The Metazoan Parasite Fauna of Grebes (Aves: Podicipediformes) and Its Relationship to the Birds' Biology
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THE METAZOAN PARASITE FAUNA OF GREBES (AVES: PODICIPEDIFORMES) AND ITS RELATIONSHIP TO THE BIRDS' BIOLOGY BY ROBERT W. STORER MISCELLANEOUS PUBLICATIONS MUSEUM OF ZOOLOGY, UNIVERSITY OF MICHIGAN, NO. 188 Ann Arbor, May 31,2000 MISCEI.I,ANEOUS PUH1.IC:ATIONS M'IJSEIJM OF %OOI.OC:Y. LJNIVERSITY OF MICIHI<;AN NO. 189 Tlic pl~blicatiorisof' the Museuni of' Zooloky, The IJnivel-sity of Michigan, consist primarily of two series-tllc Occasional Papers and thr Miscrllaneo~lsP~~l)lications. Kotli series we]-c founded by 111: Rryir~ltWnlkcl-, Mn Bradshaw H. Swales, and Dr. W. W. Newcomb. Occasionally the Mt~set~nip~~hlishes contribtrtions oc~tsidcof these series; begirir~irigin 1990 these are titled Special Pr~hlicationsand arc numbered. All sublnittcd rria~iuscriptsreceive extel-nal review. 'Tlrc Miscellaneous P~~l)lications,which include monographic siltdies, papers on field arltl I~II- scum tccll~liques,and other contributions not within the scope of the Occasional Paper-s, ar-' pub- lished separately. It is not intended th;lt they 11c grouped into volu~ries.Each riun~bel.has a title page arltl, wlicrl ~iecessal-y,a tal~leof contents. The Occasional Papers, ptlblicatio~iof' wliich was bcgun in 1913, senTeas a niediuni fol- 01-iginal stt~dicsI~ased pl-incipally upon the collectiolls ill thc MIIS~III~I.Tliry al-r isst~rdseparately. Whc~la sr~llicirnttlu~nbt-I- of' pages has been printed to make avolunie, a title page, tahlr of'contents, ;inti an i~~dexitre supplied to libraries arid individuals on tlie mailing list for the SCI-ies. A cornplctc list of publications on Birds, IFislics, Insects, Ma~r~~nals,Mollusks, Rcptilcs ii1lt1A~iipl~ib- iaris, ~u~tlother topics is availilblc. Addl.css inquiries to tlie Director, Musr~~rnof'Zoology, The Univcr- sity of' Michigan, Ann Arhor, Michiga~i48 lo!)-1075). Bailey, K.M. & (:. G;rns. 1998. 'live new syril~ra~~c.hitllishes, Monop/(~v.rro.sr,li from Pc~lills~~la~.l~ltlia iu~dM. rlurilr~c~ifrorii Sri I;ark;~. Occ. Pap. 726. 18 PI)., 7 figs $2.00. Nr~ssl)a~~rn,R.A. Xc M.E. Pfrctrdcn 1998. Revision of'tlir ASI-ican caeci1i;in genus Schis/o?nrlopclrnParkel- (hriipliibia: <;ymnol~hiori;l:(:ileciliidac). Misc. Pul~l.187. 35 pp., 15 figs., 15 tables, 2 color plates. $7.00. Wc1111, S.A. arltl li.1i. Miller. 1998. %oogonc,~ir~is~~quilc~, a ticw goocleitl Gsli ((:yp~-i~~odor~tifo~-~iies)ft-om the Alncca tlr;lin;~gcof Mcxico, iind ;I rediagnosis ofthe grnrls. Occ. I'ap. 725. 28 pp, 5 tigs. $2.00. Ntrssbatun, R.A., (::I. R;rxworthy & 0.Pronk. 191-18. Tlre ghost geckos of' Madagascar: a fitrtlitu. rcvi- sion of tlie Malagasy Icaf-toctl gcckos (Reptilia, S(l~~;imata,Gekkonidae). Misc. I't1111. 186. 26 pp,, 25 figs., 5 tahlcs. $9.00. Dyrt; R.S. 1997. Pliylogenetic Revisio~iof Atherinopsinae ('kleostci, Athcri~iopsidae),with (;otil- IIICII~S011 tl~cSystcniatics of tlie South American FI-esliwatt-I.Fish C;c~rttslIn~ilicli~bhj~,\ ((;i~.a~-tl. Misc. Publ. 185. (54 pp., 46 figs. $15.00. OCol~rlor;R.M., R.K. Colwell 8c S. Nacem. 1996. 'rlir Flower- Mires ot..Ii-inid;ld 111: 'I'lic Gc~it~sIil~ino,c(,iut (Acari: hscidae) . Misc. 1'11111. 184. 32 pp, 70 figs. $10.00. ILlr~gr,A.G. & K.A. Nt~ssbaurri. 1995. A Review Of APr-ican-Madagasci111Ckkkonid I.i/a~.tlPhylogeny A~idBiogeogl-aphy (Sqlrarnata). Misc. Plt111. 183. 20 pp, 11 ligs. $8.50. Kaxwortlly, (;.J. & R.A. Nussbaum. 1994. A review of. the Madagascan Snake Gcncl-a l's~urloxy?-hoj)z~.r, I'~~m~-l~(~rlin(~m,i11itl IIc~tmliodot~ (Squ;im;~t;t: (:olubr-idae). Misc. I'uhl. 182. 37 pp, 25 figs. $1 1.50. Gosli~~e,W.A. 1993. A sr~rvcyof ~~pprr,jawm~~scr~laturc in 1iiglic1- tclcostca~i fislics. Occ. Pap. 724. 26 pp, 9 figs. $2.20. l~~rrllrn;~n,M1.E. 8c J.A. (:arnpl)ell. 1992. Hylid frogs of tlic ge1111sI'l(~c-lrol~~lrr: Systc~ri;~tics a~ltl phylogc- nctic ~.clationsllips.Misc. 1'11111. 181. 38 1111, 2 1 figs. $9.10. I.cc (:. Bollitlgc~,(,x o//itio 1);unic.l I). Hal-ning, C:l.;ltl(l H~I\TII At~drca17iscl~e~~ NCWIII~II, AIIIIAI l)os D;~vitlA. B~-;~l~tlon,Ann i\~-ho~- OIi\rii~1'. IVI~I~I~;II-~,(;oo(i~-i( 11 S. MAI-~~II131ylo1; Grossc I'oinrc F;u-lirs 1.;1t11.c11ecB. I)citch, Rloomlicld tlills liehecc;~RI( (;owan, Ann AI-bol- I<,ithc~-i~icI<. White, AIIII R1-1,ol. The Metazoan Parasite Fauna of Grebes (Aves: Podicepediformes) and its Relationship to the Birds' Biology Robert W. Storer Bird Division Museum of Zoology The University of Michigan Ann Arbor, Michigan 48 109-1079, USA MISCELLANEOUS PUBLICATIONS MUSEUM OF Z00L2OC;Y,UNIVERSITY OF MICHIGAN, NO. 188 Ann Arbor, May 31, 2000 Tllc lilk cycle of the ccstodc, ?ic/rin hir~mls. Thc adult parasite (centel-),whicll reaches a length of approximately 2.5 Irlln, irlllabits the anterior section of the small intestine of the definitive host, herc an Ez~redGrcbc (I'odicc~j~st~ipicollis). A mean of 2,794 individual worms of this species has becn Pound in 31 individual birds. The minute (0.02 mm) eggs are passed into the wateI; where they arc presumably swept into thc moutl~ofthe intermediate host, a corixid bug (Sigma), by the insect's front legs. Grcbcs become infected by eating the bugs, which may teen1 in lakes where the birds breed. This common parasite of grebes in the Northern Ncrnisphere belongs to the fi~rnilyAmabiliidac, all br~tone ofwl~osc29 spccies are grebe specialists. Origi~ialdrawing by Jollil Mcgahan from sources listed ill thc acknowledgrnents (p. 65). ABSTRACT Slow); li. W 2000. 7'1~~M(llnzoc~17 frarc~.sil~/(~una (~gt-(,he.s (A~J~s: I'odi~ij~rdi/oi-rn(~,s) rind its ,rrlalion,ship lo th~ hir.(islbiology. ~Misc.I'r~bl. Mus. Zool., CJniv. A!lichigc~17,188:l-90, I O,figs., 7 tabl(~s.The data base of this work consists of a list oP thc n~~rlticclll~larparasites known to infect grcbcs and a list of the known spccics oS prey taken by ?ac:h specics of grcbc. Tllc formcr inclltdes information on whcrc in thc bird the parasites arc fo~~nd,the distribution of the parasite species by continents, degree of host specificity, and life cycles (whether on fresh or salt water and lists of known intermediate and paratenic hosts). These data sets are used to sl~owhow the parasite faunas arc related to the biology of the birds. The aquatic existence of grebes and the widc range of prey taken by individ~~alsand spccies of grebes are reflected in tlic wide range of helminths infecting the birds (113 species ofdigencs, 86 ofccstodes, 13 of aca~nthocephalans,and 37 of nematodes, most ofwhich have aquatic lifc cycles). One family, one subFC~rnily(each with the exception ofa singlc spccies), several genera, and 104 spccics of'hclminths are considered grcbc specialists, as at-e most ofthe specics of cctoparasitcs. Of the well-studied species of grcbc hel~ni~itlispecialists, almost all have been found in more than one gl-cbc spccies and Inany rnorc in more tllan orlc grebc genus. This is believed I-elatedto tllc fact that even grebes adapted for taking particular kinds of prey will usually take Inany other kinds. Thrrs, where several species of grcbcs arc sympatric, thcrc is a considerable ovel-lap in diet and consequently, ovcrlap in the intermediate hosts ingested. On thc other hand, most of the ectoparasites appear to be confined to single grebe species. Grebes contain large 11urnbcrsof intestinal lielminths. In a study of these parasites of 91 grcbcs from the breeding grotmds in Alberta (Stock 1985), all were found i~~fectcd,with nr~ml~ersranging from 2 to 15 specics and 112 to 33,169 individuals, the majority (96 percent) of which were cestodes. Most grebes spcrld the breeding season on eutrophic bodies ol'watcl; which have an abundant supply ofkinds and individuals ospotcntial prey for the birds. More than half of the spccies of grebcs winter on salt water where a variety of hcl~nirlthswith salt-water intermediate 1.1ostscan infect the birds, bl~ttlnfor- tl~nately,very Pew grebes taken on salt-watcr habitats have been examined for p;~r;isites. Still other kinds of'helminths may be acquired on bodies of watcr along the birds' migration routes. Most of the helm- i~lthsthat are obtained on fresh waters presumably are lost shortly after the birds' arrival on salt watcr and vi~pII(~T,S(C. What causes these changeovers in unknown, but it is suggested that the n~cchanisrnsfor this are I-eversals of the salt gradient in thc intestines. Eared Grebes coming to the highly salinc Mono Lake 011 their molt migration after the breeding season presumably soon lose their Iielminth parasites. Grebes have a u~~iquchabit of swallowing their own feathers which form pellets containing other indigestible materials and, when cast, arc believed also to rid the stomach of nen~atodcsthat Seed on the grebes' food therc. A separate plug of feathers is lodged in thc pyloric outlet of'the stomach where it acts as a sieve preventing large pieces of iildigestiblc material from passing through the intestine. This is bclicved to reduce the likelihood olpa~.asitesbeing dislodged from the wall of thc gut. Grebcs arc the only birds that do not nest on land but regularly build floating nests of matel-ial taken from the water.