Avicennia germinans ((Linnaeu))

Daniel, Thomas F., 2016, Avicennia (Acanthaceae: Avicennioideae) in North America and Mesoamerica, Proceedings of the California Academy of Sciences 63 (5), pp. 163-189 : 171-183

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.

Avicennia tomentosa Jacq., Enum. Syst. Pl. 25. 1760. Hilairanthus tomentosus (Jacq.) Tiegh, J. Bot. (Morot) 12: 358. 1898. TYPE.— See discussion.

Avicennia tomentosa Jacq. var. campechensis Kunth , Nov. Gen. Sp. 2: 229 (ed. folio); 284 (ed. quarto). 1818 (“1817”). TYPE.— MEXICO. Campeche: “Crescit prope Campeche Mexicanorum,” F. Humboldt & A. Bonpland s.n. (holotype: P-P00670135-image!).

Avicennia floridana Raf. , Atlantic J. 1: 148. 1832. TYPE.— Not designated (no specimens cited).

Avicennia floridana Gand., Bull. Soc. Bot. France 65: 64. 1918, nomen illegit. TYPE.— U.S.A. Florida: Lee County, Fort Meyers , A. Hitchcock 270 (see discussion).

Avicennia oblongifolia Chapm., Fl. South. U.S. 310. 1860. TYPE.— U.S.A. Florida: Monroe County, Oct , Key West (fide protologue); pertinent specimens, if extant, would likely be in the Chapman herbarium at NY, but none have been located.

Avicennia tonduzii Moldenke, Phytologia 1: 273. 1938. TYPE.— COSTA RICA. Puntarenas: Punta Mala [ca. 09°03ʹ54.48ʺN, 083°39ʹ04.84ʺW], zone littorale du Pacifique, Mar 1892, A. Tonduz 6776 (holotype: BRimage!, fragment of holotype at NY!; isotypes: BM-image!, M-image!, MICH-image!, US!).

Shrubs to 5 m tall or trees to 20 (–25) m tall. Young stems of reproductive shoots covered with dense ± antrorsely appressed or matted whitish and shiny trichomelike or scalelike projections to 0.05 mm long (scurfy) and sometimes also with longer appressed eglandular trichomes, the internodes infrequently glabrate. Leaves petiolate, blades narrowly to broadly elliptic (to obovate), 25– 160 (–185) mm long, 11–50 (–63) mm wide, (1.6–) 2.7–6.9 × longer than wide, acute to rounded to emarginate at apex, 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 includes ± appressed longer eglandular trichomes or sparsely to densely scurfy without longer trichomes. Inflorescences of axillary and terminal pedunculate densely bracteate (± headlike) spikes or panicles of these, inflorescence branches (if present) subtended by triangular- to oblate- to subovateconcave inflorescence bracts to 5 mm long (sometimes caducous), peduncles (5–) 10–50 mm long, scurfy, pairs of flowers congested at or toward apex of rachis, inflorescence internodes mostly 0.8– 4 mm long near midspike, rachis not or only partially or barely visible, densely pubescent with mostly antrorse eglandular trichomes to 0.2 mm long. Bracts triangular to oblate to broadly ovate to subcircular, 1–4 mm long, abaxial surface scurfy to pubescent like rachis. Bracteoles similar to bracts. Flowers 8–14 (–26) per spike. Calyx 2–4 mm long, lobes broadly ovate to subcircular, abaxially pubescent with antrorsely appressed eglandular trichomes to 0.4 mm long. Corollas (3.5–) 4–8 mm long, internally white to cream with yellow in throat, externally glabrous (proximal portion of tube) and densely pubescent with appressed eglandular trichomes to 0.5 mm long (distal portion of tube and limb), tube (1.5–) 2–3. 5 mm long, limb ± 2-labiate with 4 or 5 lobes, upper lip bifid to ± conspicuously 2-parted at apex (with division up to 3 mm long) and/or wider than lobes of lower lip, all lobes oblong to obovate, densely pubescent (at least on distal half if not throughout) internally. Stamens 4, inserted at midpoint or in proximal half of corolla tube, exserted from mouth of corolla tube, oriented in pairs near upper lip of corolla with thecae opening toward lower lip, 3–5 mm long, filaments 2. 2–4 mm long, anthers presented at ± same height, thecae 0.6– 1 mm long; pollen oblate spheroidal to euprolate, polar diameter ( P) 23–46 µm, equatorial diameter ( E) 24–29 µm, P: E = 0.92–1.90. Stigma lobes (0.2–) 0.4– 0.7 mm long. Fruit ovoid to broadly elliptic to obovoid, usually grayish (rarely black) when dry, 12–32 mm long, 9–25 mm across at widest expanse, pubescent with antrorsely appressed eglandular trichomes to 0.7 mm long (especially when less mature) and scurfy (especially when more mature).

PHENOLOGY.— Flowering: throughout the year; fruiting: April–January (especially in August, and likely throughout the year).

DISTRIBUTION AND HABITATS.— Southern North America (southeastern USA and Mexico), Bermuda, West Indies, Central America, South America ( Brazil, Colombia, Ecuador [including Galapagos Islands], French Guiana, Guyana, Peru, Surinam, and Venezuela), and tropical western Africa (see Daniel and Figuerido [2009] for distributional information on African plants, which often have been referred to as A. africana P. Beauv. ). Figure 2 shows the generalized distribution of this species in North America and Mesoamerica. Plants occur in/on tidal flats, beaches, coastal mangrove swamps (mangals), salt marshes, sand dunes, and coastal grasslands and savannas at elevations at or near sea level. Common associates include Allenrolfea occidentalis , Batis maritima , Bravaisia berlandieriana , Coccoloba uvifera , Conocarpus erectus , Distichlis litoralis , D. spicata , Echinochloa polystachia , Frankenia palmeri , Laguncularia racemosa , Maytenus phyllanthoides , Rhizophora mangle , Salicornia biguelovii , S. virginica , Scaevola plumieri , and Spartina alterniflora .

Avicennia germinans is the most widely distributed species of Avicennia in the New World. Its northern distributional limit there is in Bermuda (ca. 32°20ʹN; Lacerda et al. 2002), where another true mangrove, Rhizophora mangle L. also attains the northern extent of its distribution; these occurrences are the northernmost of true mangroves. The northernmost occurrence of A. germinans on the Atlantic Coast of North America is in northern Florida (St. Johns County, 30°06.618ʹN, 081°22.303ʹW, Williams & Eastman s.n. at USF). On the coast of the Gulf of Mexico the northern extent of the species extends as far as the Bay St. Louis, Mississippi (Hancock or Harrison county, ca. 30°2ʹN, Sanger s.n. at NY).

Of the three true mangroves that are common on the American mainland ( A. germinans , Laguncularia racemosa , and Rizophora mangle ) A. germinans is the northernmost in distribution. Its northern limit in North America appears to be determined by the frequency and duration of subfreezing temperatures, which can kill the above-water parts of the plants or cause mortality (Sherrod and McMillan 1985; Cavanaugh et al. 2014). Tomlinson (1986) noted a close correlation of the limits of the distributions of mangroves with the 24° isotherm of sea surface temperatures. A decade later Rützler and Feller (1996) indicated that like corals, mangroves cannot survive where an average water temperature falls below about 23° C. In recent decades, freeze-free winters have led to an expansion of A. germinans into salt marshes at the northern extremes of its range in the southern United States ( Guo et al. 2013). A poleward expansion of the distributional range of A. germinans in southern North America, likely correlated with warmer winter temperatures and less extreme freezes, was recently noted by Cavanaugh et al. (2014).

In the United States, plants have been documented from most of the coastal counties in Florida (except for most of those in the panhandle on the Gulf Coast in the northwestern portion of the state), rarely reported from at least two of the three coastal counties in Mississippi (e.g., Moldenke 1960; Scheffel et al. 2014; the locality cited for Mississippi by Lowe in 1921, Door Point, is in Louisiana), and documented from seven coastal parishes in Louisiana. In Texas the species has been collected along the Gulf Coast from Jefferson County near the Louisiana border intermittently southward to Cameron County on the Mexican border. Although it likely occurs (or occurred) in all coastal counties of Texas, based on specimens noted below and in literature reports ( Turner et al. 2003 and pers. comm. in 2010 from updated but unpublished maps; Rosen and Zamirpour 2014), it has been collected in Aransas, Brazoria, Calhoun, Cameron, Galveston, Jackson, Jefferson, Kleberg, Nueces, Refugio, San Patricio, and Willacy counties. Sherrod and McMillian (1981, 1985) discussed the distributional history of mangroves, including A. germinans , in Texas and in the northern Gulf of Mexico. The species has not been reported from the Gulf coast of Alabama (e.g., Kral et al. 2011).

Avicennia germinans is known from all 16 coastal states of Mexico (occurring along the Gulf of Mexico, Caribbean Sea, Pacific Ocean, and Gulf of California). The species reaches its northern extent in western Mexico along the Gulf of California in Sonora (Puerto Lobos, lat. 30°15ʹN; Felger et al. 2001) and its northern extent on the Pacific coast in Baja California Sur (ca. lat. 26°N; Turner et al. 1995). Distribution along Pacific coast of North America does not extend as far northward as that along the coasts of the Gulf of California, Gulf of Mexico, and the Atlantic Ocean likely because of upwelling of cold waters off the western coast of the peninsula of Baja California and California that limits their northward spread. Turner et al. (1995) discuss the distributions and probable environmental factors effecting them for this species in regions around the Gulf of California.

In Central America to the south of Belize, the extent of mangrove communities and the abundance of A. germinans are better developed along the Pacific than the Caribbean coast. Indeed no specimens have been seen from the Caribbean coast of Costa Rica, although A. germinans has been reported from the mouth of the Río Moín, north of Puerto Limón ( Zamora et al. 2004). In South America, as in North America, A. germinans has a broader distribution on the Atlantic than on the Pacific coast. Lacerda et al. (2002) indicated that the southernmost occurrences of A. germinans in South America are in northwestern Peru (Río Piúra, ca. 5°32ʹS) on the Pacific coast and in southeastern Brazil (Atafona, ca. 21°37ʹS) on the Atlantic coast. Like the northward limit of this species on the Pacific coast of North America, the southern distributional limit of A. germinans along the western coast of South America in Peru appears to have a similar cause due to the cold waters of the Humboldt Current and upwelling there ( Lacerda et al. 2002).

ILLUSTRATIONS.— Figure 4 ; see also Hepper (1963: 449, fig. 309); Gibson (1970: 178, fig. 32, which is based on Standley 87592 at F); Moldenke (1973: 151, fig. 1); Villiers (1973: 65, pl. 16); Correll and Correll (1982: 1253, fig. 540); Nash and Nee 1984: 13, fig. 2); Tomlinson (1986: 189, fig. B.8); Sanders (1997: 83, fig. 1); Proctor (2012: 586, fig. 218).

NOMENCLATURE.— The synonymy provided above is based on types from North America and Central America or names sometimes applied to plants from these regions. Additional synonyms were listed by Moldenke (1960, 1973). Based on information provided by Stearn (1958), authorship of the combination A. germinans often has been attributed to “( L.) Stearn” (e.g., Tomlinson 1986; Jarvis 2007; Daniel and Figueiredo 2009). In his letter to D. Ward (17 April 1962 attached to a specimen in FLAS), Stearn indicated that the correct name for the black mangrove is A. germinans ( L.) L., a case subsequently made by Compère (1963), and now widely accepted.

The lectotype of Bontia germinans at LINN also has inflorescence internodes similar to those of A. bicolor (i.e., ca. 4– 4.5 mm long near midspike). From the image of this specimen, I have not been able to discern other diagnostic characters used here to distinguish these species. Thus, I am unable to confirm to which species it pertains.

Moldenke (1960) noted that the types of A. nitida and A. tomentosa are both Herb. Jacquin s.n. specimens at BM, which he annotated as such in 1936. Images of both of these very sparse specimens have been seen, and based on what is visible and available, they likely conform to A. germinans .

Although no specimen was designated as the type of A. floridana Gand. , Gandoger’s herbarium is at LY in France, and it is possible that the holotype is there; specimens of the type number at F, MO, Cornell, NY, and US were cited by Moldenke (1960), but he did not indicate where the holotype was located .

LOCAL NAMES.— “Aili sip” (Kuna; de Nevers et al. 6554); “arbol salado” (Zizumbo & Colunga 500); “black mangrove” (e.g., Hammel 105); “cajiel” (Zizumbo & Colunga 500); “istatén” ( J. González & A. Pérez 193); “kahil” (Huave; A. Gerardo B. 4); “madre de sal” (Rico-Gray & Espejel 305); “madresal” ( Flores M. 1101; García B. 486); “madre sal” (Espejo & Hernández 2904); “mangle de sal” (Rico-Gray & Espejel 305); “mangle negro” (Chazaro B. 3299, Pennington & Sarukhan K. 9131); “mangle prieto” (e.g., DIAAPROY s.n.); “mangle pullequi” (Moran 7151); “mangle rojo” (DIAAPROY s.n.); “mangle salado” (e.g., Arteaga et al. 130; Allen 5631), “mangle senico” (Carter 2721); “palo de sal” (Elorsa C. 285); “pipi” (Salas M. & Torres B. 825); “puyece” (Ferris 5396); “puyeque” (González O. 1131; Mexia 1008); “saladillo” ( García P. & Estrada L. 1974; Magaña 1232); “sahkab luk’” (Ucán 431); and “u najil tikin xiw” ( Tapia M. 1508).

USES.— Firewood ( Mexia 1008; García B. 486); construction materials ( García B. 486); and used by natives for diseases of the throat (Choussy 1593). Additional construction, medicinal, and food uses of the species were provided by Moldenke (1960), Burkill (1985), Cheatham et al. (2000), and Alvarez C. (2008).

CONSERVATION.— The black mangrove is a widespread and often an abundant shrub or tree of tropical and subtropical shorelines and near coastal habitats where it is an important (often dominant) constituent of mangrove communities. Its conservation status was assessed as Least Concern ( LC) by Ellison et al. (2010).

DISCUSSION.— This species is readily distinguished by the combination of its scurfy young stems, densely spicate (headlike) inflorescences, bilaterally symmetric flowers with internally pubescent corolla lobes ( Fig. 3A ), styles usually conspicuously elongating near the end of anthesis or shortly thereafter as the fruits begins to form, and the grayish fruits (when dry; Fig. 3D ).

Rare individuals from Mexico (e.g., Martínez s.n. from Baja California Sur at MEXU; Rico-Gray 95 from Yucatán at UC) have remote nodes (to 7 mm distant, as in A. bicolor ) near the midpoint of the spikes, but show their affiliation to A. germinans by their internally pubescent corolla lobes, scurfy pubescent young stems, trichomes of the rachis to 0.5 mm long, and thecae 0.8 mm long.

The fragrant flowers of Avicennia germinans are commonly visited by honeybees, and they are reputed to be an excellent source of honey. Corollas are often noted to be 10–20 mm long (e.g., Moldenke 1960; Tomlinson 1986). My observations concur with those of Gibson (1970), who indicated that she had seen none more than 8.5 mm in length.

Morphological variation is considerable in this species, especially with respect to habit and leaf size and shape. For example, although it is commonly a sizable tree in portions of its range, A. germinans becomes a dwarf shrub at the northern extent of its range in the Gulf of Mexico (Sherrod and McMillan 1985). Moldenke (1960) discussed variation in this species with respect to these and other attributes.

In a letter to Paul Standley dated 19 February 1938 (attached to Brenes 12222 at F), Moldenke indicated that A. tonduzii could be “most easily” distinguished from A. germinans (as A. nitida ) by the size of the flowers. Indeed some of the specimens cited and/or annotated by Moldenke as A. tonduzii have calyces (2–2.5 vs. 2–4) and corollas (3.5–5 vs. 4–8) that are shorter than most specimens treated as A. germinans . Subsequently, Moldenke (1960, 1973) distinguished A. tonduzii from A. germinans on the basis of its short (shorter than the stamens) style (vs. style exserted from the calyx when corolla is shed, surpassing the stamens) and leaf blades mostly elongate-oblong, 3– 5 times as long as wide (vs. lanceolate or lance-oblong to elliptic or obovate, usually <3 times as long as wide). Elongate styles are evident on the holotype of A. tonduzii at BR (and on isotypes at BR and US). The other putative distinguishing characters of A. tonduzii overlap those of A. germinans when plants from throughout the ranges of both species are considered. Tomlinson (1986) considered A. tonduzii as a narrow-leaved variant of A. bicolor and treated the name as synonymous with that species. Others have followed Tomlinson’s influential account, and A. tonduzii has been recognized as a distinct species in some relatively recent literature (e.g., Lacerda et al. 2002).

In the southern Caribbean region and along the Atlantic coast of South America, the distribution and taxonomy of A. germinans become entangled with those of A. schaueriana Stapf & Leechman ex Moldenke. Stearn (1958) distinguished S. schauerana from A. germinans by the characters in the following couplet:

1a. Leaves mostly acute at tip; corolla lobes tomentose on both sides; style elongate......................................................................... A. germinans

1b. Leaves mostly rounded at tip, sometimes acute; corolla lobes glabrous internally, tomentose externally; style very short, the stigma nearly sessile.................... A. schaueriana

Leaf tips are variable in A. germinans . While length of the style might be somewhat dependent on floral stage and pollination, it appears likely that styles of A. schaueriana , like those of A. bicolor , do not elongate (or at least not to the extent) as in A. germinans . Like A. germinans , A. schaueriana has crowded inflorescences, but pubescence of the corolla lobes would appear to be a useful character for distinguishing these species.

REPRESENTATIVE MATERIALS EXAMINED.— BELIZE. Belize: St. Johns College , J. Dwyer & R. Pippin 10030 ( MO); shortcut from southern part of Belize City to Western Hwy. , A. Gentry 8560 ( F); Belize , C. Hammel 105 ( K); Belize , C. Lundell 4719 ( K); W de Tropical Park , T. Ramamoorthy et al. 3609 ( MEXU); Gale’s Point, just S of Bar River mouth, ca. 17°14ʹN, 88°18ʹW, J. Ratter et al. 6560 ( K); Caye Caulker (southern island), South Point , west coast, 17°43ʹ47ʺN, 088°02ʹ15ʺW, J. Rietsema & D. Beveridge 19906 ( NY) . Corozal: ca. 4 km SE of Sarteneja , ca. 18°21ʹN, 088°07ʹW, G. Davidse & A. Brant 32658 ( USF); ca. 2 km S of Sarteneja , 0.5 km NE of La Isla, 18°20ʹ19ʺN, 088°07ʹ25ʺW, Z. Goodwin & G. Lopez 1583 ( MO) . Stann Creek: Northeast Cay, F. Fosberg & D. Stoddart 53854 ( US); Man-O’- War Cay , W of Tobacco Cay Range, F. Fosberg & D. Spellman 54204 ( MO, US); North Silk Cay , F. Fosberg & D. Spellman 54272 ( US); Little Water Cay , F. Fosberg & D. Spellman 54343 ( F); Buttonwood Cay , F. Fosberg & D. Spellman 54421 ( US); Scipio Cay, 16°28.135ʹN–16°28.295ʹN, 088°17.793ʹW–88°17.829ʹW, D. Lentz et al. 2355 ( NY); Long Coco Caye, off Placencia, ca. 16°29.5ʹN, 88°12.7ʹW, D. Lentz et al. 2833 ( NY); Dangriga, G. Proctor 36197 ( MO); All Pines , [16°47ʹ00ʺN, 88°19ʹ00ʺW], W. Schipp 625 ( UC); South Water Cay, Spellman & Stoddart 2157 ( US); Seal Cay, Spellman & Stoddart 2481 ( MO); Rendezvous Cay, Spellman & Stoddart 2498 ( MO, US); Coco Plum Cay, Spellman & Stoddart 2567 ( MO, US); 4.5 km S of Southern Highway turnoff, D. Stevenson 1132 ( K) . Toledo: W of beach road, Punto Gorda, J. Dwyer et al. 631 ( MO) .

COSTA RICA. Guanacaste: vicinity of Playa Naranjo in P. N. Santa Rosa , F. Almeda et al. 4205 ( CAS); Playa de Coco , R . Blaisdell 300 ( FSU); P. N . Palo Verde, Valle de Tempisque, Estación Catalina , Laguna Nicaragua , 10°21ʹ10ʺN, 085°13ʹ00ʺW, U . Chavarría 1243 ( F); Puerto Jesús, Nicoya, O . Cook & C. Doyle 743 ( US); ca. 1 km E of Río Tempisque ferry, G . Crow 6116 ( F); between Playa Comchal and Playa Brasilito, L . Durkee 76-386 ( F); Port Parker, Salinas Bay , F . Elmore E1 ( DS); Sitio Ojochal , P. N . Santa Rosa , 10°47ʹ35ʺN, 85°39ʹ00ʺW, A . Fernández 315 ( K, NY); Playa de Sámara , al S de Nicoya, A. Jiménez M . 526 ( F). Puntarenas: Golfo Dulce, vicinity of delta of Río Esquinas , P. Allen 5631 ( DS, F, FSU, US); Quepos, F. Almeda et al. 3398 ( CAS, F, NY); Cantón Golfito, Península de Osa , P. N . Corcovado, Estación Sirena, Río Corcovado , 08°28ʹ50ʺN, 83°35ʹ30ʺW, L. Angulo 136 ( K); Puerto Jiménez de Osa , A. Brenes 12222 ( F); Isla de Caballo , A . Brenes 15696 ( NY); Punta Morales, J. Gomez-Laurito 9980 ( F); near mouth of Río Coto Colorado , ca. 8 km S of Golfito, 08°37ʹN, 083°09ʹW, W. Kress & L. Cablk 94-3803 ( US); between Caldera and Matalimon, E of Puntarenas , K. Lems 5026 ( NY, US); cerca de Caldera , J. León 505 ( F); near Rincon de Osa , R. Liesner 2203 ( NY); vicinity of Puntarenas , W. Maxon & A. Harvey 7841 ( US); Santo Domingo, Pittier 7109 ( K, US), 7110 ( F, K, NY, US); estuaires de Santo Domingo de Golfo Dulce , A. Tonduz 10060 ( DS, NY, US); N side of Estero, Puntarenas , I. Wiggins & D. Porter 132 ( CAS) .

EL SALVADOR. 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); 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. Castillo 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 13329 B ( 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 D 21a ( 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, 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, 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, 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 3 rd 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 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 A 1 A and FL 312, E of FL A 1 A; 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 ( R 22 E, T 23 S, S 24), 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).

Figures

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.

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.

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).

FIGURE 4. Avicennia germinans. A. Habit (composite from photos of living plants). B. Pneumatophore (Schwartz & Nickerson 9119). C. Fertile shoot (Calzada 434, Pipoly 9034, Sousa 3112). D. Inflorescence with flower in profile (Lakela 29824, Sousa 3112). E. Flower, front-view (Lakela 29824, Sousa 3112). F. Stamen (Sousa 3112). G. Gynoecium (Lakela 29824). H. Fruit with detail of surface (Lundell 7009). Drawn by Tom Davis from specimens at CAS.