ANT 154BN–14 Primate community structure

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Unformatted text preview: ANT 154BN Lecture #14: Primate community structure 22 Feb 2011 Primate communities 1. Community ecology 2. Measuring diversity 3. Primate communities: patterns Primate communities >1. Community ecology 2. Measuring diversity 3. Primate communities: patterns Ecological study at different scales Organism Group Population %"'0 Community Ecosystem Landscape Community All the organisms that inhabit a particular area An assemblage of populations of different species living close enough for potential interaction Interspecific interactions relationships among species within a community communities have collective and emergent properties Interspecific interactions competition predation herbivory symbiosis (parasitism, mutualism, commensalism) disease Interspecific interactions –/– +/– competition predation herbivory parasitism disease +/+ +/0 mutualism commensalism Community attributes Species composition what species are found there Species richness how many species are found there Species diversity number and relative abundance of species Primate communities 1. Community ecology >2. Measuring diversity 3. Primate communities: patterns Measuring diversity at different scales Alpha: diversity within a particular area or ecosystem species richness, i.e., number of species Whittaker 1972 Measuring diversity at different scales Beta: changes in species diversity between ecosystems or along environmental gradients various indices, e.g., Sorenen’s index S1, S2 = total # of species in communities 1, 2 C = number of species common to both communities Ranges from 1 = exactly the same species in each community to 0 = no overlap in species Whittaker 1972 Measuring diversity at different scales Gamma: total species over a large area or region various diversity indices calculated over a large scale Whittaker 1972 Measuring diversity at different scales Measuring diversity at different scales 5 2 Measuring diversity at different scales 6 2 Measuring diversity at different scales 10 3 Measuring diversity at different scales Diversity indices various algebraic indices (e.g., Simpson’s D, Shannon’s H) depend both on species richness and evenness less diverse more diverse for a given richness, diversity increases with evenness Diversity indices various algebraic indices (e.g., Simpson’s D, Shannon’s H) depend both on species richness and evenness less diverse more diverse for a given evenness, diversity increases with richness Diversity indices rank-abundance diagrams plot species abundance vs. abundance rank Rank-abundance diagrams NATURE A R T I C L1E S | Vol 450 | November 2007 NATURE | Vol 450 | A R T I C L E2007 1 November S a 15 10 5 Number of species 0 0 c 25 20 15 10 5 0 40 40 〈S 〉= 113.6 〈S 〉= 173.9 1 = = 〈J 〉 2,780.7 〈J 〉 8,342.0 ^ rðmÞdm~ expð{m=rÞme{1 dm 25 30 30 θ = 25.1 θ = 29.3 CðeÞde theory123,10. xOur.979 simple analytical approximations of0the exact sam=0 x = .992 20 Υ = yield Υ 0 the where d20 x/(12x) to match the first moment of the 5 20 pling theory82140.087 virtually indistinguishable fits=to.090 data, and 2 = 0.992 R R2 = 0.987 15 continuous distributions and: m 1 x 5 1 (for the metacommunity x1n 5 1). 30 where SM is the number of species in the metacommunity. Here Estimates of the ecological parameters obtained from fitting equation (5) to the RSA data of six (m)dm is the continuous probability distribution of the mean 0 ^ 0 r 0 forests. S, the number of observed s10 0 J, the local community 6 pecies; size; h, t8 biodiversity he 0 2 6 2 4 6 8 2 4 10 populations of1the 2 3 4 the metacommunity and has 4 form of species in 5 6 7 0 the 2 ~ parameter; m, the per capita immigration rate (m 5 Jm/(1 2 m)); and x, the per capita birth-toc Fisher log-series (in a singularity-free description25,26): d the familiar ~ death-rate ratio. 15 plots the above expression for m into equation plots obtains Substituting d 5 (6), one Number of species 〈S 〉= 36.9 Plot S = 〈J 〉 231.7 θ = 16.6 BCI, Panama x = 0.877225 Yasuni, Ecuador = 0 821 Υ R Pasoh, Malaysia2 = 0.995 678 Korup, Cameroon 308 Lambir, Malaysia 1,004 Sinharaja, Sri Lanka 167 45 180 species abundance of tropical forests plots b Table 1 | Relativeplots J 20 21,457 15 17,546 26,554 10 24,591 33,175 16,5 936 h 48 212 206 54 305 28 〈S 〉= 72.8 ~ m x J= 〈m〉 926.9 θ = 22.0 2,122 0.090.95891 0. x= 9,448 0.35 0 0.65 Υ= R2 = 0.93 1,999 0.07 0.994 18,551 0.43 0.57 3,281 0.09 0.91 15,633 0.48 0.52 r2 0.97 0.98 0.98 0.94 0.99 0.94 Box 2 | Relative species abundance of tropical forests a b 40 The mean number of species with n individuals in a community can be 40 written as: 30 20 10 hQn i~ SM X k~1 Pn,k ~SM ? 30 ð 0 d20^ðmÞPn,m mr 10 ð18Þ 8 coral reef communities tropical forest communities ð19Þ discrete and 0 0 where X is the total population JM ? Comparison forests (JM ~ xn ~ l ng 1{ CI t Then, on substituting equations (19) Figure~1 |hSamphix=ðof BxÞ).ropical-forest data. We plot the fits of equation n4 a Figure 1 |To understand the qualitative difference between species composition (20)sintolequation (18) and mples choy 5 r~nd/JMl,yefRm andBCI data.,Each Relative species abundance of coral-reef communities. We plot and (5) ( olid ine) to the RSA of sa defining sen m m om ro 0 the eSMRh in the coral-reef system to the tropical species abundance data the fits of equation (1) (solid line) and in coral-reefforests, we used the available obtains ws RSA for the same number of the ples withthe local abundances as one plot sho an analytical expression for sam RSA of the same in the cor for the coral-reef tree species abunda), reef (b), metacommunity (c) and plot on community: al-reef data. The RSAs are qualitatively similar to those in Fig. 1 (the relative local community (a nce data from the forest dynamics 6,7 values of m? re equal to 1, 0.84, 0.46 and 0.19 for a, b, c and d, respectively). a metacommunity (habitatsIsland (BCI), The ama to generate random samples of Barro Colorado pooled, d) . Pan bars are observed numbers of nð species binned me size 2as in the coral-reef studies.firsture 4 shows the RSA for x CðnzyÞ the sa into log abundance categories. The Fig histogram bar ~ expf{y½JM =ðmdÞ{ lnð1{xފg~ hQn i~h dy we obtain very similar values of the biological parameters to the ones 10 derived previously. The exact theory applies both under the assumption of species independence in a fluctuating community and for 5 species undergoing zero-sum0dynamics9. 2 4 6 2 4 6 10 0 Abundance category tropical between coral reefs and 8 8 10 12 10 ð1{xÞmm=JM xn ~ Cðnzmm=JM Þ Pn,m ~ 0 ~ 2 4 Cðmm=JM Þ n! 10 6 8 0 2 ~ 10 4 6 8 ð20Þ 10 Abundance category of the metacommunity Volkov et al. 2007 Rank-abundance diagrams change in grassland community with continuous fertilizer inputs Tokeshi 1993 Primate communities 1. Community ecology 2. Measuring diversity >3. Primate communities: patterns Primate distribution !"#$%&'((')&(*"+%,*-.' /%'#-)0)&'.%)&(1#&1('%2('3*-4"+% 5"(*"&*-4)%*4%#+*."&'6%,1&%*)%#-.7-)'8%-9%8*99'('4&%-(3"4*).)% *4%8*99'('4&%7"(&)%-9%&$'%:-(+8 Primate habitats !"#$%&'()*+ ,--.*/01%2"3*45671)&"38*937:8*#6;<&2(&3'*12*=03>1?&3*!6??&3'2 Comparing primate communities Suriname Kuala Lompat Ranomafama Comparing primate communities: body size Neotropical primates smaller Fleagle 1999 Comparing primate communities: activity period Abundance of nocturnal species in Madagascar Fleagle 1999 Comparing primate communities: group size Abundance of solitary feeders in Madagascar, few in America Fleagle 1999 Comparing primate communities 490 ￿. ɢ. ￿ʟ￿￿ɢʟ￿ ￿ɴ￿ ￿. ￿. ʀ￿￿￿ Tai Kibale RaleighvallenVoltsberg Manu Ketambe Morondava Ranomafana Kuala Lompat Figure 1. Map showing the location of the primate communities compared in this study. ‘‘ecological space’’ occupied by communities on different continents similar or different? Are the species in communities from some biogeographical areas more similar to one another than those of other areas? Are some ecological characteristics more consistent among communities than others? Are ecological similarities of primate communities within continental areas greater or smaller than those between primates from different continents? What is the role of phylogeny in determining the ecological characteristics of primate communities? M a t erial a n d m e t h o ds In this analysis we compare eight primate communities—two from each of the four major areas currently occupied by primates. In general, we have chosen some of the most species rich Fleagle & Reed 1996 Principal components analysis Principal components analysis Principal components analysis Principal components analysis Principal components analysis Principal components analysis characteristic 3 char act PC 2 eris ti c2 isti character c1 PC 1 component space original data space Principal components analysis 498 nocturnal, body size, diurnality, frugivory, leaping￿. ɢ. ￿ʟ￿￿ɢʟ￿ ￿ɴ￿limbing, terrestriality c ￿. ￿. ʀ￿￿￿ Nocturnality leaping Body size, diurnality, frugivory, climbing, terrestriality 3 Factor 2 (25%) 1 0 Arboreal quadrupedalism faunivory Folivory 2 folivory –1 arboreal quadrupedalism, faunivory –2 –3 –2 –1 0 Factor 1 (28%) 1 2 3 Figure 3. Plot of all the primate species on 1. Avahi laniger 25. 2. Propithecus diadema 26. 3. Propithecus verreauxi 27. 28. 4. Lepilemur microdon 5. Lepilemur ruficaudatus 29. 30. 6. Hapalemur griseus 7. Hapalemur aureus 31. 70 species, 10 variables per species the first two factors of the principal components analysis. Nycticebus coucang 49. Pan troglodytes 50. Pongo pygmaeus Nycticebus coucang 51. Hylobates syndactylus Colobus guereza Colobus polycomos 52. Hylobates syndactylus 53. Hylobates lar Piliocolobus badius Piliocolobus badius 54. Hylobates lar 55. Saguinus fuscicollis Procolobus verus Fleagle & Reed 1996 Primate communities within biogeographic regions highly congruent nocturnal, leaping body size, diurnality, frugivory, climbing, terrestriality 499 ￿￿￿￿￿ʀɪɴɢ ￿ʀɪ￿￿￿￿ ￿￿￿￿￿ɴɪ￿ɪ￿￿ 3 (a) 2 L. microdon 3 (b) A. laniger H. aureus H. simus H. griseus L. ruficauda 2 folivory E. fulvus Factor 2 (25%) P. diadema Factor 2 (25%) 1 0 –1 –2 –3 3 (c) 2 P. versus E. fulvus E. rubriventer D. madagascariensis M. rufus 1 0 M. murinus P. verreauxi P. furcifer V. variegata C. major –1 M. coquerelli C. medius –2 –1 0 1 2 3 –2 –3 3 (d) –2 –1 0 1 2 3 Factor 1 (28%) Factor 1 (28%) arboreal quadrupedalism, faunivory Factor 2 (25%) Factor 2 (25%) 1 0 G. demidoff P. badius Madagascar 2 1 0 C. polykomos C. guereza P. badius G. demidoff G. senegalensis C. ascanius C. mitis C. l'hoesti P. anubis P. troglodytes P. potto C. petaurista Fleagle & Reed 1996 C. albigena 3 (a) 2 1 0 –1 –2 –3 3 (c) 2 Factor 2 (25%) 3 (b) 2 1 0 M. nemestrina Factor 2 (25%) Factor 2 (25%) Primate communities within biogeographic regions highly congruent P. thomasi P. melalophos P. obsculra H. lar H. syndactylus H. syndactylus H. lar nocturnal,mestrina M. ne N. coucang leaping –2 –1 0 1 2 3 Factor 1 (28%) M. fasicularis P. pyg eus body size, diurnality,mafrugivory, M. fascicularis –1 N. coucang climbing, terrestriality –2 –3 3 (d) 2 Factor 2 (25%) –2 –1 0 1 2 3 Factor 1 (28%) folivory A. seniculus 1 0 –1 –2 –3 A. seniculus S. sciureus S. midas C. apella 1 0 –1 –2 –3 C. moloch P. pithecia A. paniscus S. fuscicollis S. imperator C. albifrons S. sciureus C. apella C. satanus A. paniscus A. trivirgatus –2 –1 0 1 Factor 1 (28%) 2 3 –2 –1 0 1 Factor 1 (28%) 2 3 arboreal quadrupedalism, faunivory Figure 5. Individual plots of four primate communities on the first two factors of the principal components analysis—two from Asia: (a) Kuala Lompat, and (b) Ketambe; and two from South America (c) Raleighvallen–Voltzberg, and (d) Manu. South America). It is likely that had we included a Bornean locality with a tarsier we would have seen more heterogeneity in Asian localities and more similarity to the African sites. However the presence of medium-sized, frugivorous–folivorous, suspensory gibbons in the upper right quadrant and the absence of numerous cercopithecines in the lower right quadrant also distinguish the Asian communities. America Fleagle & Reed 1996 Primate communities within biogeographic regions highly congruent nocturnal, leaping 500 3 (a) 2 1 0 M. fasicularis body size, diurnality, frugivory, climbing, terrestriality ￿. ɢ. ￿ʟ￿￿ɢʟ￿ ￿ɴ￿ ￿. ￿. ʀ￿￿￿ 3 (b) 2 P. thomasi P. melalophos H. lar folivory H. syndactylus H. lar M. nemestrina Factor 2 (25%) H. syndactylus Factor 2 (25%) P. obsculra 1 0 –1 –2 –3 3 –1 –2 –3 3 (c) 2 M. nemestrina N. coucang N. coucang M. fascicularis P. pygmaeus –2 –1 0 1 2 3 –2 –1 0 1 2 3 Factor 1 (28%) (d) Factor 1 (28%) arboreal quadrupedalism, faunivory Factor 2 (25%) 1 0 A. seniculus S. sciureus C. apella Factor 2 (25%) Asia 1 0 P. pithecia C. moloch 2 A. seniculus A. paniscus S. fuscicollis S. imperator C. albifrons S. sciureus Fleagle & Reed 1996 A. paniscus ￿￿￿￿￿ʀɪɴɢ ￿ʀɪ￿￿￿￿ ￿￿￿￿￿ɴɪ￿ɪ￿￿ 3 3 499 Factor 2 (25%) P. diadema Factor 2 (25%) Primate communities within biogeographic regions highly congruent (a) L. microdon (b) A. laniger L. ruficauda 2 1 0 H. aureus H. griseus 2 1 0 H. simus E. fulvus P. verreauxi D. madagascariensis –1 –2 –3 3 M. rufus C. major nocturnal, leaping E. rubriventer E. fulvus V. variegata body size, diurnality, frugivory, –1 climbing, terrestriality M. murinus M. coquerelli P. furcifer C. medius –2 –1 0 1 2 3 –2 –3 3 –2 –1 0 1 2 3 Factor 1 (28%) (c) 2 P. versus Factor 1 (28%) (d) 2 C. guereza folivory P. badius Factor 2 (25%) 1 0 –1 –2 –3 G. demidoff Factor 2 (25%) P. badius C. polykomos 1 0 –1 G. demidoff G. senegalensis C. ascanius C. mitis C. l'hoesti P. anubis P. troglodytes P. potto C. atys C. petaurista C. diana C. campbelli C. albigena P. potto P. troglodytes –2 –1 0 1 Factor 1 (28%) 2 3 –2 –3 –2 –1 0 1 Factor 1 (28%) 2 3 arboreal quadrupedalism, faunivory Figure 4. Individual plot of four primate communities on the first two factors of the principal components analysis—two from Madagascar: (a) Ranomafana and (b) Marosalaza Forest near Morondava; and two from Africa: (c) Tai Forest and (d) Kibale Forest. Nevertheless, in the distribution of species in the ecological space defined by the first two PCA factors, they are more similar to one another than either is to the communities from other continental regions. Compared with the Malagasy communities, the two African communities are distinguished by the abundance of medium to large, diurnal, quadrupedal, frugivorous species such as cercopithecines and chimpanzees that occupy the lower right quadrant and the Africa Fleagle & Reed 1996 Primate communities within biogeographic regions highly congruent 502 3 ￿. ɢ. ￿ʟ￿￿ɢʟ￿ ￿ɴ￿ ￿. ￿. ʀ￿￿￿ Madagascar Asia 2 Factor 2 (25%) 1 Africa 0 –1 America –2 –3 –2 –1 0 Factor 1 (28%) Figure 6. Superimposed polygons outlining the ‘‘ecological space’’ of the first two factors of the principal components analysis occupied by each of the eight primate communities. (——), South America; (– – –), Asia; (- - -), Madagascar; (· · ·), Africa. Table 2 Four measures of dispersion among individual species comprising each of the eight primate communities 1 2 3 Fleagle & Reed 1996 Factor 2 (25%) Primate communities within biogeographic regions￿￿￿￿￿ɴɪ￿ɪ￿￿ congruent ￿￿￿￿￿ʀɪɴɢ ￿ʀɪ￿￿￿￿ highly 495 1 0 (a) (b) (c) –1 –2 –3 –2 –1 0 Factor 1 (28%) 1 2 3 Figure 2. GraFihicrell6.strupiernmpotshe tpolee ons asutriesing ehe l‘oecoalgdcapersice’’ amoheg firsditwdufalctorecoestheitprn cipal p gu i u S at o i of ed hr yg me o u l n of t co ‘ giclo i is l spa on of t n in t vi o a sp s i f w hi in communities. c(omAonenosf analygisnoc(cupaed rby e acihtance efreim tcpnitroitd aommunities.ag——x,oSouthcAmeaina; .(– – –), a) p rea t pol yso , b) i ve ag e d s of th ogh e r ma e c nd (c) aver ( e ta ) nomi distr cce Asia; (- - -), Madagascar; (· · ·), Africa. Ta e Four m p complete the blat2 set. Howch eretasneyghofvdrirmermoteamongtiuntvodudasa ewias coorpoiwied because it da ev , o uresa e sy li si i n d amo ndi i f al t p c e s b mr r s ng l ea of he i t p i ate communi ies is documented that primate species demonstrate different dietary regimes at different localities, Taxonomic Taxonomic e.g., Eulemur fulvus (Richard & Dewar, 1991). Area of Centroid distance t In order to compare tComdiunetrent prpmyate comimuceities(two factcommon vsoainerlesl)l ecological he m ff i y i ol gon in a ors) (alldi arvace a b d stan n space, we created a 70 ￿ 10 data matrix based on the 70 species and ten variables. The body sizes and activity pRanernsanaf each 4p1 cies (whi8 h were 1c34ded as 1 0f·o12 diurnal, 2 for s9 e 1·3 c ·o 6r Madagascar att omaf o 46 d 15 1· e cathemeral, and 3 for noMtouondalv)a were stan0 ardized·3using z-sco3r6 s. We sp0·c2fically included c r rn a e6 i 0 Raleighvallen 149 0·79 0· 8 3 0·408 America body size in the analysis Mancause it is an14mportant72 spect of0·a8 y primat0·’471 cology and is be u i9 es e 0· a 6n 4p ·466 also readilyAsia ble for Kuolst Ltompa.t Attem32ts to ‘‘e1l·i23 inate’’ s1z2e in either0morphological or availa m a a axa m i· 5 Ket m 6 12 · ·540 ecological analyses throughbauesbe of residual29 have m·a4or flaws 1a29d usually 0amplify measuree s j n Ki al 577 1·16 1· 1 8 0·671 ment erroAfrica than Taliiminating size 5(e.g., Jung14rs et al., 11996). A Pe·arson correlation rs rather e 98 1· e · 16 0 614 matrix was calculated and this matrix was then used in the creation of factor scores using principal components analysis (Gower, 1966; Pimentel, 1979) to summarize the ecological Comparing primate communities Madagascar: abundance of nocturnal, solitary, folivorous species America: lack of folivorous, solitary species; small body sizes Asia: low species diversity; few small species; many suspensory species Africa: abundance of large frugivores Take home messages 1. Ecological communities have collective and emergent properties. 2. Species diversity is a function of both species richness and evenness. 3. Primate communities are more similar within than between broad biogeographic regions. 4. Note the general patterns in each region, summarized on the previous slide. No question to ponder today Focus on the readings; there will be things from the reading on the test. ...
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This note was uploaded on 04/05/2011 for the course ANT 154bn taught by Professor Debello during the Winter '10 term at UC Davis.

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