelula, J. García P. & E. Estrada L. 1974 (MEXU); Distr. Tehuantepec, Mpio. Salina Cruz, beach at La Ventosa, ca. 16°10ʹN, 095°09ʹW, R. Gereau & G. Martin 1921 (CAS); Laguna Superior, S of Juchitán, near Xandanl, R. King 1549 (NY, UC); Mpio. Tututepec, Chacahua, J. Magaña 1232 (XAL); Puerto Angel, C. Morton & E. Makrinius 2624 (K); La Ventosa Beach, ca. 6 mi E of Salina Cruz, A. Reznicek & D. Gregory 304 (NY); Distr. Tehuantepec, Mpio. Santiago Astata, Laguna Colorada, 4 km W de Zaachilac, 15°57ʹ39ʺN, 095°34ʹ40ʺW, S. Salas M. & E. Torres B. 825 (MEXU, XAL); Distr. Tehuantepec, Mpio. Salina Cruz, 500 m W de Salinas del Marquéz, 16°10ʹ6.9ʺN, 095°14ʹ24.3ʺW, S. Salas M. et al. 5596 (XAL); Distr. Tehuantepec, Mpio. San Pedro Huamelula, Rancho Paraiso, 15°51ʹ49ʺN, 095°50ʹ22ʺW, N. Velázquez R. et al. 265 (MEXU); Mpio. San Mateo, Huazantlán, La Salina, D. Zizumbo & P. Colunga 500 (MEXU). Quintana Roo: S de Punta Allen, 180 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 Cayo Cedros, Bahía de la Ascensión, E. Cabrera 3405 (CAS, MEXU); 10 km N de Puerto Morelos, camino a Punta Caracol, E. Cabrera & H. de Cabrera 3134 (MEXU, NY); 4 km N de la zona hotelera de Isla de Cozumel, camino a Isla de la Pasión, E. Cabrera & H. de Cabrera 13582 (MEXU); Isla Mujeres, camino al Puerto de Abrigo, E. Cabrera et al. 17200 (MEXU); Mpio. Isla Mujeres, Isla Mujeres, lado W de la marina, 21°19ʹN, 086°46ʹW, C. Chan et al. 1590 (XAL); Reserva Biósfera de Sian Ka’an, 6 km E de Ramonal, R. Durán et al. 1120 (MEXU); Mpio. Felipe Carrillo Puerto, La Laguna Xunyanche, 20°00ʹN, 087°40ʹW, J. Flores & E. Ucan 8352 (F); Mpio. Isla Mujeres, Isla de Contoy, 21°30ʹN, 086°49ʹW, J. Flores & E. Ucan 8855 (MEXU); Mpio. Othón P. Blanco, Cayo Centro en el Banco Chinchorro, 18°35ʹN, 87°20ʹW, J. Flores et al. 8959 (XAL); Cozumel Island, G. Gaumer 146 (K); Holbox Island, G. Gaumer s.n. (K); Cozumel Island, E. Goldman 653 (F, US); brecha a Punta Brava al S de Pto. Morelos, P. Moreno 843 (MEXU); Mpio. Isla Mujeres, Isla Mujeres, lado SE, 21°14ʹN, 086°46ʹW, A. Puch et al. 865 (XAL); Mpio. Cozumel, Isla Cozumel, 20°30ʹN, 086°58ʹW, A. Puch et al. 1073 (XAL); Mpio. Benito Juarez, camino a Punta Nizuc desde el entronque de la carretera Cancún-Chetumal, 21°N, 086°50ʹW, V. Rico-Gray 123 (MEXU); Isla Mujeres, N end, J. Sauer & D. Gade 3265 (MICH); Mpio. Lázaro Cárdenas, Chiquilá, 21°23ʹN, 087°23ʹW, E. Ucán 431 (XAL); Mpio. Isla Mujeres, atras del Puerto de Abrigo, zona del Sak Bajo, 21°15ʹN, 086°45ʹW, E. Ucan E. & J. Flores 1038 (MEXU, UC); Mpio. Othón P. Blanco, camino Blanco de X-Calak, rumbo a Majahual, 18°26ʹN, 087°56ʹW, E. Ucan E. et al. 620 (MEXU); Mpio. Felipe Carrilo Puerto, Vigía Chico, Reserva de la Biósfera Sian Ka’an, R. Villanueva 812 (MEXU). Sinaloa: Mpio. Escuinapa, Palmito, 9 km E al estero “Mezcal,” J. Beltrán M. 1014 (FCME); Mpio. Rosario, E edge of Mazatlán Bay, 0.7 mi W of Mex. 15, 8.6 mi N of Río del Presidio, D. Breedlove 1577 (DS); Estero Ballena, old channel of Río Fuerte (W of Los Mochis), R. Felger 8437 (CAS, MEXU); Topolobampo, A. Gibson & L. Gibson 2095 (FSU); Escuinapa, Arroyo de la Codojuiz, J. González O. 1131 (K); Mpio. Rosario, Coacoyolitos, J. González O. 6458 (CAS); 4 km W de El Toldo, 24°57ʹN, 107°57ʹW, V. Lopez S. 8.1 (MEXU); vicinity of Topolobampo, J. Rose et al. 13309 (NY); vicinity of Mazatlán, J. Rose et al. 14046 (F); N side of Topolobampo, D. Seigler & P. Richardson 11686 (MEXU); Mpio. Los Mochis, just W of Topolobampo, 25°35ʹN, 109°05ʹW, T. Van Devender et al. 2000-28 (NY). Sonora: Bahía Kino, mouth of Río de Sonora, F. Drouet & D. Richards 3542 (F); 18.5 mi N of Bahía Kino Nuevo, J. Hastings & R. Turner 64-35 (DS); Estero Tastiola, NW de Guaymas, O. Holguín s.n. (DS); Tepoca Bay, I. Johnston 3288 (CAS); beach S of Guaymas, G. Lindsay 1154 (DS); ca. 1 mi W of Puerto Lobos on narrow peninsula, C. Lowe & R. Turner 3319 (DS); Sargento, T. Mallery & W. Turnage s.n. (DS); Guaymas, W. Phillips 3483 (CAS) Mpio. Hermosillo, 2.7 km N of Punta Chueca, 20°02ʹ20ʺN, 112°10ʹW, A. Reina G. & T. Van Devender 96-635 (MEXU); Mpio. Caborca, Puerto Lobos, 30°16ʹ16ʺN, 112°51ʹ14ʺW, A. Reina et al. 97-265 (MEXU); vicinity of Guaymas, J. Rose et al. 12578 (NY); Bahía San Carlos, W of Guaymas, Weedons M-1091 (MEXU); bayshore at Empalme, I. Wiggins 6341 (DS); Isla Tiburón, Estero San Miguel, 28.968611°N, 112.20194°W, B. Wilder et al. 06-276 (CAS); Isla Tiburón, Punta Tormenta estero, 29°00ʹ51.17ʺN, 112°11ʹ54.21ʺW, B. Wilder et al. 08-329 (CAS); Isla Tiburón, Cyazim It, spit in estero at Punta Perla, 29.22442°N, 112.29345°W, B. Wilder et al. 08-377 (CAS). Tabasco: Mpio. Frontera, Playa Boquerón, 20 km E de Frontera, A. Guadarrama 876 (MEXU, NY); Mpio. Centla, Ejido Nuevo Centla, antes Playa Boquerón, 18°33ʹ10.3ʺN, 092°30ʹ48ʺW, M. Guadarrama O. et al. 6688 (MEXU, XAL); Mpio. Nacajuca, Laguna Bayazú, llegando por el Río Gonzalez, A. Hanan A. et al. 981 (MEXU); Mpio. Centla, Paso San Román, F. Ventura A. 20406 (MEXU, XAL); Mpio. Paraíso, Puerto Ceiba, Isla Dos Bocas, F. Ventura A. 20432 (XAL); Mpio. Paraíso, 4 km N de Mecoacán, S. Zamudio R. 117 (MEXU). Tamaulipas: Mpio. Altamira, Barra de Chavarria S, entrada por el Barranco, E. de Dunas 743 (XAL); 1 km W de La Pesca, cerca de la Laguna Blanca, DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 181 L. Hernández 1567 (MEXU); Mpio. Aldama, Rancho Nuevo, 23°08ʹ12ʺN, 097°46ʹ01ʺW, D. Infante et al. 445 (XAL); Mpio. Matamoros, delta del Río Bravo, 25°56ʹ56ʺN, 097°09ʹ07ʺW, D. Infante & J. Vázquez 656 (XAL); Mpio. Soto la Marina, campamento totuguero La Pesca, 23°47ʹ27ʺN, 97°44ʹ12ʺW, E. Martínez 39316 (MEXU); Mpio. Altamira, playa cerca del Puerto Industrial, A. Mora O. & J. Mora L. 5450 (MEXU); Mpio. Altamira/Aldama, Barra de Chavarria, P. Moreno C. et al. 743 (MEXU); vicinity of Tampico, E. Palmer 484 (CAS F, K, NY). Veracruz: Puente de Alvarado, J. Calzada 434 (F); Mpio. Coatzacoalcos, Laguna Ostión, camino Pajapan– San Juan Volador, 18°11ʹN, 94°36ʹW, J. Calzada 12657 (MEXU, XAL); Mpio. Panuco, alrededores de Laguna de Tamos, 22°13ʹN, 098°02ʹW, J. Calzada et al. 6268 (XAL); Mpio. Actopan, El Morro de La Mancha, La Laguna, 19°36ʹN, 096°24ʹW, G. Castillo C. 182 (F, UC, XAL); Mpio. Cosoleacaque, Polvorín, carretera Polvorín–Cosoleacaque, 17°59ʹ48ʹN, 094°38ʹ12ʺW, G. Castillo C. et al. 14838 (MEXU, XAL); Mpio. Agua Dulce, Río Tonala, cerca de Arroyo Blasillo, M. Chazaro B. 3299 (XAL); Mpio. Actopan, Estación Biológica El Morro de la Mancha, 19°36ʹ00ʺN, 096°22ʹ40ʺW, C.M.V.A. 2 (MEXU); Mpio. Cazones, Rancho Nuevo, Estero Boquilla, M. Cortés 455 (XAL); Mpio. Actopan, Laguna de la Mancha, carretera Cardel–Nautla, J. Dorantes 57 (MEXU); S de Laguna Salada, J. Dorantes et al. 1053 (F); 16 km S de Palma Sola, Laguna del Farallon [19°39ʹ19.58ʺN, 96°24ʹ40.28ʺW], J. Dorantes et al. 1171 (CAS); alrededores de Laguna Verde (SW de La Planta), Alto Lucero, J. Dorantes et al. 5132 (NY); Mpio. Alto Lucero, Lugana de San Agustín, KM 71 carretera Cardel–Nautla, 19°55ʹN, 096°31ʹW, C. Guttierez B. 1311 (MEXU); Mpio. Zempoala, Estación de Biología “El Morro de la Mancha,” INIREB, carretera Cardel–Nautla, G. Ibarra M. 3b (FCME); borde sur de la Laguna Salada, A. Lot et al. 2055 (F); Mpio. Boca del Río, Mandinga, 19°03ʹ05.52ʺN, 96°04ʹ43.56ʺW, F. Medina H. et al. 27 (MEXU, XAL); Laguna de Tampamochoco, cerca de Tuxpan, A. Mendoza s.n. (DS); Mpio. Tuxpan, 8 km de Barra de Tuxpan, L. Monroy et al. 147 (CAS, XAL); Mpio. Actopan, 50 m SW de la boca de la Laguna de la Mancha, 19°35ʹN, 096°22ʹW, A. Novelo 408 (XAL); Mpio. Alvarado, Laguna de Alvarado, 18°46ʹ13ʺN, 095°45ʹ38ʺW, R. Palestina et al. 1460 (XAL); Laguna de Sontecomapan, T. Pennington & J. Sarukhan K. 9131 (NY, K); Mpio. Coatzacoalcos, terracería La Barrillas– Laguna Ostión, 17°45ʹ30ʺN, 094°42ʹ07ʺW, A. Rincón G. et al. 1731 (XAL); Barra de Tuxpan (20°58ʹN), N of Río de Tuxpan mouth, J. Sauer & D. Gade 2981 (F); Río Coscoapan, M. Sousa 3112 (F); Mpio. Cazones, Barra Cazones, S. Vargas P. 90 (XAL); Mpio. Actopan, La Mancha, F. Ventura A. 5226 (CAS). Yucatán: Mpio. Tizimín, 55 km en el camino a Las Coloradas, cerca el Puente del Río Lagartos, J. Aguilar Z. & S. Diez M. 225 (MEXU); Mpio. Telchac, Laguna Rosa, 1 km de Puerto de Telchac, 21°20ʹN, 89°16ʹW, J. Calzada et al. 6611 (F, XAL); Peña, Chocarro & Jun 567 (BIGU); Mpio. Tizimín, 16 km E de Las Coloradas, 21°30ʹ40ʺN, 87°50ʹ15ʺW, R. Durán et al. 2575 (MICH); Mpio. Hunucmá, 7.5 km E de Sisal hacia Celestún, E. Estrada 283 (FCME); Mpio. Telchac Puerto, 2 km E de Telchac Puerto, A. Feliciano K. 325 (MEXU); Mpio. Progresso, Isla Larga de los Arrecifes Alacranes, 22°26ʹN, 089°31ʹW, J. Flores & E. Ucan 9253 (XAL); Alacran Atoll, S end of Perez Islet, F. Fosberg 41866 (NY, US); Alacran Atoll, Pajaros Islet, F. Fosberg 41904 (US); Las Bocas de Silam, G. Gaumer et al. 23340 (F, NY, US); Progreso, C. Lundell & A. Lundell 8140 (MEXU, MICH, NY); Mpio. Dzilam de Bravo, entre Santa Clara and Dzilam de Bravo, 21°25ʹN, 088°50ʹW, J. Palma & R. Allkin 300 (MEXU); 6 km W de Dzilam de Bravo, brecha a Pto. Telchac, H. Quero R. & R. Grether 2469 (MEXU); Mpio. Hunucmá, 2 km E de Sisal, 21°10ʹ15ʺN, 090°00ʹ45ʺW, E. Reyes de los Santos 607 (MEXU); Sisal, Schott 361 (F); Celestun, Schott 473 (F); Mpio. Celestún, 8 km antes Celestún, viniendo de Kinchil, 20°53ʹN, 090°20ʹW, V. Rico-Gray 60 (F, MEXU, UC, XAL); ca. 4 km de Sisal, viniendo de Hunucmá, Hunucmá, 21°13ʹN, 090°03ʹW, V. Rico-Gray 75 (MEXU); Mpio. Progreso, 1.5 km S de Chelem, camino a Progreso, 21°15ʹN, 090°20ʹW, V. Rico-Gray 87 (MEXU); 1.5 km E de Dzilam de Bravo, 21°25ʹN, 182 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 088°50ʹW, V. Rico-Gray 95 (F, UC, XAL); Mpio. Telchac Puerto, 0.5 km S de Telchac Puerto, 21°20ʹN, 089°15ʹW, V. Rico-Gray 101 (F, MEXU, UC, XAL); afueras de la cuidad de Río Lagartos, 21°35ʹN, 088°10ʹW, V. Rico-Gray 106 (F, MEXU, XAL); between Progresso and Telchac Puerto, J. Sauer & D. Gade 3207 (F); Progresso, W. Steere 3092 (MICH); Mpio. Celestún, 1 km E de Celestún, 20°51ʹ30ʺN, 090°24ʹ00ʺW, J. Tapia M. 1508 (MEXU); Tizimín, alrededores de El Cuyo, ca. 21°30ʹ45ʺN, 087°40ʹ46ʺW, M. Ventura 158 (F); Alacran Reef, Isla Perez, S end of island, B. Welch s.n. (DUKE, MEXU). NICARAGUA. León: Poneloya, ca. 12°23ʹN, 087°03ʹW, R. Haynes 8617 (NY); 7.1 km de Las Peñitas, orillas del Estero Las Peñitas, Sediles 408 (K); Estero Las Peñitas, 3.1 km de Las Peñitas, 12°20ʹN, 086°59ʹW, Sediles 413 (CAS), 443 (CAS); Estero Brasil, ca. 2 km S of Hwy. 32 on road to Velero, ca. 12°10ʹN, 086°45ʹW, D. Stevens et al. 17293 (CAS). Managua: Masachapa, J. Atwood & D. Neill AN32 (NY). Region Autonomista Atlantico Norte (northern Zelaya): Pozo Verde, 10 km NE de Puerto Cabezas, 14°06ʹN, 083°20ʹW, E. Little 25401 (F, US). Region Autonomista Atlantico Sur (southern Zelaya): Bluefields harbor, El Bluff, S. Marshall & D. Neill 6507 (USF). Rivas: Estero San Juan del Sur, 11°14–16ʹN, 085°51–53ʹW, M. Araquistain 3809 (CAS); San Juan del Sur, F. Seymour 1269 (MEXU, NY, UC). PANAMA. Bocas del Toro: Changuinola Valley, C. Cooper & G. Slater 81 (US); Water Valley, H. von Wedel 987 (US). Chiriquí: playa cerca del KM 3, J. Him 358 (US). Coclé: Aguadulce, outskirts of tidal belt, H. Pittier 4969 (US). Colón: vicinity of Colón, J. Cowell 97 (NY); vicinity of Viento Frio, H. Pittier 4116 (US). Guna Yala: trail from Cangandi to dock by Mandinga airport, [09°27ʹ39.50ʺN, 079°05ʹ0.70ʺW], G. de Nevers et al. 6554 (CAS, US). Herrera: P.N. Sarigua, 08°00ʹ41ʺN, 080°29ʹ03ʺW, I. Alvarez B4339 (US). Los Santos: Salinas de Chitre, W. D’Arcy & T. Croat 4200 (F). Panamá: Perlas Archipelago, San José Island, Naval Cove, C. Erlanson 120 (NY); Perlas Archipelago, San José Island, I. Johnston 1129 (US); Isla San José, H. Kennedy 2281 (F, US); Taboga Island, Gulf of Panama, H. Pittier 3614 (NY, US); Miraflores Locks, W. Stern et al. 50 (US). U.S.A. Florida: Brevard Co.: Merritt Island, R. Kral 4972 (FSU, UC); S end of Merritt Island, Banana River, Coquina, R. Whetstone 9116 (MO). Broward Co.: along Dania Beach Blvd., ca. 2 mi W of beach, S. Leonard 6924 (FSU). Charlotte Co., 4.2 km S of De Soto Co. along Peace River, 2.7 km W of US 17, 26°59ʹ50ʺN, 081°59ʹ10ʺW, A. Franck & B. Upcavage 1866 (USFimage!). Citrus Co.: Shell Island, near mouth of Crystal River, R. Long 1309 (USF). Collier Co.: near Naples, R. Godfrey 58071 (FSU); Everglades City, O. Lakela 29824 (NY). Flagler Co.: inland waterway in vicinity of Marineland, R. Godfrey 61686 (FSU). Franklin Co.: between St. George Sound and small tidal marsh just NE of Culpepper home on Cannonball Acres, L. Anderson 5580 (FSU). Hernando Co.: N of Aripeka on Fla. 595, J. Carlton s.n. (USF). Hillsborough Co.: Long Key, F. Lewton s.n. (NY). Indian River Co.: along Indian River Lagoon, Oslo Riverfront East Conservation Area, Oslo Road E of US 1, ca. 3 mi. S of Vero Beach, 27º35.193ʹN, 080º21.902ʹW, S. Myers 1281 (USF-image!). Lee Co.: Tarpon Bay, eastern Sanibel, W. Brumbach 7904 (NY); Little Pine Island, H. Moldenke 929 (MO). Levy Co.: causeway to Cedar Key, R. Godfrey & P. Redfearn 52828 (UC); near jct. A Street and 3rd Street, Cedar Key, K. Murray 80-43-10 (NY); Cedar Keys, I. Wiggins 19342 (DS); E side of Seahorse Key, SE of lighthouse, I. Wiggins & D. Wiggins 19430 (DS). Manatee Co.: Palmetto, G. Nash 2450 (MO); Palma Sola, vic. of Manatee, J. Simpson 80 (UC). Martin Co.: Jonathan Dickinson State Park (Girl Scout Camp), R. Woodbury & R. Roberts s.n. (USF). Miami-Dade Co.: Coral Gables, S end of Vee Lake, 25°40.5–40.7ʹN, 080°16.1– 16.5ʹW, J. Abbott 24063 (FLAS-image!); Everglades Natl. Park, Flamingo Area, 25°08.38ʹN, 080°55.88ʹW, W. Hess et al. 8579 (MO, NY); Kampong, 4013 Douglas Road, Coconut Grove, W. Judd 5602 (FLAS-image!). Monroe Co.: Long Key, D. Correll & H. Correll 40150 (MO); Rode DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 183 Harbor, Key Largo, C. Janish & J. Janish 447 (DS, MO); Big Pine Key, W. Muenscher & R. Thorne 18073 (UC); Lower Matecumbe Key, J. Pruski et al. 2826 (NY); Key West, just S of airport, 24°33.162ʹN, 081°46.024ʹW, A. Salywon 1188 (CAS); E end of Packet Key, I. Wiggins 20081 (DS). Palm Beach Co.: Jupiter Island, G. Cooley et al. 4864 (USF). Pasco Co.: just W of Port Richey along inlet canal from beach, J. Ray et al. 9980 (FSU). Pinellas Co., Boca Ciega, bayside just S of Treasure Island causeway, R. Thorne 48355 (UC). Sarasota Co.: Sarasota, Marie Selby Gardens, 27°19ʹ33ʺN, 082°32ʹ28ʺW, H. Bizet 51 (MO, NY); Historic Spanish Point, in Osprey W of US 41, 27º12ʹ15ʺN, 082º29ʹ46ʺW, M. Nolan 62 (USF). St. Johns Co.: between Matanzas and Marineland, R. Godfrey 70653 (CAS, FSU, GA, UC); Anastasia State Park, Conch Island, ca. 4.75 km NNW of jct. FL A1A and FL 312, E of FL A1A; immediately S of the St. Augustine Inlet, 29º53ʹ56ʺN, 081º17ʹ21ʺW, J. Kunzer et al. 2146 (USF-image!); Crescent Beach, D. Seigler & D. Young 10211 (MEXU); Anastasia State Recreation Area, NW Conch Island, S of Vilano Point and St. Augustine Inlet, UTM-471684, 3308353, C. Slaughter et al. 16617 (FSU); Tolomato River, 30°06.618ʹN, 081°22.303ʹW, A. Williams & S. Eastman s.n. (USF-image!). St. Lucie Co.: Hutchinson Island, Blind Creek access area, off Indian River, G. Silberhorn s.n. (USF). Taylor Co., Jug Island [29°50ʹ31.75ʺN, 83°36ʹ58.05ʺW], R. Godfrey 60403 (UC). Volusia Co.: 9 mi S Daytona Beach, near Ponce de Leon Inlet, R. Norris 541 (FSU). County undetermined: Indian River, A. Curtiss 1972 (CAS, GA, MO), s.n. (NY); Tampa Bay, P. Rolfs 248 (MO). Louisiana: Cameron Parish: Monkey Island near mouth of Calcasieu Ship Channel at Calcasieu Pass, adjacent to ferry landing, W. Vermillion s.n. (LSU-image!). Jefferson Parish: Grande Isle, J. Carlton s.n. (USF); Fifi Island, 29.255541°N, 089.978158°W, D. Atha 12910 (NY). Lafourche Parish: 0.6 mi S of Fourchon Road bridge (R22E, T23S, S24), A. Lasseigne 6146 (MEXU); S of end of La. 3090, S of Fourchon City, S of Leeville, R. Thomas et al. 103237 (MO, NY). Orleans Parish: New Orleans, Nuttall s.n. (K). Plaquemines Parish: ca. 40 mi (air) SSE of New Orleans, 29.40782, -89.79907, M. Bell s.n. (LSUimage!). St. Bernard Parish: North Islands in North Chandeleur Sound, NE of Venice, R. Thomas et al. 89768 (MEXU). Terrebonne Parish: E end of Isle Dernier, F. Givens 3733 (MO); Brush Island, F. Lloyd & S. Tracy 249 (NY). Mississippi: Harrison Co., Bay St. Louis [possibly Hancock Co.?], C. Sanger s.n. (NY); Cat Island, 30.23037°N, 89.08532°W, Scheffel et al. 2014 (living plant-image!). Jackson Co.: Ranger Lagoon, Horn Island, 30.24171°N, 88.67886°W, Scheffel et al. 2014 (living plant-image!). Texas: Aransas Co.: Redfish Bay, causeway to Port Aransas, Rte. 361, S. Hill 18296 (MO, NY); Port Aransas, B. Tharp 253 (CAS, MO, NY, UC). Cameron Co.: South Padre Island, between Old Causeway and Queen Isabella Causeway, F. Banda 81 (USF); Padre Island, just E of Port Isabel, D. Correll et al. 25539 (UC); 8 mi SW of Port Isabel, inlet crossed by FR 1792 (NY), J. Crutchfield 2985 (NY); Clark Island, near Boca Chica, C. Lundell & A. Lundell 8760 (CAS, NY, UC); Point Isabel, H. Parks 2939 (MO); bay at Boca Chica, Brazos Santiago Island, R. Runyon 2812 (NY); Point Isabel in Lower Rio Grande Valley, R. Runyon 5897 (UC); boca de Río Bravo, Schott 139 (NY); Boca Chica, G. Webster & R. Wilbur 3035 (GA). Nueces County: Aransas Pass, causeway between Aransas Pass and Port Aransas, P. Fryxell 5162 (MEXU). Refugio Co.: near Tivoli, J. Williams 415 (NY). 3. Avicennia marina (Forsk.) Vierh., subsp. australasica (Walp.) J. Everett, Telopea 5(4): 628. 1994. Avicennia tomentosa Sieber var. australasica Walp., Repert. Bot. Syst. 4: 133. 1845. Avicennia resinifera G. Forst., Pl. Esc. 72. 1786. Avicennia marina var. resinifera (G. Forst.) Bakh., Bull. Jard. Bot. Buitenzorg, ser. 3, 3: 210. 1921, nom illegit. (superfl.). TYPE.— Sheet 1460 in Thunberg’s herbarium (lectotype, designated by Everett [1994: 628]: UPS). Figures 1H–K, 2, 3B,D Shrubs to 2.3 m tall. Young stems of reproductive shoots covered with dense shiny granules or 184 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 scalelike projections to 0.05 m long (scurfy), soon glabrate. Leaves petiolate, blades ovate to elliptic (to obovate), 44–100 mm long, 19–41 mm wide, 1.7–2.8 (–3.6) times longer than wide, acute (to rounded or emarginate) at apex, subattenuate to attenuate at base, surfaces discolorous (abaxial lighter), punctate-pitted (sometimes inconspicuously so abaxially), adaxial surface lacking trichomes, abaxial surface covered with a dense scurfy layer. Inflorescences of axillary and terminal (sessile to) pedunculate ± headlike spikes, peduncles (0–) 1–40 mm long, scurfy or distally pubescent like rachis, rachis not or but barely visible, internodes near midspike 1–4 mm long, scurfy and pubescent with ± antrorse eglandular trichomes to 0.2 mm long. Bracts opposite, broadly ovate to triangular, concavoconvex, 3–4 mm long, abaxial surface scurfy and often pubescent like rachis. Bracteoles similar to bracts except smaller. Flowers mostly 4–16 per spike, sessile. Calyx 3.5–4 mm long, lobes elliptic to broadly elliptic, concavoconvex, imbricate, abaxially pubescent with antrorsely appressed eglandular trichomes to 0.8 mm long, margin ciliate with similar but spreading trichomes. Corollas 3.5–6.5 mm long, internally drying dark or blackish proximally and light brownish distally (those from Australasia are usually described as yellowish or orangish and the color is often darker in the corolla tube), externally glabrous (tube and base of lobes) and densely pubescent with appressed eglandular trichomes to 0.2 mm long (remainder of lobes), tube 1.5–2 mm long, limb actinomorphic, 4-lobed, lobes ovate-triangular to ovate-elliptic, 2–4.5 mm long, apically entire (or 1 lobe sometimes slightly bifid apically with division to 0.2 mm long), internally lacking eglandular trichomes (at least distally) but sometimes punctate-pitted (proximally). Stamens 4, inserted in distal half of corolla tube near base of lobes, exserted from mouth of corolla tube, oriented symmetrically (i.e., equally distant from one another) around corolla with thecae opening toward central gynoecium, 1.5–2 mm long, filaments 0.5–0.8 mm long, anthers presented at same height, thecae 1–1.2 mm long; pollen prolate spheroidal to euprolate, polar diameter (P) 28–41 µm, equatorial diameter (E) 24–27 µm, P:E = 1.04–1.75. Style not evident, stigma lobes 0.2 mm long. Fruit ovoid to subellipsoid, proximally blackish and distally light brownish when dry, 15–24 mm long, 10–19 mm across at widest expanse, pubescent with erect to flexuose to antrorse eglandular trichomes to 0.3 mm long (especially when less mature) and scurfy (especially evident when more mature). 2n= 64, 96 (Dawson 1989). PHENOLOGY.— Flowering: February, August–September; fruiting: February, November. DISTRIBUTION AND HABITAT.— Avicennia marina has the most extensive distribution among species in the genus; it is native to eastern Africa, southern Asia, Indian Ocean and western Pacific Ocean islands, and Australia. Subspecies australasica occurs primarily in subtropical and temperate Australasia (i.e., southeastern Australia and northern New Zealand). It is the southernmostoccurring taxon among species of Avicennia in the Old World (to 38°45’S; Duke 2006), and the southernmost-occurring mangrove in the world. In southern California, where this taxon has been introduced and become naturalized (Fig. 2), plants occur in salt marshes with Batis, Juamea, Salicornia, Spartina, and Suaeda at or near sea level. ILLUSTRATIONS.— Munir (1986: 1179, fig. 546); Duke (1991: 314, fig. 7); Clarke and Myerscough (1991: 285, fig. 1). NOMENCLATURE.— The name “A. marina var. australasica (Walp.) Moldenke” has been used for this taxon, but as discussed by Everett (1994), this combination was not validly published at this rank, and other infraspecific taxa of A. marina are currently treated as subspecies. Avicennia marina subsp. australasica and the synonyms noted above are all based on A. resinifera. See Moldenke (1960) and Duke (1991) for a full list of synonyms of Avicennia marina. LOCAL NAME.— Gray mangrove. CONSERVATION.— Avicennia marina has been assessed as a taxon of Least Concern (LC) by Duke et al. (2010). This taxon is not native in the New World, but has become naturalized locally DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 185 and is potentially invasive. On his collection 28024 made in 1979, Moran noted that plants in California had been introduced from Aukland, New Zealand about 1966–69. He also noted the presence of about 100 or more flowering-size plants plus many seedlings in the wildlife reserve where his observations were made. Initial efforts to eradicate the species were unsuccessful. DISCUSSION.— This species is readily distinguished from those native to the western Hemisphere by its actinomorphic flowers with yellowish to orangish corollas bearing ovate-triangular to ovate-elliptic lobes and its equidistant stamens that are inserted near the base of the corolla lobes, dehisce toward the center of the flower, and vary from 1.5–2 mm in length (Fig. 3B). Duke et al. (1998) provided genetic evidence that supported the morphological recognition of A. marina as a distinct species and that supported recognition of the three infraspecific taxa (treated by him as varieties, but here recognized as subspecies): subsp. marina, subsp. eucalyptifolia (Valeton) J. Everett, and subsp. australasica. The latter subspecies would appear to consist of or contain polyploids based on the reported chromosome numbers of 2n= 64 and 96 (Dawson 1989, as A. resinifera). These numbers suggest a possible base number of x = 8 or x = 16. A chromosome number of 2n= 36 was reported for an unspecified subspecies of A. marina by Subramanian (1988; without citation of voucher). These appear to be the only recent chromosome counts for both this species and for the genus. If these numbers are accurate, both polyploidy and dysploidy would appear to have played a role in the evolution of taxa in A. marina. The only other known chromosome counts for Avicennia are 2n= ca. 66 and n= ca. 33 by Raghavan and Arora (1958; with a meiotic figure showing n= 33, but without citation of a voucher) for A. alba Bl. Sanders (1997) indicated a base chromosome number for Avicenniaceae of x= 18. Although no rationale was stated for this number, his conclusion was probably based largely on Subramanian’s count of 2n= 36 for A. marina. Avicennia marina subsp. australasica was distinguished by Duke (1991) from the other two varieties of A. marina by the fully (or nearly so) pubescent calyces (vs. pubescent only near the base), and the gray, fissured (vs. green, chalky smooth, and often flaky in patches) bark of the mature trunk. SPECIMENS EXAMINED.— U.S.A. California: San Diego Co.: Northern Wildlife Preserve, North Mission Bay, tidal area 100 m W of Rose Creek, 32.7949°N, 117.2247°W, I. Kay 29 (UCR); E edge of Kendall/Frost Marsh, Mission Bay, San Diego, ca. 32˚47.5ʹN, 117˚13.8ʹW, sea level, 9 September 1979, R. Moran 28024 (CAS, GH, MEXU, NY, UC, US); same locality, 12 June 1990, R. Moran 31036 (CAS, JEPS). ACKNOWLEDGMENTS I thank Tom Davis, 2015 intern in biological illustration at the California Academy of Sciences, for the fine illustration of the black mangrove; photographers (M. Fagg and B. Petersen) for making their images available; and Judy Gibson (SD), Layla Hains (SD), Carlos Parra (COL), Rebecca Peters (CAS), Jon Rebman (SD), and Dagoberto Rodríguez (LAGU) for supplying images of pertinent herbarium specimens. The following herbaria generously provided access to their collections, which served as the basis for this study: BM, CAS, DS, DUKE, F, FCME, FLAS, FSU, GA, GH, JEPS, K, MEXU, MICH, MO, NY, P, UC, UCR, US, USF, XAL. 186 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Series 4, Volume 63, No. 5 LITERATURE CITED ALVAREZ C., N.S. 2008. Avicennia germinans (L.) L. In D. Louppe, A.A. Oteng-Amoako, and M. Brink, eds., Prota 7(1): Timbers/Bois d’oeuvre 1. Wageningen, Netherlands. <http://database.prota.org/PROTAhtml/ Avicennia%20germinans_En.htm> [accessed 31 January 2016]. AYMARD, G. 2015. Avicennia. In R. Bernal, S.R. Gradstein and M. Celis, eds., Catálogo de plantas y líquenes de Colombia. Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá, Colombia. <http://catalogoplantascolombia.unal.edu.co> [accessed 24 December 2015]. BURKILL, H.M. 1985. The Useful Plants of West Tropical Africa, vol. 1. Royal Botanic Gardens, Kew, UK. BORG, A.J., AND J. SCHöNENBERGER. 2011. Comparative floral development and structure of the black mangrove genus Avicennia L. and related taxa in the Acanthaceae. International Journal of Plant Sciences 172:330–344. BORG, A.J., L.A. MCDADE, AND J. SCHöNENBERGER. 2008. Molecular phylogenetics and morphological evolution of Thunbergioideae (Acanthaceae). Taxon 57:811–822. BREEDLOVE, D.E. 1986. Listados Florísticos de México IV. Flora de Chiapas. Instituto de Biología, UNAM, Cd. México, Mexico. CAVANAUGH, K.C., J.R. KELLNER, A.J. FORDE, D.S. GRUNER, J.D. PARKER, W. RODRIGUEZ, AND I.C. FELLER. 2014. Poleward expansion of mangroves is a threshold response to decreased frequence of extreme cold events. Proceedings of the National Academy of Sciences 111:723–727. CERóN-S., I., E.G. GONZALEZ, A.E. SCHWARZBACH, D.E. SALAS L., E. RIVERA O., N. TORO P., E. BERMINGHAM, AND W.O. MCMILLAN. 2015. Contrasting demographic history and gene flow patterns of two mangrove species on either side of the Central American Isthmus. Ecology and Evolution 5:3486–3499. CHEATHAM, S., M.C. JOHNSTON, AND L. MARSHALL. 2000. The Useful Wild Plants of Texas, vol. 2. Useful Wild Plants, Austin, Texaqs, USA. CHIVIAN, E., AND A. BERNSTEIN. 2008. Sustaining Life. Oxford University Press. New York, New York, USA. CINTRóN, M., G. AND Y. SCHAEFFER N. 1992. Ecology and management of New World mangroves. Pages 233– 258 in U. Seeliger, ed., Coastal Plant Communities of Latin America. Academic Press, San Diego, California, USA. CLARKE, P.J., AND P.J. MYERSCOUGH. 1991. Floral biology and reproductive phenology of Avicennia marina in south-eastern Australia. Australian Journal of Botany 39:283–293. COMPèRE, P. 1963. The correct name of the Afro-American black mangrove. Taxon 12:150–152. CORRELL, D.S., AND H.B. CORRELL. 1982. Flora of the Bahama Archipelago. J. Cramer, Vaduz, Liechtenstein. COSTANZA, R., R. D’ARGE, R. DE GROOT, S. FARBER, M. GRASSO, B. HANNON, K. LIMBURG, S. NAEEM, R.V. OʹNEILL, J. PARUELO, R. RASKIN, P. SUTTON, AND M. VAN DEN BELT. 1997. The value of the world’s ecosystem services and natural capital. Nature 387:253–260. DANIEL, T.F., AND E. FIGUEIREDO. 2009. The California Academy of Sciences Gulf of Guinea Expeditions (2001, 2006, 2008) VII. Acanthaceae of São Tomé and Príncipe. Proceedings of the California Academy of Sciences, ser. 4, 60:623–674. DAWSON, M.I. 1989. Contributions to a chromosome atlas of the New Zealand flora—30 miscellaneous species. New Zealand Journal of Botany 27:163–165. DUKE, N.C. 1991. A systematic revision of the mangrove genus Avicennia (Avicenniaceae) in Australasia. Australian Systematic Botany 4:299–324. DUKE, N.C. 2006. Australia’s Mangroves. University of Queensland, Brisbane, Australia. DUKE, N. 2010. Avicennia bicolor. The IUCN Red List of Threatened Species 2010: e.T178847A7625682. <http://dx.doi.org/10.2305/IUCN.UK.2010-2.RLTS.T178847A7625682.en> [accessed 25 January 2016]. DUKE, N.C., J.A.H. BENZIE, J.A GOODALL, AND H.E.R. BALLMENT. 1998. Genetic structure and evolution of species in the mangrove genus Avicennia (Avicenniaceae) in the Indo-west Pacific. Evolution 52: 1612–1626. DUKE, N., K. KATHIRESAN, S.G. SALMO III, E.S. FERNANDO, J.R. PERAS, S. SUKARDJO, T. MIYAGI, J. ELLISON, N.E. KOEDAM, Y. WANG, J. PRIMAVERA, O. JIN EONG, J. WAN-HONG YONG, AND V. NGOC NAM. 2010. Avicennia marina. The IUCN Red List of Threatened Species 2010: e.T178828A7619457. <http://dx.doi.org/10.2305/IUCN.UK.2010-2.RLTS.T178828A7619457.en> [accessed 1 February 2016]. DANIEL: AVICENNIA IN NORTH AMERICA AND MESOAMERICA 187