(DRAFT) - Taxonomy
                            Species GLOBE-BERRY, TUMAMOC
                                Species Id ESIS702036
                                   Date 13 MAR 96



TAXONOMY

NAME - GLOBE-BERRY, TUMAMOC OTHER COMMON NAMES - GLOBE-BERRY and TUMAMOC ELEMENT CODE - CATEGORY - Angiosperm PHYLUM AND SUBPHYLUM - MAGNOLIOPHYTA, CLASS AND SUBCLASS - MAGNOLIOPSIDA, ORDER AND SUBORDER - VIOLALES, FAMILY AND SUBFAMILY - CUCURBITACEAE, GENUS AND SUBGENUS - TUMAMOCA, SPECIES AND SSP - MACDOUGALII, SCIENTIFIC NAME - TUMAMOCA MACDOUGALII AUTHORITY - TAXONOMY REFERENCES - COMMENTS ON TAXONOMY - Tumamoc Globe-berry Tumamoca macdougalii Rose KINGDOM: Plant GROUP: Angiosperm DIVISION: Magnoliophyta CLASS: Magnoliopsida ORDER: Violales FAMILY: Cucurbitaceae Tumamoc globe-berry (Tumamoca macdougalii Rose) is a delicate vine; slender stems annual (bases of the largest ones become woody and perennial), arising from a cluster of shallow tuberous roots, tubers spread irregularly downward or horizontally from the plant's central axis, tubers connected by fragile slender connecting roots; stems and leaves glabrous. Seedling and juvenile plants possess triangular or pentagonal-shaped bright green leaves. Leaves of adult plants are pedately parted into three lobes, these spreading and secondarily lobed, the tips mucronate or rounded, highly variable in width of leaf (20-90 mm) and width of lobes (2-10 mm), thin and bright green when in dense shade to dark green, thickened, and leathery in full sun. Leaves of juvenile plants pentagonal in outline, with the medial (distal) division much larger than the others, bright green, 5-20 mm across, pointed. Flowers monoecious; male flowers borne on 20-150 mm long racemes in clusters of 2-6, the Taxonomy - 1 (DRAFT) - Taxonomy Species GLOBE-BERRY, TUMAMOC Species Id ESIS702036 Date 13 MAR 96 perianth lobes narrowly lanceolate, to 5 mm long, pale greenish-yellow, stamens 3, borne on the tube about one-fourth the distance below the mouth, 2 of the anthers 2-celled, the third 1-celled; female flowers similar, but are solitary and borne on pedicels from 4-25 mm long which originate in the leaf axils, calyx with very slender tube and 5 small sepals, corolla paleyellow with narrow, elongated lobes, petals 8-12 mm long. Fruits are succulent, spherical to ovoid, pale green when immature with darker green vertical stripes, becoming yellow, orange, and finally bright red at maturity, 8-10 mm in diameter, average 0.5 g (.01-1.84 g). Seeds average 2.9 (0-10) per fruit, average 22 mg, 5-8 mm long, 4-7 mm wide, 2-3 mm thick, pear-shaped (obovoid), truncate at apex, rough or tuberculate, not marginate, and dark gray to black (03,08,10,11,15,16, 17). Tumamoca is a monotypic genus of the Cucurbitaceae family (04, 13). The Tumamoc globe-berry was first collected by David T. Macdougal on July 31, 1908, on Tumamoc Hill west of Tucson, Arizona, where the Carnegie Institute Desert Laboratory was located. The specimen was sent to John N. Rose at the U.S. National Herbarium, who described the new genus and species in honor of the type locality and of the collector (04,15,17). The type specimen is housed at the U.S. National Herbarium in Washington, D.C. (#591589, 15). Tumamoca macdougalii has often been confused with Ibervillea sonorae (S. Watson) E.L. Greene, a similar plant in a related genus. Rose (15) discusses the similarities and differences between the two genera. He makes reference to the flower differences (monoecious vs. dioecious), variation in seed size, shape and markings, and to the habit of Tumamoca forming a "cluster of tuberous roots instead of a single globular root" as does Ibervillea. Anthers of Tumamoca are borne on the slender corolla tube well below its opening while those of Ibervillea are borne at the tube opening (02,10). Reichenbacher and Felger (12) state that the Tumamoca seeds are larger than those of Ibervillea and are easily separated from the surrounding bright red juicy matrix. Clay May (02), a Tucson botanist and chemist, extracted flavonoids from T. macdougalii, I. sonorae var. sonorae, and I. tenuisecta (A. Gray) D.K. Small. Ibervillea sonorae is the geographically closest relative of T. macdougalii and I. sonorae var. sonorae is the only sympatric close relative. May (02) concluded that Tumamoca was less closely related to the Ibervillea taxa than the latter were to each other, thus supporting the generic distinction. Summer monsoon showers in the Sonoran Desert of greater than one-half inch produce dramatic morphological changes in the above ground growth of T. macdougalii. The slender, leathery leaves produced during the early summer drought (termed "drought leaves) expand to more than twice their previous size (07). After substantial summer rains flower racemes produce 15-17 male flowers as long as soil moisture is adequate. During times of drought the average is 3-6 per raceme. Mills (03) found tuber shape and orientation to be influenced by soil conditions. He reports that "in loose sandy loams, where most tubers were found, tubers grew more or less perpendicularly. Those tubers that were sharply angled or nearly horizontal seemed to be in heavier clay soil. Potato-like tubers, those that ended bluntly, were Taxonomy - 2 (DRAFT) - Taxonomy Species GLOBE-BERRY, TUMAMOC Species Id ESIS702036 Date 13 MAR 96 present more often where soil below the tuber changed abruptly to heavy clay." Some larger tubers were found growing around the roots or trunks of cacti or trees. Taxonomy - 3
                                  (DRAFT) - Status
                            Species GLOBE-BERRY, TUMAMOC
                                Species Id ESIS702036
                                   Date 13 MAR 96



STATUS

Coded Status E: Federal Endangered Non-consumptive recreational Ornamental COMMENTS ON STATUS - U.S. STATUSES AND LAWS: The Tumamoc globe-berry (Tumamoca macdougalii) has been designated as Threatened pursuant to the Endangered Species Act of 1973 (50 CFR 17.12; P.L. 93-205, 87 Stat. 884; 16 U.S.C. 1531-1540), as amended. The species has this status wherever found including the State of Arizona and Sonora, Mexico. Removal and reduction to possession of any Federally listed plant from an area under Federal jurisdiction is unlawful (50 CFR 17.61 and 17.71). This species is protected by the Lacey Act (P.L. 97-79, as amended; 16 U.S.C. 3371 et seq.) which makes it unlawful to possess any wild plant (including roots, seeds, and other parts) within U.S. territorial or special maritime jurisdiction (as defined in 18 U.S.C. 7); or to import, export, transport, sell, receive, acquire, or purchase in interstate or foreign commerce any wild plant (including roots, seeds, and other parts) taken, possessed, transported, or sold in violation of any State law or regulation. It is also unlawful to import, export, transport, sell, receive, acquire, or purchase any wild plant (including roots, seeds, and other parts) taken or possessed in violation of any U.S. law, treaty, or regulation or in violation of Indian tribal law. RESPONSIBLE FEDERAL AGENCIES: USFWS -Responsible for the management/recovery, listing, and law enforcement/protection of this species. BIA -Responsible for the law enforcement/protection of this species with applicable State and Federal laws on public lands under their control. Also responsible for management/recovery on Bureau of Indian Affairs lands. BLM -Responsible for the law enforcement/protection of this species with applicable State and Federal laws on public land under their control (43 CFR 4140). Also responsible for management/recovery on Bureau of Land Management lands. DOD -Responsible for the law enforcement/protection of this species with applicable State and Federal laws on Status - 1 (DRAFT) - Status Species GLOBE-BERRY, TUMAMOC Species Id ESIS702036 Date 13 MAR 96 public land under their control. Also responsible for management/recovery on Department of Defense lands. NPS -Responsible for the law enforcement/protection of this species with applicable State and Federal laws on public lands under their control. Also responsible for conservation (Nat. Park System Organic Act - 16 U.S.C. 1, 2-3)/management/recovery on National Park Service lands. Taking, possessing, or disturbing of Federally listed species is prohibited on NPS lands (36 CFR 2.1, 2.2, and 2.3). USFS -Responsible for the law enforcement/protection of this species with applicable State and Federal laws on public lands under their control. Also responsible for management/recovery on Forest Service lands. The Forest Service is responsible for integrating management, protection, and conservation of Federally listed species into the Forest Planning process (36 CFR 219.19 and 219.20). All Federal agencies have responsibility to ensure that any action authorized, funded, or carried out by that agency is not likely to jeopardize the continued existence of the species or result in the destruction or adverse modification of Critical Habitat (50 CFR 402), and to utilize their authorities to carry out programs for the conservation of the species. STATE STATUSES AND LAWS: STATE: Arizona DESIGNATED STATUS: Protected ADMINISTRATIVE AGENCY: AZ Comm. of Agriculture & Horticulture STATE STATUTE: Ariz. Rev. Stat. Chapt. 7, Sec. 3-901B. STATE: Sonora, Mexico UNOFFICIAL LIST: Under evaluation presently. INTERNATIONAL STATUSES, TREATIES, AND AGREEMENTS: This plant is under evaluation to determine its status in Sonora, Mexico. ECONOMIC STATUSES: This species may be of value for some of its yet undetermined phyto-chemical products and it has ultural value as a component of Arizona's natural heritage of biotic diversity. It is a succulent and may be a target of collectors. 80/12/15:45 FR 82480/82481 - Notice of continued review 85/05/20:50 FR 20806/20810 - Proposed rule 85/09/27:50 FR 39526/39527 - Notice of review 86/04/29:51 FR 08681/08683 - Final rule, Endangered Status - 2
     

HABITAT ASSOCIATIONS

HABITAT - TERRESTRIAL TERRESTRIAL LAND USE - Residential Commercial and Services Transportation, communications, and Util Mixed Urban or Built-up Land Shrub and Brush Rangeland COMMENTS ON HABITAT ASSOCIATIONS - The range of Tumamoca macdougalii covers some 31,000 square miles of Sonoran Desert from just southeast of Guaymas, Sonora, Mexico, to Tucson, Arizona, west to Organ Pipe Cactus National Monument (12) and north to Pinal County, Arizona. Until recently the species was only known in the United States from Pima County, Arizona (17), but one population has been discovered just north of Pima County in Pinal County. T. macdougalii is not uniformly or randomly distributed throughout the available habitat (08). Instead, the distribution is highly contagious (=clumped), consisting of discrete isolated populations of one to several hundred plants which cover areas of less than one acre to about 400 acres (12). The species is capable of occupying a wide range of habitats from halophytic coastal scrub communities on clayey saline hardpans only a few hundred feed from the Gulf of California shoreline to rocky loamy soils derived from weathered granite at nearly 3000 ft. elevation in south central Arizona (11). Southeast of Guaymas, it occurs in a halophytic coastal scrub community on extremely salty hardpans; at Kino Bay it occurs in coastal scrub on salty sand of old barrier dunes; in Tucson it is found on hot dry, south-facing slopes of basalt. The largest population known is found in creosotebush desertscrub on gravelly loams primarily derived from weathered granites (08). The Federal Register (04) describes the habitat of T. macdougalii as palo-verde/cholla cactus scrub and creosotebush bursage/bursage desertscrub. Dominant associated species include creosote bush (Larrea divaricata), foothills and blue paloverde (Cercidium microphyllum & C. floridum), whitethorn acacia (Acacia constricta), saguaro (Carnegiea gigantea), prickly pear (Opuntia phaeacantha), cane cholla (Opuntia versicolor), mesquite (Prosopis juliflora), ironweed (Olneya tesota), and triangleleaf bursage (Ambrosia deltoidea). In Sonoran Desertscrub, Arizona Upland Subdivision, Paloverde- Mixed Cacti Associations at elevations of 450-795 meters (1476-2608 ft) it is found on well-drained coarse soils dominated by foothills paloverde, triangleleaf bursage, ironwood, buckhorn cholla (Opuntia acanthocarpa), chainfruit cholla (O. fulgida), prickly pear, barrel cactus, saguaro, creosotebush, range ratany (Krameria parviflora), whitehorn acacia, and ocotillo (Fouquieria splendens) (11). Soils range from rocky to gravelly, sandy silty, and clayey, and may be derived from granite, basalt, and rhyolite. T. macdougalii has been reported from desert riparian communities on poorly drained, finely textured soils at lower elevations in the Lower Colorado Valley Subdivision (11) Creosotebush-bursage Associations with creosotebush, triangleleaf bursage, mesquite, and white bursage (Ambrosia dumosa). In some cases wash habitats can be quite mesic and it has been found in association with a host of introduced and native weedy forbs and grasses. Habitat Associations - 1 Many botanists are under the impression that T. macdougalii is commonest along washes. Although in some areas it is frequent along washes it is unclear if this is due to the greater availablility of microhabitat, simply because there are more potentially sheltering trees and shrubs along washes, or if the plants actually derive any benefit from occasional flows in the channel. The type population and several others are on the lower slopes of hills and are not related in any way to drainageway channels. T. macdougalii is also known from Sonoran Desertscrub, Central Gulf Coast and Plains of Sonora Subdivisions (12). In the Guaymas region vegetation is dominated by mixed lowland (mainly non-halophytic trees, shrubs, and cacti) and coastal (maily halophytic shrubs) desertscrub. Typical halophytes include Suaeda moquinii, Maytenus phyllanthoides, and Frankenia palmeri. On the Tohono O'Odham Nation in Pima County, Arizona, T. macdougalii is found in communities dominated by allscale (Atriplex polycarpa) usually associated with mesquite and other common Sonoran Desert species in valley bottoms (07,11). It is believed that the plants are widely scattered but ubiquitous throughout the allscale plant communities on the reservation (09). Types of vegetation on the Tohono O'Odham Nation that T. macdougalii has been found in are Mesquite-Allscale-burrowseed association (09). Thirteen plants of T. macdougalii found northwest of Kaka on the Tohono O'Odham Nation were located immediately below bedrock outcrops at the base of a hill in limestone soils. This Pinal County location is the only population found on limestone soils (09). Vegetation was dominated by a foothills paloverde-triangleleaf bursage-creosotebush association. Popualtions in Avra Valley and the Tucson basin are between 2000 and 2680 feet elevation. Thirteen populations are found on the lower slope of small mountains and hills, but none occur on the bedrock-dominated mountain slopes. No populations have been found in the floodplains of any major drainagewasy such as the Rillito River, Brawley, Wash, or the Santa Cruz River, although many populations are adjacent to or within the floodplains of minor washes (11). The parameters of its preferred habitat are poorly known. There are no obvious environmental features limiting its distribution except that it is limited to lower elevations because of frost susceptibility, and its southern and western range limits are probably related to increased competition in the less arid Sonoran thornscrub vegetation, while its western and northern range limits are probably due to decreasing summer precipitation and lower winter minimum temperatures, respectively. In the Tucson-Avra Valley area, which is at the species' northern and eastern limits, although it occurs in a number of different habitats it is restricted to isolated discrete populations whose boundaries do not coincide with habitat boundaries (07,11). Populations thoughout the range are separated by considerable expanses of apparently suitable, but unoccupied habitat with individuals clumped within populations. Estimating the abundance of the species by searching small areas and extrapolating to the entire range is impossible without an extremely large sample size. Habitat Associations - 2
                                (DRAFT) - Food Habits
                            Species GLOBE-BERRY, TUMAMOC
                                Species Id ESIS702036
                                   Date 13 MAR 96



FOOD HABITS

TROPHIC LEVEL - AUTOTROPH Food Habits - 1
                         (DRAFT) - Environment Associations
                            Species GLOBE-BERRY, TUMAMOC
                                Species Id ESIS702036
                                   Date 13 MAR 96



ENVIRONMENTAL ASSOCIATIONS

G = General A = Adult LIM = Limiting RA = Resting Adult J = Juvenile FA = Feeding Adult RJ = Resting Juvenile BA = Breeding Adult FJ = Feeding Juvenile P = Pupae L = Larvae E = Egg RL = Resting Larvae FL = Feeding Larvae
LIFESTAGE ENVIRONMENTAL ASSOCIATIONS G G G G G G G Environment Associations - 1
                               (DRAFT) - Life History
                            Species GLOBE-BERRY, TUMAMOC
                                Species Id ESIS702036
                                   Date 13 MAR 96



LIFE HISTORY

HABIT: The Tumamoc globe-berry is a liana/vine (08,11,15). LIFE CYCLE: Tendrils and dwarfed spare lea ves begin to emerge from the subsurface tubers of 5-20 percent of the perennial plants in a population by late April or early May. A few scattered leathery, much reduced leaves, several racemes of male flower buds, and, on monoecious plants, a few female flower buds are produced. Although stems may elongate to 3 meters or more, growth remains very sparse though May and most of June. Only a fraction of the buds produced actually reach anthesis and very few fruits survive to maturity (08). New shoots emerge from the subsurface tubers, elongate while climbing through the nurse plant, and attain lengths of up to 7 meters prior to the onset of summer rains (a little less than half of the total seasonal growth) (07). Growth on the remaining plants usually does not occur at all until the first heavy summer rainfall of 0.5-1.0 inches sometime in July; then growth is explosive (07). Following a heavy summer rain growth can begin in just a few hours. Stems, leaves, and flower buds already in place develop to their fullest potential and reproduction begins within two to five days after. Some plants apparently lapse into periods of dormancy for no apparent reason and may not develop stems and leaves for more than one growing season at a time. After summer rains abate the plants revert to the conservative mode of producing only drought leaves. Resources are increasingly channeled into fruit maturation and away from vegetative growth. A few flowers are sporadically produced during this period, usually from late August through November. Juvenile plants usually possess a single herbaceous (i.e., green) stem, and adults have at least one and sometimes several white to light gray woody stems (13). Each node along the stem produces one tendril, one leaf, one male flower raceme, and one female flower bud (07). Flowering begins before the summer rains, though female buds are not produced or are quickly aborted (04). The length of the longest stem is strongly and positively correlated with tuber weight. This suggests that accurate predictions about tuber weight can be made simply by measuring the lengths of the longest stems (13). The plants begin retreating to the subsurface tubers again by mid-fall, usually mid- to late October. Growth is sporadic after that (08). Cessation of growth depends on rainfall and the date of the first hard frost. By mid- to late December frosts have usually destroyed all of the green stems and foliage. Larger adults retain a few feet of woody stem, but otherwise all of the aboveground parts of the plant wilt and gradually disintegrate (12). Plants overwinter, dormant in their underground tubers. TYPE OF REPRODUCTION: Sexual (15). Experiments on the pollination ecology of T. macdougalii indicate that its pollen is self-compatible. Fruits are, however, not produced without pollination. Fruit set is highest Life History - 1 (DRAFT) - Life History Species GLOBE-BERRY, TUMAMOC Species Id ESIS702036 Date 13 MAR 96 with cross-pollination, and, at least one site, lowest when flowers are left to natural pollination (02,10). This indicates the species has difficulty attracting and utilizing pollinators and may explain how populations are able to maintain monoecious, all female, and all male plants. REPRODUCTIVE PHENOLOGY: Germination: Germination commences within a few days after the first summer rainfall of over 0.5 to 1.0 inches. This may occur any time from late June to early September. The seedlings of any given growing season are produced from the fruits of previous years. It is not known how long the seeds may survive in the soil. Seedlings have a very low survival rate. The number of plants in each increasing size class decreases dramatically, presumably plant size is related to age. Leafing: Slender stems, tendrils, and dwarfed spare leaves begin to emerge from the subsurface tubers of 5-20 percent of the plants in a population by late April or early May (08). Although stems may elongate to 3 meters or more the growth remains very slender through May and most of June. Within 48 hours of the first heavy rains the size of the leaves more than triples (08). Budding: Male flower buds are produced before the summer rains in May and June, but only a few reach anthesis (08). About one-tenth the number of female flowers are produced. Clay May reports that not all adult plants flower every year. During periodic droughts the plant selectively aborts reproductive and vegetation components. The first components aborted are female flower buds, followed by male buds, fruits, leaves, branch tips, and finally whole branches (07). Anthesis: Anthesis occurs from late July through the rainy season which usually ends sometime in late August or early September. Flower production declines rapidly after the rains subside. Male flower buds persist, but few reach anthesis. Fruiting: Fruit production begins with one or a few fruits early in the summer, though most of these never mature. Most fruit production occurs in one burst of flowering triggered by the first heavy summer rain. This can occur at any time from late June through August (08,13). Fruits mature in four to five weeks (07). Seed/Fruit Dispersal: Fruit maturation usually does not begin until late August and continues through September. Most fruits ripen and then disintegrate on the vine. Some mature fruits fall to the ground and are taken by ants. So far there are no recorded instances of wildlife taking the fruits in the wild (08,09,10). The vast majority of seeds simply fall to the ground immediately below the maternal parent and are incorporated into the litter. SEX OR SPORE STATUS: Tumamoc globe-berry was originally reported to be monoecious by J.N. Rose (15), but it is now known to be facultatively dioecious (02,07,10,11,13) and self-compatible, while expressing towards androecy (07). The ration of male to female functions in the species is undoubtedly linked to the favorability of the precipitation regime (11). Female flower production is significantly negatively correlated with both tuber weight and length of longest stem. Plants may be Life History - 2 (DRAFT) - Life History Species GLOBE-BERRY, TUMAMOC Species Id ESIS702036 Date 13 MAR 96 predominantly female when small and then become increasingly more male as they age. This contradicts the findings of Clay May (02) who found that the smallest plants were the most male (13). It is thought that part of the plants adaptations to its unpredictably arid environment probably includes selective flower and fruit abortion. This ability helps the plant reduce moisture losses in times of drought (13). Pollination studies indicate significant inbreeding depression in self-pollinated flowers (11). Although no data have yet been obtained, it is possible that some genetic differentiation has occurred between populations due to the frequency of inbreeding (11). The pollinator(s) have not been identified yet and the data on pollen viability, effects of self-fertilization, and many other features are preliminary at this time (11). The pollinator(s) are believed to be one or more moth species. Pollination studies conducted so far indicate significant inbreeding depression in self-pollinated flowers (10). Pollinator availability appears to be very limited causing a significantly reduced fruit set in natural populations. POLLINATION, SPORE AND SEED DISSEMINATION: Pollination studies indicate significant inbreeding depression in self-pollinated flowers (11). The pollinator(s) have not been identified yet, but are thought to include at least one and possibly several species of small moth. Pollination studies indicate that pollinator availability is very limited and is partially responsible for a very low rate of fruit set in natural populations. Fruits are large and juicy. The colors change from green to yellow to bright red and seeds are relatively large and thick-shelled. These features strongly suggest that the species depends on birds for seed dissemination, yet so far there are no recorded observations of any wildlife species utilizing the fruits in the wild (12). Ants occasionally collect the fruits, but probably not frequently and the resultant dispersal would be limited. The highly contagious (=clumped) distribution suggests very strongly that seeds are inefficiently dispersed (07,08). Some obscure environmental parameter may be responsible, but this seems unlikely. It has been suggested that the original disperser was a bird species which is now, at least locally extinct, and that this is responsible for the contagious distribution (07). The fruit seems best suited for dispersal by a bird. In such a case the fruit would be swallowed whole and seeds dispersed through digestion. The juicy inner matrix is what would attract the disperser (12). The fruit wall is easily digested. In contrast Ibervillea fruits are too large to be swallowed whole and the tough rind is probably inedible. In the latter case the fruits are mostly eaten by birds with long thin bills (such as thrashers) by poking a hole in the rind and cleaning out the interior of the fruit (12). It is believed that nurse plants provide support for the delicate vine and provide a means of displaying mature fruits to potential seed dispersers (11). Seeds are thick-shelled and are believed to be able to survive in the soil for several years (08). Life History - 3 (DRAFT) - Life History Species GLOBE-BERRY, TUMAMOC Species Id ESIS702036 Date 13 MAR 96 SEED BIOLOGY: Seeds average 2.9 (0-10) per fruit and 22 mg in weight. Seeds are 5-8 mm long, 4-7 mm wide, 2-3 mm thick, pear-shaped (obovoid), truncate at the apex, rough or tuberculate, not marginate, and dark gray to black (03,08,10,11,15,16,17). Seeds generally overwinter in the soil and germinate after the following year's heavy summer rains (08). May (03) has found a very high seed and seedling mortality rate. In natural populations approximately 605 seeds are required to produce a single adult plant. Germination rates of 50-90 percent have been obtained with and without seed scarification. Tumamoca is considered relatively easy to propagate under greenhouse conditions (11). POPULATION BIOLOGY: Tumamoca macdougalii is believed to be a relatively long-lived species. A preliminary life table developed by May (02). Using data from several Tucson area populations indicates that only 1 in 20 seeds germinates and of the resulting seedlings 1 in 22 survives to adulthood. A conservative age estimate based on the number of nodes on the woody portions of stems indicates that the largest adults may be 25 years old. The plants are likely to be much older than that because stem clipping is extremely common and very old plants may, therefore, have stems appearing to be relatively young. Because seed dispersal is limited, 76 percent of seedlings were found beneath the canopies of large adults, presumably their maternal parents. In the past four years some populations have perceptibly declined in numbers while others, only a few miles away, have increased significantly. The Sabino Canyon population numbered 41 plants when first studied in 1984 (07). In 1985 the population lost several plants, but gained others with a net increase of one plant (08). This is despite the fact that photographs taken both years indicate that summer precipitation in 1985 was well below that of 1984. The Central Arizona Project (CAP) population was first documented to include 468 plants in 1984 (07), by 1985 the population had grown to at least, if not many more than, 736 individuals, and by 1986 it increased to 813 plants. A population a few miles north of the CAP population was comprised of 152 individuals in 1984 (07), but by 1986 although the site was not comprehensively resurveyed, microsites formerly supporting several dozen plants were completely devoid. This population seems to have dropped drastically in numbers. The difficulties of relocating plants once discovered, of distinguishing them from neighbors only a few milimeters away, and of accounting for dormant plants makes it nearly impossible to construct accurate life tables. The main factors affecting survival are known: 1) intensity and seasonal distribution of summer precipitation; 2) intensity, duration, and seasonal distribution of droughts; 3) date and intensity of first hard frost; 4) local predation pressure, varying from minor stem clipping to partial or complete excavation; and 5) anthropogenic habitat alteration. The recoverability of the species is limited. Some progress can and is being made towards preservation of known populations, particularly in the Tucson area. Much of the urban and agricultural Life History - 4 (DRAFT) - Life History Species GLOBE-BERRY, TUMAMOC Species Id ESIS702036 Date 13 MAR 96 growth impacting the species in this region is beyond the control of the Endangered Species Act and cannot be controlled through any state or local regulatory mechanisms. Although the Bureau of Reclamation is installing javelina proof fencing around four of the largest populations, it is unlikely that any additional progress will ever be made towards control of animal predation. The factor which really seems to be responsible for the rareness of the plant is a faulty mechanism of seed dispersal combined with a lak of pollinators. Until these problems are fully understood it will be difficult to develop a truly effective recovery plan. ECOLOGICAL/EDAPHIC FACTORS: Tumamoca macdougalii has been found to occur on a wide variety of soil types. Reichenbacher (11) has found it on well-drained coarse soils in Arizona Upland Subdivision and on poorly drained, finely textured soils at lower elevations in the Lower Colorado Valley Subdivision. The species is capable of occupying a wide range of habitats from halophytic coastal scrub communities on clayey saline hardpans only a few hundred feet from the shoreline in Mexico to rocky loamy soils derived from weathered granite at nearly 3000 feet in south central Arizona. Wiggins collected a specimen from dry gravelly soil and Reichenbacher and Felger reported the soils from the Carbo locality to be gravelly loams characteristic of major drainageway bottoms. Soils on the Tohono O'Odham Nation in the mesquite-allscale associations are sodic-saline sandy loams (09,14). Populations have been found on loams, silt loams, sandy loams, hard clays, loose saline sands, and poorly drained sodic-saline soils (01, 05 12,14). Crossman (01) describes the overall soil characteristics as well-drained soils, fine grained and friable with organic matters consistently present. Of the samples tested pH's ranged between 7-8 with virtually no EC. Parent materials range from granite, rhyolite, rhyolite/andesite mixed, andesite, andesite basalt, basalt, limestone, and from soils developed on undifferentiated valley fill (01,11). The largest population in east Pima County occurs on soils derived from weathering of amole granites in the Tucson Mountains (11). Soil texture varies from coarse to very fine. Thirteen plants found near Kaka (Pinal County) were not on the bedrock limestone of the hill itself but were immediately below bedrock outcrops at the base of the hill in limestone soils. This is the only record of the plants occurring on limestone soils (09). Mills (03) states that "tuber shape and orientation is influenced by soil conditions. In loose sandy loam, where most tubers were found, tubers grew more or less perpendicularly. Those tubers that were sharply angled or nearly horizontal seemed to be in heavier clay soil. Potato-like tubers, those that ended bluntly, were present more often where soil below the tuber changed abruptly to heavy clay." The population at Sabino Canyon (Pima County) is noteworthy in that it is the only population known from rocky terraces adjacent to a perennial stream (08). T. macdougalii seems to be remarkably non-selective in its microhabitat preferences. Plants have been found in brushy, weed-choked ephemeral stream floodplains and on dry rocky slopes of Life History - 5 (DRAFT) - Life History Species GLOBE-BERRY, TUMAMOC Species Id ESIS702036 Date 13 MAR 96 ridges and south-facing slopes (12). Several plants have been found in man-modified microhabitats. One group was found on an embankment created two or three decades ago to serve as a firing range backdrop (07). The plants are always found climbing among shrubs and trees, which provide shade and elevated humidities for delicate seedling and juvenile plants, and support for the larger adult vines (13). The problem of survival on a wide variety of soil types may be circumvented by requiring germination and establishment to occur beneath a nurse plant, where the soil environment is highly modified by shade of the nurse plant and organic material from decaying vegetation (11). Clusters of 50-100 plants have been found beneath the canopy of a single nurse plant. TROPHIC STATUS: T. macdougalii is completely autotrophic and phototrophic. It does not participate in any symbiotic or parasitic relations (15). CHARACTERISTIC DOMINANCE: In some situations T. macdougalii can be locally abundant in microsites of 100 to 300 square feet. For the most part it is a widely scattered to extremely rare component of the vegetation at any given site. In the largest population known approximately 800 plants are scattered over an area of 400 acres. The densest concentration known consists of roughly 100 individuals plants in an area of approximately 250 square feet. COMMUNITY ECOLOGY: T. macdougalii is clearly a constituent of the climax vegetation of the regions it inhabitats. The plant is long-lived and, due to its limited dispersal ability, is limited to undisturbed habitats. Out of the approximately 2500 plants known in existence only 3 have been found on sites which were clearly disturbed. These plants were obviously the result of seed dispersal from plants on undisturbed sites only a few meters away. The species produces an average of only 3-4 seeds per fruit, the seeds are large and probably long-lived; these features indicate a climax vegetation species. SPECIES INTERRELATIONSHIPS: No symbiotic relationship is known, however, T. macdougalii is usually found under trees and shrubs (nurse plants), which provide shade and protection, as well as support for the vine (04). Only two plants (one adult and one juvenile), out of approximately 2500 known individuals, have ever been found without any nurse plant shelter at all. These were growing in dried mud at the upstream foot of a dike (07). The 'nurse plants', 'host plants', or 'shelter plants' serves two functions: 1) they provide a moist warm environment with elevated local humidity required by seeds and seedlings for successful germination and establishment; 2) and they provide support for the weak stems and a means of displaying flowers to potential pollinators and mature fruits to potential seed dispersers (11). The plants do not appear to be more abundant or vigorous in association with any particular host species. They most often grow on the northeast side of the nurse plant. Life History - 6 (DRAFT) - Life History Species GLOBE-BERRY, TUMAMOC Species Id ESIS702036 Date 13 MAR 96 Observed 'nurse plants' include foothills paloverde (Cercidium microphyllum), creosote bush (Larrea divaricata), blue paloverde (Cercidium floridum), whitethorn acacia (Acacia constricta), saguaro (Carnegiea gigantea), prickly pear (Opuntia phaeacantha), ironwood (Olneya tesota), and triangleleaf bursage (Ambrosia deltoidea), buckhorn cholla (Opuntia acanthocarpa), chainfruit cholla (O. fulgida), barrel cactus (Ferocactus wislizenii), range ratany (Krameria parviflora), ocotillo (Fouquieria splendens), allscale, pencil chola, and desert willow trees (Chilopsis linearis) (04,06,07, 08,11). Some nurse plants are preferred in some areas over others. On the Tohono O'Odham Nation 80 percent of the known plants are sheltered by allscale while plants in the Tucson-Avra Valley area were mostly sheltered by ironwood, foothills paloverde, and mesquite (09). Allscale can only shelter a few plants due to its small size. From the plants perspective the problem of survival on a wide variety of soil types may be circumvented by requiring germination and establishment to occur beneath a nurse plant, where the soil environment is highly modified by shade of the nurse plant and organic material from decaying vegetation (11). Clusters of 50-100 plants have been found beneath the canopy of a single nurse plant. OTHER LIFE HISTORY DESCRIPTORS: None. Life History - 7
                           (DRAFT) - Management Practices
                            Species GLOBE-BERRY, TUMAMOC
                                Species Id ESIS702036
                                   Date 13 MAR 96



MANAGEMENT PRACTICES

RESULT MANAGEMENT PRACTICE Beneficial Maintaining undisturbed/undeveloped areas Beneficial Restricting/regulating human use of habitats Beneficial Land Acquisition Beneficial Maintaining Sperm/Seed Banks Beneficial Stocking captive-reared wild-strain animals Beneficial Transplanting wild animals Beneficial Controlling/Restricting Noncommercial Harvest Beneficial Controlling/Removing Native Vertebrates Adverse Collecting Existing Collecting Adverse Inherent Reproductive Characteristics Existing Inherent Reproductive Characteristics Adverse Predation Existing Predation Adverse Rural Residential/Industrial Areas Existing Rural Residential/Industrial Areas Adverse Highway/Railroads Existing Highway/Railroads Adverse Soil compaction by heavy equipment in mine areas Existing Soil compaction by heavy equipment in mine areas Adverse Grazing Existing Grazing Adverse Existing COMMENTS ON MANAGEMENT PRACTICES - The U.S. Fish and Wildlife Service determined, effective May 29, 1986, that Tumamoca macdougalii (Tumamoc globe-berry) is an endangered species (51 FR 15906) (04). The species is threatened with habitat destruction from increased agricultural development, urbanization, a proposed Central Arizona Project (CAP) canal, grazing, and collection. The growth of urban centers and associated features, primarily in eastern Pima County, and agricultural development in Arizona and Sonora (especially Avra Valley, Arizona) make Tumamoca one of the few endangered species directly in the path of a rapidly expanding metropolitan area (11). The Bureau of Reclamation is working with the Fish and Wildlife Service to determine the status of Tumamoca macdougalii on the CAP route. Known populations as well as potential habitat on Bureau of Land Management administered lands may be impacted by land imprinting and seeding projects or by the possibility of land transfers from the Bureau of Land Management to the State of Arizona or to private interests (04). In recent history Tumamoca has never been abundant, but the reasons for its decline are largely unknown. It is speculated that limited seed dispersal to suitable habitat may be responsible for the sparse and unpredictable distribution (07). Except for ants, wildlife do not appear to be utilizing the fruits (07,12). Management Practices - 1 (DRAFT) - Management Practices Species GLOBE-BERRY, TUMAMOC Species Id ESIS702036 Date 13 MAR 96 Animal predation on the leaves and tubers has been observed many times (08,11). Javelina are known to forage extensively on the subterranean tubers and devastate the plant (07,11). Some plants are incompletely excavated and tuber remains often resprout, though the plant may be greatly reduced in size. Approximately 60 percent of the plants known at one site on the Tohono O'Odham Nation were partially or completely excavated by javelina. One or more plant(s) were severely damaged by javelina in 8 of 42 Tucson area populations. Leaf damage by the larva of an as yet unidentified moth and stem clipping by lagomorphs and/or rodents has also been observed (11). Stem clipping is extremely common and affects 10-30 percent of the plants in some populations. Clipping ranges from removal of small stem segments to abscission of all above-ground parts of the plant. The impact of livestocking in Tumamoca habitat is not known. Livestock have not been observed to eat the vine itself, but pruning of shrubs, devastation of areas surrounding watering holes, and reduction of groundcover may reduce the area of suitable germination and establishment habitat in any given area (11). Approximately six plants are believed to have been vandalized in 1983 and 1984 (07,11). One such plant occurred on the CAP route and 2-5 near a street along the base of Tumamoc Hill. CAP construction will result in substantial habitat destruction and population displacement. Five of the 41 populations found in eastern Pima County have been bisected by roads or by the CAP aqueduct (11). Eleven populations are so close to roads or some other development that they will be or have been substantially reduced. Two populations are close enough to soem sort of development and may be impacted in the future. The CAP canal route bisects the largest population known (523 plants) (11). A conservation program is in progress to relocate and monitor affected plants (13). Of the 468 plants documented at the CAP in 1984, 106 were transplanted, 73 of these survived and presumably 363 carried through. Of the 272 plants discovered during the 1985 location and marking project, 177 were relocated and 61 survived. Of the 75 plants discovered or created (by inadvertently breaking tuber clustersf) during excavations in 1986, 27 survived. Thus, as far is known, the total number of plants still living at the CAP population is 524 plants (13). This should be considered a rough estimate, since nothing is known about the fate of the 363 plants not involved in the transplant operation and it is not known if the 197 transplanted plants which were not growing in 1986 are dead or dormant. A total of 54 plants have been found in Sonora, Mexico (12). One Sonora population has been eliminated by conversion of habitat to buffelgrass pastureland. This may have also impacted other, currently undocumented populations. It is not known if the populations near Guaymas and at Kino Bay have been reduced by the transportation and agricultural developments which have impacted much of these areas. Conservation measures are being implemented at this time. It has been proposed by Reichenbacher (11) to 1) purchase land adjacent to the aqueduct to preserve portions of the population not affected by the project, 2) collect seeds from the transplanted plants to use in developing a reserve of live plants to offset mortality of the transplanted plants, 3) develop a plan to transplant all of the Management Practices - 2 (DRAFT) - Management Practices Species GLOBE-BERRY, TUMAMOC Species Id ESIS702036 Date 13 MAR 96 affected plants into the adjacent preserve (03,13), 4) and purchase for preservation in perpetuity, three populations on private lands elsewhere in Avra Valley. The City of Tucson agreed to avoid two large populations which were in the path of its proposed delivery line for CAP water. The Tohono O'Odham Nation is working with the Fish and Wildlife Service and Bureau of Land Management to avoid impacts of a proposed agricultural development on several small populations in western Avra Valley. The U.S. Forest Service is monitoring a population in Sabino Canyon and plans to attempt to establish new populations nearby (08,11). At the suggestion of the Fish and Wildlife Service and concerned citizens organizations, the Arizona Commission of Agriculture and Horticulture added Tumamoc globe-berry to its list of plants protected under the Arizona Native Plant Law (11). The Bureau's transplantation study has a 49.5 percent survival rate in 1987. If all plants with missing tags that showed no evidence of growth are included then the survival rate is only 36 percent (18). UNAPPROVED PLAN: U.S. Fish and Wildlife Service. 1987. Recovery Plan for Tumamoc Globe-berry (Tumamoca macdougalii) (subintegra). Tech. Draft. U.S. Fish and Wildl. Serv., Albuquerque, NM. 47 pp. The primary objective of the recovery plan is to maintain, protect, and enhance natural populations of Tumamoca macdougalii. Populations in the United States and Mexico should be protected by: 1) enforcing existing laws and regulations (Endangered Species Act, CITES, Lacey Act, and the Arizona Native Plant Law (ARS 3-901B, 2) conducting additional surveys on Tohono O'dham Nation lands, Organ Pipe and Saguaro National Monuments, State and private lands in east Pima County, 3) obtaining an agreement from the Bureau of Land Management not to dispose any lands supporting populations, 4) seeking cooperation of private landowners for preservation of existing populations on private land, and 5) developing a program of property acquisition and preservation with The Nature Conservancy. The Tumamoc globeberry needs to be studied in its natural habitat. Information is needed on: 1) its soil needs, 2) moisture requirements as related to phenology and life history characteristics, 3) nurse plant relationships, preferences, and optimal shading, 4) seed and seedling germination requirements in the wild, 5) demographic trends, Management Practices - 3 (DRAFT) - Management Practices Species GLOBE-BERRY, TUMAMOC Species Id ESIS702036 Date 13 MAR 96 6) life history, 7) pollination biology, 8) gene flow (electrophoretic and pollination biology) between disjunct population centers, 9) the effect of javelina foraging on globe-berry populations, and 10) moth species whose caterpillar feeds on the leaves of Tumamoc globe-berry. Propagation and transplant techniques should be developed to provide nursery stocks and to establish non-extant populations. Efforts to develop public awareness, appreciation, and support for the preservation of Arizona agave are also needed. Ongoing Recovery Activities: Bureau of Reclamation has or will purchase(d) land 1) adjacent to the Tucson Aqueduct to preserve parts of the population not effected by construction and 2) three populations on private lands elsewhere in the Avra Valley. They also collected seeds and transplanted plants. Surveys, monitoring, and studies by different agencies and individuals are continuing. The Arizona-Sonora Desert Museum has found it relatively easy to propagate T. macdougalii. Management Practices - 4
                                   (DRAFT) - References
                               Species GLOBE-BERRY, TUMAMOC
                                  Species Id ESIS702036
                                      Date 13 MAR 96



     

References

***** REFERENCES FOR ALL NARRATIVES EXCEPT N-OCCURRENCE ***** 01 Crossman, L.H. 1986. Pers. comm. U.S. Bur. of Reclamation, Supervisory Soil Scientist, Phoenix, AZ. 02 May, C. Unpublished data. Tucson, AZ. 03 Mills, G.S. 1986. Report on the transplantation of Tumamoca macdougalii from the Phase B corridor of the Tucson Aqueduct of the Central Arizona Project. SWCA, Inc., rept. to U.S. Bur. Recl., AZ Project Off., Phoenix, AZ. 04 U.S. Fish and Wildlife Service. 1986. Endangered and threatened wildlife and plants; determination of Tumamoca macdougalii to be endangered. Fed. Reg. 51:8681/8683. 05 Phillips, A.M., M.A. Bernzott, and B.G. Phillips. 1986. Special status plant inventory on certain BLM lands in south-central Arizona. Final rept., order no. AZ-95-PH6-0396 to the Bureau of Land Management, Phoenix Dist. Office, AZ. 06 Reichenbacher, F.W. 1984. Rare plants of the Central Arizona Project Tucson Aqueduct Phase B. The Ariz. Nature Conservancy rept. to U.S. Fish and Wildl. Serv., Albuquerque, NM. 07 Reichenbacher, F.W. 1985. Status and distribution of the Tumamoc globe-berry (Tumamoca macdougalii Rose). F.W. Reichenbacher and Assoc. rept. to Ariz. State Univ., Dept. of Botany & Microbiol., Tempe, AZ. 08 Reichenbacher, F.W. 1986. Field surveys of Tumamoc globe-berry in the Coronado National Forest with observations on growth, phenology, transplantation, and propagation techniques. F.W. Reichenbacher and Assoc. rept. to U.S. For. Serv., Coronado Nat. For., Tucson, AZ. 09 Reichenbacher, F.W. 1987. Tumamoc globe-berry surveys on the Tohono O'Odham Nation, Pima and Pinal Counties, Arizona. Rept. to U.S. Fish and Wildl. Serv., Albuquerque, NM. 10 Reichenbacher, F.W. 1987. Pollination experiments on natural populations of Tumamoc globe-berry. F.W. Reichenbacher and Assoc. rept. to U.S. Fish and Wildl. Serv., Albuquerque, NM. 11 U.S. Fish and Wildlife Service. 1987. Recovery Plan for the Tumamoc Globe-berry (Tumamoca macdougalii J.N. Rose). Tech. Draft. U.S. Fish and Wildl. Serv., Albuquerque, NM. 47 pp. 12 Reichenbacher, F.W., and R.S. Felger. 1985. Field investigations of the Tumamoc globe-berry in Sonora, Mexico, October, 1985. F.W. Reichenbacher and Assoc. rept. to U.S. Fish and Wildl. Serv., Albuquerque, NM. 13 Reichenbacher, F.W., and R.H. Perrill. 1987. Monitoring of transplanted Tumamoca macdougalii, Tucson Aqueduct, Phase B. Central Ariz. Project. F.W. Reichenbacher and Assoc. rept. to U.S. Bureau Recl., Ariz. Proj. Off., Phoenix, AZ. 14 Robinette, D. 1986. Pers. comm. U.S. Soil Conservation Service. 15 Rose, J.N. 1912. Tumamoca, a new genus of Cucurbitaceae. Contr. U.S. Natl. Herbarium, Vol. 16, Wash., D.C. 16 Shreve, F., and I.L. Wiggins. 1964. Vegetation and flora of the Sonoran Desert. 2 Vols., Stanford Univ. Press, Stanford, CA. 840 pp. References - 1 (DRAFT) - References Species GLOBE-BERRY, TUMAMOC Species Id ESIS702036 Date 13 MAR 96 References - 2 (DRAFT) - References Species GLOBE-BERRY, TUMAMOC Species Id ESIS702036 Date 13 MAR 96 17 Toolin, L.J. 1981. Status report: Tumamoca macdougalii. Rept. to U.S. Fish and Wildl. Serv., Albuquerque, NM. 18 ERO Resources Corporation. 1987. Draft report "Second year monitoring study of transplanted Tumamoca macdougalii Tucson Aqueduct Phase B Central Arizona Project" to Bureau of Reclamation, Phoenix, AZ. ***** REFERENCES FOR N-OCCURRENCE NARRATIVE ONLY ***** 01 Brown, D.E., and C.H. Lowe. 1980. Biotic communities of the Southwest. U.S. Forest Serv., Rocky Mount. Forest & Range Exp. Stn., Gen. Tech. Rept. RM-78, Tempe, AZ. 02 Reichenbacher, F.W., and R.S. Felger. 1985. Field investigations of the Tumamoc globe-berry in Sonora, Mexico, October, 1985. F.W. Reichenbacher & Assoc. rept. to U.S. Fish and Wildl. Serv., Off. of End. Species, Albuquerque, NM. 03 Reichenbacher, F.W. 1985. Status and distribution of the Tumamoc globe-berry (Tumamoca macdougalii Rose). F.W. Reichenbaher & Assoc. rept. to Arizona State Univ., Dept. of Botany & Micro., Tempe, AZ. 04 Reichenbacher, F.W. 1985. Rare plan survey selected areas of the Schuk Toak and San Xavier Districts of the Papago Indian Reservation. F.W. Reichenbacher & Assoc. rept. to Franzoy Corey Engineering Co., Tempe, AZ. 05 Reichenbacher, F.W. 1986. Field surveys of Tumamoc globe-berry in the Coronado National Forest with observations on growth, phenology, transplantation and propagation techniques. F.W. Reichenbacher & Assoc. rept. to U.S. Forest Serv., Coronado Nat'l. Forest, Tucson, AZ. 06 Reichenbacher, F.W. 1986. Rare plants of the San Xavier Farms, San Xavier District, Tohono O'Odham Nation. F.W. Reichenbacher & Assoc. rept. to Franzoy Corey Engineers & Architects, Phoenix, AZ. 07 Reichenbacher, F.W., and R.H. Perrill. 1987. Monitoring of transplanted Tumamoca macdougalii, Tucson Aqueduct, Phase B. Central Arizona Project. F.W. Reichenbacher & Assoc. rept. to U.S. Bureau of Recl., Ariz. Proj. Off., Phoenix, AZ. 08 Reichenbacher, F.W. 1987. Tumamoc globe-berry surveys on the Tohono O'Odham Nation, Pima and Pinal Counties, Arizona. F.W. Reichenbacher & Assoc. rept. to U.S. Fish and Wildl. Serv., Off. Endang. Species, Albuquerque, NM. 09 U.S. Fish and Wildlife Service. 1987. Recovery Plan for the Tumamoc Globe-berry (Tumamoca macdougalii J.N. Rose). F.W. Reichenbacher & Assoc. for U.S. Fish and Wildl. Serv., Albuquerque, NM. 47 pp. 10 Robinette, D. 1984. Pers. comm. U.S. Soil Conservation Service, Tucson, AZ. 11 Tierra Madra Consultants & Cornett & Associates. 1985. 1985 San Xavier planned community biological survey and assessment. Tierra Madra Consultants & Cornett & Assoc. rept. to U.S. Bureau of Indian Affairs, Phoenix, AZ. 12 Toolin, L.J. 1981. Status report: Tumamoca macdougalii. Rept. to U.S. Fish and Wildl. Serv., Albuquerque, NM. References - 3 (DRAFT) - References Species GLOBE-BERRY, TUMAMOC Species Id ESIS702036 Date 13 MAR 96 References - 4