THREE NEW SPECIES OF GALIUM (RUBIACEAE) FROM BAJA CALIFORNIA LAURAMAY T. DEMPSTER Dempster, Lauramay T. (Department of Botany, University of California, Berkeley). Three new species of Galium (Rubiaceae) from Baja California. Bfittonia 22: 184-190. 1970.--The species described as new are apparently narrow endemics, all belonging to that section of Gallum having fruits with long straight specialized hairs. Galium wigginsii and G. moranii are of the widespread G. wrightii-G, parlshii complex, whereas G. carterae is most like the rarely collected G. oresbium Greenman of northeastern Mexico.

The following new species of Galfum are named for three botanists who have greatly expanded our knowledge of Baja Californian botany: Ira Wiggins, Annetta Carter, and Reid Moran. It is unfortunate that we have only one collection of Galium carterae and only three of G. moranii, but G. wigginsii is abundant in an area which, although not accessible by car, is nonetheless more easily visited than many other parts of Baja California. ~ have therefore had an opportunity to study this species in the: field, and to count its chromosomes. The single collection of G. carterae clearly shows it to be strikingly different from any other known species. The three collections of G. moranii differ among themselves only in minor respects, and as a group are well differentiated from their nearest relatives. All three species belong to that group which has dry fruits with long straight (i.e., not hooked) specialized hairs. I t is possible that further exploration may show these species to be more widely distributed than our present information indicates (Fig. 1). Much of Baja California is still difficult of access and, furthermore, Galium is inconspicuous and less likely to be collected than larger or more colorful plants. Although G. carterae and G. moranii may be truly narrow endemics, it seems probable that G. wigginsii ranges somewhat farther north and south than its present known occurrence. G a l i u m c a r t e r a e Dempster, sp. nov.

(Fig. 2).

Planta perennis humilis conferta cinerea hirtella, caulibus fragilibus; folia elliptica carnosa uninervia 3-7 mm longa, internodia aequantia, marginibus plerumque reflexis; stomata depressa, in pagina superiore deficientia; corollae extra hirtellae; fruetus sicci, carpellis reniformibus pills Iongibus rectis obtectis. TYPE: M E X I C O : ]3A3-A CALIFORNIA SUR" Sierra de la Giganta, head of S branch of Cafiada de Teojo, in vicinity of Portezuelo de Peloteado (SW of Notri), dense brittle perennial, forming a small clump on vertical cliff, ca. 850 m, fruiting in October, Carter & Medellin-Led 4656. (HOLOTYPE: UC 1334279). The type collection is the only one, but the plant is so unusual that there can be no doubt of its specific rank. The collector tells me that she saw several individuals, but was able to reach only one. The stems, especially the lower portion, are remarkable for the great expansion of their "angles," which overhang and nearly conceal the reduced and sunken "sides." This condition is even more extreme than the similar one previously observed in the G. angustifolium complex (Dempster & Stebbins, in press). The stems in G. carterae, furthermore, are sharply constricted at the nodes, resulting in a BPdTa'O•IA 2 2 : 1 8 4 190. April-June, 1970.

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2r ./ -7

Q = G. w ig_ginsii O=G. moranii I~=G. carte

&

rae

FIG, 1. Map of Baja California, showing location of three new species. series of spool-like internodes which break apart very easily. As a consequence, the type specimen, although carefully handled, broke into many pieces before it reached me. The new species seems most nearly related to G. oresbium Greenman, a mainland species which has been collected apparently only twice, the first time in Nuevo Le6n by Pringle (2542), the second time in Coahuila by C. H. Muller (3184). Galium oresbium is, however, a much more elongated plant, with much smaller, as well as relatively narrower, leaves, and the stems are: not brittle. The leaves of both species are fleshy and strongly revolute, but those of G. oresbium, unlike G. carterae, have heavily collenchymatous midribs and are densely set below with exceedingly minute hairs. I have seen no. proper flowers of either species, but a few late and depauperate buds were found. Although it is impossible to be certain of the sexual system in either species, the presence of mature fruits on the same plant with hairless ovaries of depauperate flowers suggests that both G. oresbium and G. carterae are polygamous. The leaves of G. carterae are clearly and conspicuously one-nerved. Cleared and

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FIc. 2. Galium carterae. A, middle portion of stem, showing fleshy, congested, revolute leaves, X 4; B, cross section of leaf, X 25; C, thinner, nearly plane upper leaf, X 4; D, upper leaf cleared and stained to show venation, X 7; E, detail from middle portion of edge of same leaf, X 55; F, lower portion of stem with one remaining leaf, to show thickened internodes and constricted nodes, X4; G, cross section of stem, showing greatly enlarged "angles" and sunken "sides," N25; H, mature carpel, XT. (Carter 4656, type). .-> Fio. 3. GaIium wigginsii. A, branch of medium size, X unopened perfect flower and 6 staminate flowers, X 7 (Hardham & Dempster 17007); D, leaf with leaf bases, X 7 (Moran 15480) ; E, detail from middle portion of edge X 55; F, somatic metaphase, X 2500 (Dempster 4226).

1 (Breedlove 16307) ; B, cymule with 1 (Moran 15510); C, single node, X 4 cleared and stained to show venation, of same leaf to show minute pubescence,

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stained preparations, however, show the presence of two additional and much weaker traces connecting directly to the basal vascular ring. In the strictest sense, therefore, the leaves are 3-nerved. G a l i u m wigginsii Dempster, sp. nov.

(Fig. 3).

Planta perennis polygama erecta 15-35 cm alta, nodis basalibus lignosis; caules foliaque velutini; folia uninervia, anguste vel late elliptica vel ovata, apicibus acutis sensim angustatis; folia inferiora conferta 2.5-5 mm longa, internodia aequantia, superiora remota ad 11 mm longa, quam internodia 2-4-plo breviora; stomata, depressa, in pagina superiore deficientia; cellulae glandulosae subapicales veI saepe deficientes; inflorescentia tantummodo bracteata, plerumque angusta, ramis brevibus sed interdum patentioribus longioribusque; corollae rotatae glabrae plerumque rubrae sed interdum viridi-flavae, apicibus prolongatis; fructus sicci, pills rectis hispidi; chromosomatum numerus, 2n: = 22. TYPE: M E X I C O : BAJA CALIFORNIA: Sierra San Pedro MSrtir, meadow at La Encantada, 2200 m, Wiggins & Demctree 4879. (~O*.OTYPE: UC 562794; ISOTYI'ES: DS, F, GH, NY, US). Baja California: Sierra San Pedro MSrtir, 1650 to 3150 m, in meadows and on cool slopes among granite boulders, with open forest of Pinus jeffreyi, P. lambertiana, and Libocedrus. Additional specimens examined: BAJA CAHrORNIA: Sierra San Pedro M~rtir, Santa Rosa, Moran & Thorne 14,399 (SD); Sierra San Pedro M/trtir National Park, Goldman 1225 (F, US), Breedlove 16,307, Moran 11,262 (DS, SD, US), 15,057, 15,289, 15,480, 15,510, 15,632 (all SD) Moran & Thorne 14,124 (SD), Robertson 13 (UC), Wiggins 16,597 (DS), Wiggins & Demaree 5034 (DS, GFI, US), Dempster 4222~ 4223, 4226, Hardham & Dempster 17,007 (all JEPS). MEXICO:

Friedrich Ehrendorfer in 1952 annotated specimens of this taxon in several herbaria (DS, F, UC, US) as Galium wrightii Gray subsp, wigginsii Ehrendf. Although this name was never published, I have chosen to adopt the epithet for the new species, selecting as the type the specimen designated by him. Galium wigginsii is clearly a part of the G. wrightii-G, parishii complex, but within the group it is distinctive as well as geographically isolated. The very fine pubescence resembles that of G. parishii of California and Nevada, as well as some scattered forms of G. wrightii fro.m Arizona and New Mexico. The thin and flaccid elliptical leaves are in rather sharp contrast to the usually fleshy and much broader leaves of G. parishii, and the usually stiffer as well as much narrower leaves of G. wrightii. Their gradual acute apices are very different from the abruptly apicnlate apices of G. pariskii. The change from small, congested leaves to larger remote leaves occurs much higher on the stein and is more abrupt than in any form of G. parishii or G. wrightii. T h e absence of stomata from the upper leaf surfaces is in contrast both to G. parishii and to most of G. wrightii, including the latter as found in the Sierra Juitrez of Baja California. The sunken stomata are in agreement with G. parishii. The inflorescences of G. wigginsii resemble those of G. wrightii, but are usually much narrower. They are, however, much broader, less leafy and tess congested than those of G. parishii. Although the leaves are obviously 1-nerved to the naked eye, it can be demonstrated by clearing and staining that they are often 2- or 3-nerved (average 1.3 nerves per leaf). A similar situation exists in G. wrightii, wherein, however, the number of nerves is substantially higher (average per leaf 2.5 nerves). The diploid chromosome count is in agreement with all counts made so far in the G. wrightii-G, parishii complex.

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FIG. 4. Galium moranii. A, portion of branch, X 1; B, branchlet with mature fruits: 1 bicarpellate, the other three each with 1 aborted carpel, X 2; C, corolla, N 7; D, cleared, stained and dissected monocarpellate fruit, shown without hairs, X 7; E, cleared and stained leaf, with leaf bases, to show venation, X 7; F, detail from middle portion of edge of same leaf, X 55. (Carter & Sousa 5172, type).

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[VOL. 22 (Fig. 4).

Planta perennis scandens flaccida laxa hirtella, inflorescentiis laxis paucifloris; caules foliaque molliter pubescentes; caules ad 0.75 m longi, internodiis plerumque 2-7 cm longis, quam foliis 1.5-7-plo longioribus; folia anguste usque late elliptica 7-14 mm longa, in pagina superiore sine stomatibus; cellulae glandulosae deficientes vel aliquot subapicales; pedicelli quam fructus 1-4-p[o longiores; corollae rotatae extra hirtellae, viridi-flavae vel albae vel sub-roseae; fructus uni- vel bi-carpellati, carpellis fere sphaericis, multis pills rectis tenuissimis obtectis; endospermium maturitate ut videtur haud indurescens. TYPE: M E X I C O : BAJA CALIFORNIA SUR: Sierra de la Giganta, headwaters o f main canyon of Arroyo Hondo, N side of Cerro Giganta, 840 m, Carter & Sousa 5172. (HOLOTYPE: UC 1334278). Trailing on shaded boulders and steep north-facing slopes below cliffs; Sierra de la Giganta at 780-10.35 m. Collections studied: Sierra de la Giganta: Cerro del Barreno, Carter & Moran 5331 ; north side of Cerro Giganta, Carter & Sousa 5163. This species, like the last, must be referred to the G. wrightii-G, parishii complex, from which it is, however, even more divergent than G. wigginsii. Outstanding differences include the weak stems, the very thin, elliptical, and relatively large leaves, the long pedicels, and the soft endosperm. The last-mentioned character can be seen only in the type collection, since the other two. have no mature fruits. The character, which may not be consistent throughout the species, is striking. Although I have examined thousands of specimens, I have never heretofore seen long-haired Galium fruits of any age which, like the apparently mature fruits of the G. moranii type material, are actually extruded when pressed; i.e., although the fruits will flatten under pressure when young, they always remain hard and round when mature. The fruits of G. maranii are not, as appears at first, fleshy like those of G. cali]ornicum H. & A., and G. nuttallii Gray, in which the ovary expands and becomes juicy. Rather, it is the endosperm that fails to become hard and resistant to distortion. When extruded, it is sticky and adheres to the paper in which the plant is pressed, much after the manner of the flesh of the truly baccate species. The term "pedicel" as used in the description needs to be qualified. For practical purposes these are pedicels, in the sense that they are leafless stem-like organs supporting each a single flower. However, with this, as with other Galium species, closer examination reveals in many, or even most, instances a minute bract, or at least some indication of a node, showing that the true pedicel is only a fraction of the apparent pedicel. Often it is virtually impossible to say what is truly a pedicel, and what is a combination of pedicel and branchlet. Bractlets marking nodes on apparent pedicels may be fairly large to minute, caducous, or perhaps entirely nonexistent. The three present collections show slight variation among themselves, but it is evident that more intensive study is needed in the area of the Sierra de la Giganta, and perhaps elsewhere, to demonstrate the full range of variation. ACKNOWLEDGMENTS I am grateful to Lawrence Heckard, Curator of the Jepson Herbarium, who generously provided laboratory space for m y work. Thanks are due also to Rimo Bacigalupi for critically reading the manuscript and editing the Latin.