NEW (THYMELAEACEAE) A OF DIRCA SPECIES FROM THE NORTHEASTERN MEXICO OF SIERRA GUYL NESOM MARKH. MAYl .D and El PI . Depart we Botany of ;;/ Umversi Texas ty of TX Amttn, 78715, U.S. A. A newly scovered po pulation of Dmv?:from the Siern Madre One.ntalofTam di 1 mexkana D. nov.Ite xpands the vlexico, de^ illustrated here as sp. IS ^ Me The new nown liree species and It TfiiSt of the genus k to occui in xico. s] : d .imilar to D. pa/»sim of the eastern Unite States in it: long c alyx tube with . i ncted D. kring limb, ,.a 1 Dmc )f the San Fnmcisco Bay region in west-ce ntral Californ la. / Illt'Xlanu, resem atter in its bi th whitish pubesa.nee, sessile flc)wers an d fruiits, distinct ' oung and twigs abaxi surfaces. ;alyces, .ersistenc pt ai id al le;if p from D. na ger, nar rowe r tube, shot .pecies differs Una pobla.cion de Di ra, reci enteimente Jescub ieria en la Sie ( mexk. Tamauli Mexico, d escribe iconografiElaquic:omo D. ,pas, se e numero gen ;ndo esta pr eleva a trese d< especi es er el ero, si( la 1 - I Lan ecea de Mexico. ueva especu D. pal.'turn. este los ;lel ( mbo bade con anado, pero su caliz ab li 1 con restnngi,da D. :ciden tain de la reg ion de la Bahia es y 0, 1: G de parece a esta u i-oeste yem; con pubescenci a blanq la, flores y frut las is iLiecir [ y la pub escen ciapersisterite en laIS ranlas jove nes y eia el enves'de las mn mas tubo largo ocadentalis flores y I i masc imen mas cortos Los dos sisten- lobulos ;ortos, y esti lo y fib tos . masde25C Dtna separadi pot iO kilometre; mexica na esta alejada :)s s. , Dma The genus has previously included only two species, D. palustris L. United and adjacent scattered but widespread in the eastern States L., Canada, and D. occidentals A. Gray, endemic to six counties of the San & (McMinn Bay Forderhase 1935; region west-central California Francisco in On NevUng Vogelman 1962; various manuals; Fig. a 1953; floristic 1). botanical reconnaissance September 1994 Madre in in the Sierra Oriental of Tamaulipas, Mexico, we observed a population of low shrubs that we recognized as Dtrca, but the plants were and not sterile identifiable to spe- March we cies. In early 1995 were able to study the population peak at flowering and beginning of The Mexican most fruit. plants closely resemble D. but occidentalis, there are several consistent morphological differences between them and they are geographically disjunct by more than 2500 kilometers (California/Tamaulipas, The known see Fig. closest popula- 1). tions o{ Dirca palustns, at the southwestern corner of range in eastern its Louisiana, Arkansas, and southeastern Oklahoma, more than 1100 are ki- We lometers disjunct from the Mexican plants. believe that justifiable it is and desirable to recognize this Mexican population as a species distinct from both more of northern its relatives. & Nesom exicana Mayfield, sp. (Fig. 2) . Woody dm 6-20 shrubs dm, averaging 16 from tall, ca. a single t branched near the base, with spreading, flexuous branches, the stem cm wide 3(-5) near the bcise of the plant, the ultimate branches 2-3 wide, bark smooth and gn grayish reddish-brown, to the current year's opened gle flower, 462 SiDA 16(3) 1995 persistently loosely and sparsely pubescent, glabrous below. Leaves deciduous, alternate, entire, at maturity broadly elliptic to slightly ovate, mm 4-8 cm basally rounded, long, 2.5-6.0 wide, 1.3-1.7 times longer than wide, the largest distally situated on the branchlets, glabrous above, the lower surface persistently sparsely strigose-sericeous on the lamina and mm along the veins, petioles 1-2 long. Buds covered by the enlarged peti- mixed (Rowers and ole base, leaves), the apical apparently falsely terminal, bud the 4 scales whitish-sericeous, forming a foliaceous, deciduous involu- cre to the dowers. Flowers appearing before or concurrently with the leaves, sessile in axillary and apical fascicles, deflexed to somewhat more nodding at full anthesis, the axillary fascicles invariably producing 3 flowers, the sometimes apical apparently "twinned" with fascicles (5-)6 flowers; corolla mm absent, not represented by infracalycular structures; calyx (7-)8-10 mm long, petaloid, the tube narrow but widening 5-7 slightly distally, mm long, abruptly expanded into a flaring limb 2-4 long, the tube/limb mm ratio (of length) (1.5-)1. 7-3.0, the 4 calyx 1.5-3.0 lobes long, the and limb lobes yellow, the tube lighter yellow and drying creamy; stamens filaments inserted within the calyx tube 8, essentially at a single level (at mm mm the throat, 2.0-3.5 below the lobe apex), the longest exserted 2-3 mm above the calyx lobes, those inserted below the lobes 1-2 longer than mm below those inserted the sinuses, the thecae 0.4-0.8 long, basifixed; "hypogynous Heinig disc" or "disc" (sensu 195 a ring of connate, irregu- 1) mm lar, slightly fleshy scales ca. 0.2-0.4 high and wide, basally adnate to the inner surface of the calyx base; stigma minutely capitate, above the level of the pre-dehiscent anthers on a fully elongated style, equalled or slightly surpassed by the dehiscing anthers. Fruits drupaceous, -seeded, 1 pyriform when to ovoid, sessile, green young, mature fruits not observed. MEXICO. Typf,: Tamaulipas, Municipio. Hidal^i;(K mountainous road from al.,n,^ Sra. Nombres Engracia (Tamaulipas) Dnlces (Nuevo to LL'on)/Arro\o Ohscuro, 2.0 road mi NE of Los Caballos toward Canada El Mimbre, 15.0 road mi w'of lowermost crossing of Mimbre, arroyo El 23"59'()9"N, 99"28'37"W, 1800 m, Mar 7863 ca. S 1995, G/iy Nesoiii with Mark Mayfieldand Greg Anderson MEXU; AAU, ANSM, ARIZ, (hoi.otyfi;: isotvpfs: ASU, BH, BRIT, CAS, CHAP, COLO, CONN, DAV, DUKE, ENCB, FSU, FTG, GA, F, GH, GUADA, KANU, lEB, K, LSU, M, MICH, MO, MSC, NCU, NLU, NY, OBI, OKL, RM, TENN, UARK, OS, OSC, USA, TEX, UAT, UC, UCR, UNL, VDB, US, WIS, P, S, MEXICO. exammed: jllection Tamaulipas: type locality, [sterile, le m, Nesom MayfieU/2()85 with (TEX). — The which Dina of the locality. site at mexuzana was obser m (Arroyo Obscuro) a steeply sloping, nooh-facing, is I ding rocky The a watercourse. area limes tone with kar is ^he elevation near the road 1800 met but plant is ca. ers, a Nesom and Mayfield, new species of Dirca from Mexico 463 Dirca extend 20 meters (elevation) above the road and to at to at least ca. we may 80 meters below They well occur over a broader area than least it. where investigated, particularly downslope, but this the only locality is from Engracia Dirca was encountered along the 35 kilometer road Sta. to Dulces Nombres. Indeed, we found no other site along this road with a and similar physical floristic definition. The dominant canopy trees at the site are Carya ovata (P. Miller) K. & Koch, Pmuspatula Schlecht. Cham., Pseudotsuga menziesii (Mirb.) Franco, & 20-25 and Quercus launna H. B. Large trees of these reach about meters height, with Pseudotsuga perhaps reaching 30 meters. Judging from in tall stumps in the area, particularly large individuals (presumably o^ Pseudotsuga and Pmus) have been removed by logging. Large trees oiCarpinus caroliniana Uqmdambar the but do not Walt, and styraciflua L. are scattered at locality The woody understory includes the reach the height of the canopy. full Garrya Muell.-Arg., Cornus urbimana Rose, Croton virletianus following: macrophylla Benth., rubra Wats., Litsea pringki Bartlett, Persea Ilex aff. S. Most Hu, and Taxus Schlecht. pododaema Blake, Philadelphus caldcolus globosa bud and were beginning break of the canopy species are deciduous just to predominance evergreen the understory includes a of species. at this site; conspicu- Arboreal bromeliads, including three species of Tillandsia, are among the Ferns form a major part of the herbaceous flora at this site; ous. common most are species of Adiantum, Woodivardia, Botrychium, common and Polystkhum. Other herbs include Phanerophlebta, Polypodium, and Chmaphila, Chiropetalum, Goodyera, Sisyrinchium, Stachys. species o^ — We and observed population biological observations. Characteristics of the 800-1000 plants (by estimate) at the Dirca site, where they occur for about 300 meters along the road and adjacent slopes on the east side of the water- more open-canopy They microsites, tend be densely clustered in course. to We more did not unearth any plants, singly and scattered over a larger area. A but there was no indication that they reproduce clonally. report of rhi- & (McMinn zome Forderhase 1935) has not production D, in occidentalis more (Spongberg comm.). been corroborated by recent observations pers. synchronous The population of Dirca mexicana strongly in entire is we would flowering. Had we arrived 3 or 4 days later, it is likely that have appear wither quickly been unable to observe floral features, as the flowers to The appear be weakly with the onset of fruit maturation. flowers to may protogynous and there evidence that they also be self-compatible. is before development of the calyx, the style (with apical stigma) Initially, full At time, the stigma usually elongated past the level of the anthers. this is we appears slightly moist and presumably receptive; also observed that many at this stage appear to have pollen attached. At anthesis, the full filaments have raised the open anthers to the level of the stigma or slightly beyond Further, there time separating it. is little intial receptivity of the stigma and dehiscence of the anthers, and distance between the stigma little and open anthers. mm Visiting the flowers were one species of bee (individuals 7 in ca. and length) at least four species of butterflies. Pollination effective, judg- is from ing the apparently ubiquitous deposition of pollen on the stigmata of pre-staminate flowers. Bees and were probing butterflies inside the flowers, and although seems highly likely that nectar was available them (prob- it to ably produced by hypogynous the production was disc), not evident its in numerous flowers that we examined. Nor could we any detect fragrance (D. ocddentalis was noted by Howell [1970} to be fragrant). Fruit matura- tion was beginning only on a few plants and we were unable make an to estimate of the success of fruit and seed production. We did not determine what feature or features account for the remark- able o^ Dirca rarity mexicana. All of the other species yet identified from — Arroyo Obscuro are found in other sites in the same general area none Dina except the could be considered (but comments below rare see regard- ing Viburnum). however, Rarity, also is a feature of both other species of the genus, particularly D. D'ma occtdentalts (Stebbins 1942; Johnson 1994). uncommon palustns widespread but of occurrence. is — Morphological amipansons. Dirca mexicana resembles D. most ocddentalis in of the features that have been used to distinguish the from D. latter palustris (Vogelmann 1953; see key below): vestiture, presence or absence of pe- a duncle and pedicel, and the morphology The distal of the calyx. flowers of D. mexicana, however, more are similar in general configuration to those of D. palustris: both have a relatively long and narrow tube with a shorter, abruptly widening limb, the staminal filaments are inserted above the middle of the calyx, and the and style anthers are exserted for a relatively short length. In D. occidentalism the tube shorter than the broadly funnel- is form limb, the staminal filaments are inserted below the middle of the calyx, and the and stamens Vogelman style are long-exserted. (1953, 80) p. emphasized the taxonomic usefulness of the of filament which level insertion, turn in is indicative of the throat position and flower shape: "In most in- stances this character alone sufficient to distinguish the two is species." The comparative illustrations furnished by Vogelmann, however, do not accurately represent this difference, nor does the detailed illustration of D. Holm Cronquist palustris in (1981, p. 635); those by (192 are more simi- 1) own lar to our observations. Differences between the new species and D, Bud wn own young twigs nd both with bro or reddish-br piibescence; scales ai whole leaf surfaces complfitely glabrous; 3wers ind fruirs pedicellate, th.e fl. I ma pedun merely crenulate-undulate with- cluster often .culate; calyx rgin ;, United D, palu. out distinct lobes;.=astern St:ates... Bud with whitish pubescence youn, twigs and abaxial leaf:surfaces scales g ; and anc[deeply persistently pubesci;nt; flowers fruits se ssile; calyx distinctly Bay region of Califori or northeastern Mexic iia nm 4- imb Calyx tube 2-4 long, broade ned a broadly funnelform 2. inl:o r 1 mm 6 long; stanamal filaments uiserred below the middle of the calyx. mm and 3_4(-5) at)ove th flower at maturity; st yle .e mm Bay stigma (l-)2-4 above the nthers maturity; San Franci SCO a at — seem point provide Bwgeographk does not possible at this to pattern. It among morphologically based hypothesis of relationship the three spe- a ofDirca, because the genus appears to be relatively isolated, its closest cies (and thus the evolutionary polarity of character states) difficult to relatives Domke Nevhng surprising to find a 1934; 1959). specify (see Still, it is view between D. mexicana and D. in greater overall similarity occidentalism between of the well-known pattern of close relationship and disjunction species of the eastern and southeastern United States and the sierra of north- & Graham immediate Mexico (Miranda Sharp 1950; 1973). In the eastern from numerous disjunct their area of Dirca mexicana are species that are Carya United Carpinus caroliniana, primary range the eastern States in (e.g.): W. Desmodium (Muhl. ex Barton, glutinosum Chimaphila umbellata ovata, (L.) and Willd.) Wood, Liquidambar styraciflua, Pedicularis canadensis L., and (Michx.) Schott. In addition, there a rare yet Polystichum acrostichoides is (Nesom undescribed species oi Viburnum in prep.) in the close vicinity that V most obovatum Walt, of the southeastern apparently closely related to is from same United Another recently recognized, disjunct species the States. Nesom (May and submitted), area of Tamaulipas, Scrophularia sp. nov. field and closely similar to marilandica L. of the eastern U.S. species S. S. is & Cham. Taxus one of the Schlect. of the Pacific region. globosa, californica widely separated understory species at the Dirca site, also has close relatives and western United the eastern States. in Biotic disjunctions between the eastern and western United States, similar common known well though not particularly (e.g., to that in Dirca, are Wood added Sharp 1951; 1970), and other examples can easily be (e.g., This pattern emphasized by the recent discovery of Sericocarpus, lonactis). is ( 466 SiDA 1995 16(3) a second species ofNeu^isia in California (Shevock et 1992), both species al. extreniely rare. In contrast, clearly established disjunctions of extant plants from the western United States to the Sierra Madre Oriental of Mexico are unknown Numerous to us. disjunctions exist between the Sierra Madre Oriental and S.M. Occidental, but most or all of these appear to be deriva- McVaugh tives of the general pattern described by where (1952), the evo- A lutionary antecedents occupy a more southern position. few Mexican species have a broken but somewhat continuous from still distribution the through eastern sierra northern Coahuila into the mountains of southwest- New ern Texas and southeastern Mexico Populus Michx., (e.g., tremuloides Nesom Pseudotsuga menziesii [Mirb.} Franco, and see 1993). Dm-^ In the broadest view of the origin of mexicana, the simplest hy- pothesis that belongs the element with is it to floristic closest evolu- its tionary ties to the flora remaining in the southeastern United States, the geographic continuity between them probably established during the middle Miocene (Graham The to late 1973). disjunction between D. and palustns may D. ocadmtaln be considerably older, as a vegetation probably includ- ing these species was spread across North America through much of the Tertiary, beginning as early as the Eocene (Graham 1972, 1993). In a con- Axelrod trasting view, (1975) hypothesized that the closely related dis- juncts found in eastern Mexico, the Appalachians, and the West coast of United the States represent remnants of continuous a forest earlier spread Mexico and more into simultaneously fragmented as a result of a spreading dry climate in the mid-Oligocene. Evaluation of the theory latter diffi- is cult at present because of the lack of evidence northern temperate for ele- ments American in Latin prior to the Late Tertiary. These elements have not been rec covered from the Oligo-Mi ocene Sim ojove Group of Chiapas, 1 M Mexico; appe Panam; Q^uereus first ars in a in the 10-Pli ocene, aiid in South America arrives by about one mi Uion ago nd A//,ins years a Q^uen.us not until Many Ame about 340, )()() years age east(;rn North rica disj uncts were ). Mc present in eastern _^xico by th< middle Plio cene, nd global a ; M paleotemper ature history suggests that cooling in the middle iocene may have been appropriate time ar for principal introduction. thei: 1 r We thank Greg Anderson help with for collecting, observations on floral new We biology of the species and his comments on the manuscript. are grateful to Billie Turner, Steve Spongberg, Alan Graham, and Simpson Beryl comments for on the manuscript, John Gene Wofford and Bob to Taylor, Krai, and Sidney McDaniel information on Oklahoma, for Dirca Ten- in and nessee, Mississippi, Linda Vorobik respectively, to for the illustration, and to Jaime Hinton for his encouragement and company on and spring fall fDircafromMex and example the of illecting trips for his nuing vigor. New York Press y 1934 Untersucliun^en die stem itischeundgeographischeGhederun^ Liber s> 1972 Outhnt ot tlu oiuin in historit d reLo^ninon of flonstic iffinitie: 1 AmeiRi A Crihim Asii ind eastern Noirh In (cd rionstics ind paieo- ) _ 197^ Historv the irboitscuu tempLrite element in the northern Latir ol md bioti In A Grihim cd Vci^etuion Ve^et ition History ol Northerr nLeuh JOHNSC \ lU, 199 PliUuolthe\Ve-,tL. 1 M A new 4LSOM Submitted ind ,) I [ , Mex ^'Trom > Dma McMiN ^ [OR DLRHAS, 19.5 on ouidentalis N u.d Jotes 1 } 1 1 I I Midiono 120 Gra> > Vm M< tP,u.u sttd phyiot^env n'.'h'AmuK Hut 1 M A Ch^ru 1950 Mi'iMN, 'ind Rr tenstic 1 ) 199'' ' NLs'ni "(Vj' nirc e species ot Astc I A^ste ! Coalnull Mexico Fhyi ologia 74 296-3 04 onofthegenusD^/;^#i/ m 196/ The tlie ' Dina (Thymeleaceae). ouidentalts J the southern Appalachians and ;n