540 Biodiversity and CiJ1fJervarton 10: 541-550.2001. Q 2001 Xluwer Academic Publishers. Printed in the NetherlandJ. Taylor JS, Church KE and Rusch DH (1999) Microhabitat selection by nesting and brood-rearing northern bobwhite in Kansas. Journal of Wildlife Management 63: 686-694 Terborgh 1. Estes lA. Paquet P. Ralls K, Boyd-Heger D. Miller 81 and Noss RF (1999) The role of lOp car. nivores in regulating lerreslrial ecosystems. In: Soule ME and Terborgh (OOs) Continenlal Conservation, Elevational distribution of restricted range forest tree pp 39-64. Island Press, Washington. DC Torger.;en TR and Bull EL (1993) Toe atrophy caused by carpenter ants in Vaux's swiflS. AvocetL'\ 17: taxa in eastern Tanzania •• 209-210 • Torgmen TR and Bull EL (1995) Down logs as habitat for forest-dwelling ants -the primary prey of pileated woodpeckers in northeastern Oregon. Northwest Science 69: 294-303 JON C, LOVETIJ", G. PHILIP CLARKEJ,2, ROBIN MOOREJ Tremblay JP. Gauthier G, Lepage 0 and Desrochers A (1997) Factors affecting nesting success in greater snow geese: effects of habitat and association with snowy owls. Wilson Bullettn 109: 449-461 and GILBERT H, MORREy3 Uchida H (1986) Passerine birds nesting close to the nests of birds of prey. Japanese Journal of Ornithology J Environmenl Department, Universiry of York, York YOlO 5DD, UK; 1PnsenJ address: The Hennirage, 35; 25-32 [in Japanese with English summary] Crewkeme, Somuut TAl8 8ET, UK; 336 CONUlughrRoad, Nunthorpe, MiddJesborough TS708S, UK;

Uela M (1994) Azure-winged magpies. Cyanopica C)'Q1Ul. 'parasitise' nest defense provided by Japanese •Author for correspondence (e- mail: [email protected];fax: +44-/904.431998) lesser sparrowhawks. Accipirer guJaris. Animal Behaviour 48: 871-874 Ueta M (1998) Azure-winged magpies avoid nest predation by nesting near a Japanese lesser spalTow- Received 7 July 1997; accepted in revised form 4 May 2000 hawk's nest. Condor 100: 400-402 Vlilinlinen VM (2000) Predation risk associate1 with nesting in gull colonies by two Ayrhya species: observalions and an experimental test. Jouma.! of Avian Biology 31: 31-35 Vermeer K (I %8) Ecological aspects of ducks nesting in high densities among lands. Wilson Bulletin 80: Abstract. The forests of eastern Tanzania are a globally important biodiven>ity hotspot. In this study 361 7B-B3 eastern Tanzanian restricted range forest tree taxa were assessed. Of these taxa, 223 occurred in the Eastern Vermeer K (1970) Breeding biology of California and ring.billed gulls: a study of ecological adaptation to Are, 150 in Coastal forests. 17 in Northern forests and 21 in the Lake Nyasa forests. The majority of the the inland habitat. Canadian Wildlife Service Report Series 12 taxa had restricted e1evational ranges with 76.3% occurring in no more than two 200 m e1evalional bands Werner HE (1992) Individual behavior and higher-order species interactions. American Naturalist 140: out of a total potential elevation range of 3000 m. The majority of the taxa occupied a small area in the SS-S32 eastern T~nzanian f~rests, with 201 taxa being only found at a few sites. In determining priority areas for Werner EE and Anholt BR (1996) Predator-induced behavioral indirect effects: consequences to competi. conservation. selection of taxon definitions can have imponant effects. For example. tree size varies with tive inleractions in anurnn larvae. Ecology 77: 157-169 elevation, so if only large trees are used then site selection will be biased towards particular areas, Wheelwright NT, Lawler JJ and Weinstein JH (1997) Nest.site selection in Savannah sparrows: using gulls as scarecrows? Animal Behaviour 53: 197-208 Key words: elevation range, forests, rarity, Tanzania, trees Whittaker 18 (l991) Effects of ants on temperate woodland trees. In: Huxley CR and CUller OF (eds) , Ant-ph:m! Interactions. pp 67-79. O:tford Univeristy Press. Oxford . Wiklund CG (1982) Fieldfare (Turdus pi/oris) breeding success in relation to colony size. nest position and Introduction , association with merlins (Fulco columburius). Behavioml Ecology and Sociobiology II: 165-172 Wilcove OS (1985) Nest predation in forest tracts and the decline of migralOry songbirds. Ecology 66: 1211-1214 Working on the assumption that resources for conservation are scarce, large scale Witmer GW. Bucknall JL, Fritts TH and Moreno DG (1996) Predator management to protect endangered compilation, of distribution patterns have, been used to highlight global priori,¥, ar- avian species. Transactions of the Nonh American Wildlife and Natural Resources Conference 61: 102- JOB eas on the basiS of thetr bIOlogical diverSity (e.g. Mittenneier et al. 2000. Williams WOOllon JT (l993) Indirect effects and habitat use in an intertidal community: interaction chains and and Gastnn 1994; Williams et a!. 1996), Whilst global conservation priOri:ies can 6e inleracLion modifications. American Naturalist 141: 71-89 identified by the use of such spatially relatively coarse data, conservation is ultimately Wunderle JM and Pollock KH (198:5) The bananaquit-wasp nesting association and a random choice model. Ornithological Monographs 36: :595-603 effected by)ocal implementation. which requires a fine grained analysis pinpointing Young BE, Kaspan M and Manin TE (1990) Species-specific nest site selection by birds in anHlcacia key areas or habitat types_ trees. BiotrOpica 22: 31o-3l5 The forests on the mountains and coastal plains of eastern Tanzania are recognised as one of the world's mnst important biodiversity hotspnts (Myer! 1990; Myers et al, 2000), The forests contain a remarkable number of pl,ant species of restricted geo- graphical distribution (Burgess et a!. 1998; Lovetl 1988; Lovett et a!. 2000; Polhill 1968) that appear to be both recently evolved neo-endemics and ancient relictudh •••• palaeo~endemics (Lovett and Friis 1996). Such geographically rare species may also be locally rare, for example they may have a limited elevational range and low abun- dance (Gaston 1994), a fact that will affect management for conservation within the biodiversity hotspot. ,

542 543

Once global conservation priorities have been determined, knowledge of the local ecology and distribution of geographical rare species are needed 10 formulate effec- tive management plans., For example, the Tanzanian forests occur in an area where there are sharp topographic. climatic and edaphic gradients that may affect local spe- cies distributions. They are highly fragmented, with many small patches covering • Fo •• " •••• ..". only a few hectares that can be rich in restricted range taxa, Moreover, the Tanzanian forests are still poorly known biologically with many new species still be- ing discovered. Consequently, knowledge of general ecological patterns of restricted range taxa may help to formulate management plans that are relevant to as yet undis- covered species. In this study we compile information on as many restricted range trec taxa as possible. dividing them into categories based 00 taxonomic rank, geographical distribution, elevational dislributioo and habit. The aim is to ascertain if taxa defined as rare in the sense of having restricted geographical ranges at a continental scale, also have reslricted local geographical and elevational distributions. The results are discussed in the context of specific management strategies.

Methods

The list of restricted range forest taxa in eastern Tanzania was initially compiled from information contained in the Flora of Tropical East Africa (FTEA) (Turrill et al. o tOO 200 1952-), for taxa defined as a 'tree' in growth habit. This covered from a few Idlomeltat; metres to over 60 m tall. Our own field observations were used to supplement this information. The eastern Tanzanian list was originally complied for the World List of Threatened Trees (Oldfield et al. 1998). The list was hroken down by a number f'gure J. Location of the Northern, Eastern Arc. Lake Nyasa and Coastal forests in Tanzania (after Lovett of variables for analysis: height, taxonomic rank, habit, distribution and habitat. 1990). Eastern Tanzanian forests were defined as closed canopy evergreen or semi-evergreen vegetation formations east of the arid corridor (sensu Wergcr 1978) and divided into but whi~h we considered t~ be forest plants were also included. Taxa were dividect.fto Eastern Arc forests, Coastal forests, Northern forests and Lake Nyasa forests (sensu three height classes aceordmg to the tallest record of the tree cited in the FfEA or from Lovett 1990; Figure I). The Eastern Arc forests accuron a range of ancient crystalline our own field observations: <10 m tall, 10-20 m tall and >20 m tall. Four levels o'f mountains that run from the Taita hills, Kenya to the Makambako Gap in the southern taxonomic rank were considered: , species, subspecies and variety. 1\\'0 growth Udzungwa mountains, Tanzania. The Coastal forests (in Tanzania) are predominatelyon habits were recognised: 'shrubby trees' (i.e. trees which have also been recorded as the sedimentary rocks and derived formations between the Eastern Arc and the Indian shrubs) and trees that have only been recorded as trees. Four patterns of restricted Ocean coastline. The Northern forests are on the northern Tanzanian volcanics and "range distribution were used: restricted to a few specific localities in Tanzania (F); associated mountains on the rift valley. The Lake Nyasa forests are in the high rainfall occurring in several to many localities in Tanzania (T); Oecumng i~Tanzania and any area of northern Lake Nyasa south of the Makambako Gap and are predominately one of the neighbouring countries (T + 1); and a widely 4isjunct population of a taxon on Pliocene volcanics or volcanic soils overlying crystalline basement rocks. The occurring elsewhere (D). The elevational range of each taxon in 200 m elevation bands maximum elevation of each forest type is: Eastern Arc forests 2400 m. Coastal forests over a total elevational range of 0 to 3000 m (IS bands) was also recorded. ., ••• 1100 m (but usually much lower), Northern forests 3000 m, and Lake Nyasa forests P:~blems encountered included the desc"ription of a number of FfEA collecting 2850 m. Some species occurring in Coastal forests also occur in Eastern Arc forests, and localitIes as woodland, even where field observations suggest that it meets the criteria so have an elevation range above that of Coastal forests as a whole. Restricted range in standard definitions for forest (e.g. that of White 1983). Fo~ example, vegetation trees that occur in vegetation derived from forest (e.g. thicket and coastal bushland), on the shores "of Lake Lutamba in SE Tanzania, is described in collection notes as l

544 545 wa/d by the German botanist Schlieben. This is translated in the FTEA as woodland, Table J. Frequency of restricted range taxa of differ- perhaps because 'wald' is more commonly used for forest than 'forst' in German. ent laxonomic ranks in each of the four geographical categories and two habit categories. Field observations revealed that most, if nal all, of the restricted range species in the area would have been collected in forest. Thus the 16 endemic species known only EA C N LN T ST from the location cited as 'Lake Lutamba' are forest dependant rather than occurring • • Genus 15 6 0 t lJ 6 in woodland as cited in the FTEA (Clarke 1995). Species 153 105 II S 136 108 .. Problems of definition also arose with laXa of restricted range. which have been Subspecies 26 18 3 4 12 31 Variety 29 2t 3 S 12 42 divided into subspecific entities. For example the genus Ophrype/alum is monotypic Total 223 ISO 17 21 IS? and restricted to SE Kenya and eastern Tanzania, and so presents the problem as 173 to whether it should be regarded as two endemic subspecies or an endemic genus. EA - Eastern Arc; C - Coastal; N - Nonhero; LN - Similar examples include the species Polyscias s/uhlmannii, which is only known Lake Nyasa; T - tree; ST - shrubby tree. from NE Tanzania and SE Kenya, but is represented by two endemic varieties, and and 80% in no more than two bands. For three out of four different distribution cate- Drypetes usa mba rica, which includes five varieties within its restricted range in SE gories recognised (F, T, D) there is no relationship between frequency and elevation Kenya and NE Tanzania. Taxa that have two or more disjunct populations, especially in elevation bands below 2000 rn (Table 2). Tanzania and anyone of the neighbouring where these populations are widely separated (e.g. between west-central and eastern countries category show a relationship with elevation, mainly because the frequencies Africa) are especially prone to the effect of differing taxonomic biases (Lovett 1993). an: high for the first three elevation classes. In tenns of percentages of taxa in each The disjunct populations might be considered as disjunct species by some taxono- elevation class, the percentage of taxa that occurs in a few specific localities in Tan- mists but not others, for example: Pterocarpus mildbraedii, Mesogyne insignis and zania (F) increases with elevation, whereas the T + 1 class percentages decrease with Mammea a/rieana (Lovett 1993). These oUlliers may be morphologically (pheno- elevation. typically) similar to other taxa in the disjunction, but the possibility exists that they The relationship between distribution category and area category is presented may contain differing genotypes due to their long isolation, and hence have different in Table 3. A X2 test was used to see if the two classifications were independent. ecological requirements. For the purposes of this study, the lowest taxonomic rank was used as an operational data unit, and currently accepted was followed. . Although the list of restricted range taxa is fairly comprehensive, it was not possible . to obtain information on some taxa, for example the list of restricted range taxa in the Sterculiaceae is not complete, particularly for the genus Cola. A list of taxa analysed is available from the first author by request.

Results -o" A total of 361 restricted range taxa in 41 families were recorded. The restricted range U taxa were mostly at the taxonomic level of species (244) with 19 genera, 43 subspe- cies and 55 varieties. The frequency of each category of restricted range taxon in the four different geographical categories and two habit categories is given in Table 1. Ten families accounted for 85% of the restricted range taxa, with the most being in the (132) followed by Fabaceae (64), Euphorbiaceae (31), Annonaceae (24), Ebenaceae(12), Sapindaceae (12), Sapotaceae (11), Rutaceae (9), Verbenaceae •••••••• (7) and Clusiaceae (6). 2 3 45678 9 10 The majority of taxa had restricted elevational ranges, with 76.3% occurring in Number of 200 m bands no more than twO 200 m elevational bands. No taxa occurred in more than 10 bands Figure 2. Histogram of the frequency distribution of the nwnber of elevational bands occupied by the 361 (Figure 2). If the taxa are placed in 400 m bands, then 49% occur in only one band eastern Tanzanian restricted range tree l.a.la. .. l

546 547

Tub/e 2. Frequencies of reslricted runge taXa Tab/I! 4. Size of restricted range tree taxa in each of in four claues of geographical distribution in lhe distribution and area categories. each 200 m elevalional band. Tree height (m) Elevation (m) F T T+I 0 <10,. 10-20 >20 Total 0-200 31 16 62 3 200-400 40 19 '9 , Few sites 119 '2 26 197 400-600 48 13 44 , Tanzania only 25 12 12 49 600-800 36 II 26 , Tanzania + t 52 32 18 102 800-1000 44 15 24 6 Disjunct 4 5 4 13 1000-1200 25 19 18 4 Total 200 101 60 361 1200-1400 29 21 17 3 1400-1600 36 17 20 3 Eastern Arc III 63 49 223 1600-1800 36 14 21 3 Coastal 91 41 18 150 1800-2000 40 12 18 3 Nonhem 8 7 2 17 2000-2200 23 8 10 2 Lake Nyasa 14 3 4 21 2200-2400 12 , 7 2 Total 224 114 73 411 2400-2600 , I 3 0 2600-2800 I I I 0 2800-3000 I I 0 0 There is no relationship between distribution category and size of trees (Table 4, F - restricted tu a few specific localities in X 2 = 9-13, P = 0.17). This relationship remains not significant if the low frequen- Tam-.ania; T - occurring in several to many lo- cies of the disjunct class are combined with Tanzania and anyone of the neighbouring calities in T.anzania; T + 1 - occulTing in Tanza- 2 nia and anyone of the neighbouring countries; countries class (X = 6_88, P = 0.14)_ Similarly, there is no relationship between o - disjunct population occurring elsewhere. geographical area category and tree height, indicating that the mixture of tree heights is not significantly different for different areas (Table 3, Northern and Lake Nyasa 1able J. Frequencies of four different categories of re- area are combined to avoid small frequencies, X2 = 7.48, P = 0.11). There is a stricted range taxa in each geographical area recognised. significant decline in tree height with elevation at the 5% level (T~~le5, X2 b 18.47, The totals for the dislributiOli classes do nol agree with P 0.048), though this relalionship is not simple. for example there are substantially those in the main [able because some lrees occur in Cwo = of the area categories. grealer than expected frequency trees of >20 tall in the 800-1200 m band_

F T T+ I 0 Total

Eastern Arc 130 37 45 II 223 Discussion Coastal '7 17 72 4 150 Northern 6 2 8 1 17 Lake Nyasa 8 4 9 0 21 The results show three clear patterns. Firstly, the majority of the restricted range tree Total 201 60 134 16 411 taxa occur in the Eastern Arc and Coastal forests; secondly the taxa occur throughout

FQr footnotes see Table 2. • Table 5. Size of restricted range tree taxa in 400 m elevation bands. Expecled frequencies given by a Because of low frequencies in the Northern and Lake Nyasa areas, these categories X 2 test are given in brackets. i were combined_ The classifications are not independent (X2 = 35.07, P » 0.0001). Elevation band (m) The observed Eastern Arc frequencies for those species restricted to a few specitic calegories (F) and Tanzania only (T) are larger than expected, while those for the Tree height (m) 0-400 400-8.00 800-1200 1200-1600 1600-2000 >2000 Tor: ••. Coast are smaller than expected. The opposite is true for those taxa occurring in 20 20 (27) 24 (22) 31 (19) 18(17) 13(16) 4 (10) 110 -. areas, whereas those in the Coastal forests also occur in an adjacent country. Total 154 125 110 100 92 '7 608

------..

548 549

the elevations] range of the forests; and thirdly most of the taxa have local distri- W. Mziray var. hanangensis (Rutaceae) cited in FfEA may refer to a collection that butions. Each of these points is discussed below in the context of management for has subsequently been assigned to Vepris barenensis (M.G. Gilbert) W. Mziray. If conservation. this is the case it win only be found in four 200 m bands. The monotypic northern That most of the restricted range taxa occur in the Eastern Arc and Coastal forests Lake Nyasa endemic Temnocalyx nodulus,.characteristically occurs in upper montane (Tables I and 3), with fewer in the Lake Nyasa and Nonhern foreslSsuppons pre"ious forest and so the lower elevation records cited in flEA arc unusual; and Mildbraedia observations (Burgess et aL 1998; Lovett 1988) and confirms the boundaries of the earpinifolia (Pax) Hutch. var. earpinifalia (Euphorbiaceae) is a forest edge plant with eastern African biodiversity hotspot (Myers et al. 2000). Lower elevation forests tend wide ecological tolerances. to nave more restricted range taxa in Tanzania and anyone of the neighbouring coun- In summary, the forests of eastern Tanzania are exceptionally rich in tree taxa tries class. This is because the endemic rich Coastal forests extend from Tanzania of restricted distribution, with most of the restricted range taxa being concenlrated into south eastern Kenya, whereas the Eastern Arc forests are only represented in in the Eastern Arc and Coastal forests. In addition to being of restricted range on Kenya by remanent outliers on the Taila and Shimba Hills. There are however some a continental scale, many species also occur only in a few sites and have a narrow notable narrow range endemics in the other areas, for example Temnocalyx nodulus elevational range. In order to conserve as many taxa as possible, a full elevational Robyns (Rubiaceae) is in a monotypic genus and is only known from the Poroto and range of forest needs to be protected together with the many small patches of forest Rungwe mountains of the Lake Nyasa forests. In addition the Lake Nyasa forests which contain rare plants, which may have an area of occupancy as small as a few have 21 restricted range taxa and the Northern forests have 17. There is a danger that hectares. This latter point is quite important because empirical evidence from else- such taxa may be overlooked because of the exceptional importance of the adjacent where shows that tropical forest fragmentation results in a loss of species (Turner Eastern Arc and Coastal forests, but which, if they occurred in a less endemic rich 1996; Turner et aL 1996). In Tanzania, small forest fragments may be an imponant country, might be a focal point of conservation attention. means of survival for species lhat occur only in a few sites. A factor that does cause Although the mixture of tree heights is similar in the different areas (Table 3), in loss of narrow range endemics is disturbance, which has been noted in both Coastal the contex.tof prioritising areas for conservation on the basis of numbers of restricted forest (Mwasumbi et aL 1994) and Eastern Arc forest (Lovett 1999). This indicates range taxa, selection of definitions can have an important effect. For example, if a that restricted range taxa are not tolerant of a wide range of ecological conditions and lower height limit of 20 rn is used, then Eastern Arc forests become proportionally that management for conservation in the eastern Tanzanian forests should minimise much more important than Coastal forests, and subrnontane forests in the elevation disruptive interventions such as forest product extraction. I band of 800-1200 m become the most imponant forest type (Tables 4 and 5). Like- .. wise, in terms of genetic variation, most of the restricted range genera and species are trees, but most of the subspecies and varieties are shrubby trees (Table 1). In Acknowledgements practical management terms, if exploitation is a threat, then by using large size limits for the definition of a tree, it becomes biased towards those used in commercial timber The World Conservation Monitoring Centre supponed the initial data gathering exploitation and ignores the fact.that the preferred size of buHding wood in Tanzania through the Conservation and Sustainable Management of Trees project fundedv the is small diameter poles for traditional housing (Hall and Rodgers 1986), and that most Government of the Netherlands and managed by Sara Oldfield and Charlotte Lusty. Y'e domestic fuel is derived from small diameter trees. gratefully acknowledge comments from Neil Burgess on an earlier draft of this paper. Restricted range taxa occur throughout the elevational range of the forests (Table 2), indicating that the full range of altitudinal forest types would need to be References conserved in order to protect as many restricted range taxa as possible. Numbers of

taxa fall with elevations over 2000 m, a pattern which coincides with reduced forest Bursess NO, Clarke GP and Rodgers WA (1998) Coasull forests of eastern Africa: status. endemism pat- area and the upper tree line. A management strategy of conserving a full altitudinal terns and their potential causes. Biological Journal of me Linncan Society 64:.337-367 range of forest is made an the more important by the fact that most of the restricted Clarke GP (1995) Status Reports for 6 Coastal Forests in Lindi Region. Tanzania. Frontier-Tanzania Tech. nical Report No. 18. The Society for Environmental Exploration and the University of Dar es Salaam. range taxa occur in relatively few elevational bands (Figure 2) and at few sites in LondonIDar es Salaam """'"••••. the forests (Tables 2 and 3). This observation fits with the general pattern that taxa Gaston KJ (1994) Rarity. Chapman & Hall. London which are rare in terms of being of restricted distribution on a continental scale are Hall JB and Rodgers WA (1986) Pole cutting pressure ill Tanzanian forests. Journal of Forest Ecology and Management 14: 133-140 also locally rare (Gaston 1994). Only three species occur in nine or ten 200 m eleva- Lovett JC and Friis I (1996) Some patterns of endemism in the tropical north east and eastern African I L tional bands. Of these, the lower elevation record of Vepris hanangensis (Kokwaro) woody flora. In; van der Mat.sen WG. van der Burgt XM and van Medenbach de Rooy 1M (cds) The l, ,

, '

550 Biodiversiryand Conservation 10: 551-566,2001. @ 200 I K/uwer Academic Publishers. Primed in the Nerherlands. Biodiversity of African Plants. Proceedings XIVlh AETFATCongress 22-27 August 1994, Wageningen. The Netherlands. pp 582-601. Kluwer Academic Publishers, Dordrecht. The Netherlands Lovett JC (1988) Endemism and affinities of the Tanzanian montane forest ftora. Monographs in System- atic Botany from the Missouri Botanical Garden 25: 591-598 Non-apoid flower-visiting fauna of Everglades Lovett JC (1990) Classification and status of the moist forests of Tanzania. Mitteilungen aus dem Institut rur Allgemeine BOIani" Hamburg 238: 287-300 National Park, Florida •• Lovett JC (1993) Eastern Arc moist forest flora. In: Lovett JC and Wasser SK (eds) Biogeography and Ecology of the Rain Forests of Eastern Africa, pp 33-55. Cambridge University Press, Cambridge .Lovell JC (1999) Tanzanian forest tree plol diversity and devation. Journal of Tropical Ecology 15: 689- JOHN B. PASCARELLA"', K.D. WADDINGTON and P.R. NEAL 694 Department of Biology, University o/Miami, P.O. Box 149//8. Coral Gables. FL 33124-042/. USA; Lovett le. Rudd S, Taplin J and Fridmodt-M~ller C (200) Panems of plant diversity in Africa south of the J Present address: Department 0/ Biology, Valdosta State University, Valdosta, GA 3/698, USA; Sahara and their implications for conservation management. Biodiversily and Conservation 9: 37-46 •AUlhor jorcorresponderu:e (e-mail: [email protected] ..jOJ:: +/-9/2-333-7389) Mittermeier RA, Myers N and MiUermeier CG (2000) HotspOis. Earth's Biologically Richest and Most Endangered Terrestrial Ecoregions. Cemex, Mexico Mwasumbi LB. Burgess ND and Clarke OP (1994) The Vegetation of Pande and Kiono Monsoon Forests, Received 25 August 1999: accepted in revised form g May 2000 Tanzania. Vegetatio 113: 71-81 Myers N (1990) The biological challenge extended: extended hot-spots analysis. Environmentalist 10: 243-256 Abstract. The non-apaid flower-visiting fauna of Everglades National Park (ENP), Rorida. was surveyed Myers N, MiUermeier RA, Miuermeier CG, da Fonseca GAB and Kent J (2000) Biodiversity hotspots f(lf conservation priorities. Nature 403: 853-858 during 1995--t991 as part of a community pollinator survey. One hundred and lhirty one sampling mps were made to four areas of Everglades National Park (Shark Valley. Chekika. Long Pine Key (LPK). and Oldfield S, Lusty C and MacKinven A (1998) World List of Threatened Trees. World Conservation Press, Aamingo). Species-monlh curves indicate that lhe sampling effort resulted in capture of most of the Cambridge flower-visiting animal species in the. park. A total of 143 insects and I bird species were recorded. Diptern Polhill RM (1968) Tanzania. In: Hedberg I and Hedberg 0 (eds) Conservation of Vegetation in Africa were the most diverse group (55 spp.), followed by Lepidoptera (42 spp.) and non-apoid Hymenoptera (34 South of lhe Sahara, pp 166-178. Acta Phytogeographica Suecica. Stockholm spp.). The majority of species were rare (56% of species were found on fewer man five trips). The highest Turner 1M (1996) Species loss in fragments of tropical rain forest: u review of the evidence. Journal of Applied Ecology 33: 200-209 diversity of species was found from January to May during the peak flowering period in some plant communities. The greatest total diversity was found in Long Pine Key and Shark Valley had the lowest Turner 1M. Chua KS, Ong JSY, Soong BC and Tan HYW (1996) A century of plant species loss from an diversity. Chekika and Aamingo were intennediate in diversity. AnimaJs visited 178 plant species ••....26% of isolated fragment of lowland tropical rain forest. Conservation Biology 10: 1229-1244 the potentially animal pollinated Angiospenn diversity of the park. 1\venty-five species of plants had only Turrill WB. Milne-Redhead E. Polhill RM, Beenlje H (eds) (1952-) The Aora of Tropical East Africa. non-apoid flower visitors; the majority of these species had only visits by Lepidoptera. Potedtially important A.A. Balkema, Rouerdam pollinator species include members of the Syrphidae. Coleoptera. and LepidD~ However, many of the Werger MJA (1978) The KaITOO-Namib Region. In: Werger MJA (ed) Biogeography and Ecology of South- flower-visiting species may not be effective pollinators. This study will be useful for designing sampling ern Africa, pp 231-299. Dr. W. Junk Publishers, The Hague protocols for including invertebrates in assessments of ecological restoration undClWaY in lhe Everglades White F (1983) The Vegetation of Africa. A Descriptive Memoir to Accompany the UNESCOI ecosystem and for more detailed studies of the importance of non-apaid ftower-visitors as effective pollinators. UNSO/AETFAT Vegetation Map of Africa. UNESCO, Paris Williams PH and Gaston KJ (1994) Measuring more of biodiversity: can higher-taxon richness predict wholesale species richness? Biological Conservervation 67: 211-217 Key words: Biodiversity. Everglades National PaIt, Ronda, pollinators, Sytphidae Williams PH, Gibbons D, Margules C, Rebelo A. Humphries C and Pressey R (1996) A comparison of richness hotspots, rarity hoispots and complementary areas for conserving diversity of British birds. Conservation Biology 10: 155-174 Introduction

Everglades National Park (ENP) comprises a large area (1.5 million acres) of sub•. tropical wilderness in southern Florida, USA (Davis and Ogden 1994). In southeast. ern Florida, ENP contains the largest remaini[]g patch of upland pine rockland, as well as mangrove, hardwood subtropical forest, cbasllIl prairie, and various wetland habitats. Although many vertebrate species and apimal communities of ENP have been well studied (Robertson and Frederick 1994), we know little about the im. portance of invertebrates in community (!ynamics a[]d ecosystem functions (Sax.d'8l' et al. 1990). Although the flora is well known (Avery and Loope 1996), there have '- been few studies of plant-pollinator interactions in ENP. Invertebrates play imp'onant roles as pollinators of native plants and may also be important in the reproduc- _ live success of invasive non-indigenous plant species, several of ~hich threaten the