I Spermo/ep/s (Ap'iaceae). Phytoneuron 2012-87: 1^9. TAXONOMY OF APIASTRUM, AMMOSELINUM, AND SPERMOLEPIS (APIACEAE) Guy L. Nesom 2925 Hartwood Drive Fort Worth, Texas 76109 guyTtesom@sbcglobal.net ABSTRACT A taxonomic summary is given for the three closely related and primarily North American genera Apiastrum (1 species), Ammoselinum (3 species), and Spermolepis (11 species). Ap iastru m includes the single species A. angustifolium, which occurs in California, Baja California, and Baja California Sur. Ammoselinum includes A. rosengurtii (endemic to Uruguay) and the North American A, butleri and A, popei, Spermolepis includes one species endemic to the Hawaiian Islands — S. hawaiiensis, one species endemic to Argentina — S. castellanosii, and nine species native to North America: S. echinata, S. mermis, Spermolepis organensis Nesom, sp. nov. (Dona Ana Co., New Mexico), Spermolepis laevis Nesom, sp. nov. (central Texas to south-central Oklahoma), S. divaricata, Spermolepis (Leptocaulis) diffusa (Nutt. ex DC.) Nesom, comb, nov., Spermolepis (Ammoselinum) gigantea (Coulter & Rose) Nesom, comb, nov., Spermolepis lateriflora Nesom, sp. nov. (California, Arizona, New Mexico, Texas, Chihuahua, and Sonora), and Spermolepis infernensis Nesom, sp. nov. (San Diego Co., California). Species descriptions and keys to the genera and species are provided and a discussion of inflorescence architecture points out distinctions within and among the genera; most of the species are illustrated by photos of specimens. An epitype is designated for Spermolepis hawaiiensis; leetotypcs are designated for Apiastrum latifolium (a synonym of Apiastrum angustifolium), Ammoselinum popei. Ammoselinum castellanosii Ammoselinum sect. Hesperoselinurn, the genus Leptocaulis (a synonym of Spermolepis), and Spermolepis (Leptocaulis) diffusa, KEY WORDS: Apiastrum, Ammoselinum, Spermolepis, Apiaceae, inflorescence architecture The genera Apiastrum, Ammoselinum, and Spermolepis are very similar among themselves and are all placed in tribe Selineae (Downie et al. 2010) of subfamily Apioideae. The species are primarily North American but Ammoselinum and Spermolepis each include a South American species and Spermolepis hawaiiensis is endemic to Hawaii. The monospecific Apiastrum is restricted to Pacific coastal region of Mexico (Baja California and Baja California Sur) and California. Plants of Ammoselinum and Spermolepis are annuals with narrow, characteristically linear to filiform leaf segments, narrow involucel bractlets but lacking an involucre, white petals with straight apices, laterally compressed fruits, narrowly conical stylop odium, and styles absent, the sigmas divergent Apiastrum is similar but apparently is specialized in its lack of sepals, stylopodium, and involucel bractlets and its branching-inflorescence architecture. Loss of peduncles has occurred in some species of all three genera. 1. Medial and distal leaves appearing opposite; schizocarps dqiressed-ovoid, 1-1.5 mm; sepals absent; stylopodium obsolete, styles filiform; involucel bractlets absent Apiastrum 1. Leaves alternate; schizocarps broadly ovoid to ovoid-oblong, urceolate- ovoid, or urceolate- oh long, 1.5-5 mm; sepals small but present; stylopodium present, styles obsolete; involucel bractlets present. 2. Schizocarp ovoid-oblong to urceolate-ovoid or broadly ellipsoid, ribs sparsely to densely scaberulous with single-celled papilla-like projections Ammoselinum 2. Schizocarp broadly ovoid to ellipsoid or elliptic-ovoid, ribs and intervals variously hairy or at least tuberculate Spermolepis Nesom: Taxonomy of Apiastrum, Ammoselmum, and Spermolepis In maintaining Ammoselinum and Spermolepis as separate genera in the present manner, the definition of Ammoselinum is narrowed to only the three species with mericarps scabrous-ribbed, otherwise glabrous, and with corky-expanded, appendage-like lateral ribs (Fig 1). Those species without expanded lateral ribs and with hairs or at least tubercles on both the ribs and intervals are referred to Spermolepis. Considerable variation in fruit shape and vestiture also exists among the species of Spermolepis as accepted here (Fig. la-q). Spermolepis gigantea and S. castellanosii appear to be distinct as a pair on the basis of the relatively elongate fruits (compared to other species of Spermolepis), but S. gigantea is unique in its hispid-hirsutulous fruit vestiture with long, sharp- pointed hairs without tuberculate bases; hairs of S. castellanosii are similar to those of S. infemensis. Spermolepis divaricata and S. diffusa are distinct as a pair on the basis of their short-ellipsoid fruits with tiny upcurved hairs; they also are distinct from the rest of the genus in their relatively smooth epidermis (vs. minutely "bubbly'' in the others). Spermolepis echinata, S. hawaiiensis, S. lateriflora, S. infemensis, and S. inermis are similar in their broadly ovoid fruits with multicellular, tuberculate trichome bases and the first three species are echinate-brrstly with apically hooked hairs. Figure la-u. Mature or near-mature fruits of Apiastrum, Ai but not exactly the same scale (see descriptions for measure Ammoselinum pope i, (a) Ammoselinum rosengurtii, (d)Amj loselinum, and Spermolepis. At approximately 2nts). {a) Apiastrum angustifolium, (b) selinum butleri. i: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 3 Figure le, f, g, h. (e) Spermolepis echinata, (f) S. lateriflora, (g, h) variants ofS. ham n: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 4 e ] i, j, k, 1, m, n. (i) Spermolepis echi text (j) S. echinata x 5. inermisf?, Cory 48' ■is , (m) S. inermis, (n) 5. /aevw. Nesom: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 5 Figure lo, p, q, r, s, t. (o) Spermolepis laevis variant with slightly rugulate ribs and intervals - Wolff '2102, (p) S. laevis variant with a few tubercles, some with short, blunt-tipped hairs - Whitehouse 18439, (q) S. divaricata, (r) S. diffusa, (b, t) S. castellanosii Nesom: Taxonomy of fyfatfrun, Ammosefaium, aid Spermdepe . . t 4**J& ■ u «s Figure lu. ifyxrmolepis gigantea (at much larger scale than the other fruit photos). Among the genera of tribe Selineae in the account by Downie et al. (2010), Oligzclaaus Chodat & Wilczek is the only other American genus in the tribe outside of the "Arracacia Clade" and the "Perennial Endemic Norm American Clade" — except for Ammoselimim and Spermolepis, all other genera are Eurasian. Oligzclaatis includes only the single species O. pala^micas (Speg.) Perez- M or. (synonym = O. anehmts Chodat & Wilczek), which apparently is restricted to Argentina. Mainly because of its dorsally compressed fruits (flattened parallel with the plane of the commissure) with numerous oil tubes on the broad commissural face, M ami as and Constance (1950) eliminated Oligzclaatis as a possible congener or even close relative when considering the generic placement of their new species Ammoselimim rosengirtii (see illustrations of O. amtmis in Chodat and Wilczek 1902, pp. 527—528, vs. lateral compression in the Apiastnim-Ammoselimim-Spermolepis group, flattened perpendicular to the plane of the commissure). The base chromosome number of the Apiastrum, Ammoselimim, and Spermolepis group appears to be x = 11, as it appears in all three genera, with reductions to x = 10 and x = 8. Among the uncinate-bristly species of Spermolepis, three dysploid levels exist: 2n = 22 (S hcrwaiiensis), 2n = 20 (S ecHnald), and 2n = 16 (S lateriflora). If the uncinate-bristly species represent a single clade, men the dysploid changes appear to be more indicative of individual speciation events that consistent : Taxonomy of Apiastrum, Ammoselinum, and Spermolepis indicators of cladistic relations hips. Interesting research remains to be done with regard to chromosome numbers (see comments following S. divaricatd). A close ifclduonsh,} at u.»ij ihe 1 3 species of Apiastrum, Ammoselinum, and Spermolepis is suggested by morphological similarities as well as their general geographic coherence in a broad region (North America, South America) where other potentially related species apparently do not exist. It would not be indefensible to treat all 13 species within a single genus. Branching pattern and umbel architecture In all species of Ammoselinum except one and in two species of Spermolepis, umbels are borne on ebracteate peduncles mat appear to originate only at leaf axils (Fig. 2A). Growth is indeterminate, as upward vegetative growth is continued even at the distalmost node. In these plants, however, the peduncle appears to be the extension of the primary ("pr") stem axis. Continued upward stem growth and production of additional umbels continues from growth of the axillary ("ax") bud. In this interpretation, the umbels actually are produced as terminal structures, rather than axillary ones. In Ammoselinum butleri, Spermolepis lateriflora, and S. infernensis, the peduncle is absent (presumably suppressed) and the umbellet rays appear to arise from the leaf axils (Fig. 2C). The axillary bud in these plants apparently is suppressed ;it the distalmost node, so that growth may be characterized as determinate. Peduncle suppression is complete in A. butleri but in S. lateriflora, apparently over its whole geographic range, some plants produce pedunculate umbels from nodes below the distalmost (see Figures 7 and 8 and examples cited below, under the species). In Spermolepis divaricata, S. diffusa, and S. inermis, the primary axis at each node forms a peduncle and compound umbel, but on the branch arising from the distalmost axillary bud, both the terminal leaf and the axillary bud are suppressed (Fig. 2B). In effect, each branch terminates in two compound umbels and growth may be characterized as determinate. In Apiastrum angustiflolium, leaves appear to be opposite and two branches and two sessile (compound) umbels arise at each node (Fig. 2D). In the interpretation hers, this arises from (a) suppression of the peduncles, (b) complete foreshortening of the distal internode that would constitute the axillary axis, and (c) duplication of the axillary bud in order that growth continues from two upward branches. Axillary buds apparently are suppressed at the distalmost node and growth may be characterized as determinate. Nesom: Taxonomy of Apiastrum, Ammoselinum, and Figure 2. Variation in inflorescence architecture in Ammoselinum., Apiastrum, and Spermolepis. A. Pedunailaie-axillary-in determinate: Ammoselinum popei, Ammoselinum rosengurtii, Spermolepis eciunaM, S. hawaiiensis, & castelkmosii. B. Pedunculate- axillary-determinate: Spe}tiv:Aepi.s divaricata. S. diffusa., S inermis, S. laevis, S. orgaiiensis, S. gigantea. C. Sessile-axil lary-deteraiin ate: Ammoselinum butleri, Spermolepis lateriflora, S. infernensis. D. Sessile-dichotomous- determinate: Apiastrum angustifoUum. "ax" = axillary brand], arising from the axillary bud. "pr" = primary branch, continuing from the main branch from below. In determinate arrangements, rise axillary bud is suppressed at the distaimost node. : Taxonomy of Apiastrum, Ammosdinum, and Spermolepis APIASTRUM Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1: 643. 1840. TYPE: Apiastrum angustifolium Nutt. ex Torr. & A. Gray Herbs, annual, with odor, 0.4-5 dm, glabrous; taproot slender. Leaves appearing opposite to subopposite medially and distally; basal and cauline 3-4 ternately compound; blades broadly ovate to ovate in outline, herbaceous; leaflets divided, ultimate divisions linear to narrowly oblong, margins entire; petioles scarious margined at base, distal petioles foreshortened and scarious-margined along whole length. Umbels compound, loosely convex, axillary, sessile (rays appearing to arise from leaf axils), peripheral flowers not different; involucral bracts absent; involucel bractlets absent. Pedicels present. Flowers bisexual; sqjals absent; petals white, margins, entire, apices slightly inflexed; stylopodium depressed-reduced, nearly obsolete; styles evident, filiform, 0.2 mm, arching-divergent. Schizocarps depressed-ovoid [mericarps reniform], laterally compressed, not beaked, splitting, ribs 3. barely raised, mostly delimited by line of papillae, oil tubes 1 per interval, filiform not filling the interval, surface shallowly tuberculate, otherwise glabrous; carpophore bifid the whole length. Base chromosome number, x = 11. 1. Apiastrum angustifolium Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1(4): 644. 1840. TYPE: USA. California. [San Diego Co.:] St. Diego, N Cal, April, T. Nuttall s.n. (probable holotype: GH 00075076; isotypes: K digital image!, PH 01015720 digital image!). Apiastrum angustifolium var. tenellum Nutt. ex Torr. & A Gray, Fl. N. Amer. 1(4): 644. 1840. TYPE: MEXICO. Baja California. Cerro [Cedros] Island, Mar 1889, E. Palmer s.n. (probable holotype: PH 743994 digital image!). Torrey and Gray did not indicate that they saw a collection; their concept of the taxon was from Nuttall's manuscript. The plant in the PH collection matches Torrey and Gray's brief description ("stem dichotomous from the base; leaves less divided; rays of the umbel very slender; umbellets 1-2- flowered; seed morerugulose"). Apiastrum latifolium Nutt. ex Torr. & A Gray, Fl. N. Amer. 1(4): 644. 1840. LECTOTYPE (designated here): USA. California. [Santa Barbara Co.:] "St. Barbara, N Cal." [on PH sheet], no other collection data, T. Nuttall s.n. (GH 00075075; isolectotype: PH 01044838 digital image!). Torrey and Gray cited "Nuttall! Douglas!" Helosciadium leptophyllum var. ? latifolium Hook. & Arn., Bot. Beechey Voy., 347. 1838. No collection was cited (pp. 347-348). The protologue gave only this: "The specimens are only in young fruit, and the segments of the upper leaves are considerably broader than in any form we have yet seen, while even the lower ones are broader than in H. laciniatum, DC, which we consider a mere variety of this species." As synonym of Apiastrum fide Mathias and Constance (1945). Stems 4-50 cm. Leaves: blades 1-5 cm, ultmutw segments 5-25 mm; petioles 20-40 mm. Peduncles absent. Umbels axillary only: involucre bracts absent; involucel bractlets absent; fruiting rays 2-5 per node (2 umbels per node), (0-)7-25(-50) mm (central umbellet sessile); umbellets 3-7 flowered; fruiting pedicels (0-)2-8(-15) mm (central l-2(-3) flow^ers sessile), unequal, spreading. Schizocarps 1-1.5 mm. In = 22 (San Diego Co. - 2 counts, Bell & Constance 1957; Baja California, Constance et al. 1976). Map 1. Figure 3. Flowering Mar-Apr, Chaparral, coastal sage scrub, blue oak savanna, rock outcrops, granite slopes, shale slopes, serpentine soil, steep slopes, recently burned areas, grassy openings, roadsides; 0-400(-1500) m. Calif.; Mexico (Baja California. Baja California Sur). n: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis \Q (k\ / j ~~j I ftjKAiLip^Xj ^Oo^ :{a p-r— __ • ^-\A\ J A \ | \> / &A~ ~~~~ff — ^ \ A /A S 1 La 7v^^^ V ) \ I AaC \ Apiastrum angustifolium .it iiS^lmJ 4000 ft, 28 Apr 2010, Heil & McClain 32322 (SJC); San Simon Valley between Rodeo and Arizona-New Mexico line, fine textured soils of the valley bottom in desert grasslands, 4100 ft, with Hilaria mutica, Scleropogon brevifolius, Bouteloua eriopoda, Ephedra trifurca, Prosopis, Gutierrezia sarothrae, 10 April 1978, Moir 1 01 (NMC). Luna Co. : ca. 1 1 mi NW of Florida Station, among rocks in bed of dry creek, 29 Apr 1947. McVaugh 8128 (SMU, TEX). Collections examined. MEXICO. Chihuahua. Extreme NW corner [of the state], about 50 m E and 1.5 mi. S of U.S. border, in silty, heavily overgrazed bottom of draw, with Yucca elata, Sporobolus airoides, Astragalus wootonii, 13 May 1980, Spellenberg & Ward 5525 (NMC). Sonorsi. 6.7 mi b\ i j i.l (Mexico li) N ol \Ii Jj1u>« rocky hill- desert skubs 7 *m i'*i3 Eelger 17451 (ARIZ); vicinity of Cerro Pelon, ca. 5 mi SE of Desemboque, 21 Apr 1968, Felger 17894 (ARIZ); near El Guayabo on road 18 km E of Alamos. 27° 0' 20" N, 108° 47' 10" W, 250 m, 16 Mar 1989, Ferguson s.n. (ARIZ); along Hwy 89, 9.2 mi S of Arroyo Los Ajos, 7.4 mi N of Mututucachi, roadside in oak savanna, 212 Apr 1995, Fishbein 2242 (ARIZ); Rio Mayo, San Bernardo, arroyo, Lower Sonoran, margin of a wash, 22 Feb 1935, Gentry 1339 (ARIZ.); Carbo, 50 km N of Hermosillo, wash near Mex Hwy 15, 6 Apr 1975, Helmkamp s.n. (UCR); Rancho El Aguilar Noria, N of Ures and Santiago, 29° 33' N, 110° 25-26' W, open, broad drainage, Sonoran desert scrub, on mesic N slopes, ca. 500 m, common, 21 Apr 1991, Joyal 1995 (TEX); Dto. de Altar, Picu Pass, 23 Mar 1926, Long 7a (ARIZ); 18.7 mi W of Rte 19, along turnoff 3.2 mi N of Esqueda, oak grassland, mesquite bottomlands, 27 Mar 1970, McGill 64048 (ASU); 4.6 mi S of Cucurpe, cliff-face along San Miguel River and road, 29 Mar 1970, McGill & Pinkava 6519 (ASU); Near El Guayabo on road 18 km E of Alamos, 250 m, 16 Mar 1989, Martin & Ferguson s.n. (ARIZ); Mpio. Soyopa, E side of Mex 16, 1 km S of Rio Yaqui Bridge, 28° 31' 30 " N 109° 32 W, 180 m, 14 Mar 19SS,Martin, Ferguson, &Moore s.n. (NMC); NE side of Rio Yaqui bridge on Mex 16, just S of Tonichi, 200 m, in sand, 18 Feb 1997, Reina G. 97-44 (ARIZ); Arroyo La Quema, near Tepoca, tropical decid. forest on slopes, rocky stream canyon, 560 m, 21 Mar 1998, Reina G. 98-378 (ARIZ); Mpio. Benjamin Hill, 3.5 km SW of Benjamin Hill on road to Palo Alto, Sonoran desert scrub, 2408 ft. locally very common annual on disturbed rocky soil, 1 Jan 2003, Reina G. 2003- 286- A (ASU); 0.8 km N of Mex 16 on road to San Antonio de la Huerta, 28° 34' 16" N, 109° 34' 52" W, very open thornscrub, 299 m, locally uncommon on flats, 15 Mar 2005, Reina G. 2005-256 (TEX); Agua Prieta, Hwy 2, ca. 26 mi E of Agua Prieta and 24 mi W of the state line at Puerto San Luis, desert scrub of Juniperus, Acacia. Prosopis, Yucca baccata, Gutierrezia, etc. on rocky volcanic hills, 1300 meters, 19 Mar 1984, Sanders 4712 (UCR); Alamos, Rio Mayo Region, Rancho La Huerta, ca. 2 mi NW of Alamos on the road to San Bernardo, north of the Alamos airstrip, weedy disturbed areas near buildings and roads, 420 meters, 15 Mar 1993, Sanders 13152 (UCR); Alamos, Rio Mayo region, roadside c. 12 km W of Alamos on the road to Navojoa, in vicinity of Canon Agua Marina, at the foot of the Sierra de Alamos, burned roadside in hilly country 1640 ft, 15 Mar 1993, Sanders 13180 (UCR); Alamos, Parque Chalaton and along canyon bottom above, SW edge of Alamos in foothills of the Sierra de Alamos, tropical deciduous forest and cleared areas, 420-450 meters, 17 Mar 1993, Sanders 13365 : Taxonomy of Apiastrum, Ammosdinum, and Spermolepis (UCR); Dist. Alamos, near Cerros, 4 Mar 1933, Shreve 6167b (ARIZ); 8 mi S of Estacion Llano, 3 Apr 1935, Shreve 7323 (ARIZ): Palm Canyon, 17 mi SE of Magdalena, in Sierra Babiso, (= Cerro Cinta de Plata), stream bed,13 Feb 1977, Van Devender s.n. (ARIZ); 4 mi of El Ocuca on Mex 2, 21.4 mi E of Altar, annual in wash, 10 Mar 1977. Van Devender s.n. (ARIZ); 17 mi SE of Magdalena on road to Cucurpe, Palm Canyon, Cerro Cinta de Plata, 15 May 1979, Van Devender et al. s.n. (ARIZ); Alamos, Rio Mayo region; Arroyo Mentidero at the crossing of El Chinal Road, near Rio Cuchujaqui, 11.5 km (by air) S of Alamos, tropical deciduous forest, 240 meters, 10 Mar 1993, Van Devender 93-97 (ARIZ, UCR); Alamos, Rio Mayo region; La Huerta, l.SkmNNE of Alamos on San Bernardo Road, 410 meters, 9 Mar 1993, Van Devender 93-216 (ARIZ, UCR); La Huerta, 1.8 km NNE of Alamos on San Bernardo Road, common annual in yard, 410 meters, 9 Mar 1993, Van Devender 93-21 6 (ASU); 0.4 mi E of Punto Cirio, Sierra Bacha, Sonoran Desert desert scrub, 40 m, 24 Mar 1995. Van Devender 95-210 (ARIZ. UCR); El Llano de Curea, foothills thornscrub, locally uncommon annual, 514 meters, 19 Mar 2004, Van Devender 2004-161 (ASU); 2.2 km SE of Rancho Las Borregas headquarters on road to Nogales, SE tributary of Arroyo Planchas de Plata, sycamore- oak canyon, 1187 m, 22 Apr 2004, Van Devender 2004-250A (ARIZ); 7.9 mi N of Esqueda, 11 May 1948, Wiggins 11 777 (TEX). 2. Spermolepis echinata (Nutt. ex DC.) A. Heller, Contr. Herb. Frankl. & Marsh. Coll. 1: 73. 1895. Leptocaulis echinata Nutt. ex DC, Prodr. 4: 107, 1S30. Avium echinatum (Nutt. ex DC.) Benth. & J.D. Hook, ex S. Wats., Bibl. Index N. Amer. Bot, 412. 1878. TYPE: USA. Arkansas. "In Amer. bor. ad Red River" [protologue], T. Nuttall s.n. (holotype: BM digital image!; isotype: PH digital image!). De Candolle noted ("v.s. ") that he had seen the Nuttall collection. Stems 5^10 cm. Leaves: blades broadly ovate in outline, 0.7-2.5 cm, 3-pinnately compound, ultimate divisions filiform, 2-18 mm x 0.5-1 mm; petioles 3-20 mm. Peduncles (l-)2-5(-6.5) cm. Umbels axillary mostly at distal nodes, al! pedunculate; invoiucel bractiets l-3(-4), linear. 1-3 mm, margins scabrous-toothed; fruiting rays 5-9(-12), (0)1-15 mm (central umbellet sessile to subsessile), unequal, suberect and evidently clustered; umbellets (l-)3-9-flowered; fruiting pedicels l-6(-7) mm (central flowers short-pedicellate). Schizocarps 1.5-2 mm, densely echinate-bristly. 2n = 20 (Constance et al. 1976; Prairie Co., Arkansas, Demaree 61921, duplicate SMU!). Figure 8. Flowering (Mar-)Apr-May(-Jun). Sand, gravel, silt, sandy clay, sandy roadsides and flats, disturbed areas, ditches, disturbed sites, pastures, rocky slopes, shell banks, sandstone outcrops, beaches, creek bottoms, lake shores, prairies, post oak woods, live oak woods, oak-inesquite woodland, desert shrub; (0-)100-300(-1500) m; Ala., Ark, Fla., Ga., 111., Iowa, Kans., Ky., La, Miss, Mo, NY, N.C., Okla, S.C, Term., Tex, Va.; Mexico (Coahuila, Tamaulipas). Specimen examined. MEXICO. Coahuila. Piedras Negras, Pringle 8309 (fide Villarreal 2001, voucher not seen in present study). Tamaulipas. 20 mi W of Reynosa, desert scrub in clayish soil, 28 Feb 1944, Painter & Barkley 14378 (TEX). Attributions of Spermolepis echinata to Arizona, California, and New Mexico have been based on collections identified here as S. lateriflora. In Texas, typical S. echinata reaches as far west as Brewster, Culberson, Jeff Davis, Pecos, and Presidio counties, but it does not extend to El Paso Co. at the easternmost extension of the distribution of S. lateriflora. No confirmed records of S. echinata exist from New Mexico. A plant collected in north-central Texas has the habit and inflorescence of Spermolepis echinata and echinata-like fruits (densely tuberculate-hairy) but with the hairs relatively short and without an uncinate apex. Wilbarger Co.: 14.5 mi W of Electra, Waggoner pastures, turn W 0.6 mi S on Hwy 85, tall grass in draw, mesquite savanna, sandy loam, 12 May 1945, Whitehouse 9828 (SMU). : Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 3. Spermolepis hawaiiensis H. Wolff, Repert. Spec. Nov. Regni Veg. 17: 440. 1921. TYPE: USA. Hawaii. Kauai, Weimea, [no date], Hillebrand s.n. (holotype: probably B, Wolff material at B mostly extant fide HUH online database). EPITYPE (designated here): USA. Hawaii. (Kauai Island): Koai'e Canyon, just below "the fingers" near the ridge and above N-facing cliffs W of Lonomea Camp and Kawai'iki and E of Hipalau, 704 meters, 21 Apr 2004, N. Tangalin 47 (PTBG 043006, digital image on JSTOR!; Fig. 9). This collection was made near the type locality. Stems 5-20 cm. Leaves: blades oblong to ovate in outline, 1-4 cm, 3-pinnately compound, becoming sessile, smaller, and less divided distally, ultimate divisions linear to linear-lanceolate, 3-6 mm; petioles 10-30 mm. Peduncles 1-3 cm. Umbels at distal nodes, axillary, all pedunculate; involucre bracts absent; involucel bractlets (0-)l-5, linear-lanceolate. 1-6 mm; fruiting rays 2-7, (0- )5-15 mm (central umbellets sessile to short-pedicellate), unequal, spreading-ascending; umbellets 2- 8-flowered; fruiting pedicels (0-)2-6 mm (central flower sessile to short-pedicellate), unequal, spreading-ascending to ascending. Schizocarps 3-4 mm, densely echinate-bristh', hairs arising from multicellular tuberculate bases. 2« = 22 (Wagner et al. 2005). Figure 9. Flowering (Dec, Feb-)Mar-Apr. Steq3 mesic forests, gulch slopes and ridge tops in dry forest, shrub lands, steep to vertical cliffs, cliffs bases, ridges in coastal dry cliff vegetation, N-facing slopes, ridges on bare rock, open, rocky, goat-ravaged area, a'a lava; 50-700 m; endemic to the Hawaiian Islands — Hawaii, Kauai. Lanai, Maui, Molokai, Oahu. Information from NTBG (2012) and USFWS (2010). 4. Spermolepis infernensis G.L. Nesom, sp. nov. TYPE: USA. California. San Diego Co.: Hell Hole Canyon near Borego, 5-7 Apr 1932, C. Epling & W. Robison s.n. (holotype: RSA!; isotype: UC! digital image!). Similar to Spermolepis lateriflora in its epedunculate (sessile) umbels but different in its sparse fruit vestiture of apically straight, blunt-tipped hairs. Stems 7-13 cm, branching from the base. Leaves: blades broadly ovate in outline, mostly 1- 2 cm, finely ternately dissected, ultimate divisions linear to oblong, mostly 2-6 mm; petioles 10-20 mm. Peduncles usually absent, occasionally present and 2-3.5 cm. Umbels axillary, sessile at distal nodes, sometimes pedunculate at proximal nodes; involucre bracts absent; involucel bractlets (l-)2-4, linear, 1^1 mm, margins entire, herbaceous, without scarious margins; fruiting rays 4-5 per node, (0- )3-10 mm (inner umbellets sessile or subsessile to short-pedicellate), unequal, spreading; umbellets (2-)4-7 flowered; fruiting pedicels 1-5.5 mm (inner flowers short-pedicellate). Schizocarps 1.5-2 mm, ribs rounded, sparsely to moderately tuberculate to hispidulous-spinulose on the angles and intervals with pustulate multicellular mounds, each pustular mound with a straight, erect, blunt-tipped, unicellular, hairlike cell, 0.1-0.2 mm or some mounds without a hair, lateral ribs not strongly differentiated; oil tubes 3 per interval, barely evident between the thickened ribs. Chromosome number not reported. Map 5. Figures 10, 11. Flowering Mar-Apr. Desert shrubland; 600-700 m; California (San Diego Co.). In deriving the concept and description of Spermolepis infernensis, I have seen only the two sheets of the type collection, which include 9 separate plants of consistent morphology. The type collection is from the area of the Hellhole Canyon Preserve, which is northeast of Escondido at elevations of about 1800-2000 feet elevation. The reference by Epling and Robison to "Borego" apparently meant Borrego, since Hellhole Canyon is in the general vicinity of the Borrego Valley and the Anza-Borrego Desert State Wilderness Park (see comments above in connection with S. lateriflora). : Taxonomy of Apiastrum, Ammosdinum, and Spermolepis Spermolepis infernensis and S. lateriflora perhaps are sister species, in view of their similarity in inflorescence structure and distribution in the western USA It is possible that S. infernensis is a recent derivative of S. lateriflora, with a reduction in the density of the fruit vestiture and a developmental change that results in truncation of the hairs. The difference in appearance is striking, however, and the effect on dispersal potential surely must be significant. 5. Spermolepis inermis (Nutt. ex DC.) Mathias & Constance, Bull. Torrey Bot. Club 68: 124. 1941. Leptocaulis inermis Nutt. ex DC, Coll. Mem. 5: 39, plate 10, fig. B. 1829. Spermolepis patens var. inermis (Nutt. ex DC.) Mathias, Brittonia 2: 243. 1936. TYPE: USA. Arkansas. "In Amer. bor. ad Red River" [protologue], T. Nuttall s.n. (holohpt BM digital innge'. isotype: PH digital image!). De Candolle noted ("v. s.") that he had seen the Nuttall collection. Leptocaulis patens Nutt. ex DC, Prodr. 4: 107. 1830. Apium patens (Nutt. ex DC.) S. Wats, Bibl. Index N. Amer. Bot, 413. 1878. Apiastrum patens (Nutt. ex DC.) Coulter & Rose, Rev. N. Amer. Umbell, 110. 1888. Spermolepis patens (Nutt. ex DC.) B.L. Robins, Rhodora 10: 34. 1908. TYPE: USA. Arkansas. "In Amer. bor. ad Red River" [protologue], T. Nuttall s.n. (holotype: BM digital image!; isotypes: NY digital image!, PH digital image!). De Candolle noted ("v.s. ") thai he had seen the Nuttall collection. Stems 8-80 cm. Leaves: blades oblong-ovate in outline, 3-5 cm, 3-pinnately compound, ultimate divisions filiform, 3-30 mm x 0.1-1 mm; petioles 4-15 mm. Peduncles 2-7 cm. Umbels terminal and axillary, all pedunculate; involucre bracts absent; involucel bractlets 1-4, linear to linear-lanceolate, 2-5 mm, margins scabrous-toothed; fruiting rays 5-11, suberect and evidently clustered, unequal, 1-13 mm (central umbellet subsessile); umbellets 2-7-floweied; fruiting pedicels (0-)l-6 mm (central flower sessile to short-pedicellate). Schizocarps 1.2-2 mm, tuberculate, without trichomes; oil tubes 1 per dorsal interval. Chromosome number not known (reported in error as In = 22 by Bell & Constance 1957; voucher from Florida identified here as S. divaricata). Map 3. Figure 13. Flowering Apr-May (-Jun). Sand, river silt gravelly soil, clay loam, sand prairies, blackland prairies, shale glades and barrens, rocky ridges, granite outcrops, limestone crevices, oak-juniper woodland, ditch banks, woods edges, roadsides, fields; 30-200(-900) m; Ala, Ark, 111, Ind, Iowa, Kans, La, Md, Minn, Miss, Mo, Nebr, N.Mex, Okla, Term, Tex.; Mexico (Coahuila). East of the Mississippi River in the southeastern USA Spermolepis inermis occurs only in widely disjunct localities in Mississippi, Alabama, and Tennessee; in Louisiana and Alabama, the scattered populations occur mostly in prairie patches. Bonafide records seen in the present study are documented here. Mississippi. Monroe_Coj ca. 3 mi SSW of Prairie along Hwy 25, 2.4 mi S of jet with Hwy 382, 7 May 1996, McDonald 9364 (VDB); ca. 1.5 mi SW of Aberdeen, remnant prairie roadside at jet state hwys 25 and 382, 23 May 1996, McDonald 9466 (BAYLU). Oktibbeha Co. : Botanic Garden of the South, ca. 1.5 mi S of Sessums on Sessums Rd, Black Prairie region, 21 May 1997, Leidolf 1512 (VDB). Alabama. Sumter Co. : Blackland prairie between Gainesville and Alabama 17, 20 May 1975. Krai 55614 (VDB), Tennessee. Warren Co.: 1 mi NE of Morrison, ballast of railroad adjacent swamp and Hwy 55, 18 May 1987, Patrick & Wofford 2029 (SMU). Spermolepis inermis probably is not native in Maryland, where it has been reported (Brown & Brown 1984). Radford et al. (1968) noted its occurrence in New Hanover Co, North Carolina, but no voucher exists in the NCU herbarium (Alan Wealdey, pers. comm.). Some Alabama collections at VDB previously identified as S. inermis are instead S. divaricata (Baldwin Co, belong 7712: Dallas Co, Sessler 1 1 78; Geneva Co, Krai 90807), and this mistaken identity probably also has been the case for the Maryland and North Carolina reports. : Taxonomy of Apiastrum, Ammosdinum, and Spermolepis Additional specimens examined (outlying populations). USA. New Mexico. Eddy Co.: Carlsbad Caverns Natl. Park, 1.8 mi WSW of E boundary via sewage lagoon road, 0.7 mi E of sewage lagoons, small arro}~o lined with shrubs, on bajada escarpment, 3650 ft, 19 Apr 1977, Burgess 4496 (TEX); Carlsbad Caverns Natl. Park, Walnut Canyon, ca, 0.2 mi S, 0.2 mi W of BM 3901, gravel alluvium dominated by Brickellia herniate:, 17 May 1977, Burgess 4556 (ARIZ, TEX). MEXICO. Coahuila. Rio Grande, Tule Canyon, on Coahuila side above Upper Madison Falls, calcareous gravelly soil, Dasylirion, Yucca. Molina, Rhus, Acacia, 475 m. 10 Apr 1973, Johnston et al. 10615 (LL); San Rosendo Canyon (flows into Rio Grande opposite Brewster Co.), 500-700 m, calcareous gravelly loam, Dasylirion, Yucca, Molina, Larrea, Acacia, Quercus, Prosopis, 9 Apr 1973, Johnston et al 10597B (LL); Musquiz, spring 1935, Marsh 113 (TEX); Sierra de Santa Rosa. Canon El Puerto, Rancho El Puerto, 28 24 N, 101 54 W, matorral de Acacia coulteri, Pithecellobium pollens, Zanthoxylum fagara, Yucca thompsoniana, y Opuntia lindheimeri, 900-1000 m, 6 Jun 1991, Villarreal 5963 (BRIT): Mpio. Musquiz, Hacienda La Rosita, 26 Jun 1936, Wynd & Mueller 295 (ARIZ). Fruits of a collection from south-central Texas, identifiable as Spermolepis inermis in every other way, have some tubercles producing very short, blunt-tipped hairs, similar to those of S. organensis and S. infernensis. Wilson Co. : 8 mi S of Elmendorf, 4 May 1945, Cory 48795 (SMU). Spermolepis inermis, S. divaricata, S. diffusa, S. laevis, and S. organensis appear to be closely related among themselves. If the ancestral species of Spermolepis had hairy fruits with (e.g., 5. echinata, S. gigantea), then S. inermis can be understood as derived through loss of the hairs. Spermolepis laevis probably is a derivative of 5. inermis, through further loss of the surface ornamentation of the fruits. Spermolepis divaricata and S. diffusa perhaps are sister species, through reduction of the inflorescence from an ancestor shared with A', inermis (maintaining the potential to produce fruit hairs). Spermolepis organensis is possibly a peripheral isolate of S. inermis; its short, stubby fruit hairs are like those of 5. infernensis and rare populational forms of S. inermis, perhaps through partial derepression of the hair formation. 6. Spermolepis laevis G.L. Nesom, sp. nov. TYPE: USA. Texas. Llano Co.: Enchanted Rock, granitic sandy soil, 15 May 1933, E. Whitehouse 11292 (holotype: SMU) Similar to Spermolepis inermis in general appearance, especially its strictly pedunculate, terminal and axillary umbels and its relatively short, suberect and evidently clustered fruiting rays; different in its completely smooth fruit surface, with evident ribs but lacking tubercles on the ribs or intervals. Stems 8^18 cm. Leaves: blades ovate to broadly ovate in outline, 1-4 cm, 3-pinnately compound, ultimate divisions filiform, 4-15(-25) mm; petioles 2-15 mm. Peduncles 2-5 cm. Umbels terminal and axillary, ail pedunculate; involucre bracts or very rarely 1-2; involucel bractlets 1-4, linear to linear-oblong or linear-lanceolate, 1-3C-5) mm, margins scabrous-toothed; fruiting rays 3-8, suberect and evidently clustered, unequal, (0-)2-9(-13) mm (central umbellet sessile to subsessile); umbellets (l-)3-8-flowered; fruiting pedicels (0-)3-5 mm (central flowers sessile to subsessile). Schizocarps 1-1.2 mm, minutely beaked, surface smooth, dorsal ribs 3, oil tubes 1 per dorsal interval. Chromosome number not reported. Map 3. Figure 14. Flowering Apr-May(-Jun). Granite outcrops, granitic gravel, limestone gravel, sandy fields, oak-cedar slopes, live oak savannas; 200-500 m; Okla., Tex. Additional collections examined. Oklahoma. Johnston Co. : 10 mi N of Tishomingo, near Wapanucka Road jet, grazed prairie along State Hwy 99, sandy granitic soil from nearby granite knobs, 29 May 1948 [fruit], Bobbins 3064 (UC). Texas. Bell Co.: nature prairie near Little River, 5 : Taxonomy of Apiastrum, Ammosdinum, and Spermolepis Jun 1930, Wolff 2201 (VDB); near Little River, field, 11 May 1930, Wolff 2102 (BRIT); Temple, from around Substation #5, 11 May 1930, Wolff '21 02 (SMU, perhaps duplicate of the BRIT sheet of same number). Burnet Co. : Granite 'hills'[?], sandy soil, 29 May 1922, Thorp s.n. (TEX); near Burnet, sandy soil, 26 Apr 1931, Whitehouse 11291 (SMI I): Inks Lake State Parle, ca. 1 mi S of Hwy 29, granite outcrop, 1 May 1947, Whitehouse 18439 (SMU). Gillespie Co. : ca. 12 mi N of Fredericksburg, dry soil on oak-cedar slope, 29 Jun 1957, Cor r ell & Johnston 17269 (LL). H amilton Co. : [no other locality data], 12 Jun 1941. Thorp s.n. (TEX). Llano Co. : [no other locality data], 11 Jun 1930, Thorp s.n. (TEX). Mason Co. : Mason Mountain Wildlife Management Area, in Middle Pasture, 0.2 mi N of Mile-O-More Lake, 23 Apr 2001, Sanchez 2355 (BAYLU); MMWMA, in Middle Pasture, near the Lodge, near Una Branta Lake, 19 Apr 2003, Sanchez 3224 (BAYLU); MMWMA m West Pasture, downstream from Comanche Lake, near the Beaver Dam, sandy soil, 15 Apr 2005, Sanchez 3697 (BAYLU); Mason Mountain Wildlife Management Area, in Middle Pasture, 0.9 mi NE of gate into Headquarters Pasture, sandy soil, 29 May 2005, Sanchez 3879 (BAYLU, BRIT); MMWMA in Middle Pasture, 0.3 mi NE of Headquarters Bids., live oak savanna, sandy soil, 17 May 2008, Hansen 5922 (BAYLU, TEX); granite outcrops on S side of RM 1222, 2.6 km E from intersection of US Hwy 87 and RM 1222 at Camp Air, frequent in dry crevice, 17 May 1979, Walters 301 (SMU). Tarrant Co. : near Fort Worth, in old field, 15 Jul 1924, Ruth 1107 (SMU); gravel road N of Crowley under Santa Fe bridge, limestone gravel soil, Grand Prairie. 21 Apr 1946, Whitehouse 15491 (SMU). The Oklahoma collection (Johnston Co.) consists of 5 plants — 3 are typical Spermolepis echinata but 2 are typical S. laevis. the fruits completely glabrous. From Bell Co., Texas, Wolff 2201 has perfectly smooth fruits; fruits of Wolff 2102 (Fig. lo) are very slightly tuberculate but better identified as Spermolepis laevis than S. inermis; Wolff 657 (BRIT) from Bell Co., however, is clearly S inermis. Both taxa have been collected in Bell, Burnet, Gillespie, and Tarrant counties. Field study toward a better understanding of the distribution of Spermolepis laevis and its biology, especially its possible interaction with S. inermis, will be interesting and useful. 7. Spermolepis organensis G.L. Nesom, sp. nov. TYPE: USA. New Mexico. Dona Ana Co.: Organ Mts., Rock Springs Canyon, NWNW Sec 34, T22S, R4E, common on gravelly loamy granitic soil on 5 deg N-facing slope, with Quercus arizonicus, Juniperus deppeana, Garrya wrightii, Cercocarpus montanus, Rhus trilobata, 5400 ft, 6 Jun 1995, L Mcintosh 3106 (holotype: NMC!). Similar to Spermolepis inermis in its strictly pedunculate, terminal and axillary umbels; different in its shorter fruiting peduncles and in its corky fruit surface with vaguely formed tubercles, some tubercles producing short, straight hairs, some without hairs. Stems ca. 20 cm, purple. Leaves: blades broadly ovate in outline, 1.5-2 cm, 2-3 ternately compound, ultimate divisions filiform, 4—11 x 0. 1 mm; petioles 2-8 mm. Peduncles 0.9-3.5 cm. Umbels terminal and axillary, all pedunculate, indeterminate; involucre bracts absent; involucel bractlets 1-4, margins smooth, 1-3 mm; fruiting rays 4-7, suberect and clustered, unequal, (0-)4-10 mm (central umbellet sessile); umbellets (3-)4-6-flowered, fruiting pedicels (0.2-)l-4 mm (central flowers sessile to subsessile). Schizocarps 1.2-1.5 mm, dorsal ribs 3, rounded, often somewhat obscured by the corky epidermis, lateral ribs similar to dorsal, oil tubes 1 per dorsal interval; surface with vaguely formed multicellular tubercles and' or rounded lateral ridges, tubercles sometimes with a straight, erect, blunt-tipped unicellular hairlike cell. Chromosome number not reported. Map 3. Figure 12. ly of Apiastum, Ammoselinum, and Spermolepis 24 , 1800 m; N.Mex., known Flowering May-Jun. Granitic gravelly loam, oak-juniper s only from the type collection. The distinctive features of Spermolepis organensis might be interpreted as derived from those of S. inermis. The peripheral geographic location of S. organensis (Map 3) also suggests that this might be true but other outlying populations of S. inermis exist. Recognition of 5. organensis as a species, albeit it weakly defined, emphasizes the apparent, at least partial degression of the fruit liair formation, the corky fruit surfaces, and the relatively short fruiting peduncles. If S. organensis were interpreted as of hybrid origin from extant parents, the only oilier species of Spermolepis currently known from the Organ Mountains is S. lateriflora. The nearest known occurrence of 5. inermis is to the east in Eddy Co., that also somewhat of a geographic isolate. 0# Spermolepis inermis Spermolepis organensis A Spermolepis laevis : Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 8. Spermolepis divaricata (Walt.) Raf., Bull. Bot., Geneve 1: 217. 1830. Dctucus divaricatus Walt, Fl. Carol., 114. 1788. Ammi divarication (Walt.) Pers., Syn. PI. 1: 308. 1805. Aethusa divaricata (Walt.) Spreng., PI. Umbell. Prodr., 22. 1813. Sison divaricaium (Walt.) Spreng., Sp. Umbell., 113. 1818? Leptocaulis divaricatus (Walt.) DC, Coll. Mem. 5: 39. 1829. Babiron divaricaium (Walt.) Raf., New Fl. 4: 24. 1836. Apium divaricaium (Walt.) A.W. Wood, Amer. Bot. Fl., 140. 1870, NEOTYPE (Ward 2008, p. 475): USA. South Carolina. Charleston Co.: Wadmalaw Island, S of Charleston, 27 May 1988, D. Boufford & E. Wood 23862 (GH; isoneotypes: MO, NY). Walter did not cite a specimen or locality; Ward (2008) noted that "Spm. 40-C, a wispy fragment., was labeled "Daucus" by Fraser, and annotated as "divaricatus Walt." by A. Gray, though one wonders what he saw that was recognizable." Sison pusillum Michx., Fl. Bor.-Amer. 1: 168. 1803. Ligusticum pusillum (Mchx.) Pers., Syn. PI. 1: 315. 1805. TYPE: USA. [protologue] "In sabulosis aridis Carolinae," A Michauxl (holotype: P?). Babiron dichotomum Raf, New Fl. 4: 24. 1838. TYPE: USA. Florida. Rafmesque did not cite a specimen, noting only "Florida," As synonym of S. divaricata fide Mathias and Constance (1944). Babiron pusillum Raf, New Fl. 4: 23. 1838. TYPE: USA. Alabama or Georgia. Rafmesque noted "sent me from Alabama, and by Dr. Torrey from Georgia as the Daucus pusillus\ see 788." As synonym of S. divaricata fide Mathias and Constance (1944), Stems 7-40 cm. Leaves: blades oblong to oblong-ovate in outline, 0.5-5 cm, 3-pinnately compound, ultimate divisions linear, 3— 10(— 15) x 0.2-1 mm; petioles 1-30 mm. Peduncles 17-40(- 50) mm. Umbels terminal and axillary, all pedunculate; iff olucre bracts absent; involucel bractlets 1-3, narrowly lanceolate, 0.5-1 mm, the margins usually callous-toothed; rays 3-6, divaricately spreading, unequal, 5-17 mm; umbellets (3-)4-6-flowered; fruiting pedicels (0-)2-9 mm (central 1-2 flowers subsessile to sessile). Schizocarps 1.5-2 mm, scabrous with minute, upcurved hairs not arising from a tuberculate base. Chromosome number: see comments below. Map 4. Figure 15. Flowering Mar-Apr(-May). Sandy woodlands (longleaf pine-turkey oak, pine-oak, oak scrub, flatwoods, evergreen scub vA ) pia-tK iannant-, sandy peat, sandy roadsides, fields, pastures, and clearings, lawns, abandoned gardens, orange groves, moist ditches, swamp and salt marsh edges, shell mounds, sand ridges, sandhills. *atsd prairies, sandy peat of bogs; 0-200 m; Ala., Fla., Ga., La., Mss., N.J., N.C., S.C., Tex., Va. Confusion exists regarding chromosome numbers of Spermolepis divaricata, S. inermis, and X echinata. Numbers of 2n = 22 and 2n = 16 apparently both are based on vouchers both identified as S. divaricata. Vouchers for both counts were collected in Florida, so at least it is clear that neither could have been the basis of a count for S. inermis. The count of In = 20 for S. echinata w r as made from an Arkansas plant (see description of S. echinata) securely identified as that species. While it seems unlikely that S. divaricata has two such distinct dysploid numbers, the possibility opens an interesting evolutionary study. In = 22 (Bell & Constance 1957; Florida. Okaloosa Co.: roadside banks just E of bridge 3.6 mi W of Crestview, 17 Apr 1954, Bell 1470, NCU [fide A Weakley], VDB!, "Voucher for chromosome count of n = 11" on specimen). Initially identified by Bell as Spermolepis divaricata; annotated as S. inermis by Mathias & Constance and reported in publication as S. inermis with n = 11; later annotated by H.E. Ahles and by Alan Weakley as S. divaricata; confirmed by Weakley (pers. comm.) as S. divaricata. 2» = 16 (Bell & Constance 1957; Florida. Escambia Co.: sandy roadside along Fla. Hwy 297, 5 Apr 1955, Bell 1514, NCU [fide A. Weakley], "Voucher for chromosome count of n = 8" on specimen). : Taxonomy of Apiastrum, Ammosdinum, and Spermolepis Initially identified by Bell as Spermolepis echinata and rq>orted in publication as S. echinata with n = 8 in the Bell and Constance publication. Specimen later annotated by Mathias & Constance and by Alan Weakley as S. divaricata; confirmed by Weakley, pers. comm., as S. divaricata. Attributions of the species to New Mexico have been based on mis identifications of Cyclospermum leptophyllum. PLANTS Database attributes Spermolepis divaricata to New Jersey, based on an unpublished "Chrysler Herbarium Checklist" ( by J. Meyer from 1990. The voucher, correctly identified in the checklist, is this : New Jersey. [Cam den Co. ] : Camden, ballast, 26 Jun 1866. C.F. Parker s.n. (CHRB digital image!). The species is regarded here as a waif in New Jersey and not a permanent member of the state flora. The plants of Spermolepis divaricata from Acadia Parish. Louisiana, from prairie remnants along a railroad right-of-way, have pedicels in the upper range of length (mostly 5-9 mm) for the species but the central flowers of each umbellet are subsessile and fertile. A cadia Par. : along RR ca. 3.5 mi SW of Crowley, 7 May 1966, Lemmon 1168 (LSU digital image!). Two collections of Spermolepis divaricata are recorded here for Texas. Austin C o. : Industry, 1895, Mr. H. Wurzlow s.n. (BRIT). Liberty Co. : along Co. Rd 2252 W of the Davis Hill Baptist Church and N of Hwy 105 E of Cleveland, 26 Apr 1997, Brown 20285 [without fully mature fruits but the pedicels are short and the umbellets have sessile central flowers with evidently maturing fruits] (NLU). The difference between Spermolepis divaricata and S. inermis can be subtle but is nevertheless real. Before full fruit maturation, outgrowths of the ovary surface of S. divaricata can look like developing tubercles of S. inermis although they usually have an antrorse orientation. This similarity perhaps was the basis for the Mathias & Constance annotation of Bell 1470 (as S. inermis; see comments above about chromosome numbers), which lias short hairs and some tuberculate bases without hairs. In S. inermis. the central rays are consistently very short and there is a relatively small angle of divergence, overall giving the umbels a congested appearance. In S. divaricata, the rays (including the central) mostly ars equal to subequal in length and diverge at a relatively greater angle, giving the umbels a more open appearance. Spermolepis divaricata and S. diffusa characteristically produce a determinate inflorescence - - the axillary bud and terminal leaf are suppressed at the distalmost node (Fig. 2B). In some plants from Florida, however, this apparent specialization is not expressed (de-repressed?) and the pattern is determinate (Fig. 2A). These apparently are populational variants. Examples: Alac hua C o.: Dunn 525 (FLAS), Rabat 488 (FLAS, 9 plants. 2 determinate. 9 indeterminate), Scudder 1491 (FLAS). Baker Co.: West & Arnold s.n. (FLAS). Hernando Co.: Nee et al. 2798 (FLAS). Highlands Co. , Baltzell 1 799 (FLAS, 2 plants, 1 determinate, 1 indeterminate). 9. Spermolepis diffusa (Nutt. ex DC.) G.L. Nesom, comb. nov. Leptocaulis dijfusus Nutt. ex DC, Prodr. 4: 107. 1830. LECTOTYPE (designated here): USA. Arkansas. "In Amer. bor. ad Red-River" [protologue], T. Nuttall s.n. (BM 001042SS4 diaiul image! isi lectotypes: BM 001042883 digital image!, NY digital image!, PH-2 sheets digital images!). De Candolle noted ("vs. ") that he had seen the Nuttall collection. Stems 15-75 cm. Leaves: blades oblong to oblong-ovate in outline, 0.5-5 cm, 3-pinnately compound, ultimate divisions linear, 3-15 x 0.2-1 mm; petioles 1-30 mm. Peduncles 20-50 mm. Umbels terminal and axillary, all pedunculate; involucre bracts absent; involucel bractlets 1-3, linear- lanceolate, 0.5-1 mm; fruiting rays 2—M-6), divaricately spreading, subequal, 15-33 mm; umbellets 2-4(-5)-flowered; fruiting pedicels (8-)14-32 mm (all flowers of each umbellet with subequal pedicels, none sessile or subsessile). Schizocarps 1.5-2 mm, scabrous with minute, upcurved hairs not arising from a tuberculate base. Chromosome number not reported. Map 4. Figure 16. Flowering Apr-May(-Jun). Sandy clay, sand, roadsides, fencerows, fields, pastures, dunes and sandy hills, thin soil under oak-juniper, sandy soil in longleaf pine, pine, oak-pine, oak-hickory, post oak, and post oak-blackjack oak w r oods, lake shores; 50-500 m; Ark, Kans., La., Mo., Okla., Tex. Aptly named Leptocaulis diffusus has long lain in synonymy of Spermolepis divaricata but its morphological distinction from typical S. divaricata is easy to discern. The long pedicels and rays of S. diffusa usually provide ID-at-a-glance; if the pedicels are in the shorter part of the range, the lack of sessile or subsessile flowers provides a second criterion. The range of S. diffusa is entirely west of the Mississippi River. The two species appear to be sympatric in central Louisiana and southeastern Texas. Vouchers documenting the occurrence of both species in Natchitoches and Vernon parishes (Louisiana) are atNLU. Plants in a few scattered Texas collections have abnormally short pedicels: Anderson Co., pedicels 8-11 mm, Bridges & Kindscher 13731 (BRIT, TEX):. Comanche Co., pedicels 6-10 mm. Skinners 20078 (SMU). In the Anderson Co. collection, a low percentage of the umbellets produced a single central, subsessile flower that did nor develop a mature fruit. These plants are within the geographic range of S. diffusa and are regarded here as populational variants of that species, perhaps reflecting ancestal (S. divaricata-ake) characteristics. : Taxonomy of Apiastrum, Ammosdinum, and Spermolepis The doss similarity and probable sister relationship of Spermolepis diffusa to S. divaricata might be emphasized in treating the two as eonspecific varieties. The course here emphasizes their distinction in morphology and geography (and by inference, ecology). Pointed field observations in their region of sympatry would be interesting, and a chromosome count for S. diffusa might provide evidence for an internal isolating mechanism, especially in view of the apparent lability in chromosome number within the genus. 10. Spermolepis castellanosii Perez-Mor., Lilloa 5: 32, fig. 1. 1940. LECTOTYPE (designated here): ARGENTINA. Prov. Rio Negro. San Antonio Este, 21 Nov 1928, A. Castellanos (BA 28/1184). Perez -Moreau cited three other collections — two from Neuquen (leg. Ragonese, Perez-Moreau) and one from Mendoza (leg. Ruiz Leal). Stems ca. 4-8 cm tall, simple or few-branched mostly at nodes above the base. Leaves: blades broadly ovate in outline, 25^0 mm, ultimate divisions 4-8 mm, scaberulous on margins and nerves; petioles 7-10 mm, searious-margined at base. Peduncles 20-50 mm. Umbels axillary, all pedunculate; involucre bracts absent or 1; involucel bracelets 3^1, linear-lanceolate, entire, 1.5-7 mm, unequal; fruiting rays 3-5, 1-15 mm (inner 1-2 umbellets short-pedicellate), spreading; umbellets 3- 5-fiowered; fruiting pedicels (1— )7— 9 mm (inner flower short-pedicellate), loosely convex to irregular. Schizocarps oblong-ellipsoid, attenuate toward the apex, 3.2-5.3 mm, hispid-hirsutulous on the angles and intervals with narrowly triangular, straight, blunt-tipped hairs, dorsal ribs 3, rounded, lateral ribs not expanded; oil tubes 1 (rarely 2-3) per interval, 2 on the commissural face. 2n = 64 (Constance et al. 1976; Hunziker 12523, Cordoba, Argentina). Figure 17. Spermolepis castellanosii apparently is endemic to west-central Argentina. Perez-Moreau (1940) cited collections from the provinces of Rio Negro, Neuquen, and Mendoza. The hexaploid chromosome count by Constance et al. (1976) was from Prov. Cordoba. Photos on Flickr by Joseph Fourier (2009) are from Prov. San Luis. 11. Spermolepis gigantea (Coulter & Rose) G.L. Nesom, comb. nov. Ammoselinum giganteum Coulter & Rose, Contr. U.S. Natl. Herb. 7: 89. 1900. TYPE: USA. Arizona. Maricopa Co.: mesas near Phoenix, 17 Jun 1882, C.G. Pringle 28 (holotype: GH; isotypes: fragment JEPS digital image!, NY digital image!, US digital image!). Ammoselinum occidentale Munz & Johnston, Bull. Torrey Bot. Club 52: 224. 1925. TYPE: USA. California. Riverside Co.: "Hayfields" [Hayfield pumping plant locality], Chuckwalla Valley, Colorado Desert, locally abundant in heavy soil of a dry basin under shrubs and in the open, 500 ft, 13 Apr 1922. P.A. Munz & D.D. Keck 4930 (holotype: POM digital image!; isotypes: BM digital image!, JEPS!, UC!). Stems 8-26 cm, simple or branching from the base. Leaves: blades obovate to broadly ovate in outline, 12-25 mm, ultimate divisions linear, (1— )4— 13 mm; petioles 3-25(3-0) mm. Peduncles 2- 5.5 cm or penultimate umbel sometimes sessile (see NY isotype, Fig. 19. Umbels terminal and axillary; fruiting rays 4-10, (0-)2-22 mm (inner umbellet sessile), unequal, spreading; umbellets 1- 10 flowered; pedicels (0-)2-8 mm (inner flower sessile to subsessile); involucre bracts (0-)l-3, linear or sometimes 3-fid; involucel bractlets 1-6, linear to linear-lanceolate, entire or less commonly 2-3-fid, 2-12 mm, sometimes searious-margined at base. Schizocarps narrowly elliptic-ovate to urceolate-oblong or ovoid-oblong, 3-4 mm, ribs low-rounded, hispid-hirsutulous on the ribs and intervals with sharp-pointed, 1-2-celled hairs arising from a conical, non-pustulate base, dorsal ribs 3, cordlike and thickened, lateral ribs flattened and broad, nearly obscuring the commissural sulca; dorsal oil tubes 3 per interval, commissural oil tubes 2. 2n = 38 (from label of McKay 64: "n = 19 Chromosome vouchers cultivated in University of California Botanical Garden, C-775"). Map 5. Figures 18, 19, 20. n: Taxonomy of Apiastrum, Calif. Flowering Mar-Apr. i, sandy flats, desert shrubland with Larrea; 200-800 m; Ariz., ire and represented in Arizona only by the collections cited here from Maricopa, Pima, and Pinal counties; in California it is known only from the type of Ammoselinum occidentale, collected in eastern Riverside County (Map 4). Wolff (1 927) and Mathias and Constance (1945) identified Pringle 8314 from Coahuila as Ammoselinum giganteum, but that collection, far-distant from the main range of that species, is identified here as typical Ammoselinum popei (see citation above). Collections examined. USA. Arizona Maricopa Co. : 39 mi NW of Wmtersburg, sandy flat among Larrea, 22 Mar 1936, Wiggins 8431 (UC). Pima Co. : ca. 10 miE of Tucson onNogales Road, On open roadside, 2500 ft, 27 Apr 1945, Gould 3064 (ARIZ). Pinal Co. : near Casa Grande, 3 Apr 1937, Darrow s.n. (ARIZ), Casa Grande, 1400 ft firs faintly Pastinaca-scented, 2 May 1965, McKay 64 (ARIZ); Eloy, 25 Mar 1 930, Peebles, Harrison, & Kearney 6496 (ARIZ-2 sheets). Spermolepis gigantea is characterized by these features: umbels terminal and axillary, all pedunculate; involucel bracelets long, s carious -margined on the proximal 1/3-1/2; involucre bracts usually present, often ternate; peduncles, rays, and pedicels minutely hispidulous on the angles; and fruits relatively long, narrowly elliptic-ovate, hispid with non-pustulate-based hairs. The broad lateral ribs are perhaps the reason that is has been treated as a species of Ammoselinum. ©Spermolepis gigantea •Spermolepis infernensis Map 5. Distribution of Spermolepis gigantea and S. infernensis. : Taxonomy of Apiastrum, Ammosdinum, and Spermolepis Munz & Johnston (1925) included Ammosdinum giganteum and A. occidentale as the two members of Ammosdinum sect. Hesperosdinum Munz & Johnston, but Mathias and Constance (1944, p. 104), without comment, placed A occidentals in the synonymy of A. giganteum. where it has since resided. Munz and Johnston observed that Ammosdinum occidentale differed from A giganteum in its "lower more compact habit, unbranched stems, smaller more compact umbels, pubescent (rather than conspicuously callous-toothed) smaller carpels, and twice as many commissural oil tubes." In the observation here, however, the fruit vestiture is the same in both taxa, and Mathias and Constance (1944) treated A. occidentale as a synonym of A giganteum. Mathias and Constance described the lateral fruit ribs of A giganteum as having "corky appendages," but in the observation here, they are elaborated hardly more than the dorsal. ACKNOWLEDGEMENTS I am grateful for loans (to TEX) from ARIZ, FLAS, NMC, RSA, SJC, SRSC, and UC-JEPS and for hospitality at BAYLU, BRIT-SMU-VDB, NLU, and TEX-LL while studying there. Thanks to Mirta Arriaga at BA for providing close-up photos of the fruit of Spermolepis castellanosii, to Liora Spitz at CURB for providing a digital image of the early collection of Spermolepis divaricata from New Jersey, to Sarah Taylor and Jochen Schenk at MACF for digilal images of the McFadden collection of Spermolepis lateriflora from Los Angeles County, to John Pruski for a digital image and close examination of the fruits of Pr ingle 8314 (MO), to Tim Flynn and David Lorence at NTBGfor providing the image of Spermolepis hawaiiensis, to Jon Rebman and Mary Alice Kessler for observations on the fruit vestiture of SD collections (confirming their identity as Spermolepis lateriflora), to Carol Ann McCormick and Alan Weakley for information on NCU specimens, to Sam Kieschnick at BRIT for providing scanned images of several specimens, and to Stephen Downie for early comments on the taxonomy. This research has been supported in large part by the Flora of North America Association. LITERATURE CITED Bell, C.R. and L. Constance. 1957. Chromosome numbers in Umbelliferae. Amer. J. Bot. 44: 565- 572. Boufford, D.E. 1977. Ammosdinum butleri (Umbelliferae), new to North Carolina. Sida 7: 220. Brown, RG. and M.L. Brown. 1984. Herbaceous Plants of Maryland. Port City Press, Baltimore, Maryland. Bryson, C.E. 1991. Tw ? o weedy species, Ammoselinum butleri (Umbelliferae) and Lepidium austrinum (Cruciferae), new to Mississippi. Sida 14: 506-508. Calflora. 2012. The Calflora Database. Information on California plants for education, research and conservation, based on data contributed by dozens of public and private institutions and individuals, including the Consortium of Calif. Herbaria. Berkeley, Chodat, R and E. Wilczek. 1902. Contributions a la flore de la Republique Argentine. Bull. Herb. Boissier ser. 2. 2: 281-296, 475^190, 521-544. Constance, L. 1993. Spermolepis. Pp. 165, in J.C. Hickman (ed.). The Jepson Manual: Higher Plants of California. Univ. of California Press, Berkeley. Constance, L. and M. Wetherwax. 2012. Spermolepis. Pp. 199, in B.G. Baldwin, D.H. Goldman, D.J. Keil, R Patterson, and T.J. Rosatti (eds.). The Jepson Manual: Vascular Plants of California (ed. 2). Univ. of California Press, Berkeley. Constance, L., T.I. Chuang, and C.R. Bell. 1976. Chromosome numbers in Umbelliferae. V. Amer. J. Bot. 63: 608-625. Coulter, J.M. and J.N. Rose. 1900. Monograph of the North American LTmbelliferae. Contr. U.S. Natl. Herb. 7: 5-256. Coulter, J.M. and J.N. Rose. 1909. Supplement to the monograph of North American Umbelliferae. Contr. U.S. Natl. Herb. 12: 441-451. : Taxonomy of Apiastrum, Ammosdinum, and Spermolepis Downie, S.R., K. Spalik, D.S, Katz-Downie, and J. -P. Reduron. 2010. Major ciades within Apiaceae subfamily Apioideae as inferred by phylogenetic analysis of nrDNA ITS sequences. PI. Divers. Evol. 128: 111-136. Fourier, J. 2009. Spermolepis castellanosii. Flickr. Two photos, 1 Nov 2009, from Prov. San Luis, Argentina, Gonzalez, A 2010. Fotos de flora nativa de Uruguay y adventicias. Flickr. Keener, B.R. 2007. Noteworthy collections : Alabama. Castanea 72: 47-48. Mathias, M.E. and L.Constance. 1944. Apiastrum. Genus 10. North American Flora 28B: 71. Mathias, M.E. and L. Constance. 1944. Spermolepis. Genus 11. North American Flora 28B: 71-73. Mathias. M.E. and L. Constance. 1944. Ammoselinum. Genus 21. North American Flora 28B: 103— 104. Mathias M.E. and L. Constance. 1950. Four new American Umbelliferae. Bull. Torrey Bot. Club 77: 133-139. Munz, P.A. and I.M. Johnston. 1925. Miscellaneous notes on plants of southern California - IV. Bull. Torrey Bot. Club 52: 221-228. Munz, P.A (in collaboration with D.D. Keck). 1959. A California Flora. Univ. of California Press, Berkeley. National Tropical Botanic Garden (NTBG). 2012. Herbarium Database, Herbarium PTBG, National Tropical Botanic Garden, Kalaheo, Hawaii, Perez-Moreau, R.A 1940. Una nueva especie de Umbelliferae. Lilloa 5: 31-34. Radford, A.E., HA Ahles, and C.R Bell. 1968. Manual of the Vascular Flora of the Carolinas. Univ. of North Carolina Press, Chapel Hill. SEINET. 2012. Southwest Environmental Information Network Managed at Aizona State Univ., Tempe. TENN. 2012. Univ. of Tennessee Herbarium website, Torrey, J.G and A Gray. 1840. Leptocaulis. Fl. N Amer. 1: 608-609. USFWS. 2010. Spermolepis hawaiiensis, 5-Year Review, Summary and Evaluation. U.S. Fish and Wildlife Service, Pacific Islands Fish and Wildlife Office. Honolulu, Hawaii. Villarreal Q., J. A 2001. Listados floristicos de Mexico. XXIII. Flora de Coahuila. Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Mexico, D.F. Wagner, W.L., D.R Herbst, and D.H. Lorence. 2005. Flora of the Hawaiian Islands website. Ward, D.B. 2008. Thomas Walter Typification Project, V: Neotypes and epitypes for 63 Walter names, of genera D through Z. J. Bot. Res. Inst. Texas 2: 475^186. Wolff, H. 1927. Umbelliferae- Apioideae- Ammineae-Carinae, Ammineae Novemjugatae et Genuinae. Pp. 1-398 in A Engler (ed). Das Pflanzenreich, Heft 90 (IV. 228). W. Engelmann. Berlin. m: Taxonomy of Apiastrum, Ammosdinum, and Spermolepis 3! Figure 3. Apiastrum cmgustifolium, representative plant (ASU). Nesom: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 33 sj(£ ^u .tfj w%-% %# A \ L i: Taxonomy of Apiastum, Ammoselinum, and Spermolepis 34 Figure 5. Ammoselinum rosengurtii (holotype, UC Nesom: Taxonomy of Apiastrur, Figure 6. Spermoiepis lateriflora, representative collection (AEJ.Z). i: Taxonomy of Apiastrum, Ammosdinun, and Spermolepis 36 Figure 7. Spermolepis lateriflora, representative plant, bottom left of Fig. 5 (ARIZ). Nesom: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 37 Figure 8. Spermolepis echincita, representative collection (FSU). i: Taxonomy of Apiastrur 11 4 J«™-™ — .S^OS jf~ ^S-^5SES~ ^ »"™™v srasKsssssastsreffisi ~— ™~'~"™"™" E2S2 SS.S"™'s"r j «™. rao ._»«„«__i™ Nesom: Taxonomy of Apiasfrun nfernensis, holotype (RSA). : Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 40 Figure 11. Spermolepis infernensis, representative plant (from holotype, RSA). n: Taxonomy of Apiastrum, Ammose/inum, and Spermo/epis 41 APIACEAE 3106 SPERMOLEPIS E Coll: 6 JUN 1995 NEW MEXICO, Dona Ana Co., Figure 12. Spermolepis organensis, holot\pe (NMC). i: Taxonomy of Apiastum, Figure 13. Spermolepis inermis, representative plant (SMU). Figure 14. Spermolepis laevis, representative plant (SMU). and Spermolepis iiKiiiiiim 168420 FLORIDA STATE UNIVERSITY Figure 15. Spermolepis divaricata. representative plant (FSU). Nesom: Taxonomy of Apiasfrur i Spermolepis 45 Figure 16. Spermolepis diffusa, representative plant (MO). i: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 46" Figure 1 7. Spermolepis caste Ikmosii, from the original illustration (Perez -Moreau 1940). Nesom: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 47 Figure IS. Spermolepis gigctntect, m: Taxonomy of Apiastrum, Ammoselinum, and Spermolepis 4§ Figure 19. Ammoselinum gigonteum isotype (NY) = Spermolepis gigcmtea. i: Taxonomy of Apiastum, Ammoselinum, and Spermolepis 49