Endangered Species

UPDATEIncludingEndawered a Reprint Species of theTechnical latest USFWS Bulletin

THE UNIVERSITY OF MICHIGAN March 1988 Vol. 5 No. 5 School of Natural Resources

In this issue: The Longleaf Pine Habitat Loss Landscape of the Threatens Two Mid- Southeast: Almost Gone western Plants and Almost Forgotten Service Decides Not To List Spotted Owl The Longleaf Pine Landscape the Southeast: Almost Gone and Almost Forgotten

by Reed F. Noss

Escaping bleak northern winters, was once an open pine savanna with a ent, and had lower lire frequencies. perhaps for the rest of their lives, hordes dense carpet of wiregrass (Aristida Hammocks usually occurred on slopes, of visitors to the southeast are greeted stricta) and wildflowers became a and on islands, peninsulas, and other by a mass of green as they arrive on the thicket of hardwoods. Today, intensive areas protected from fire (Harper 1911). coastal plain. Many northern visitors forestry (even on public lands such as In the coastal plain, very subtle changes express amazement that so much of this national and state forests) and urban and in elevation (sometimes on the order of region is still "wilderness." Unfortu- agricultural development threaten the a few centimeters) can have profound nately, what the unfamiliar see as ver- few remaining semi-natural stands. effects on plant species composition dant wilderness is actually a severely This article summarizes some unique (e.g., Abraharnson et al. 1984). Slopes degraded landscape. The dense pine ecological aspects of the longleaf pine greater than 15 degrees are very effec- plantations and second-growth hard- landscape, and suggests a strategy to tive firebreaks (Wate et al., in press ). woods that appear natural to the untu- In Florida and small portions of ad- tored eye are artificial assemblages, -- joining states, another upland commu- vastly different in structure and comp nity, "scrub," occurred usually as is- sition from the vast, open piney woods The dense pine plantations lands of often dense sand pine (Pinus that stretched in presettlement times and second-growth hard- clausa) and short-statured oaks from Virginia to Texas. (particularly Quercus myrtifolia, Q. When the first European settlers woods that appear natural geminata, and Q. chapmanii ) in the came to the southeastern coastal plain, to the untutored eye are open sea of longleaf (Myers, in press). they were impressed by the open, park- artificial assemblages, The sharp discontinuities between like character of the "pine barrens* vastly different in structure scrub and longleaf communities, once which dominated this region (Bartram and composition from the thought to reflect soil differences 1791, Williams 1837). William Bar- (Laessle 1958), appear to be largely a tram wrote that "this plain is mostly a vast, open piney woods result of fire history, in part controlled forest of the great long-leafed pine that stretched in presettle- by the vegetation itself (Myers 1985). (Pinus palustris Linn.); the earth cov- ment times from Virginia Whereas Florida longleaf pine commu- ered with grass, interspersed with an to Texas. nities bum lightly at 2 to 5 year intervals infinite variety of herbaceous plants...." under natural conditions, scrub bums At this time, longleaf pine communities catastrophically every 10 to 40 years or covered at least 60 or 70 million acres, longer. Soil differences are determined over 60% of the upland area of the by the vegetation, rather than vice versa restore and maintain viable examples of coastal plain (Wahlenberg 1946, (Kalisz and Stone 1984). As uplands, Croker 1979, Ware et al., in press). The this landscape in perpetuity. both scrub and longleaf communities presettlement distribution of longleaf are readily developable and do not have pine communities is shown in Fig. 1. Ecology the regulatory protection afforded to Today these communities cover less Longleaf pine communities, as the wetlands in Florida (Hart 1987, Noss than 10 million acres (Croker 1979), dominant vegetation of the southeastern 1987a). and almost all of this acreage is second- coastal plain, formed the matrix in Longleaf pine communities can be growth and degraded by logging, which other plant communities were divided into two general types: turpentining, grazing, and disruption of embedded. The longleaf pine landscape sandhills (also called "high pine," in- the natural fire regime. comprised several associations domi- cluding sites with more clayey soils, or Fire was the natural force that main- nated by longleaf pine in the overstory "clayhills") and flatwoods. Christensen tained longleaf pine communities. and wiregrass in the ground layer, with (in press) divides the sandhill commu- When fire frequencies and spread of strips and patches of other community- nity into three subgroups: pine-turkey fires were reduced by deliberate sup- types such as hammocks (mesic hard- oak (Quercus luevis) sandridge, fall- pression and by artificial firebreaks, wood forests) and various wetlands. line sandhill, and Florida sandhill asso- such as roads and developments, the These communities were below long- ciations. Pine-turkey oak sandridge for- community changed radically. What leaf pine on the slope-moisture gradi- ests are found in austere habitats such as

Vo1. 5 No. 5 Endangered Species UPDATE I Endangered Spades UPDATE

A forum for information exchange on endangeredspccies issues

March 1988 Vd. 5 No. 5

Kathryn Kohn ...... Editor Miclwl Soule ...... Faatlry Advisor Yu Man Lee...... Produdion Asrict.

I~~~huctionsfor Authors: Figure 1: Distribution of Longleaf Pine Communities, adapted from The Endangered Species UPDATE Ware et al., in prw welcomes micler related to specia pmtection in r wide mnge of areas in- cludiig but not limited to: iswrch and ridge tops, sand rims of the California pines dominated the mesic flatwoods, mmagement activities for endrngered coastal bays, and relict dune ridges on the most common flatwoods commu- rpecier, themtical rpprorcha to rpecia lower coastal tenaces. These communi- nity, but have been largely replaced by consemtion, and habitat pdonmd pesetve design. Bodr reviews, editaid ties have a characteristic herb layer of the faster-growing slash pine (Pinus el- oanments, and um0un-e of lichens. The fall-line sandhills, a mostly liottii) by foresters. Mesic flatwoods are cunent events and publicrtiona are also continuous band of rolling hills more open than other flatwoods com- welcome. extending hmsouthern North Caro- munities. Saw palmetto (Serenou re- Readers include a broad nnge of lina to Alabama, vary with topography pens) is the most common shrub, but professionals in bxh scientific and and associated soil differences some well-burned stands have virtually policy fields. Articles should be written (Christensen, in press). Florida sandhill no shrub layer. Lower, wet flatwoods, in an easily understandable style for a associations have three phases, co- and those near the coast, are usually knowledgeable audience. Manuscripts should be 7-10 double dominated by turkey oak on the driest dominated by slash pine, with thick spaced typed pager. For fwther sites, bluejack oak or sand post oak patches of saw palmetto, gallberry (Ilex information please contact Kathryn (Quercus incana or Q. margaretta, re- glabra or I. coriacea), fetterbush Kdvn at the number listed below. spectively) on finer-textured, more fer- (Lyonia lucida), wax myrtle (Myrica tile soils, and southern red oak (Quercus cerifera) and other shrubs. The wettest Subscription Infowmtwn: falcata) on calcareous soils, grading flatwoods are generally dominated by The Endangered Species UPDATE is into the southern mixed hardwood for- pond pine (Pinus serotina) with loblolly pblished monthly by the Shod of est (Monk 1968, Christensen, in press). bay (Gordonia lasianthus) and a dense Natural Resources at The University of The clayhill community in the red hill shrub layer. Michigan. Annual rubmiptions arc S 15 each ($18 outside the U.S.). Send cfieclc country of southern Georgia and adja- The two characteristic species of a money ader (made payable to The cent Florida (Harper 1906, Means, in virtually all longleaf pine communities University of Michigan) to: press) is essentially equivalent to the are longleaf pine and wiregrass. Wire- Endangered Species UPDATE bluejack oak-sand post oak phase of grass is a bunch grass that grows ex- School of Natural Resources Florida sandhill. tremely dense under natural conditions 'he University of Michigan In contrast to the various sandhill or of frequent fire. Competition with the Ann Arbor, MI 48109-1 115 high pine associations, flatwoods oc- overlapping roots of wiregrass plants (31 3)763-3243 cupy low, flat sites which can be fairly may be one important mechanism of arid but are poorly-drained in wet peri- hardwood exclusion in longleaf pine Cmc ods. Flatwoods are the most extensive communities (Clewell 198 1). Wire- vegetation type in Florida, occupying grass, like many herbaceous plants of Red-Cockaded Woodpecker some 5Wo of the state (Davis 1967, longleaf communities, normally flow- (Picoidcsborealip) Abrahamson and Hartnett, in press). In ers only after summer fues, and viable Photo by Reed Nosc contrast to sandhills, there is often a seeds are rarely produced; no seedlings well-developed shrub layer. Longleaf have ever been reported in the field.

Vol. 5 No. 5 Endangered Species UPDATE 2 Wiregrass may be a climatic relict, with adequate to account for the dominance tors such as rat snakes. The average age present clumps existing vegetatively for of pyrophytic vegetation (Komarek of cavity ttees is 95+ years today perhaps 5,000 years (Clewell 1981). 1968, Abrahamson et al. 1984). (Jackson et al. 1979), and 103+ years in Hence, once the native wiregrass The high incidence of fues in long- the largest remaining population of this groundcover has been eliminated, such leaf pine communities is related to their species in the Apalachicola National as by site preparation techniques of in- high flammability, a consequence of the Forest, Florida (Hovis and Labisky tensive silviculture or by long-term fire volatile oils and resins in longleaf pine 1985). Because longleaf pines can live suppression, community restoration is needles and wiregrass. Mutch (1970) to 400 years, the average, and probably difficult. suggested that the flammable properties optimal, age of cavity trees was cer- The diversity of species in natural of some plant species have been favored tainly much older in the past, Modem longleaf sites is extremely high. Cle- by natural selection as a means of reduc- forestry operations, using short rota- well (1981) recorded from 59 to 67 ing competition with other species. tions, do not allow the pines to reach an species of herbays plants in stands age and size preferred by red-cockaded sampled by 30 1-m quadrants. Total- Endangered Species woodpeckers. An odd and tragic coinci- number of species in 8 stands ranged Table I* lists some endangered, dence is that this woodpecker, like its from 66 on xeric sites to 133 in flat- threatened, and otherwise "special" fellow community member, wiregrass, woods adjoining grass-sedge savannas. species associated with longleaf pine never has been documented to success- Animal diversity is also high in these communities in Florida. The list is not fully colonize a new site or a fonner site communities. Although a general prin- complete, as we are still learning about from which it has been eliminated. ciple of ecology is that structurally the distribution of many rare species, Another rare species that is declin- complex habitats contain more species, particularly plants and invertebrates. A ing rapidly in the coastal plain is the fox Engstrom et al. (1984) found more more complete list of longleaf-asmi- squirrel (Sciurus niger), again because breeding bird species in longleaf pine ated rare species throughout the range old-growth longleaf pine-turkey oak forests than in more complex beech- of longleaf pine must await further forest is the preferred habitat (Wood magnolia forests. Conversion of natural research. 1985). Sherman's fox squirrel (S. n. sh- longleaf stands to pine plantation re- Many of the plants on this list are emni),a subspecies endemic to north- duces the species richness, diversity, essentially ecotonal, either restricted to central Florida, is particularly threat- density, and biomass of birds (Repen- or achieving their greatest abundance in ened (Ehrhart 1979). An endemic ning and Labisky 1985). the emtones between longleaf pine and scarab beetle, (Ataenius sciwus) ap- Longleaf pine is one of the most fite- other community-types @. Hardin, pears to be restricted to fox squirrel adapted of all plants. It is the only tree in pers. comm.). The herb bog (also called nests (Franz 1982). Habitat destruction this region with seedlings (the so-called pitcher plant bog, seepage slope, or and fire suppression have eliminated "grass-stage," a dense tuft of needles at savanna) community is an ecotone be- fox squirrel populations in many areas, ground-level, arising from a long, thick tween flatwoods and shrub bogs or and small pop~onsisolated on re- tap-root) adapted to survive fire (Bruce other wetlands (Clewell 1981). Fires maining sandhill fragments may suffer 1951). Apical growth begins after 3 to sweeping down from the flatwoods from genetic and demographic prob- 15 years in the grass stage (Croker and prune back encroaching shrubs and lems. Although Sherman's fox squirrel Boyer 1975). Because natural longleaf maintain the open herb bog community, has been a candidate for listing by the stands are open and savanna-like, fires which can sometimes be hundreds of U.S. Fish and Wildlife Service for sev- are frequent but of low intensity, usu- meters wide (Means, in press). This eral years, no action has been taken, and ally burning only the ground layer. Such unique community was characteristic of the state of Florida still has an open fres, which in Florida longleafcommu- the Gulf coastal plain of panhandle season on fox squirrels with a bag limit nities occur naturally at 2 to 5-year in- Florida, Alabama, and Mississippi, but of 2 per day. Based on new evidence in- tervals but at longer intervals such as 3- approximately 97% of its former area dicating severe decline, I have recently 10 years in some other areas (Chris- has been destroyed (Folkerts 1982). (Nov. 21,1987) fded a formal petition tensen, in press), expose bare mineral The red-cockaded woodpecker with Interior Secretary Hodel to list this soil which longleaf pine seeds require in (Picoides borealis), lately the focus of species as Federally Threatened. order to germinate (Chapman 1932). In much controversy concerning national Finally, the gopher tortoise presettlement times, single fires some- forest management in the south, is de- (Gopherus polyphemus) is one of the times burned for weeks and covered pendent upon old-growth longleaf pine most characteristic animals of the areas the size of several counties forests. These woodpeckers excavate sandhills, and probably the most impor- (Means and Grow 1985). Although nesting and roosting cavities in old, liv- tant grazing herbivore of these commu- Indians may have increased fire fre- ing pines infected with red-heart dis- nities (Means, in press). This fully ter- quencies in some areas (Myers and ease. The "resin wells" they peck restrial turtle is a keystone species, with Peroni 1983), the high frequency of around the cavity entrance produce a over 80 species of commensal inverte- lightning strikes in this region is shiny slick of sap which repels preda- brates and vertebrates using its bur- (* Table 1 may be found on Update p.6)

vol. 5 No. 5 Endangered Species UPDA7E 3 rows, which may be 30 feet long and 15 I examples of longleaf communities. A potential solution to the problem feet deep (Eisenberg 1983, Cox et al Furthermore, the typical species-level of restoring and preserving longleaf 1987). Many of these species are strictly approach of specifying minimum vi- pine systems would be to recognize obligate commensals (e.g., scarab able populations, minimum required them as endangered landscape-types beetles of the genera Aephodius, foraging areas (as for the red-cockaded (see Noss 1987b). This designation Copris, and Onthophagus), whereas woodpecker), and other sub-optimal would acknowledge the mosaic of inter- others, such as the Federally Threat- requirements does little to foster true acting communities that occur in natu- ened eastern indigo snake (Drymarchon recovery even at the species level. The ral landscapes, the ecologically critical corais couperi), find optimal habitat extreme manipulations of habitat often gradients and ecotones which are often there. The gopher tortoise is threatened recommended in species recovery plans ignored in traditional vegetation analy- by habitat destruction, fre suppression evoke images of outdoor zoos, not of sis, and the regional variation in species (which reduces herbaceous food wild, natural ecosystems. composition among sites (gamma di- plants), and hunting and poaching for its Recognizing the longleaf pine sys- versity). The goal shifts from preserv- meat. The state of Florida permitted tem as an endangered community ing separate species and communities hunting of gopher tortoises through (Means and Grow 1985) might do more as "living museums" to preserving en- 1987, although public pressure finally to protect higher-order patterns and tire regional landscapes of intact, inter- eliminated the open season in 1988. The processes. But by standard classifica- acting ecosystems. Instead of species U.S .F.W.S. listed western populations tions, longleaf pine is part of many dif- diversity per se, we are interested in of gopher tortoises (extending from the Tombigbee and Mobile Rivers in Ala- bama to southeastern Louisiana) as Threatened in the July 7,1987 Federal Register. A focus on rare species, with their narrow and patchy distributions, is not working to protect intact examples Can We Bring Back the of longleaf communities. Lungleqf Pine Landscape? A rational answer to this question would probably be "no." But effective conservation never has been guided by ferent associations, each with a unique native diversity at all scales (Noss and pure rationality, as the writings of John pattern of species composition and Hanis 1986). Muir, Aldo Leopold, and other great abundance. Moreover, community- Opportunities to restore and main- conservationists vividly attest. The few level evaluation procedures, such as tain viable examples of longleaf pine remaining near-natural stands of long- those applied by heritage programs of landscape-types exist on our large pub- leaf pine are majestic enough to inspire The Nature Conservancy, may not as- lic lands, especially the national for- determined action in any sensitive per- sign longleaf pine communities high ests. Some 62% of known red-cock- son who has walked through their dense value. For example, the Florida Natural aded woodpecker colonies (an indica- wiregrass carpets and heard the wind Areas Inventory (FNAI 1987) ranks tor of old-growth longleaf pine) occur singing through the 8 to 18"-long sandhill as '%4/S3," meaning that it is on national forests (Jackson 1978). needles of the pines. apparently secure globally but some- Unfortunately, these national forests Biologists are beginning to realize what vulnerable at a state scale, and are not being managed in a way that the limitations of the species approach ranks mesic flatwoods as "G5/S5" or will allow recovery of this species, or to biological conservation (e.g., Noss demonstrably secure at both global and of natural landscape-types with the and Harris 1986, Hutto et al. 1987). state scales. The problem here is that characteristic spatial pattern of com- Although Table 1 shows that many ranking criteria do not adequately ac- munities. To many biologists who threatened and endangered species are count for the quality of remaining ex- have reviewed them, the present round associated with longleaf pine communi- amples. Also, although longleaf pine of 50-year forest plans are an ecologi- ties, other major vegetation types in communities are still not as rare as cal disaster (Jackson 1986, Means Florida (e.g., sand pine scrub, tropical many community-types, the fact that 1987, Noss 1987~). hardwood hammock) also have large this once-dominant vegetation of the If we are to bring back the longleaf lists. The large lists relate to a high level coastal plain has been reduced by some pine landscape, large public land areas of endemicity - about 385 piesof 85% (or by much more, perhaps 99.98, must be dedicated to this purpose. plants are endemic to Florida, more than if we consider old-growth examples) These "wilderness recovery areas" any state besides California and Hawaii should be cause for concern. Perhaps must be distributed throughout the (Gentry 1'986). A focus on rare species, "extent of anthropogenic decline" is a natural range of longleaf pine and asso- with their narrow and patchy distribu- more ecologically appropriate criterion ciated communities, in order to repre- tions, is not working to protect intact than rarity. sent the full spectrum of native species (Continued on Update p. 5)

Vd. 5 No. 5 Endangered Species UPDAE 4 composition, abundance patterns, and New Orkms,La. Presses of Florida, Gaiie. inter-population genetic variation. Davis, J.H. 1967. Gmml map of the luaurl McDiad,R.W., ed. 1979. Rare and Endan- Broad habitat canidors between sites vegurion d Florida. Agr. Exp. Su, IFAS, gered Biotr of Florida. VoL 3, Amphiiiand Univ. Fla., Gdnelville. Repiled. University Prerses of Florida, are needed to allow for movement of Man, LM. 1979. Shemum's fox aquimL Gaimdle. animals and dispersal of pmpagules. Pages 17-18 in JN. Layne, ed. Ram adEn- Means, D.B. 1987. Impaus on diversity of the Considerable ecological research will dangered Biota of Florida, Vd. I, Manmals. 1985 Lmd and RuKxaa Manageanent Plan for be necessary to determine optimal Univ. Presses of Floda, Gainesville. National Fonsts m Florida. Unpublished Re- Eisenbea 1.1983 'Ibe gapher mtoisc aa a key pon to ?he Wildemers Society. methods for restoring and managing the stone "peck. Roc 4th Ann. Mtg. Gopher Merns, D.B. In press. Tmesrrial Habitats, and constituent populations, communities, Tortoise Ccunc., Valb,Go.: 1-47. hshwatcr Habitats. Gap. 5 & 6 in S.H. and the landscape as a whole. One Engstrun, T., R.L. Crawford, and W.W. Baker. Wolfe, J.A. Reidenauer, and D.B. Me~na.An immediate priority for longleaf sites is 1984. Breeding bid popuhom in relath to Ecological Characteriwion of the Florida Pan- changing forest structure following fue exclu- handle. USDI Fish & Wid. Sew., Washing- to reestablish the natural regime of sion: a 15-yur study.Wilaa~Bull. %:437-450. ton, D.C. summer fires. Above all, we will need a Florida Natural Areas Inventory 0.1987. Mas, D.B., and G. Grow. 1985. The endan- new attitude of respect and humility in Lists of Special Plants, Special Animals, and gered longleaf pine canmunity. ENFO Report order to repair the damage we have Natural Communities. Tabhassee. 85(4): 1-12. Folkerts, G.W. 1982 The Gulf Coast pitcher Monk, C.D. 1968.Successional and envircm- done. A land ethic (Leopold 1949) is plant bogs. Am. Sci 70: 260-267. mental relationships of the forest vegetation of long overdue. Frsnz, R., ed. 1982. Rare and Endangered Biota nonh central Florida. Am.Midl. Nat.79: 441- of Florida. Vol. 6, Invedmtes. University 457. Presses of Florida, Gamesville. Mutch,R.W.1970. Wil'dlpndfires andecosystems Reed Noss is a wildlife dogist, consultant, Gentry, A.H. 1986. Endemism in tropical versus a hypothesis. Ecology 51: 1046-1051. freelance writer, and lecturer living in tanpcm plant communities. Pages 153-181 Myers, RL. 1985. Fiand the dynamic relation Gainesville, Florida. He has authored many tech- in ME. Soule, ed. Conservation Biology: The ship between Florida srndhill and sand pine nical papers on conservation biology and land- Science of Scarcity and Diversity. Sinauer As- saub vegetation. Bull. Tomy Ba Club 112: scape emlogy topics. sociates, SunderIand, Maas. 241 -252. Hlrper, R.H. 1906. A phytogeog@ical sketch Myen, RL. In press. Scrub and high pineland. In ofthe Altamaha Grit region of the coastal plain R.L Myers and J.J. Ewel, eds. Ecosystems of References: of Georgia. Ann. N.Y. Ad.Sci. 7: 1-4 15. Florida. Academic Press, Orlando, Fla. H~rpa,R.M. 1911 .The relation of climax vege- Myen,RL,andP.A. Peroni. 1983. Approaches Abrahamron, W.G., A.F. Johmm, J.N. Lpyne, tation to islands and peninsulas. Bull. Tomy to determining aboriginal fire use and its im- and P.A. Peroni. 1984. Vegetation of the Arch- Bot Club 38: 515-525. pact on vegetation. ESA Bull. 64: 217-218. bold Biological Station, Florida: An example HaR. 1987. The dark side of protecting Noss, R.F. 1987a. Prol~ingnatural areas in of the Southern Lake Wales Ridge. Fla. Sci. wetlands. Palmetto 6(3): 10-11 fragmented landscapes. Nat Areas J. 7: 2-13. 47: 209-250. Hovis, J.A., and R.F. Latisky. 1985. Vegeutive Noss, R.F. 198%. Fran plant communities to Abrahamson, W.G., and D.C. Hartnett. In press. associations of red-cockrded woodpecker landscaper in conservation inventories: a look Pine flatwdand dry prairies. In RL Myers colonies in Florida. Wildl Soc. Bull. 13: 307- at The Nature Conservancy (USA). Biol. and J.J. Ewel, eds. Ecosystems of Florida. Aca- 3 14. ~XN.41: 11-37. demic Preu, Orlando, Fla. Hum, R.L, S. Reel, and P.B. Ludres. 1987. A Noss, R.F. 1987c. Florida's national forests: cur Baltram, W. 1791. The Tmela of William critical evaluation of the species appach to last chance. ENFO Repon 87(6): 1-14. Bamm. (M. van Doren, ed, 1955). Dover biological conservation. End. Sp. UPDATE Noss, R.F., and L.D. Harris. 1986. Nodes, net- Ribl., NY. 4(12): 1-4. works, and MUMS: preserving diversity at all Bruce, D. 1951. Fi resistance of longleaf pine Jaclwon, J.A. 1978. Analysis of the distribution sales. Environ. Manage. 10: 299-309. seedlings. J. For. 49: 739-740. and population status of the redcodtaded Repenning, R.W., and R.F. Iabisky. 1985. Ef- Chapman, H.H. 1932. Is the longleaf type a woodpecker. Pages 247-252 in R. Tiian, ed. fects of even-age timber management on bird climax? Ecology 13: 328-334. Roc FiNatioaal Symposium on Environ- communities of the longleaf pine forest in Christensen, N.L. In press. The vegetation of the mental Concems m Rights-of-way Manage- north Florida. J. Wildl. Manage. 49: 1088- coastal plain of the southeastern United States. ment. Miss. State Univ., Miss. State. 1098. Chap. 11 in M.G. Barbour and W.D. Billings, Jaw,J.A. 1986. Biopolitics, managment of Wahlenberg. W.G. 1946. Longleaf Pine: Its Use, eds.Vegetotion of North America. Cambridge fedeml lands, and the consavation d the Red- Ecology, Regeneration, Growth, and Manage- Univ. Press, NY. Cockded Woodpecker. Am.Birds, 40: 1162- ment. Charles Lathrop Pa& Forestry Found., Clewell, A.F. 1981. Natural setting and vegeta- 1 168. Washington, D.C. tion of the Florida Panhandle. USACE, Con- Jackson, J.A., M.R. Lermam, and R.G. Hooper. Ward, D.B., ed. 1979. Rare and Endangered tract No. DACW01-77C-0104Mobile, Ala 1979. Tree age and cavity initiation by red- Biota of Florida. Vd. 5. Plants. University Clewell, A.F. 1985. Guide to the Vascular Plants cockaded woodpeckers. J. For. 77: 102-103. Pmses of Florida, Gainesville. of the Florida Panhandle. Florida State Univer- Kale, H.W.. ed. 1979. Rare and Endangered Biota Ware, S., C.C. Frost, and P. Doem. In press. sity Press, Tallahassee. of Florida. Vol. 2, Birds. University Presses of Southern mixed hardwood forest (the former Cox, J. D. Inkley, and R. Kautz 1987. Ecology Florida, Gainesville. longleafpine forest). Chap. 11 in W.H. Martin, and habitat pmtection needs of gopher tortoise Kalisz., P.J., and E.L Stone. 1984. The longleaf ed. Biotic Communities of the Southeastern (Gopherus polyphemus) popdatiolls found on pine islands of the Ocala National Forest, Flor- United States. Vol. 1, Temstrial Communities. lands slated for large-scale development in ido: a soil study. Ecology 65: 1743-1754. John Wiley & Sons, NY. Florida. Nongame Wildlife Program Tech. Komad, E.V. 1968. Lightning and lightning Williams, J.L. 1837. The Tenitmy of Florida. Rep. No. 4. Florida Game and Fresh Water fires as ecological forces. Pm.Tall Timbers A.T. Goodrich, N.Y. Fish Comm., Tallahassee Fla. Fire Ecdogy Conf. 8: 169-197. Wood, G.W. 1985. Sightings of fox squimls in Croker, T.C. 1979. The longleaf pine story. J. For. Laessle, A.M. 1958. The origin and successional South Carolina. For. Bull. No. 46.,Clemson Hist., Jan. 1979: 32-43. relationship of sandhill vegetation and sand- Univ., Georgetown. S.C. Cmker, T.C.,& W.D. Boyer. 1975. Regenerat- pine scrub. Ecol. Monogr. 28: 361-387. Wunderlin, R.P. 1982. Guide to heVascular ing longleaf pine naturally. USDA For. Sew. Layne, J.N., ed. 1979. Rare and Endangered Plants of Central Florida. University of South Res. Pap. SO-105. Southern For. Exp. Sta., Biota of Florida. Vol. 1, Mammals. University Florida, Tampa.

Vd 5 No. 5 Endangered Species UPADE 5 Table 1: Species associated with Florida longleaf pine communities and listed by the U.S. Fish & Wildlife Service, the state of Florida, and/or the Florida Natutal Areas Inventory (FNAI). Longleaf pine communities are not necessarily the primary habitat of every species listed and many species are ecotonal. Data are from Kale (1979), Layne (1979), McDiarrnid (1979), Ward (1979)' Franz (1982), Wunderlin (1982), Clewell (1985), FNAI (1987), and Deborah White, FNAI @en.comm.).

Status Codes:

FWS: E-listed Endangered, T = listed Threatened; C1= candidate for listing, with substantial evidence in support; C2 = candidate for listing, but with substantial evidence of vulnerability and/or threat lacking; 30 = no longer considered for listing because taxon does not meet ESA definition of "species;" X = proven to be more widespread or abundant than previously believed

STATE: E = listed Endangered; T = listed Threatened, SSC = listed Species of Special Concern.

FNAI: G1 (or S 1) critically imperilled globally (or in state); G2 (or S2) imperilled globally (or in state); G3 (or S3) = very rare or local throughout range (or in state); G4 (or S4) = apparently secure globally (or locally) but may be rare in parts of range; G5 (or S5) = demonstrably secure globally (or locally); GH (or SH) = of historical occurence; G#? (or S#?) = tentative rank; G#G# (or S#S#) = range of rank; G#T# = rank of taxonomic subgroup; G#Q# = rank of questionable species; U = no rank assigned due to lack of information.

Species Common MIS State FNAl Name Status Status Status

Plants Agalinis purpurea var.carteri ..Carter's Large Purple Foxglove...... C2 ...... ,..... GSTUQBU Agalinis stenophylla ...... Narrow-leaved False Foxglove ...... C2 ...... GHQ/SH Ascepias viridula ...... Southern h4ilkweed ...... C2 ...... T ...... G2/S2 Aster plumosus ...... Plumose Aster ...... 3B ...... G2Q/S Aster spinulosus ...... Pine-woods Aster ...... C2 ...... T ...... G 1/S 1 Baptisia hirsuta ...... Hairy Wild-Indigo ...... C2 ...... E ...... G2Q/S2 Baptisia simplicifolia ...... Scare-Weed ...... C2 ...... T ...... G2G3D2S3 Calamintha ashei ...... Ashe's Savory ...... C1 ...... T ...... G3/S3 Calamintha dentata...... Toothed Savory ...... C2 ...... G2G3/SZ3 Callirhoepapaver...... Woods Poppy-Mallow ...... T ...... G3G5/S2 Clitoriafragrans...... Pigeon-Wing ...... C 1 ...... T ...... G3/S 3 Conradina glabra ...... Apalachicola Rosemary ...... C2 ...... T ...... Gl/S 1 ConradiM grandif2ora...... LargeFlowered Rosemary ...... C2 ...... G2/S2 Dicerandra frutescens ...... Scrub Mint ...... E ...... E ...... G 1/S 1 Eriogonwnfloridanum...... Scrub Buckwheat ...... C2 ...... T ...... G3Q/S3 Euphorbia telephioides ...... Telephus Spurge ...... C2 ...... G l/S 1 Gentianapennelliana ...... Wiregrass Gentian...... 3C ...... E...... G2?/S2 Gymnopogonfloridanus ...... Florida Beardgrass ...... C2 ...... G2?/S2 Hartwrightiafloridana...... Hartwrig htia ...... C2...... 2...... T ...... G2G3/S2S3 Hedeoma graveolens...... Mock Pennyroyal ...... C 1 ...... E ...... G2/S2 Lecheadivaricata ...... Pine Pinweed...... C2 ...... pin...... G2B2 Liatisprovincialis ...... Godfrey's Blazing-Star ...... C2 ...... E ...... G2B2 Liliurn catesbaei ...... Southern Red Lily ...... T ...... G4G5/S2

Vd. 5 No. 5 Endangered Species UPDA TE 6

Species Common MIS State FNAi Name Status Status Status

Notophthalmusperstriatus...... Striped Newt ...... G3/S3 Rana areolata ...... Gopher Frog ...... SS ...... G5/S 3 Reptiles

Crotalus horridus ...... Canebrake Rattlesnake...... G5/S3 Drymarchon corais couperi .....Eastern Indigo Snake ...... T ...... T ...... G4T3IS3 Eumeces egregius lividus ...... Blue-Tailed Mole Skink ...... T ...... T ...... G4?T'/S2 Gopherus polyphemus ...... Gopher Tortoise...... C2 ...... SSC ...... G262 Lampropeltis calligaster...... Mole Snake...... G562S3 Lampropeltis getulus goini ...... Apalachicola Common Kingsnake ...... G5W2 Neoseps reynoldsi ...... Sand Skink...... T ...... T ...... G262 Pituophis melanoleucas mugitus ...... Florida Pine Snake ...... C2 ...... SSC ...... G5T3?/U Sceloporus woodi ...... Florida Scrub Lizard ...... C2 ...... G3/S3 Stilosoma extenuatwn ...... Short-Tailed Snake...... C2 ...... T ...... G3/S3 Birds

Falco sparveriuspaulus ...... southeastern American Kestrel ...... C2 ...... T...... G5T3T4/S3? Haliaeetus leucocephalus...... Bald Eagle ...... E ...... T ...... G3/S2S3 Picoides borealis ...... Red-CocWed Woodpecker ...... E ...... T ...... G262 Picoides villosus ...... Hairy Woodpecker ...... G5/S3?

Mammals

Felis concolor coryi ...... Florida Panther ...... E ...... E ...... G4TllS 1 Mustela frenata olivacea ...... Southeastern Weasel ...... G5T4/S3? Mustela frenata peninsulae ...... Florida Weasel ...... C2...... G5T3/S3? Podomysfloridanus ...... Florida Mouse ...... C2 ...... SSC ...... G363 Sciurus niger shermani ...... Sherman's Fox Squirrel...... C2 ...... SSC ...... G5T2/S2 Ursus americanusfloridanus..... Florida Black Bear ...... C 2...... G5T3/S3

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EndangeredSpedes UPDATE 8 Vol. 5 No . 5 Book Review

The Last Extinction edited by Les Kaufman and Kenneth Malbry

The last Extinction grew out of a timeas a means of drawing a lessans for can begin the days of stewardship, we public lecture series entitled the current plight of many of theearth's must first end and leave behind the age "Extinction: Saving the Sinking Ark," species. Of these lessons, perhaps the of exploitation. We can only pray that held in Boston, Massachusetts, at the most important is that "when the rate of the passage comes soon and that it is a New England Aquarium during the fall extinction strongly outpaces the evdu- peaceful one. . ." of 1984. The theme around which the tion of new species, as it is doing with a six essays in the book have been organ- vengeance today, the stage is set for ized is that it is both possible and critical extreme changes . . . Destruction and to slow the mass extinction episode that extinction may be rapid, but recovery is is now in progress. In the preface, the painfully slow." editors note that although many people 'Ihe next two chapters are case stud- know that we are rapidly losing the ies. Ghillean Prance, a tropical bota- diversity of life on Earth, they are not nists, outlines the biology and ecology doing anything about it, either because of the amazon rain forest and discusses the idea is only an abstraction or be the nature and causes of the massive cause they think the problem is too vast humancaused devastation that is now for individuals to have any effect. being imposed upon the land Jim Wil- Hence the overall goal of the book is to liams and Ronald Nowak of the United "bring extinction down to earth" and to States office of Endangered Species , provide readers with a better under- complement Prance's chapter with an standing of what extinction is, what it is assessment of species extinction and not, and how it might be abated. To protection in North America Both bring the extinction issue home, several chapters underscore the need to pre- case studies are reviewed from the tem- serve critical habitats as a primary perate zones as well as the tropics. means to protecting the species that In general, the book is geared more depend on them. toward a lay audience than technical ex- Finally, the last chapters wrestle perts. Nevertheless, its clear explana- with future options and conservatian tions of the issues and challenges before needs. Thomas Foose, Cansewation us, as well as its valuable presentation Coordinator for the American Associa- of information ranging from a catalogue tion of Zoological Parks and Aquariums The editors also provide a sampling of of species that have gone extinct in (AAZPA) explores the limits and op- professional and membership organiza- North America since humans first ar- portunities for captive propagation and tions involved in issues relating to spe- rived on the continent to a discussion of reintroduction of endangered species. cies and habitat survival or related is- mass extinctions revealed in the fossil David Ehrenfeld, a biologist and con- sues. The information fathis section is record will be of interest to profession- servationist, takes a long view of the drawn in large part from the Conserva- als and students familiar the literature kind of world we might be living in if tion Directory 1986 and from the report on the topic. we do, or do not make our best efforts at "Grass Roots Conservation of Biologi- The book is divided into three gen- preserving biological diversity. He cal Diversity in the United States," pre- eral sections. The first two chapters, begins by noting that the "future is shy. pared for the Office of Technology written by Les Kaufman and David If you want to catch a glimpse of it, you Assessment. The latter report is avail- Jablonski respectively, set the context have to sneak up on it from behind. So able through the U.S. Government for the book. In the first chapter, the place to start for a look into the Printing Office. the Kaufman examines philosophical, sci- future is past." Ihe papetback version of The Last Extinction entific, and practical stumbling blocks Ehrenfeld then appropriately con- cost $7.95. It is published by: to biological diversity conservation. cludes the Last Extinction by describ- 'Ihe ha Press Jablonski, a paleontologist, reviews ing the situation which we now face in Mass. Institute of Technology mass extinctions throughout geological simple and honest tenns: "Before we Cambridge, Mass 02142

Vol. 5 No. 5 Endangered Species UPDATE 9 Technical NOte~ Produced by 7-h. centw tor cmsennhn &byat ~t~b~dUniversw

Local Extinction, Metapopulations and Endangered Species by Susan Harrison

Although the word "extinction" has tendency for the species to be absent cies now reduced to a single population a negative connotation for most conser- from many apparently suitable habitats. or a few isolated ones by drastic human vationists, population ecologists con- An illusaative example is the Karm modification of their habitat. These sider local extinction to be a normal and blue butterfly (Lycaena melissa species are endangered not only by even frequent process in the population samuelis ), a New York State endan- habitat destruction, but also by the ob- dynamics of certain species. No local gered species. The Karner blue inhabits struction of the essential flow of mi- population of a species is immortal, and the pine bmens, a post-fire succes- grants between the remaining habitats in some organisms, populations tend to sional community. Its larvae feed on by dams and water withdrawals. be rather short-lived. Habitat succes- Lupinus perennis, a fire-dependent How can species with natural sion, occasional catastrophes, plagues shrub. While periodic fire is necessary extinction and recolonization dynamics of biotic enemies (diseases, preclators, to maintain its habitat and its host plant, be protected? A few very general prin- competitors), and periodic imbalances the Karner blue cannot withstand fire in ciples can be advanced which provide in resource bases are among the factors any of its life stages. When a large area a sharp contrast to the current single that predispose some species to local- of pine barrens existed in a fire-succes- population emphasis of conservation ized population crashes. sional mosaic, the Karner blue (despite biology. First, one population is not Frequent local extinction is more being a rather weak flier) survived by enough; indeed, the more populations common among small, short-lived or- colonizing recently burned areas fnrm and habitats that can be conserved, the ganisms such as annual plants, rodents, populations on adjacent unburned habi- longer a metapopulation may be ex- and invertebrates than among long- tat. Habitat reduction and fire suppres- pected to persist. Second, empty habi- lived organisms such as trees and large sion have combined to make coloniza- tats may be as important to a species' mammals. Extinction is also prevalent tion increasingly difficult, and have led persistence as the ones that it occupies among species with specialized and to the buttefly's decline. since the species' distribution at any patchy habitat associations (certain soil Other examples from the conserva- moment is partially a matter of chance. or vegetation types, treefalls, ponds), tion literature illustrate the great variety Third, unobstructed dispersal between which cause local populations to be of metapopulational pattems that exist, habitat patches is critical to the survival isolated from frequent immigration. as well as how these pattems relate to of metapopulations. If natural recoloni- The existence of "natural" extinction conservation problems. The bay check- zation cannot keep pace with hurnan- bears implications for species conserva- erspot butterfly (Euphym editha accelerated extinction, artificial intro- tion strategies. A species can only per- bayed ), a federally listed threatened ductions may be a valuable alternative. sist regionally as long as new popula- species, is confmed to patches of grass- The development of viability analysis tions are founded at a rate which bal- land on serpentine soil. Because of the methods that take all these factors into ances local extinctions. "Metapopula- configuration of its habitat, this butter- account remains a major challenge for tion" refers to a system of conspecific fly forms a metapopulation consisting conservation biologists. populations which exchange immi- of a very large "source" population sur- grants with one another, and metapopu- rounded by numerous small and m- lation dynamics are the changes in dis- sient "satellite" populations. The The Technical Notes page is &- tribution and abundance of a species source population is the key to the re- signed as a forum for thoughts resulting from extinction and coloniza- gion-wide survival of the species. The and information on research is- tion. Metapopulation dynamics are Concho water snake, also federally sues related to endangered spe- most significant in species that are vul- listed, inhabits riffles and rapids along cies protection. Contributions nerable to local extinction and are not the Concho and Colorado River. Local are welcome. Material should be exceptionally good disbursers. In such populations of this snake form a linear sent to: species, populations may appear and "shifting mosaic," soon to be disrupted disappear in what has been called the by a proposed dam along the lengths of Kathy Freas "blinking Light" or 'shifting mosaic" the two rivers. Other species endemic Center for Conservation Biology pattern. The symptoms of this mode of to desert riverine environments, includ- Stanford University persistence are a distribution that is spa- ing certain pupfishes (Cyprinodon Stanford, CA 94305 tially and temporally variable, and a spp.), epitomize metapopulational spe-

Vd. 5 No. 5 Endangered Species UPDATE 10 Bulletin Board

New Newsletter on Monarchs being developed in the United States McKelvey, U.M. Professor of Mathe- & Insect Conservation issues and around the world to address these matical Sciences and Editor of Natural critical problems. The video and ac- Resources Modeling will be the work- A new newsletter produced in Califor- companying printed literature are avail- shop director. For further information nia, will serve to disseminate informa- able through the National Wildlife Fed- and application forms, contact: Biologi- tion about the Monarchs and their Cali- eration for $20. For more information cal Resource Modeling Workshop, fornia wintering colonies, as well as and order forms contact: Noel Gerson, Wildlife Biology/Foresy, University provide information on the latest issues National Wildlife Federation, 14 12 of Montana, Missoula, MT 59812 in insect conservation. The first issue, 16th Street N.W., Washington D.C. (Phone 406-243-5272). which was published in February 1988, 20036; (202) 637-3776. featured articles on migration, protec- WWF Special Report on tion, and management of California Biological Diversity wintering colonies as well as a listing of Workshop on Modeling Meth- recent literahue on the Monarch. Fu- ods In Biological Resource Last December, World Wildlife Fund ture issues will include information on Management published an end-of-year Special Re- the biology and ecology of the Mon- port on Biological Diversity which esti- archs, brief biographies of people N.S.F. is sponsoring a workshop on mates 1,000- 10,000 species are becom- studying and working on the conserva- Modeling Methods in Conservation ing extinct each year due to various tion of butterflies, and summaries of a Biology at the University of Montana in threats, such as tropical deforestation. tagging study. To receive the Danaus, Missoula on July 18 - August 12,1988. The report contains articles on species send a self-addressed stamped (56 The workshop is intended to bring to- conservation strategies, likely cents) envelope to Walt Sakai, Life gether a group of 25 college teachers in extinctions of birds, insects, and plants, Sciences, Santa Monica College, 1900 the mathematical sciences, ecology, and a list of threatened animal species Pico Blvd., Santa Monica, CA 90405. population biology, and biological re- which are dependent on tropical forests. source management fields to participate In the Fall of 1988, WWF will launch a Video on Tropical Rain in a 4-week summer program. The major campaign to raise funds and to Forests program includes lectures, computer stimulate political action for the conser- modeling labs, and field trips, exploring vation of biological diversity. The re- The National Wildlife Federation has scientific and socioeconomic issues in port may be obtained free by writing produced an 18-minute video about conservation biology. N.S .F. funding WWF International Information Divi- tropical forests. The video first focuses will cover participants' living expenses sion, 1196, Gland, Switzerland. on the numerous pressures causing plus a $250 weekly stipend, during the rapid tropical deforestation, and then 4-week session this summer and the 1- Bulletin Board information provided by Jane examines some of the solutions that are week follow-up in June 1989. Robert Villa-Lobos, Smithsmian Institution

Non-Profit Organization U.S. POSTAGE PAID Ann Arbor. MI UPDATE Perm~tNo. 144 School of Natural Resources The University of Michigan Ann Arbor, MI 481 09-1 115