-, and F. paudflora(0\eaceae).

TAXONOMY OF THE WATER ASHES:
FRAXINUS CAROLINIANA, F. CUBENSIS, AND F PAUCIFLORA (OLEACEAE)

Guy L. Nesom

2925 Hartwood Drive

Fort Worth, TX 76109, USA

www.guynesom.com

ABSTRACT

Water ash has commonly been identified as a single species, Froxinus caroliniana Miller, but
is here recognized to comprise three distinct entities each treated at specific rank: (1) F. caroliniana
sensu stricto, from northern Florida west along the Gulf Coast to Texas and Oklahoma and north
along the Atlatic coast to Virginia; (2) F. cubensis Griseb., from Cuba and the southern two-thirds of
the Florida peninsula; and (3) F. paucijlora Nutt, from northern Florida and southern Georgia. All
three are similar in gross morphology and occur in swampy or riverine habitats but the ranges of F.
caroliniana and F. cubensis are essentially parapatric. while the range of F. paucijlora lies mostly
within that of F. caroliniana and overlaps with F. cubensis. Samaras of F. caroliniana sensu stricto
are broad, often rhombic-elliptic, (ll-)13-19-nerved pel' side, and 3-winged at a low frequency,
while those of F. cubensis are narrower, 5-9-nerved per side, and consistently 2-winged. Samaras of
F. paucijlora are similar in shape to those of F. cubensis, but the abaxial leaf surfaces are covered by
a cuticular micro-reticulum similar to that characteristic of the F. americana complex. County-level
distributions are mapped and a key, descriptions, and typifications are provided. A lectotype is
designated for Fraxinus hybrida Lingelsheim.
KEY WORDS: Fraxinus caroliniana, Fraxinus cubensis, Fraxinus paucijlora, Florida

The name Fraxinus caroliniana Miller is currently commonly used in southeastern USA
herbaria and floristic accounts to identify various expressions of water ash — all with samara bodies
flat and winged nearly to the base. This broad view has been adopted by Miller (1955), Long and
Lakela (1976), Duncan and Duncan (1988), Godfrey (1988), Wallander (2008), and Weakley (2010)
and by Wunderlin and Hansen (2003, 2010) even to include another swamp species, F. profunda
(Bush) Bush, in synonymy. Brown and Kirkman (1990) and Nelson (1995) separated F. profunda but
identified all water ash as F. caroliniana without citing synonyms. Long and Lakela (1976, p 687)
were particularly emphatic: "This species [F. caroliniana] is extremely variable, but variations
scarcely deserve taxonomic recognition." Other students of the woody flora of northern and central
Florida, however, have maintained F. profunda as a separate species and recognized a distinction
between F. caroliniana and F. paucijlora Nutt. (e.g., Sargent 1894, 1902; Small 1913; Kurz &
Godfrey 1962; Clewell 1985; Nesom 2010), using characters similar to those in the present study.
Fraxinus cubensis Griseb. of southern Florida apparent!)' has never been recognized in the USA but
it has been treated as F. caroliniana var. cubensis (Griseb.) Lingelsh. in the Cuban flora (Leon &
Alain 1957).

In working out a taxonomic treatment of Fraxinus for the developing FNANM volumes, it
became evident that "water ash" includes three entities. In addition to the trees with characteristically
broadly elliptic to rhombic and often 3-winged samaras (= F. caroliniana sensu stricto), others
produce narrower, consistently 2-winged and differently veined samaras. Then, after becoming
familiar with the distinctive abaxial leaf surface of F. americana L. (white ash), I saw that some trees
with the more narrowly 2-winged samaras have an abaxial foliar morphology similar to F. americana
and stand apart from the other water ashes. Field observations confirm the distinctiveness of these
(identified as F. paucijlora, termed here as "swamp white ash"). In overview, each of the three

Nesom: Taxonomy of v

expressions of water ash is recognized here at specific rank: F. caroliniana, F. pauciflora, and F.
cubensis.

Trees of these three taxa are generally similar in vegetative features and all grow in swampy
or riverine habitats, often or characteristically with their bases submerged. The key below separates
them on the basis of features of the samaras and abaxial foliar epidermis. Fraxinus profunda is added
to the key since it has sometimes been confused with water ash in Florida. All are contrasted with F.
americana, a species of upland woods with a different samara morphology.

1. Samara bodies plump-subterete, wings 3-7 mm across at widest point, lateral veins 3-5 on each
side of the wing, all originating from near the top of the body, essentially parallel with the wing

margins; upland habitats Fraxinus americana

1. Samara bodies flattened, wings 8-22 mm across at widest point, lateral veins 5-9 or 1 1-19 on each
side of the wing, originating in sequence from along the major vein paralleling the body edge, angling
upward; swamp and riverine habitats.

2. Large trees; leaflets (7-)9-15(-25) cm x (2.5-)3.5-7(-ll) cm, bases often rounded, less
commonly obtuse to acute or acuminate, samaras 1 35-)40-70(-75) mm, w r ings gradually expanded

from near the base of the body to proximal 1/2, (5.5-)6-10(-12) mm wide Fraxinus profunda

2. Shrubs to small trees; leaflets mostly (4-)5-ll(-12) cm x (1.5-)2-5(-6) cm, bases obtuse to
acute or abruptly attenuate; samaras (25-)30-50 mm, wings expanded gradually from near base,
(10-)12-20(-22) mm or 6-10(-12) mm wide.

3. Leaflet abaxial surfaces micro-foveolate-papillose (use 20^10X lens) with a cuticular
reticulum overlaying and obscuring the epidermal surface, minute glandular peltate scales not
evident Fraxinus pauciflora

3. Leaflet abaxial surface without a cuticular overlay, the epidermal surface visible, minute
glandular peltate scales usually evident and abundant.

4. Samaras 35^6 x (10-)12-20(-22) mm, rhombic to broadly elliptic, elliptic-obovate,
oblong-obovate, or rhombic-ovate, wings 2-3, lateral veins (11-) 13-19 on each side of the
wing; fruit body usually half or more than half the length of the samara

Fraxinus caroliniana

4. Samaras (25-)30-50(-54) x 6-10(-12) mm, obovate-oblanceolate to narrowly obovate or
narrowly elliptic-obovate, wings 2, lateral veins 5-9 on each side of the wing; fruit body half
or less than half the length of the samara Fraxinus cubensis

FRAXINUS CAROLINIANA Miller, Gard. Diet., ed. 8. 1768. Calycomelia caroliniana (Miller)
Kostel., Allg. Med.-Pharm. Fl. 3: 1004. 1834. Leptalix caroliniana (Miller) Raf,
Alsographia Amer. 36. 1838. Fraxinus americana var. caroliniana (Miller) D.J. Browne,
Trees America 398. 1857. Fraxinus nigra subsp. caroliniana (Miller) Wesmael, Bull. Soc.
Roy. Bot. Belgique 31: 113. 1892. TYPE: USA. "Raised from seeds which were sent from
Carolina in the year 1724, by Mr. Catesby" (K).

Fraxinus platicarpa Michx., Fl. Bor.-Amer. 2: 256. 1803. Calycomelia platicarpa (Michx.) Kostel.,
Ind. Hort. Bot. Prag. 26. 1844. Fraxinus caroliniana var platicarpa (Michx.) Lingelsh., Bot.
Jahrb. Syst. 40: 221. 1908. TYPE: USA. "HAB. in Carolina" (P presumably).

Fraxinus pallida Bosc, Mem. CI. Sci. Math. Inst. Natl. France 1808: 201. 1809. Leptalix pallida
(Bosc) Raf, Alsogr. Amer. 33. 1838. Calycomelia pallida (Bosc) Kostel., Ind. Hort. Bot.
Prag. 26. 1844. As synonym of F. caroliniana fide Sargent (1894).

Nesom: Taxonomy of v

Fraxinus caroliniana Mills' var. latifolia Roemer & Schultes, Syst. Veg. 1: 279. 1817. TYPE: USA.
Protologue: "A Pensylvania ad Carolinam, rarius." As synonym of F. caroliniana based on
geography.

Fraxinus tnptera Null, Gen. N. Amer. PI. 2: 232. 1818. Samarpses triptera (Nutt.) Raf., New Fl. 3:
93. 1838 ("1836"). Fraxinus platicarpa Michx. var. triptera (Nutt.) AW. Wood, Class-Book
Bot, ed. 1861. 598. 1861. Fraxinus americana var. triptera (Nutt.) D.J. Browne, Trees
America 399. 1857. TYPE: USA. South Carolina. (Holotype: PH?). Protologue: "8.
triptera. Leaflets (about 7?) obovate, entire, subsessile, beneath tomentose, oblique at the
base; samara very broad, elliptic-obovate, most 3-winged! attenuated at the base. HAB. In the
oak-forests of South Carolina. Fruit at first sight almost similar to Halesia, more rarely 2 than
3 winged, the seed also 3-sided. Points of the leaves obtuse, the under side paler and softly
villous, the common petiole and nerves beneath smooth." Treated by Nuttall as a different
species than his "5. caroliniana" and "6. platycarpa."

Nuttall later added comments (1849, p. 63): "I observed fruit of this curious species
many years ago, in winter, in the oak forests of South Carolina, and as 1 thought, the leaves of
the same; but I am now in doubt whether the leaves than collected actually belonged to the
same plant with the fruit. I must therefore leave the species in the same imperfect manner I
than found it, as I have never since seen any other specimen, [paragraph] The fruit is the
mint cuuous of anj in the genus at rust sight jlmo*.i similar to tlk.t ot .in Halesia, being
nearly of the same breadth; the samara, in fact, appeared to be more rarely 2 than 3 winged,
the seed itself was also 3-sided, at the base the fruit is attenuated into a very slender
peduncles without being at all terete. Perhaps it is merely a variety of F. platycarpa."

Fraxinus curvidens Hoffsgg., Verz. Pfl.-Kult, Nachtr. 2: 29, 118. 1826. As a synonym of F.
caroliniana fide Sargent (1894).

Fraxinus cordataRal, Alsographia Amer. 34. 1838. TYPE: Florida. "139. Fr. L. [Leptalix] cordata
Raf. foliolis petiol. ovatis ellipt. integris, basi subcordatis, apice attenuatis obtusis. supra
luicul U.ibn-, luadv- subtus glauus pubescens — very distinct sp. from Florida, disc, by Kin,
large folioles 4 to 6 inches long 2 or 3 broad." As synonym of F. caroliniana fide Wunderlin
and Hansen (2010), but Rafinesque's description of the leaflet bases as "subcordate,"
emphasized by the epithet, does not correspond to known native ashes of Florida.

Fraxinus platicarpa Michx. var. oblanceolata M.A. Curtis, Amer. I Sci. Arts 57: 408. 1849.
Fraxinus caroliniana var. oblanceolata (M.A Curtis) Femaid & Schub., Rhodora 50: 188,
1948. TYPE: USA. South Carolina. Berkeley Co.: Santee Canal, Santee swamp, [no date],
H.W. Ravenels.n. (holotype: CHARL?; isotypes: GH, NY!).

Fraxinus platicarpa Michx. var. pubescens M.A Curtis, Amer. J. Sci. Arts 57: 408. 1849. Fraxinus
caroliniana var. pubescens (MA. Curtis) Fernald, Rhodora 39: 442. 1937. Fraxinus
caroliniana forma pubescens (M, A Curtis) Fernald & Schubert, Rhodora 50: 188. 1948.
TYPE: USA. South Carolina. Berkeley Co.: Santee Canal, Santee swamp, [no date], H.W.
Ravenels.n. (holotype: CHARL?; isotypes: GH, NY!).

Fraxinus nuttallii Buckley, Proc. Acad. Nat, Sci. Philadelphia 12: 444. 1860. TYPE: USA. Alabama.
Wilcox Co.: In swamps, no date, Buckley s.n. (PH?). For this small swamp tree with 3-
winged fruits, Buckle}' noted that "As Nuttall had not material for a complete description,
none can tell what is meant by his Fraxinus triptera; but as possibly lie may have intended
the tree now described, I call it Nuttall's Ash." As synonym of F. caroliniana based on
morphological description and geography.

Fraxinus rehderiana Lingelsh. in Engler, Pflanzenr, 4(Heft 72): 42. 1920. Fraxinus caroliniana var.
rehderiana (Lingelsh.) Sargent, J. Arnold Arbor. 2: 173. 1921. TYPE: USA. Virginia. Isle of
Wight Co.: Banks of Blackwater River near Zuni, 19 Aug 1908, A Rehder s.n. (holotype: B?;
isotype: GH). Lingelsheim made this observation: "Species fortasse hybrida, e F.
pennsylvanica et F. caroliniana orta. forma parallela F. hybridae." Placed here as a synonym

Nesom: Taxonomy of v

of F. caroliniana based on its morphological description and geography and on Sargent';

Fraxinus caroliniana Miller forma hypomalaca Fernald & Schubert, Rhodora 50: 189. 1948. TYPE:
USA. Virginia. Greensville Co.: Fontaine Creek, SW of Haley's Bridge, cypress swamp,
wooded bottomland, 9 Jul 1946, M.L. Fernald & H.E. Moore 15139 (holotype: GH!; isotype:
GH!, PH).

Fraxinus caroliniana Miller forma lasiophylla Fernald & Schubert, Rhodora 50: 189. 1948. TYPE:
USA. Virginia. Southampton Co. : Darden's Pond N of Courtland, upper border of sandy and
peaty shore, 15 Sep 1946, M.L. Fernald. B.H. Long, and I.D. Clement 15335 (holotype: GH!:
isotype: GH!, PH).

Trees or shrub-like trees with several leaning trunks and often with buttressed bases, 2.5-
10(-15) m; twigs terete; bark light gray, with thin appressed scales, not deeply furrowed; w r inter buds
brown. Leaves deciduous, pinnate, 12-40 cm, leaflets 5-7(-9), glabrous on both surfaces or sparsely
pubescent abaxially along the veins, rarely over the surface, dirk green adaxially, paler or whitish
abaxially, not scaly-punctate, not papillose, blades 4-12(-15) cm x (l-)2-3(-6) cm, lanceolate to
ovate-lanceolate, elliptic, or elliptic-oblong, lateral vein pairs (5-)7-9(-12), apex obtuse to acute or
acuminate, base obtuse to truncate or rounded, margins shallowly serrate to crenulate on distal 4/5-
2/3, lateral petiolules 2-3(-15) mm, not winged; rachis (2-)3-9(-12) cm; petiole bases not raised; leaf
scars shallowly hemispheric to depressed obovate, apex very slightly convex or notched. Flowers
unisexual (species dioecious), appearing before leaves, wind-pollinated; pistillate calyx present,
persisting or not; petals absent. Samaras 35-46 mm, rhombic to broadly elliptic, elliptic-obovate,
oblong-obovate, or rhombic-ovate, body flattened and abruptly acuminate to stipitate at the base,
usually half or more than half the length of the samara, wings 2(-3), arising from the base or proximal
1/4 of body, (10-)12-20(-22) mm wide; lateral veins (1 1-)13-19 on each side of the wing.

Flowering Feb-Apr(-May). River and stream banks, alluvial woods, flatwoods, cypress-gum
swamps (often in water), overcup oak-water locust-w^ater elm, swamp and pond margins, depressions,
ditches and canals, roadside swales; 3-60(-100) m; Ala., Ark, Fla., Ga., La., Miss., N.C., S.C., Tex.,
Va. Water ash, pop ash.

FRAXINUS CUBENSIS Griseb., Cat. PI. Cub. 170. 1866. Fraxinus caroliniana var. cubensis
(Griseb.) Lingelsh., Bot. Jahrb. Syst. 40: 221. 1908. Fraxinus caroliniana subsp. cubensis
(Griseb.) Borh., Bot. Kozlem 58: 176. 1971. Fraxinus pennsylvanica subsp. cubensis
(Griseb.) E. Murray, Kalmia 13: 6. 1983. Fraxinus pennsylvanica var. cubensis (Griseb.) E.
Murray, Kalmia 13: 6. 1983. TYPE: Cuba, Protologue: "Cuba occ, in depressis pr.
Zarabanda," 1865, C. Wright 45 (holotype: GOET; isotype: K digital image!)." Collections
at GH and NY (2 sheets) are possible type material — Cuba, without other collection data,
C. Wright 3624 — if, as in Wright collections from the southwestern USA field numbers
and published collection numbers are inconsistent. The GOET and K labels, however,
specify "Wright 45." Another possible type specimen (S digital image!) has a Charles
Wright label but is unnumbered.

Small trees or shrubs 3-14 m, sometimes multi-trunked from the base; twigs terete; bark
grayish to gray-brown, morphology; winter buds reddish brown to brown. Leaves deciduous,

pinnate, 12-23 cm. spaced along branches; leaflets (3-)5(-7), subcoriaceous. glabrous on both
surfaces except hirsute-pilose along abaxial midvein and lateral vein axils, rarely puberulent-hirsute
on il axial linuiui green to yellowish green, scaly-punctate abaxially, not papillose, blades (4-)4.5-
10.5(-13) cm x (1.5— )2— 5(— 5.5) cm, narrowly to broadly elliptic to elliptic-obovate or obovate, apex
acute to acute-acuminate, lateral vein pairs 7-11, base acute to obtuse, margins shallowly serrate to
denticulate on distal 1/2-2/3, lateral petiolules 4-10 mm, not winged; rachis 4.5-10 cm, not winged;
petiole bases not raised; leaf scars transversely elliptic and shallow concave, 3-5 mm wide. Flowers

Nesom: Taxonomy of v

unisexual (species dioecious), appearing before the leaves, wind pollinated; pistillate calyx present
and persisting at base of samaras; petals absent. Samaras (25-)30-50(-54) mm, obovate-
oblanceolate to narrowly obovate or narrow h eliiptic-obot ate, body flattened, half or less than half
the length of the samara, wings 2 (very rarely 3), arising from middle/ proximal 3/4 of body, 6-10(-
12) mm wide, lateral veins 5-9 on each side of the wing.

Flowering Feb-Mar. Hardwood hammocks, swamp forests and low woods, apparently most
commonly along river and creek banks and pond and lake edges; 0-5 m; Fla.; West Indies (Cuba).
Cuban water ash

I have seen collections of Fraxi 'mm cubensis from the following Florida counties: Brevard,
Citrus, Collier, Glades, Highlands, Hillsborough, Lee, Manatee, Marion, Martin, Monroe,
Okeechobee, Orange, Palm Beach, Polk. Sarasota, and Sumter. Various localities on the shores of
Lake Okeechobee are in Glades, Martin, Okeechobee, and Palm Beach counties.

Specimens examined from CUBA: Prov. Santa Clara. Peninsula de Zapata, on the edge of
Cienaga de Zapata, in swampy forests, small tree, J Feb 1924, Ekman 1S368 (NY); Prov. Matanzas,
Mpio. Marti, Cienaga del Majaguillar al NO de Marti, Cienaga de Gonzalito cerca del Canal de
Blanquizal, 5 m, bosque secundario de cienaga, 20 Feb 2009, Greuter 27019 (NY); Cayo la Lisa,
Cienaga de Zapata, Las Villas 24 Feb 1941, Leon et al. 19562 (NY); Los Sabados, Hacienda El Jiqui,
Peninsula de Zapata, 16 Jul 1920, Roig & Gemata 2145 (NY); Prov. Matanzas: Cienaga de Zapata,
along road between Australia and Playa Larga, close to Calal de La Laguna, evergreen or semi-
evergreen forest on limestone, temporarily inundated, 1 3 Apr 1995, Rova et al 2261 (GOET digital
image!, DNA voucher). All are localities in western Cuba, just east of Havana.

Fraxinus cubensis is similar to F. caroliniana in most features except fruit morphology' (I
have not seen F. cubensis in nature, however, to provide a comparison of its growth habit). Samaras
of Fraxinus caroliniana appear to be far more variable than those of F. cubensis (Figs. 4 and 5), and
although narrower samaras of F. caroliniana may approach of those of F. cubensis in outline, the
distinction in venation is consistent (as in the key above). It might seem justifiable to treat the two as
conspecific, but the remarkably sharp divide in geographical distribution would appear to mark some
kind of biological isolation and I have not observed unequivocal intermediates in samara morphology.
It will be interesting to investigate the area of apparent parapatry toward evaluation of their apparent
reproductive isolation, as hypothesized here.

Samaras of Fraxinus caroliniana are 3~winged (3-carpellate) at a low frequency (ca. 5-20%
on an individual) over the whole range of the species and it is more common than not to find 3-
winged fruits on any single specimen. Samaras of F. cubensis are consistently 2-w ? inged, but I have
encountered 2 collections with 3-winged fruits: Manatee Co., Palmetto, 21-23 Aug 1895, Nash 2431
\* iFK Sumtei Co >umtet Cilius \\ jlilrfw \lan.'_ement \ica \^ of Bushnell, Withlacoochee River
bottoms, 13 Aug 1958, Krai 7847 (GH).

FRAXINUS PAUCIFLORA Nutt, N. Amer. Sylva 3: 61, plate 100. 1849. TYPE: USA. Georgia.
[Charlton Co.:] "collected in Georgia in the neighborhood of 'Trader's Hill,' by the late
indefatigable and excellent botanist, Dr. Baldwyn," no date (holotype: PH; possible isotype:
PH).

Fraxinus platicarpa Mchx. var. floridana Wenzig, Bot. Jahrb. Syst. 4: 185, plate II, 11. 1883.
Fraxinus floridana (Wenzig) Sargent, N. Amer. Silva 14: 39. 1902. TYPE: USA. Florida:
"Cabanis! in lib, Berol." (holotype: B?, fragment GH). "Samoa c has dtntihu' 11.1ms
triangularibus saepe destructis, lanceolata, alae margo in longitudinem lateris partis inferioris
latior quam in Fr. quadrangulata Mchx,. angustior quam in Fr. platycarpa, apice obtusissima
emarginata basi acuminato-attenuata. 0,054 m. longa, ala 0,040 m. lata, pars inferior 0,027 m

Nesom: Taxonomy of v

long, plana, longitudinaliter sulcata." A GH collection from 1830 by E.F. Leitner is noted to
be a possible type, but the protologue indicates that the type was collected by Jean Louis
Cabanis (1816-1906 ) — see bibliographic notes by Sargent (1902, p. 39).

Fraxinus hybrida Lingelsh., Bot. Jahrb. Syst. 40: 220. 1908. LECTOTYPE (designated here): USA.
Florida. Duval Co.: swamps, near Jacksonville, Aug [1890?], Curtiss 2321 (NY!;
isolectotypes: GH-2 sheets!). Lingelsheim cited as syntypes "Curtiss 2321!, Nash 941!,
1698!" — Nash 941 (E digilal image!) and Nash 1698 (E digital image!, GH!). Curtiss made
a series of collections of this species in July and August 1894, numbered as 4536, 4536A
4536B, and 4536C (see citations below). Label data of the three iectotype ^pcumens do not
specifiy the year of collection, but a Curtiss collection of F. caroliniana sensu stricto also
numbered "2321" was made in 1880 (Georgia, Decatur Co., Oct 1880, Curtiss 2321, GH!).

Fraxinus profunda \ar. ashei E.J. Palmer, J. Arnold Arbor. 13: 417. 1932. TYPE: USA. Florida.
[Suwannee Co.:] near Hildreath, growing in the water, 15 May 1929, W.W. Ashe s.n.
(probable holotype: NCU, digital image!). Palmer (1932) did not specifically cite NCU as the
herbarium of deposition of the type but he noted that "My attention was first called to this
tree by Mr. W.W. Ashe, for whom the variety is named, and on whose notes and collections I
have drawn to supplement my own made this season in the region where it grows." There is
no specimen at GH of Ashe's collection from near Hildreath and it seems evident that Palmes'
had a loan of his material from NCU The NCU sheet is annotated by hand as "type,"

Trees 4— 10(— 15) m tall, usually single-stemmed and erect from the base; twigs terete; winter
buds dark brown. Leaves deciduous, pinnately compound, 15-30 cm. leaflets (3-)5(-7), terminal
leaflet rarely absent, petioles 3.5-9 cm, bases not raised on pedestals, petiole plus rachis 6-15 cm,
petiolules 5-11 mm, blades oblong-elliptic, elliptic, elliptic-lanceolate, or ovate-lanceolate, terminal
(5.5-)7-13(-16) cm x (2-)3.5-7 cm, lateral usually slightly smaller, dark green and glossy adaxially,
olive abaxially, apices acute to abruptly or gradually acuminate, bases rounded to obtuse or attenuate,
lateral veins 7-11 pairs, margins shallowly dentate-serrulate to serrulate on the distal 2/3; petiole base
shallowly notched adaxially, leaf scars apically shallowly convex to subtruncate, 3-4(-6) mm wide.
Flowers unisexual (species dioecious), appearing before the leaves, wind-pollinated; pistillate calyx
ca. 1 mm, persisting at base of samara; petals absent. Samaras 32^1-7 mm, elliptic-oblanceolate to
narrowly obovate, bodies (16-)20-26 mm, flattened, usually channeled along the middle, half or less
than half the length of the samara, wings 2, gradually expanded from near itiQ base to middle of the
body, (7-)9-10(-12) mm across at widest point, lateral veins 8-12 on each side.

Flowering Mar-Apr. Sloughs, swamps, usually in standing water; 2-20 m; Florida, Georgia.
Swamp white ash, pop white ash, w r ater white ash.

Additional collections examined. Florida. Alac hua C o.: S of Gainesville, Paynes Prairie
State Preserve, hammock NE of Alachua Sink, northern portion of hammock, 10 April 1982,
Easterday 858 (FLAS); along Newnans Lake, Gainesville, 13 July 1936, West & Evers s.n. (FLAS).
BakerCa: along Cedar Creek. 5.5 miles N of Glen Saint Mary, 8 Jun 1950. Kurz s.n. (FSU). Bay
Co. : bank of Econfma River, E side, 0.8 mile W of Youngstown, 24 Apr 1951, Kurz s.n. (FSU). Clay
Co. : Hibernia, Mar 1869, Canby s.n. (NY-3 sheets). DjxieCa: California Creek, 7.5 mi SSWof
Cross City, hammock, partially underwater, limestone undersurface, 8 m tree in water, 13 May 1966,
McDaniel 7489 (FSU). Duval Ca : swamps, near Jacksonville — 27 Aug 1894, Curtiss 4536 (NY),
27 Aug 1894, Curtiss 4536A (NY), 2 Jul 1894, Curtiss 4536B (NY). 1894, Curtiss 4536C (NY); 7.5
mi E of Baldwin or 4 mi W of Marietta on U.S. Rte 90, swamp, 7 Jul 1949 Kurz s.n. (FSU); 1-2 mi W
of Atlantic Beach, 6 Sep 1949, Kurz s.n. (FSU); 7.8 mi E of Jacksonville city limits on US Rte. Al A
swamp, 7 Jun 1950, Kurz s.n. (GH, VDB); 2.4 miles E of Jacksonville city limits on S side of road at
AlARte bridge, swamp, 8 Jun 1950, Kurz s.n. (FSU). Flagl er Co . : cypress swamp near Andalusia,
18 April 1940.' West & Arnold s.n. (FLAS). Jefferson Co. : Jefferson Co. : Wet hammock, 0.5 mi Wof
Aucilla River, by US Rt. 98, E of Newport, 14 June 1983, Godfrey 80693 (FLAS); slough between

Nesom: Taxonomy of v

Newport and Aucilla River, growing in 2 feet of standing water. 2 May 1956, Krai & Godfrey 2379
(FLAS, FSU, LL, NY); at junction of the Wascissa River and Aucilla River, in swampy area', small
tree (15 feet), [no date], Williams s.n. (FSU). Lafayette Co. : Lafayette Co.: Mallory Swamp
Conservation Area, SW of Branford, highly degraded mosaic of bottomland forest and basin swamp,
4 Sept 2004, Abbott 191 77 with Heaney (FLAS); on E side of Fla. Rte 51, 9 mi N of U.S. Rte 19, N
of Mayo, in water, 5 Sep ] 949, Kurz s.n. (FSU, GH). L ake Co. : along W side of the St. Johns River
along Fla. Rte 40. 22 Jul 1950, Kurz s.n. (FSU, NY); Tavares, low- hammock, 13 June 1940. Murrilt
s.b. (FLAS); vicinity of Eustis, in water, Lake Norris, small tree 12-15 ft high, 16-25 Aug 1894, Nash
1698 (AA NY); vicinity of Eustis, cypress swamp at Eldorado, tree 30-50 ft high, 16-25 Aug 1894,
Nash 1698 (NY). Levy" Co.: Waccasassa Bay State Preserve, well west of Jack's Creek; coastal
hydric hammock, with scattered freshwater depressions and scattered areas of mixed hardwoods, 9
June 1997. A bbott 1 0489 with Judd (FLAS); swamp ca. 6 mi N of Cedar Key, 22 Jul 1 960, Godfrey
60102 (SMU). Liberty Co. : floodplain woods along the Ochlockonee River, Rte 20, W of
Tallahassee, 2 Jun 1950, Kurz s.n. (NY, VDB); N of Fla. Rte 20, in W floodplain of Ocklockonee
River, 2 Jun 1950, Kurz s.n. (FSU); Liberty Co. : Telogia Creek, access road to boat ramp off Co. Rd.
67A 1.8 air mi ENE of Telogia, common small trees along sloughs and in swamp, ca. 10-25 feet tall,
trunks slender, erect, and single, none in fruit, Fraxinus caroliniana common (not collected) and in
fruit along sides of creek, 12 Jul 2010, Nesom 2010-F3 (FLAS, GH, NY); W side of Hwy 65 0.6 mi S
of jet with Hwy 267 S of Hosford, ca. 50 yards N of bridge over Telogia Creek, edge of swamp, tree
40 feet tall, fruits out of reach and not collected, 12 Jul 2010, Nesom 2010-F4 (FSU, GH, NY, TEX);
W side of Hwy 65 0.6 mi S of jet with Hwy 267 S of Hosford, ca. 50 yards N of bridge over Telogia
Creek, edge of swamp, small, singe-trunked trees in immediate vicinity of Fraxinus profunda (Nesom
F4), not in fruit, 12 Jul 2010, Nesom 2010-F5 (FLAS, GH, NY, TEX). Mario n Co. : swamp along
Rainbow River, Dunellon, in water, small tree with several trunks from near the base, bases of trunks
enlarged, 1 May 1 976. Godfrey 74890 (FSU, VDB); riverine forest beneath the Oklawaha River
bridge ("Eureka Mountain") along C-316, just E of Eureka, 24 Jul 1985, Hansen &. Wunderlin 10456
(TEX); Ocala National Forest, along Mormon Branch, a tributary of Juniper Run, moist forest along
creek, 30 July 1994, Judd 8036 (FLAS); W edge of Oklawaha r" swamp, dominant species, with N.
sylvatica var. biflora also abundant, 25 June 1975, Martin & Cooper 735 (FLAS); 1/4 mi E of
Olawaha River, Eureka, riverbottom swamp, frequent, from upper branches of uprooted tree 45 ft.
high, 6 in. diameter, 17 May 1963, Ward&Laessle 3484 (FSU), Nassau Co. : White Oak Plantation
in the wedge formed by the jet of the Little St. Mary's River and St. Mary's River, ca. 8-10 mi NW of
Yulee, swamp forest 100-200 yards NW of the Pavilion Bldg along the St. Mary's River, tree 6 m tall
in standing water, 23 Jul 1997, Wilbur 68108 (BRIT. FLAS). Orange Co. : 5 mi SE of Fort Christmas,
in a swamp, 19 Sept 1929. O'Neill s.n. (FLAS). OjLceola_Qx: deep swamps, near Kissimmee, tall
slender tree, 2 .Apr 1931, Palmer 38356 (AA, NY). Seminole Co. : Seminole/Lake County, along
banks of Wekiva River, dominant, 30 Jun 1990, Bacchus 5492 (FLAS, FSU); 2 mi NW of Sanford on
U.S. Rte 17, 22 July 1950, Kurz s.n. (FSU). Putnam Co. : along Oklawaha River, Cedar Landing, 7
July 1935, Arnold s.n. (FLAS). St. Johns Co. : bank of St. Johns River, 27 March 1910, Hood 72069
(FLAS); riverside, Tocoi, 7 June 1940, West & Arnold s.n. (FLAS). Suwannee Co. : 5 mi NW of Fort
White off of US 27 in Ichetucknee Springs State Park, ca. 0.2 mi W of Old South entrance, 0.8 mi S
of US 27 bridge, floodplain forest and swamp, 6 May 1992, Herring 709 (FLAS). Wakulla Co. :
swamp by East River, by first bridge after entering St. Marks National Wildlife Refuge on Lighthouse
Rd, single-stemmed tree ca. 9 m tall, 17 Jul 1981. Godfrey ''9020 (FLAS, FSU, VDB); river swamps,
Aucilla River, 5 mi above mouth, 10 Apr 1934, Griscom 21597 (AA); St. Mark, low woods, 29 Apr
1914, Harbison 1499 (AA); N side of Wakulla River, along U.S. Rte 319, 27 Sep 1949, Kurz s.n.
(FSU); E side of Wakulla River, swamp, 14 Jun 1950, Kurz s.n. (FSU); St. Marks NWR, Lighthouse
Road (Hwy 59), area of double bridge over East River Pool, ca. 1.8 road mi S of the Refuge
Headquarters and Visitor Center', common small trees in standing water, few in fruit, trunks erect,
single, 13 Jul 2010, Nesom 2010-F6 (FLAS, GH, NY); St. Marks NWR, Lighthouse Road (Hwy 59).
area of double bridge over East River Pool, ca, 1.8 road mi S of the Refuge Headquarters and Visitor

Nesom: Taxonomy of v

Center, single tree at edge of swamp, above water level, trunk single, erect, ca. 40 feet tall, profusely
fruiting, 13 Jul 2010, 2010-Nesom F7 (FSU, GH); W side of Wakulla River, swamp along N side of
Hwy 319/95, ca. 3 air mi NW of town of St. Marks, common frees, trunks single, erect, 10-40 feet
tall, few in fruit and fruits high and out of reach, 13 Jul 2010, Nesom 2010-F8 (FLAS, FSU, GH,
NCU, NY, TEX); near Rd 59 ca 6 mi N of the St. Marks Lighthouse, cypress-hardwood swamp,
abundant, free up to 20 feet tall, 20 Jun 1950 Redfearn 2510 (FSU, GH). Walton Co. :
Choctawhatchee River delta, 12 April 1975, Suttkus 75-6-12 (FLAS). Washington Co. : E edge of
Caryville, 20 Aug 1950, Kurz T-3 (FSU). Georgia. Charlton Co. : along St. Mary's River, Trader's
Hill section. 15 Apr 1940, Kelley 50 (AA); St. Mary's River swamp, below Trader's Hill, 12-15 Jun
1895, Small s.n. (AA, NY); St. Mary's River swamp, below Trader's Hill, 24-26 Jul 1895, Small s.n.
(AA, NY). Decatur Co . : at easternmost bridge at head of Florida Sterile Hospital Pond, near Jink's
Siding, NE of Chattachoochee, in creek swamp, 19 Aug 1975. Gholson 4562 (FLAS). Dougherty
Co. : along the Flint River at Albany, 24-28 May 1895, Small s.n. (AA, NY). Citations of collections
from FLAS are provided by Walter Judd, who identified the specimens.

In the Trader's Hill area of southeastern Georgia, whence the type of Fraxinus pauciflora,
collections also have been made of F. caroliniana. J.K. Small collected both species there in June of
1895. USA Georgia. Charlton Co. : St. Mary's bottomland, Thompson's Landing, 21 Jul 1958,
Cypert 113 (GA); branch of St. Mary's River, Trader's Hill, snail tree, 3 Apr 1918, Harbison 4
(AA); St. Mary's River, Cedar Landing, river swamp, 31 May 1930, Harper 231 (GH); in the St.
Mary's River swamp, below Trader's Hill, 12-15 Jun 1985, Small s.n. (AA).

The abaxial leaf surface of Fraxinus pauciflora has a distinctive morphology otherwise

known only in the F. americana group (F. americana, F. biltmoreana, F. smallii, and F. albicans).
This is easily observed with a dissecting microscope (40X) and once the nature of the character is
understood, a 10-20X hand lens may be sufficient to make the observation.

Lingelsheim (1908) was fully aware of the distinctive abaxial leaf morphology of Fraxinus
americana and upon studying specimens of F. pauciflora from Florida recognized the similarity in
this feature of these water ash trees to white ash. His explicit speculation that they were hybrid in
origin between F. americana and F. caroliniana was reflected in his choice of epithet.

Collections at NY of Fraxinus pauciflora studied by Gertrude Miller were annotated by her
(on 25 May 1951) as 'Fraxinus americana complex" or in a few cases as 'Fraxinus americana L. —
caroliniana Mill." Earlier, at GH in 1950, she had annotated some collections of F. pauciflora as "F.
floridana Sarg.?" She was aware of the epidermal distinction and in her monograph (Miller 1955, p.
25) she observed that these trees, which she knew at the time only from Lake County, Florida, "seem
to be the result of a rare or chance hybrid between a member of the red ash complex and one of the
white ash complex. ... This hybrid has persisted at or near the point where the original hybrid
appeared, but has not been able to extend its range, or only very slightly. Until more evidence
becomes available, and largely for convenience, in this study the hybrid is arbitrarily called F.
caroliniana Mil. - c

Sargent (1902, p. 39, plate 717) also separated Fraxinus caroliniana andF. pauciflora (using
the name F. floridana for the latter), observing that "A small Ash-free which grows in ponds and deqj
river-swamps in eastern and western Florida and ill southern Georgia and vhich has usualh been
considered a form of the Water Ash, Fraxinus Caroliniana, varies constantly from that species in the
form of the fruit. It is desirable that a plate of this second species of Water Ash should appear in a
Silva of North America, and although the foliage and winter -buds do not afford characters by which
the two trees can be readily distinguished in the herbarium, it is convenient to treat them as species
rather than as varieties." Sargent's note that collections of F. floridana were made "near Jacksonville,

Nesom: Taxonomy of v

Eustis, and Apalachicola, Florida, and in Charlton County, southern Georgia" leave little doubt that it
is the same species identified here as F. pauciflora.

In limited field study in northern Florida (two days in Franklin, Liberty, Leon, and Wakulla
counties), I saw Fraxinus caroliniana only along the immediate sides of creeks and rivers. Fraxinus
pauciflora apparently is much more common in that area, occurring within and along the edges of
swamps. Only a small percentage (ca. 10-20%) of the F. pauciflora trees were in fruit. In other
areas, however, label data indicate that F. caroliniana sensu stricto has a broader ecological range.

ACKNOWLEDGEMENTS

I am grateful to Terry Peacock, Refuge Manager, St. Marks National Wildlife Refuge, for
permission to make collections on the federal property, to stalls at BRIT-SMU, FSU, GH, NY, to Eva
Wallander for comments and digital images of Fraxinus cubensis, Daniel Ward for discussions and
perspective on Florida Fraxinus and his comments on the manuscript, and Walter Judd for identifying
FLAS collections of water ashes and providing the specimen data for F. pauciflora (as cited here), to
TEX-LL for hospitality during visits, and to GA and GH for loans to TEX for my study there. This
study was done in part under contract for the Flora of North America Association, in conjunction with
preparation of the FNANM treatment of Fraxinus.

LITERATURE CITED

Brown, C.L. and L.K. Kirkman. 1990. Trees of Georgia and Adjacent States. Timber Press,

Portland, Oregon.
Clewell, AF. 1985. Guide to the Vascular Plants of the Florida Panhandle. Florida State Univ.

Press, Tallahassee.
Duncan, W.H. and M.B. Duncan. 1988. Trees of the Southeastern United States. Univ. of Georgia

Press, Athens.
Fernald, M.L. and B.G. Schubert. 1948. Studies of American types in British herbaria. Rhodora 50:

186-190.
Godfrey, R.K. 1988. Trees, Shrubs, and Woody Vines of Northern Florida and Adjacent Georgia

and Alabama, Univ. of Georgia Press, Athens.
Kurz, H. and R.K. Godfrey. 1962. Trees of Northern Florida. Univ. of Florida Press, Gainesville.
Leon, H. andH. Alain. 1957. Flora de Cuba. Vol. IV. Contrib. Ocas. Mus. Hist. Nat. La Salle 16: 1-

556.
Lingelsheim, A. 1908. Vorarbeiten zu einer Monographic der Gattung Fraxinus. Bot. Jabrb. 40:

185-223.
Lingelsheim, A 1920. Oleaceae-Oleoideae-Fraxineae. Pflanzenr. IV 243: 1-61.
Little, E.L., Jr. 1952. Notes on Fraxinus (ash) in the United States. J. Wash. Acad. Acad. Sci. 42:

Long, R.W. and O. Lakela. 1976. A Flora of Tropical Florida. Banyan Books, Miami, Florida.
Mller, G.N. 1955. The genus Fraxinus, the ashes, in North America, north of Mexico. Agric. Expt.

Si., \ k-ni >>\'\.iikII I im . llh.K.1. . ',
Nelson, G. 1995. The Trees of Florida. Pineapple Press, Inc., Sarasota, Florida.
Nesom, G.L. 2010. Notes on Fraxinus profunda (Oleaceae). Phytoneuron 2010-32: 1-6.
Nuttall, T. 1849. The North American Sylva , vol. 3. Smith & Wistar, Philadelphia.
Palmer, E.J. 1932. Leaves from a collector's note book. J. Arnold Arb. 13: AY1-M1.
Sargent, C.S. 1894,1902. Silva of North America. (1894. Fraxinus caroliniana, Vol. 6, pp. 55-56

plates 274, 275; 1902. Fraxinus floridana, Vol. 14, p. 39, plate 717.
Small, J.K. 1913. Florida Trees: A Handbook of the Native and Naturalized Trees of Florida.

Published by the author, New York.
Wallander, E. 2008. Systematics of Fraxinus (Oleaceae) and evolution of dioecy. Plant Syst Evol

273: 25^9.

Nesom : Taxonomy of water ashes 1 Q

Weakley, A.S. 2010. Flora of the Southern and Mid-Atlantic States. Working draft of 8 March

2010. Univ. of North Carolina Herbarium (NCU), Chapel Hill.

<http://www.herbarium.unc.edu/WeakleyFlora20 1 0Mar.pd£>
Wenzig, T. 1883. Die GdXiung Fraxinus Tourn. Bot. Jahrb. 4: 165-188.
Wunderlin, R.P. and B.F. Hansen. 2003. Guide to the Vascular Plants of Central Florida (ed. 2).

Univ. Press of Florida, Gainesville.
Wunderlin, R.P. and B.F. Hansen. 2010. Atlas of Florida Vascular Plants. [S.M. Landry and K.N.

Campbell (application development), Florida Center for Community Design and Research]

Inst, for Systematic Botany, Univ. of South Florida, Tampa, <http://www.plantatlas.usf.edu/>

Figure 1. Distribution of Fraxinus caroliniana and F. cubensis. Fraxinus cubensis also o<
western portions of Cuba (see text).

Figure 2. Distribution of Frax

n : Taxonomy of water ashes 1 2

Figure 4. Representative samaras of Fraxinus cubensis.

Nesom : Taxonomy of water ashes 1 3

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Figure 5. Representative samaras of Fraxinus caroliniana.